Painted by J. J. Masquerier.         Engraved by W. T. Fry.



William Spence, Esq^r., F.L.S.

Published by Longman & C^o. London, July 1825.

[Pg i]

AN

INTRODUCTION

TO

ENTOMOLOGY:

OR

ELEMENTS

OF THE

NATURAL HISTORY OF INSECTS:

WITH PLATES.

By WILLIAM KIRBY, M.A. F.R. and L.S.

RECTOR OF BARHAM,
AND

WILLIAM SPENCE, Esq. F.L.S.

IN FOUR VOLUMES.

VOL. IV.

FIFTH EDITION.

LONDON:

PRINTED FOR

LONGMAN, REES, ORME, BROWN, AND GREEN,

PATERNOSTER ROW.

1828.

[Pg ii]

PRINTED BY RICHARD TAYLOR,

RED LION COURT, FLEET STREET.

[Pg iii]

CONTENTS OF VOL. IV.

[Pg 1]

AN INTRODUCTION TO ENTOMOLOGY.

LETTER XXXVII.

INTERNAL ANATOMY AND PHYSIOLOGY OF INSECTS.

SENSATION.

Having given you this full account of the external parts of insects, and their most remarkable variations; I must next direct your attention to such discoveries as have been made with regard to their Internal Anatomy and Physiology: a subject still more fertile, if possible, than the former in wonderful manifestations of the power, wisdom and goodness of the Creator.

The vital system of these little creatures, in all its great features, is perfectly analogous to that of the vertebrate animals. Sensation and perception are by the means of nerves and a common sensorium; the respiration of air is evident, being received and expelled by a particular[Pg 2] apparatus; nutrition is effected through a stomach and intestines; the analogue of the blood prepared by these organs pervades every part of the body, and from it are secreted various peculiar substances; generation takes place, and an intercourse between the sexes, by means of appropriate organs; and lastly, motion is the result of the action of muscles. Some of these functions are, however, exercised in a mode apparently so dissimilar from what obtains in the higher animals, that upon a first view we are inclined to pronounce them the effect of processes altogether peculiar. Thus, though insects respire air, they do not receive it by the mouth, but through little orifices in the sides of the body; and instead of lungs, they are furnished with a system of air-vessels, ramified ad infinitum, and penetrating to every part and organ of their frame; and though they are nourished by a fluid prepared from the food received into the stomach, this fluid, unlike the blood of vertebrate animals, is white, and the mode in which it is distributed to the different parts of the system, except in the case of the true Arachnida, in which a circulation in the ordinary way has been detected, is altogether obscure.

In order that you may more clearly understand the variations that occur in insects, and in what respects they differ amongst themselves, and from the higher animals, in the vital functions and their organs, I shall consider them as to their organs of sensation, respiration, circulation, nutrition, generation, secretion, and muscular motion.

Organs of Sensation.—The nervous system of animals is one of the most wonderful and mysterious works of[Pg 3] the Creator. Its pulpy substance is the visible medium by which the governing principle[1] transmits its commands to the various organs of the body, and they move instantaneously—yet this appears to be but the conductor of some higher principle, which can be more immediately acted upon by the mind and by the will. This principle, however, whatever it be, whether we call it the nervous fluid, or the nervous power[2], has not been detected, and is known only by its effects. The system of which we are speaking may therefore be deemed the foundation and root of the animal, the centre from which emanate all its powers and functions.

Comparative anatomists have considered the nervous system of animals as formed upon four primary types, which may be called the molecular, the filamentous, the ganglionic, and cerebro-spinal[3]. The first is where invisible nervous molecules are dispersed in a gelatinous body, the existence of which has only been ascertained by the nervous irritability of such bodies, their fine sense of touch, their perceiving the movements of the waters in which they reside, and from their perfect sense of the degrees of light and heat[4]. Of this description are the infusory animals, and the Polypi. The nervous molecules in these are conjectured to constitute so many ganglions, or centres of sensation and vitality[5]. The second, the filamentous, is where the nervous system consists of nervous threads radiating from the mouth, as in the[Pg 4] Radiata, or star-fish and sea-urchins[6]. The third, the ganglionic, is where the nervous system consists of a series of ganglions connected by nervous threads or a medullary chord, placed, except the first ganglion, below the intestines, from which proceed nerves to the various parts of the body. This system may be considered as divisible into two—the proper ganglionic, in which it is ganglionic with the ganglions arranged in a series with a double spinal chord. This prevails in the classes Insecta, Crustacea, Arachnida, &c., and the improper ganglionic, in which it is ganglionic with the ganglions dispersed irregularly, but connected by nervous threads, as in the Mollusca[7]. In the fourth, the cerebro-spinal, the nervous tree may be said to be double, or to consist of two systems—the first taking its origin in a brain formed of two hemispheres contained in the cavity of the head, from which posteriorly proceeds a spinal marrow, included in a dorsal vertebral column. These send forth numerous nerves to the organs of the senses and the muscles of the limbs. The second consists of two principal ventral chords, which by their ganglions, but without any direct communication, anastomose with the spinal nerves and some of those of the brain, and run one on each side from the base of the skull to the extremity of the sacrum. This system consists of an assemblage of nervous filaments bearing numerous ganglions, from which nervous threads are distributed to the organs of nutrition and reproduction[8]. Its chords are called the[Pg 5] great sympathetic, the intercostal, or trisplanchnic nerves[9]. While the first of these two systems is the messenger of the will, by means of the organs of the senses connects us with the external world, and is subject to have its agency interrupted by sleep or disease[10]; the latter is altogether independent of the will and of the intellect, is confined to the internal organic life, its agency continues uninterrupted during sleep, and is subject to no paralysis. While the former is the seat of the intellectual powers, the latter has no relation to them, but is the focus from whence instincts exclusively emanate: from it proceed spontaneous impulses and sympathies, and those passions and affections that excite the agent to acts in which the will and the judgement have no concern[11].

It is probable, though the above appear to exhibit the primary types of nervous systems, that others exist of an intermediate nature, with which future investigators may render us better acquainted[12]: but as our business is solely with that upon which insects in this respect have been modelled, without expatiating further in this interesting field, I shall therefore now confine myself to them.

We have before seen[13] that the nervous system of insects belongs to the ganglionic type: but it requires a more full description, and this is the place for it. It originates[Pg 6] in a small brain placed in the head, and consisting almost universally of two lobes, sometimes extremely distinct. It is placed over or upon the œsophagus or gullet, and from its posterior part proceeds a double nervous chord, which embracing that organ as a collar dips below the intestines, and proceeds towards the anus, forming knots or ganglions at intervals, in many cases corresponding in number with the segments of the body, and sending forth nerves in pairs, the ramifications of which are distributed to every part of the frame. In the perfect insect the bilobed ganglion of the head or the brain is usually of greater volume than in the larva, and the ganglions of the spinal chord are fewer, which gives a more decided character of centricity to the whole nervous system[14]. This may be considered more particularly with respect to its substance and colour; its tunics, and parts.

I. Substance and Colour.—The nervous apparatus of insects is stated by those who have examined it most narrowly, though consisting of a cortical and medullary part, the latter more delicate and transparent than the former, to be less tender and less easy to separate than the human brain[15]. It has a degree of tenacity, and does not break without considerable tension; in general, it is clammy and flabby, and under a microscope a number of minute grains are discoverable in it, and when left to dry upon glass, it appears to contain a good deal of oil, which does not dry with the rest[16]. That of the ganglions differs from the substance of the rest of the spinal[Pg 7] chord, in being filled with very fine aërial vessels, which are not discoverable in the latter[17]. With regard to colour, Lyonet states that the chords of the spinal marrow in the larva of the great goat-moth are of a blueish gray, and have some transparence[18]; Malpighi and Swammerdam observed that the cortical part of the ganglions of that of the silk-worm and the hive-bee had a reddish hue, while the medullary part was white[19]; Cuvier relates that the brain and the third ganglion in Hypogymna dispar, with us a scarce moth, differed in colour from all the rest, being quite white, while the others were more or less tinted, and examined under a lens appeared variegated by reddish sinuous markings, resembling blood vessels as they are seen in injected glands[20].

II. Tunics.—The coats that inclose the various branches of the nervous system in insects seem analogous to those of vertebrate animals. The first thing that strikes the eye, when these parts in a recent subject are submitted to a microscope, is a tissue of very delicate vessels, which ramify beyond the reach of the assisted sight; these are merely air-vessels or bronchiæ derived originally from the tracheæ of the animal: but besides these is an exterior and an interior tunic; the first corresponding with the dura mater of anatomists; and the other, which is[Pg 8] the most delicate and incloses the cortical and medullary parts, with the pia mater[21].

III. Parts.—The nervous system of insects consists of the brain; the spinal marrow and its ganglions; and the nerves.

i. Brain.[22] Linné denied the existence of a brain in insects, and most modern physiologists seem to be of the same opinion. A part however, analogous to this important organ—at least in its situation, and in its emission of nerves to the principal organs of the senses, in which respect it certainly differs very materially from the upper cervical ganglion, which Dr. Virey regards as its analogue[23]—is certainly to be found in them; and as Messrs. Cuvier and Lamarck distinguish this part by the name of brain, we may continue to call it by that name without impropriety. The brain of insects, then, is distinguished from the succeeding ganglions of the spinal chord by its situation in the head, the middle of the internal cavity of which it occupies, and by being the only ganglion above the œsophagus. It is usually small, though in some cases larger than they are[24]. It consists of two lobes, more or less distinct and generally of a spherical form. In Oryctes nasicornis and Pontia Brassicæ the lobes are separated both before and behind[25]; while in the larva of Dytiscus marginalis, but not in the imago, in which there are two large hemispheres separated by a furrow, the brain is undivided[26]. Cuvier mentions the[Pg 9] larva of a saw-fly in which this part is formed of four nearly equal spherical bulbs[27]: in the Scorpion (to judge by the figure of Treviranus[28]) the two lobes represent an equilateral triangle, the exterior angle of which terminates in several lesser spherical bulbs; in Acrida viridissima, Nepa cinerea, Clubiona atrox, and the common Louse, the lobes are pear-shaped[29].

ii. The spinal marrow and its ganglions[30]. From the posterior part of the brain of insects, but in the ground and water beetles (Eutrechina and Eunechina) from its sides below[31], issue two chords which diverging embrace the œsophagus, and dipping below it and the intestines,—a situation they maintain to the end of their course,—and in their further progress uniting at intervals and dilating into several knots or ganglions, compose their spinal marrow. This part is so named, from a supposed analogy to the spinal marrow of vertebrate animals, which however admits of some degree of doubt; yet, since it mixes the functions of that organ with those of the great sympathetic nerves, the denomination is not wholly improper, and may be retained. Though this chord is usually double when it first proceeds from the brain, and surrounds the œsophagus like a collar, yet in some insects it may be called a single chord. This is the case with that of the common louse, in which Swammerdam could perceive no opening for the transmission of the part just named[32]; if he was not mistaken in this, the brain, as well as the rest of the spinal marrow in[Pg 10] that animal, would be below the intestines; from the figures of Treviranus it should seem that the spiders, at least Clubiona atrox, are similarly circumstanced[33]; in the cheese-maggot, which turns to a two-winged fly (Tyrophaga Casei), the chord is also single, but it has a small orifice through which the gullet passes[34]. At the union of the chords in other cases below that organ, a knot or ganglion is usually formed, and an alternate succession of internodes and ganglions commonly follows to the end. The internodes also may generally be stated to consist of a double chord, though in many cases the two chords unite and become one, or are distinguished only by a longitudinal furrow, and even where they are really distinct and separable, in the body of the insect they lie close together[35]. In the rhinoceros beetle (Oryctes nasicornis) and Acrida viridissima &c. all the internodes consist of a double chord[36]; but in many other insects numerous variations in this respect occur.—Thus in the stag-beetle the last internode is single[37]; in the caterpillar of the cabbage butterfly (Pontia Brassicæ) the five first are double, and the six last single[38]; in that of the great goat-moth (Cossus ligniperda) the three first only are double, but the others terminate in a fork[39]; in the cockroaches (Blatta) the four first, in Hydrophilus piceus the three first, and in Eristalis tenax the two first only are double, the rest being all single[40]. A singular variation takes place in Hypogymna dispar; all the internodes are[Pg 11] single, except the second, the chords of which at first are separate, and afterwards united[41]; and, to name no more, in Clubiona atrox there is only one internode, which is single, with a longitudinal furrow[42]. In some, as in the louse, the grub of Oryctes nasicornis, and the cheese-maggot, there are no internodes, the spinal marrow being formed of knots separated only by slight or deep constrictions[43].

I must next say something of the ganglions[44]. Lyonet has observed that, in the caterpillar of the great goat-moth, these in one respect differ remarkably from the chords that connect them; in the latter the air-vessels or bronchiæ only cover the outside of the tunic, while in the former they enter the substance of the ganglion, which is quite filled with their delicate and numberless branches[45]. Every ganglion may be regarded in some degree as a centre of vitality or little brain[46], and in many cases, as well as the brain, they are formed of two lobes[47]. I shall now consider them more particularly as to their station, number, and shape.

1. With regard to the first head, their station, they are most commonly divided between the trunk and abdomen; but in some cases, as in Hydrophilus piceus and Acrida viridissima, the first ganglion is in the head[48]; in others, as in the louse, the water-scorpion, and the grub of the rhinoceros-beetle, they are confined to the trunk, their functions in the abdomen being supplied[Pg 12] by numerous radiating nerves[49]; in others again, as in the scorpion, they are all abdominal. The ganglions vary also in their situation with respect to each other. Thus in some, as in the larva of the Chamæleon-fly (Stratyomis Chamæleon), they are so near as to appear like a string of beads[50]; in that of the ant-lion (Myrmeleon) the two ganglions of the trunk are separated by an interval from those of the abdomen, which are so contiguous as to resemble the rattle of the rattle-snake[51]. In others the internodes are longer, and the ganglions occur at nearly equal intervals, as in the larva of the Ephemeræ[52]; but in the majority they are unequal in length: thus in the scorpion the three first ganglions are the most distant[53]; in the hive-bee the third and fourth[54]; and in the spider the last[55].

2. The ganglions also in different species, and often in the same insect in its different states, vary in their number. Thus in the grub of the rhinoceros-beetle the whole spinal marrow appears like a single ganglion divided only by transverse furrows[56]; in the water-scorpion there are two[57]; in the louse there are three[58]; in the rhinoceros-beetle there are four[59]; five in the stag-beetle[60];[Pg 13] seven in the hive-bee and some Lepidoptera[61]; eight in the grub of the stag-beetle[62]; nine in the great Hydrophilus[63]; ten in Dytiscus[64]; eleven in the grub of the great Hydrophilus[65]; twelve in the grub of Dytiscus and the caterpillars of Lepidoptera[66]; thirteen in the larva of Æshna[67]; and twenty-four in Scolopendra morsitans[68]. You must observe that, generally speaking, the number of ganglions is less in the imago than in the larva. With regard to the distribution of these knots to the different primary parts of the body, the following table will exhibit it, as far as I am acquainted with it, at one view. I omit those in which the ganglions are only in one of these parts.

3. I am next to say a few words upon the shape of the ganglions. Most commonly it approaches to a spherical figure, but in many instances, as I said before, they, as[Pg 14] well as the brain, consist of two lobes: they are, however, seldom all precisely of the same shape. In the Dytisci, and Carabi, the last is marked with a transverse furrow, which seems to indicate the reunion of two[73]; in the stag-beetle, the first ganglion is oval or elliptical, the second hexagonal; the third and fourth shaped like a crescent, and the last like an olive[74]; in the caterpillar of the great goat-moth the first is oblong and constricted in the middle, and the seven last are rhomboidal[75]; in the great Hydrophilus the second, and in the silk-worm all the ganglions are quadrangular[76]; in Hypogymna dispar the third is heart-shaped[77]; the great ganglion which forms the spinal marrow of the cheese-maggot is pear-shaped[78]; that of the grub of the rhinoceros-beetle is fusiform[79]; and in the scorpion all the ganglions are lenticular[80]. But the most remarkable in this respect are those of a spider (Clubiona atrox): in this insect the brain sits upon a bilobed ganglion of the ordinary form, which is immediately followed without any internode by another bilobed one, terminating on each side in four pear-shaped processes or fingers, which give it a very singular appearance[81].

iii. The nerves[82] of insects, as of other animals, are white filaments running from the brain and spinal marrow to every part of the body which they are destined to animate; and their numerous ramifications, when delineated,[Pg 15] form no unpleasing picture[83]. In the caterpillar of the goat-moth the accurate Lyonet counted forty-five pairs of them, and two single ones, making in all ninety-two nerves; whereas in the human body anatomists count only seventy-eight[84]. From the brain issue several pairs, which go to the eyes, antennæ, palpi, and other parts of the mouth: sometimes those that render to the mandibles issue from the first ganglion, as in the larva of Dytiscus marginalis, the stag-beetle, &c.[85]; those both of mandibles and palpi in the great Hydrophilus[86]; and in Blatta some which act also upon the antennæ[87].

The optic are usually the most conspicuous and remarkable of the nerves. In some insects with large eyes, as many Neuroptera, Hymenoptera, and Diptera, their size is considerable; in the hive-bee they present the appearance of a pair of kidney-shaped lobes, larger than the brain[88]; in the dragon-flies, whose brain consists of two very minute lobes, these nerves dilate into two large plates of a similar shape, which line all the inner surface of the eyes[89]; in the stag-beetle they are pear-shaped, and terminate in a bulb, from which issue an infinity of minute nerves[90]; it is probable that this takes place in all cases, and that a separate nerve renders to every separate lens in a compound eye[91]; the optic nerve in Dytiscus and Carabus is pyramidal, with the base of the pyramid at the eye and the summit at the brain[92]; in Eristalis tenax it[Pg 16] is very large, cylindrical, and of a diameter equal to the length of the last-mentioned part, upon the side of which it is supported; it terminates in a very large bulb corresponding to the eye[93]: in Scolopendra morsitans the optic nerves divide into four branches long before they arrive at the eyes, and in this insect the nerves which render to the antennæ are so thick as to appear portions of the brain, which they equal in diameter[94]. Swammerdam discovered in the grub of the rhinoceros-beetle and in the caterpillar of the silk-worm, a pair of nerves which he regarded as analogous to the recurrent nerves in the human subject, and therefore he distinguishes them by the same name[95]: they issue from the lower surface of the brain, or that which rests on the œsophagus, and at first go towards the mouth, but afterwards turn back, and uniting form a small ganglion; this produces a single nerve, which passing below the brain follows the œsophagus to the stomach, where it swells into another ganglion, from which issue some small nerves that render to the stomach, and one more considerable which accompanies the intestinal canal, producing at intervals lateral filaments which lose themselves in the tunics of that tube[96]. Lyonet afterwards discovered these nerves in the caterpillar of the goat-moth[97], and Cuvier in other insects[98].

The other nerves which issue from the brain exhibit no remarkable features. Those which originate in the spinal marrow are mostly derived from the ganglions, and[Pg 17] are sometimes interwoven with the muscles, as the woof with the warp in a piece of cloth[99]; those from the three or four first commonly rendering to the muscles of the legs, wings, and other parts of the trunk, and those from the remainder to the abdomen. After their origin they often divide and subdivide, and terminate in numerous ramifications that connect every part of the body with the sensorium commune. A pair of nerves is the most usual number that proceeds from each side of a ganglion[100]; but this is by no means constant, since in the louse, the hive-bee, and several other insects, only a single nerve thus proceeds[101]; and in the larva of Ephemeræ, while two pairs issue from the six first ganglions, only a single one is emitted by the five last[102]. In the spinal marrow of the rhinoceros-beetle, both larva and imago, the nerves consist of simple filaments which diverge like rays in all directions[103]: the same circumstance distinguishes the cheese-maggot, only some of the nerves appear to branch at the end[104]: in the louse, the last ganglion sends forth posteriorly three pairs of nerves which render to the abdomen[105]. Sometimes, though rarely, nerves originate in the internodes of the spinal marrow. Cuvier indeed has asserted that in invertebrate animals all the nerves spring from the ganglions, and never immediately from the spinal marrow; but Swammerdam, in describing those of the silk-worm, mentions and figures four pairs as proceeding from the four anterior internodes, excluding[Pg 18] the first[106]; and at the same time he gives it as his opinion, that all the nerves in insects really originate from the marrow itself, and not from the ganglions, which he asserts are of a different substance, and are inclosed in the marrow for the sake of giving it greater firmness[107]. In this opinion, however, he seems singular[108]. Those remarkable nerves described by Lyonet under the name of spinal bridle (bride épinière) also take their origin, not from the ganglions, but from a bifurcation of the spinal marrow. Of these, in the caterpillar of the goat-moth there are ten, the first issuing from the bifurcation of the internode between the fourth and fifth ganglions, and the remainder from the succeeding ones. After approaching the succeeding ganglion, these nerves form a pair of branches that diverge nearly at right angles from the bridle, and producing several lesser branches, lose themselves in the sides of the animal[109]. Besides the nerves above mentioned, two generally issue from the posterior part of the last ganglion, diverging in opposite and oblique directions: some of these render to the parts of generation; and in the silk-worm, and probably other species, the innermost pair is perforated for the passage of the vasa deferentia[110].

[Pg 19]

After duly considering this general outline of the nervous system of insects, the question will continually occur to you,—is then what you have called the brain the sensorium commune of these animals, in the same manner as it is in those with warm blood? To this query a negative must be returned. In the latter, the brain is the common centre to which, by means of the nerves and spinal marrow, all the sensations of the animal are conveyed, and in which all its perceptions terminate. The nerves and spinal marrow are merely the roads by which the sensations travel; and if their communication with the brain, by any means be cut off at the neck, the whole trunk of the animal becomes paralytic, evidently proving that the organ by which it feels is the brain. This, however, is so far from being the case in insects, that in them, if the head be cut off, the remainder of the body will continue to give proofs of life and sensation longer than the head: both portions will live after the separation, sometimes for a considerable period; but the largest will survive the longest, and will move, walk, and occasionally even fly, at first almost as actively without the head, as when united to it. Lyonet informs us, that he has seen motion in the body of a wasp three days after it had been separated from the head; and that a caterpillar even walked some days after that operation; and when touched, the headless animal made the same movements as when intire[111]. Dr. Shaw has observed—an observation confirmed in Unzer's Kleine Schriften,—that if Geophilus electricus be cut in two, the halves will live and appear vigorous even for a fortnight afterwards; and what is more remarkable, that the tail part always survives the[Pg 20] head two or three days[112]. The sensorium commune of insects, therefore, does not, as in the warm-blooded animals, reside in the brain alone, but in the spinal marrow also. It was on this account probably that Linné denied the existence of a brain in insects, regarding it merely as the first ganglion of the spine.

Cuvier and other modern physiologists, from the ganglionic structure of this organ, are of opinion that it is not the analogue of the cerebro-spinal system of vertebrate animals, but rather of their great sympathetic nerves. Indeed, considering solely the external structure of the nervous system of insects, a great resemblance strikes us between it and these nerves; for besides its general ganglionic structure, there is also in them an upper ganglion in the neck, seemingly corresponding with what we have named the brain of insects, from which the nervous chord dips to the lower part of the neck, where it forms a second ganglion, which appears to correspond with what we have considered as their second ganglion[113]. We may observe, however, that at least in one respect there is even an external resemblance between the brain of insects and that of vertebrate animals:—it most commonly consists, as has been stated, like them, of two lobes, often very distinct; a circumstance which not unfrequently distinguishes the other ganglions[114], and is not borrowed from the ganglions of the great sympathetics. With respect[Pg 21] to the internal structure of the ganglions and spinal marrow of insects, we know little to build any theory upon, except that the internal substance of the former is filled with air-vessels; at least so Lyonet, as has been already observed, found in the goat-moth, while only the tunics of the latter are covered by them. Taking the above resemblance to the brain of vertebrates into consideration, there appears ground for thinking that the nervous system of insects, like some of their articulations[115], is of a mixed kind, combining in it both the cerebro-spinal and the ganglionic systems; and this will appear further if we consider its functions.

That learned and acute physiologist Dr. Virey, assuming as an hypothesis, that the structure of the system in question is simply ganglionic, and merely analogous to the sympathetic system of vertebrate animals, has built a theory upon the assumption, which appears evidently contradicted by facts. Because, as he conceives after Cuvier, insects are not gifted with a real brain and spinal marrow, he would make it a necessary consequence that they have no degree of intellect, no memory, judgement or free will; but are guided in every respect by instinct and spontaneous impulses,—that they are incapable of instruction, and can superadd no acquired habits to those which are instinctive and inbred[116]. This consequence would certainly necessarily follow, was their nervous system perfectly analogous to the sympathetic of warm-blooded animals. But when we come to take into consideration the functions that in insects this system confessedly discharges, we are led to doubt very strongly[Pg 22] the correctness of the assumption. Now in these animals the system in question not only renders to the nutritive and reproductive organs, which is the principal function of the great sympathetic nerves in the vertebrates; but by the common organs maintains a connexion with the external world, and acquires ideas of things without, which in them is a function of the cerebral system: from the same centre also issue those powers which at the bidding of the will put the limbs in action, which also belongs to the cerebral system. That insects have memory, and consequently a real brain, has been before largely proved, as also that they have that degree of intellect and judgement which enables them to profit by the notices furnished by their senses[117]. What can be the use of eyes,—of the senses of hearing, smelling, feeling, &c. if they are not instructed by them what to choose and what to avoid? And if they are thus instructed—they must have sufficient intellect to apprehend it, and a portion of free will to enable them to act according to it. With regard to the assertion that they are incapable of instruction, or of acquiring new habits; few or no experiments have been tried with the express purpose of ascertaining this point: but some well-authenticated facts are related, from which it seems to result that insects may be taught some things, and acquire habits not instinctive. They could scarcely be brought from their wild state, and domesticated, as bees have been so universally, and both ants and wasps occasionally[118], without some departure from the habits of their wild state; and the fact of the corsair-bees, that acquire predatory habits[Pg 23] before described[119], shows this more evidently: but one of the most remarkable stories to our purpose upon record, is that of M. Pelisson, who, when he was confined in the Bastile, tamed a spider, and taught it to come for food at the sound of an instrument. A manufacturer also in Paris, fed 800 spiders in an apartment, which became so tame that whenever he entered it, which he usually did bringing a dish filled with flies but not always, they immediately came down to him to receive their food[120].

All these circumstances having their due consideration and weight, it seems, I think, most probable, that as insects have their communication with the external world by means of certain organs in connexion with their nervous system, and appear to have some degree of intellect, memory, and free will, all of which in the higher animals are functions of a cerebral system, and at the same time in other respects manifest those which are peculiar to the sympathetic system,—it is most probable, I say, as was above hinted, that in their system both are united.

I must bespeak your attention to a circumstance connected with the subject of this letter, which merits particular consideration: I mean the gradual change that takes place in the nervous system when insects undergo their metamorphoses; so that, except in the Orthoptera, Hemiptera, and Neuroptera Orders, in which no change is undergone, the number of ganglions of the spinal chord is less in the imago than in the larva. There seems an exception indeed to this rule in the case of the rhinoceros-beetle,[Pg 24] in the larva of which there is only one ganglion, while in the imago there are four[121]. But as this one ganglion occupies the whole spinal marrow, it is really of greater extent than the four of the imago; so that even in this case there is a concentration of the cerebral pulp. In some cases, as in Dytiscus marginalis, and Hydrophilus piceus[122], the imago has only one ganglion less than the larva, but more generally it loses four or five. Dr. Herold has traced the gradual changes that take place in the spinal marrow of the common cabbage-butterfly (Pontia Brassicæ), from the time that it has attained its full size to its assumption of the imago. Of these I shall now give you some account.

In the full-grown caterpillar, besides the brain there are eleven ganglions, the chords of the four first internodes being double, and the rest single: from each ganglion proceed two pairs of nerves, one from each side. In this the lobes of the brain form an angle with each other[123]. In two days the double chords mutually recede, so as to diminish the interval between the ganglions, and the single ones have become curved: thus the length of the spinal marrow is shortened about a fourth, and the fourth and fifth ganglions have made an approach to each other[124]. On the eighth day, when the insect has assumed the pupa but remains still in the skin of the caterpillar, the flexure of the internodes is much increased; the first ganglion is now united to the brain, and the fourth and fifth have joined each other, though they are still distinct; the spinal marrow has now lost considerably[Pg 25] more than a third of its length[125]. On the fourteenth day, the internodes, except the double ones, have become nearly straight again; the fourth and fifth ganglions have coalesced so as to form one, and the sixth and seventh have each lost their pairs of nerves[126]. Shortly after this, these last ganglions have nearly disappeared, and the chords of the three first internodes have again approached each other[127]. The next change exhibited is the absorption of the first ganglion by the brain, the union of the chords of the first internode, which is now straight, the approximation of the second and third ganglions, and the enlargement of the one formed by the union of the fourth and fifth, at the expense perhaps of the sixth and seventh, which have now intirely disappeared, and in their place is a very long internode. These united ganglions retain the pairs of nerves they had when separate[128]. Just before the assumption of the imago, the direction of the lobes of the brain becomes horizontal, the second and third ganglions unite, and the internode between the third and fourth is shortened[129]. Lastly, when the animal is become a butterfly, the second and third ganglions have coalesced, and are joined to that formed by the union of the fourth and fifth; a short isthmus or rather constriction, with an orifice, being their only separation: each of these united ganglions send forth laterally four pairs of nerves[130]. In his figure, Dr. Herold has not represented the orifice for the passage of the gullet, but doubtless one exists, which for an animal that imbibes only fluid food is probably[Pg 26] very minute. In Hypogymna dispar, we learn from Cuvier, this orifice is of that description, and of a triangular shape[131].

It can admit of no reasonable doubt that one of the principal intentions of these changes is to accommodate the nervous system to the altered functions of the animal in its new stage of existence, in which the antennæ, eyes, and other organs of the senses, as well as the limbs and muscles moving them, and the sexual organs, being very different from those of the larva, and if not wholly new, yet expanded from minute germs to their full size, may well demand corresponding changes in the structure of the nervous system by which they are acted upon.

But are these changes also concerned, as Dr. Virey conjectures, in producing that remarkable alteration which usually takes place between the instincts of the larva and imago? In order to answer this question, it will be requisite first to quote the ingenious illustration with which this able physiologist elucidates his ideas on this point. "The more readily," he observes, "to comprehend the action of instinct, let us compare the insect to one of those hand-organs in which a revolving cylinder presents different tunes noted at its surface, and pressing the keys of the pipes of the organ, gives birth to all the tones of a song: if the tune is to be changed, the cylinder must be pulled out or pushed in one or more notches, to present other notes to the keys. In the same manner let us suppose that nature has impressed or engraved certain determinations or notes of action, fixed in[Pg 27] a determinate series in the nervous system and the ganglions of the caterpillar, by which alone she lives, she will act according to a certain sequence of operations; and, so to speak, she will sing the air engraven within her. When she undergoes her metamorphosis into a butterfly, her nervous system being, if I may so express myself, pulled out a notch, like the cylinder, will present the notes of another tune, another series of instinctive operations; and the animal will even find itself as perfectly instructed and as capable of employing its new organs, as it was to use the old ones. The relations will be the same; it will always be the play of the instrument[132]."

This illustration is doubtless at the first glance very striking and plausible: but a closer examination will, I think, show, that, as in so many other instances in metaphysical reasoning, when fanciful analogies are substituted for a rigid adherence to stubborn facts, it is satisfactory only on a superficial view, and will not stand the test of investigation; and as this is a question intimately connected with what I have advanced on the subject of instinct in a former letter, I must be permitted to go somewhat into detail in considering it.

To prove his position, Dr. Virey ought at least to be able to show that, whenever a change takes place in the instincts of insects in their different states of larva and imago, a corresponding change takes place in the external structure of the nervous chord. But what are the facts? In three whole orders, viz. Orthoptera Hemiptera, and Neuroptera, as mentioned above[133], the structure[Pg 28] of the nervous chord is not changed; and yet we know that many tribes of these orders acquire instincts in their imago state altogether different from those which directed them in their state of larvæ. A perfect Locust, for instance, acquires the new instincts of using its wings; of undertaking those distant migrations of which so many remarkable instances were laid before you in a former letter[134]; and, if a female, of depositing its eggs in an appropriate situation. But if such striking changes in the instinct of these tribes can be effected without any perceptible alteration in the structure of the nervous chord, it is contrary to the received rules of philosophical induction to refer to this alteration the changes in the instincts of other tribes where it is found. Is it not far more probable that this alteration has in fact no connexion with the changes of instinct, but is solely concerned with those remarkable changes in the organs of sense and motion, which occur in the larva and imago states of the orders in which it is observed? In a common caterpillar, the form of the body, the legs, the eyes, and other organs of the senses, all strikingly differ from those of the imago; whereas, with the exception of the acquisition of new wings, a perfect locust differs little from its larva: so that we may reasonably expect a corresponding change, such as we find it, in the structure of the nervous chord of the lepidopterous insect, not called for in that of the neuropterous species, in which accordingly it does not take place.

This reasoning, in opposition to Dr. Virey's theory, that the changes of instinct depend on the altered structure of the nervous system, becomes greatly strengthened[Pg 29] when we advert to the higher classes of animals, which surely in any investigation of the nature of instinct ought to be closely kept in view; for the faculty, though often less perfect in them than in insects, is still of the same kind, and may consequently be expected to follow the same general laws. In a young swallow, for example, all its instincts are not developed at once any more than in an insect. The instinct which leads it to migrate does not appear for some months after its birth, and that of building a nest still later. But we have not the slightest ground for believing that these new instincts are preceded by any change in the structure of the great sympathetic nerve, or of any other portion of the nervous system: and the same may be said as to the sexual instincts developed in quadrupeds some years subsequent to their birth. If, then, these remarkable changes in the instinct of the higher classes of animals can take place independently of any visible change in the nerves, what substantial reason can be assigned why they may not also in the class of insects?

On the whole, I think you will agree with me, that there is nothing in Dr. Virey's hypothesis which should lead me to alter the opinion I have already so strongly expressed in a former letter[135], as to the insufficiency of the mechanical theories of instinct hitherto promulgated, adequately to explain all the phenomena; and unless they do this they are evidently of small value. Such theories as I have there adverted to may often seem to be supported by a few insulated facts, but with others, far more numerous, they are utterly at variance; and, to omit many other instances, I am strongly inclined to[Pg 30] doubt the possibility of satisfactorily explaining the variety of instincts exercised by a bee[136], or the extraordinary development of new ones in particular circumstances only[137], on any merely mechanical grounds.

And after all, even suppose it could be demonstratively shown that every instinct is as clearly dependent on secondary causes, as I have formerly admitted that some doubtless seem to be, yet what would this teach us as to the essential nature of instinct? We have advanced indeed a step; but still, as I have before observed in referring to the theories of Brown and Tucker, we have only placed the world upon the tortoise, and instinct, as to its essence, which is what we want to detect, is as mysterious as ever: just as, though we can clearly prove that the mind is acted upon by the senses, yet this throws no light upon the essential nature of the mind, which we are forced to admit is inscrutable, as if to teach us humility, and prevent our vainly fancying, that though allowed to discover some of the arcana of nature, we shall ever be able to penetrate into her inmost sanctuaries.

That Dr. Virey should regard instinct in insects as purely mechanical was the natural consequence of his denying them any portion of intellect; but his opinion cannot I think be consistently assented to, if it be the fact, as I have just shown[138], that they are not wholly devoid of the intellectual principle. Whatever is merely mechanical, must, under similar circumstances, always act precisely in the same way. An automaton once constructed, whilst its machinery remains in order, will invariably perform the same actions; and Des Cartes, when[Pg 31] he had constructed his celebrated female automaton, imagined that he had irrefragably proved his principle, that brutes are mere machines. But if, instead of losing himself in the wilds of metaphysical speculation, he had soberly attended to facts, he would have seen that the instinct of animals can be modified and counteracted by their intellect, and consequently cannot be regarded as simply mechanical. Though the instinctive impulse of an empty stomach powerfully impel a dog to gratify his appetite, yet, if he be well tutored, the fear of correction will make him abstain from the most tempting dainties: and in like manner a bee will quit the nectary of a flower, however amply replenished with sweets, if alarmed by any interruption. The ants on which Buonaparte amused himself with experiments at St. Helena, though they stormed his sugar-basin when defended by a fosse of water, controlled their instinct and desisted when it was surrounded with vinegar[139]: and in the remarkable instance communicated to Dr. Leach by Sir Joseph Banks, the instinct of a crippled spider so completely changed, that from a sedentary web-weaver it became a hunter[140]. There is evidently, therefore, no analogy between actions strictly mechanical and instincts, which, though they may often seem to be excited by mechanical causes, are liable to be restrained or modified by the connexion of the instinctive and intellectual faculties[141]; and while we are ignorant how this connexion takes place, it is obviously impossible to reason logically on the subject.

In thus denying that any existing mechanical theory of instinct is satisfactory, I by no means intend to assert[Pg 32] that instinct is purely intellectual. I have already given you my opinion[142], that it is not the effect of any immediate agency of the Deity; nor am I prepared to assent to the doctrine of a writer, who has in some respects written ably on the subject in question, who says, that "the Divine Energy does in reality act not immediately, but mediately, or through the medium of moral and intellectual influences upon the nature or consciousness of the creature, in the production of the various, and in many instances truly wonderful, actions which they perform[143]." The same objection applies to this as to so many other metaphysical theories, that it is not adequately supported by facts; and all theories not so supported are injurious to science in proportion as their plausibility is greater, by leading the student to relax in that observation of nature and attentive study of the instincts of animals, on which alone sound hypothesis on this subject can be ultimately founded.

I shall conclude these remarks on the nature of instinct with a few observations as to the circumstances in which insects may be supposed to be guided by this faculty, and those in which intellect seems to direct them. The bee, when it takes its flight to a field where flowers abound, is governed by intellect in the use of its senses; for these are given to it as guides: and when it arrives there, they direct it to the flowers, and enable it to ascertain which contains the treasures it is in search of; but having made this discovery, its instinct teaches it to imbibe the nectar and load its hind legs with pollen.—Again: its senses, aided by memory, enable it to retrace[Pg 33] its way to the hive, where instinct once more impels it in its various operations. So that when we ascribe a certain degree of intellect to these animals, we do not place them upon a par with man; since all the most wonderful parts of their economy, and those manipulations that exceed all our powers, we admit not to be the contrivance of the animals themselves, but the necessary results of faculties implanted in their constitution at the first creation by their Maker. I may further repeat, that the mere fact of being endowed with the external organs of sense, proves a certain degree of intellect in insects. For if in all their actions they were directed merely by their instinct, they might do as well without sight, hearing, smell, touch, &c. but having these senses and their organs, it seems to me a necessary consequence, that they must have a sufficient degree of intellect, memory, and judgement, to enable them advantageously to employ them.

There is this difference between intellect in man, and the rest of the animal creation. Their intellect teaches them to follow the lead of their senses, and make such use of the external world as their appetites or instincts incline them to,—and this is their wisdom; while the intellect of man, being associated with an immortal principle, and being in connexion with a world above that which his senses reveal to him, can, by aid derived from heaven, control those senses, and bring under his instinctive appetites, so as to render them obedient to the το ἡγεμονικον, or governing power of his nature: and this is his wisdom.

I am, &c.

[Pg 34]

LETTER XXXVIII.

INTERNAL ANATOMY AND PHYSIOLOGY OF INSECTS CONTINUED.

RESPIRATION.

"Life and flame have this in common," says Cuvier, "that neither the one nor the other can subsist without air; all living beings, from man to the most minute vegetable, perish when they are utterly deprived of that fluid[144]." The ancients, however, not perceiving insects to be furnished with any thing resembling lungs, took it for granted that they did not breathe; though Pliny seems to hesitate on the subject[145]. But the microscopic and anatomical observations of Malpighi, Swammerdam and Lyonet, and the experiments of more modern physiologists, have incontestably proved that insects are provided with respiratory organs, and that the respiration of air is as necessary to them as to other animals. They can exist indeed for a time in irrespirable air; and immersion in hydrogen or carbonic acid gases is not, as I have often ascertained, so instantly fatal to them as it would be to vertebrate animals; but like them, they[Pg 35] speedily perish in air altogether deprived of its oxygen, or placed in situations to which all access to this essential element is excluded. Their respiration too of atmospheric air produces the same change in it with that of the vertebrate animals, the oxygen disappearing, and carbonic acid gas being produced in its place. Boyle had long since ascertained, that when bees, flies, and other insects were placed under an exhausted receiver, they often perished[146]: and the same effect was even observed by the ancients to ensue, when their bodies were by any means covered with oil or grease, which necessarily closed the orifices of their respiratory organs[147].

But for the first series of experiments ascertaining the necessity of a supply of air to insects, and their conversion of it into carbonic acid, we are indebted to the illustrious Scheele[148]; and his experiments have been repeated and confirmed by Spallanzani, Vauquelin, and other chemists. The former found, that when caterpillars and maggots were confined in vessels containing only about eleven cubic inches of atmospheric air, though furnished with sufficient food, they soon died, and sooner when the space was more confined[149]. He ascertained too, that a larva weighing only a few grains consumed, in a given time, as much oxygen as an amphibious animal a thousand times as voluminous[150]. A male grasshopper (Acrida viridissima) in six cubic inches of oxygen lived but eighteen hours, and the female placed in eight cubic inches of atmospheric air, only thirty-six hours. The[Pg 36] usual tests in both instances detected the conversion of the oxygen present into carbonic acid[151]. Precisely the same result was obtained by Sorg and Ellis, who, having placed a number of flies in nine cubic inches of atmospheric air, found them all dead by the third day, the oxygen intirely vanished, and a quantity of carbonic acid nearly equal in bulk produced[152].

It is ascertained too, that insects like other animals require in the process of respiration not merely oxygen, but such a mixture of it with nitrogen or azote as composes atmospheric air: for Vauquelin found that a grasshopper placed in six cubic inches of oxygen lived only half as long (eighteen hours) as another placed in eight inches of atmospheric air; its breathing was much more laborious, and it died when not more than one-twentieth of the oxygen had been converted into carbonic acid[153]. That a large quantity of oxygen penetrates all parts of insects, is evident also from the acid prevalent in the fluids of most of them, as likewise from the wonderful power of their muscles. That azote is also received, seems probable from the ammonia which has been extracted from the fluids of many, and from the rapid putrescence of these animals[154].

The mode, however, in which the respiration of insects is carried on, differs greatly from that which obtains in the higher animals. They have no lungs, no organs confined to a particular part of the body, by means of which the whole of the blood is regularly exposed to the[Pg 37] action of the inspired air. They do not breathe through the mouth, but through numerous orifices called spiracles, and the respiratory vessels connected with these are conducted to every part of the body. In some indeed, that we have included under the denomination of insects, as the Arachnida, an approach is made to the branchial respiration of fishes.

The respiratory apparatus of insects may be considered under two principal heads:—viz. the orifices or spiracles, and other external organs by which the air is alternately received and expelled; and the internal ones, by which it is distributed. Each of these is well worthy of your attention.

I. The external respiratory organs of insects may be divided into three kinds. Spiracles; Respiratory plates; and branchiform and other pneumatic appendages.

i. Spiracles[155] (Spiracula), or breathing pores, are small orifices in the trunk or abdomen of insects, opening into the tracheæ, by which the air enters the body, or is expelled from it[156]. They may be considered principally as to their composition and substance; shape; colour; magnitude; situation; and number.

1. Composition and substance. Perhaps you may not be aware that the structure of these minute apertures is not so simple as at the first view it may seem; but when you recollect that by them the insect breathes, you will suspect that provision may be made for their opening and shutting. A spiracle therefore, speaking analogically,[Pg 38] may be regarded in numerous cases as a mouth closed by lips. In caterpillars and many other insects, the substance of the crust where it surrounds the spiracle, is elevated so as to form a ring round it. The lips, properly speaking, are formed of a single cartilaginous piece or platform, with a central longitudinal cleft or opening, when closed often extending the whole length of the piece[157]; but in some appearing always open and circular: of the former description are those covered by the elytra in the common cockchafer; and of the latter, those that are not so covered: in some, as in the antepectoral pair of the mole-cricket, there appear to be no lips, the orifice being merely closed with hairs[158]. Though the aperture is usually in the middle of the platform, in the female of Dytiscus marginalis, it is nearer the posterior side, the anterior or upper lip being the longest. In the majority, the mouth or cleft is nearly as long as the spiracle; yet in the puss-moth (Cerura Vinula) it is shorter[159]. Some spiracles, however, are unilabiate, or have only one lip. This is the case with Gonyleptes and perhaps others[160]. The lips are usually horizontal, but sometimes they dip so as to make the spiracle appear open.

With regard to the substance of these organs, it is more or less cartilaginous, and probably elastic; the surface frequently appears to be corrugate or plaited; this is very distinctly seen in the stag-beetle and the cockchafer: in the last insect, under a powerful magnifier, we are told that the lips appear to consist of parallel cartilaginous processes, separated by a cellular web[161]. In some species[Pg 39] of Copris the corrugations form a perplexed labyrinth; in the caterpillar of the puss-moth the plaits are so narrow as to look like rays[162]; and in some Dynastidæ the lips approach to a lamellated structure. Again, in Hydrophilus caraboides the upper lip, and in Dytiscus circumflexus, both lips seem formed of elegant plumes[163]: a similar ornament distinguishes the inner edge of the lips in the caterpillar of the great goat-moth (Cossus ligniperda) and others[164]. In the grub of the rhinoceros-beetle (Oryctes nasicornis) the margin of the lower or inner lip is decorated by pinnated rays, which enter the cellular membrane that covers the upper lip[165]: in this larva, and that likewise of the cockchafer, the two lips are formed of different substances; in the last the upper or outer one consists of a perforated cellular membrane, through which the air can pass, while the lower or inner one is a cartilaginous valve that closes the orifice[166]: in the former this valve is surmounted by a boss[167]. In the pupa of Smerinthus Populi, a hawk-moth not uncommon, and of some dragon-flies (Libellula depressa), the margin of the two lips is crenated, probably with notches which alternate, that the mouth of the spiracle may shut more accurately[168]. The substance is unusually thick in the spinose caterpillars of butterflies; and in the pupa of one, Uria Proteus, it is villose.

Under the present head I may observe, that in some cases, as in the puss-moth, and the larva of the common[Pg 40] water-beetle (Dytiscus marginalis), the spiracles are closed by a semifluid substance, which however, according to Sprengel, is permeable to the air[169]. The animal, where these organs are furnished with lips, has doubtless, by means of a muscular apparatus, the power of opening and shutting them: this is done, we are told, by elevating and depressing, or rather by contracting and relaxing them. Sorg counted in one case (Oryctes nasicornis) twenty, and in another (Acrida viridissima) fifty, of these motions to take place in little more than two minutes[170]: but the quickness and force of this motion is not always uniform; for the same physiologist observed, that in Carabus auratus, when feeding or moving its body rapidly, the contraction of the spiracles took place at very short intervals; but when it was fasting, and its motions were slow, the intervals were longer[171]: it is probable also, that the temperature may accelerate or retard the motion. In the summer I examined a specimen of Phyllopertha horticola, that had indeed been somewhat injured, with this view: the pulses of the abdomen, which alternately rose and fell, were at about the rate of the pulse of a man in health, sixty in a minute, and the spiracles appeared to me to keep pace with this motion: later in the year, when the temperature was lower, as I was walking, I took a specimen of some grasshopper (Locusta).[Pg 41] Upon viewing it under a lens, I observed one of the convex pectoral spiracles open and shut, and the interval between two breathings appeared nearly half a minute.

2. With regard to their shape, spiracles vary considerably. In general we may observe that the abdominal ones are usually flat, while those of the trunk are often convex[172]. Sometimes they are very narrow and nearly linear, as in many pupæ of Lepidoptera, and those in the metathorax of the sand-wasps (Ammophila) and affinities; at others they are wider and nearly elliptical, as in Lucanus and many Lamellicorn beetles: again, in Copris they are circular; in Cordylia Palmarum ovate; in Dytiscus oblong[173]; in Goerius olens lunulate; in Gonyleptes nearly of the shape of a horse-shoe[174]; and probably many other forms might be traced, if a thorough investigation with this view were undertaken.

3. The colour of spiracles will not detain us long. In the caterpillars of Lepidoptera this is often so contrasted with that of the rest of the body, as to produce a striking and pleasing effect. Thus when the body is of a dark colour, they are usually of a pale one[175]; or if the body is pale, they are dark[176], or surrounded with a dark ring[177]. This contrast is often rendered more striking by their position with regard to the partial colours that often ornament caterpillars: in those whose sides are decorated by a longitudinal stripe, the spiracles are often planted in it[178]; or just above it[179]; or between two[180]: in[Pg 42] some hawkmoths the intermediate ones are set in white or pale spots, which gives great life to the appearance of the animal. In general, in perfect insects the most prevalent colour is buff, or reddish-yellow. In the larva of the great water-beetle these organs resemble the iris of the eye, being circular with concentric rings alternately pale and dark[181].

4. The size of spiracles varies considerably. Those in the larva last mentioned are so minute as to be scarcely visible except under a lens, while those behind the fore-legs in the mole-cricket are a full line in length, and those in the pleura of Acrocinus accentifer, a Brazilian Capricorn beetle, are more than twice as long. In the same species they are often found of different sizes;—thus the anal pairs in the water-beetle lately alluded to, I mean in the perfect insect, are much larger than the rest[182], probably that the animal may imbibe a larger quantity of air when it rises to the surface of the water, where it suspends itself by the tail. In those Lamellicorn beetles in which the terminal part of the abdomen is not protected by the elytra, the covered spiracles are the largest.

5. Under the next head, the situation of spiracles, I shall not only consider the part of the body in which they are situated, but likewise their position in the crust; to which last, as it will not detain us long, I shall first call your attention. Their position in this respect is most commonly oblique: but in the abdomen of the above water-beetle they are transverse, and in a larva I possess, probably of an Elater, they are longitudinal. In spinose[Pg 43] caterpillars these organs are generally planted between two spines, one being above and the other below. The lateral line of the body most commonly marks their situation; but in many cases they become ventral, and in others dorsal. The most important circumstance, however, connected with the present head is their appropriation to particular segments or parts of the body, for, like the ganglions of the spinal marrow, they are distributed to almost every segment. Let us take a summary view of their arrangement in this respect.

No insect has any spiracle in the head; but in caterpillars and many other larvæ there is a pair in the first segment of the trunk. This is also to be found in the other states, but is not easily detected in the pupæ of Lepidoptera: in the Coleoptera order, in the grub of the Lamellicorn beetles, it is extremely conspicuous, and planted in the side of the first segment[183]; in other Coleopterous grubs it is not so readily found, but probably its station is somewhere behind the base of the arms, where it is very visible in that of the Staphylinidæ. In the imago of insects of this order, this antepectoral spiracle has been overlooked, and indeed is not soon discovered: to see it clearly, the manitrunk should be separated from the alitrunk; and then if you examine the lower side of the cavity, you will see a pair of, usually, large spiracles planted just above the arms, in the ligament that unites these two parts of the trunk to each other: in the common rove-beetle, however, (Goerius olens)you may easily see it without dissection[184]. In the Orthoptera it is situated behind the arms, as in Gryllotalpa: or between them[Pg 44] and the prothorax, as in Blatta: in the Hemiptera and Neuroptera probably the situation is not very different. In the Lepidoptera this pair of spiracles is planted just before the base of the upper or primary wings[185]: a similar situation, I suspect, is appropriated to it in the Trichoptera, but covered by a tubercle or scale. Something similar has been noticed by M. Chabrier, in the same situation and circumstances, in the collar of Hymenoptera[186]. In numerous Diptera this breathing pore is planted on each side between the collar and the dorsolum above the arms[187], and in Hippobosca in the collar itself[188].

In Lepidopterous, Coleopterous, and some other larvæ, the two segments of the body corresponding with the alitrunk in the perfect insect, are without spiracles, neither have they in this state, though pneumatic organs have been discovered[189], any real ones in that part: but not so the remaining orders, all of which have these organs in that section of the trunk. To begin with the Orthoptera:—in Blatta there seems to be a long narrow one behind the intermediate leg; in the Gryllotalpa there is one in the posterior part of the pleura; and in Locusta, above both the intermediate and hind legs[190]. It is probable, that in general those that have no spiracles in the manitrunk have four in the alitrunk, which seems the natural number belonging to the trunk. In many of the Heteropterous Hemiptera in the parapleura[Pg 45] there is an open spiracle without lips[191], to which, as in that beautiful bug Scutellera Stockeri, a channel sometimes leads. The space in which this spiracle is planted in other genera of bugs (Pentatoma &c.) is covered with a kind of membranous skin, often much corrugated[192]. In the aquatic insects of this section, and many terrestrial ones, as Reduvius, &c. this spiracle is obsolete. There is another circumstance, possibly connected with their respiration, relating to many of the bugs, which may be mentioned here. If you examine Pentatoma rufipes, a very common one, you will find between the scapula and parapleura a long orifice or chink; this upon a closer inspection, under a good magnifier, you will see completely filled with minute stiff hairs or bristles, which fringe the posterior margin of the scapula[193]. In a Brazilian species of Lygæus (sexmaculatus K. M. S.) with incrassated posterior thighs, these hairs are replaced by lamellæ which have the aspect of gills. A red, vertical, convex spiracle, with its orifice towards the head, and terminating posteriorly in a kind of conical sac, is situated towards the hinder part of the pleura in the giant water-scorpion (Belostoma grandis[194]); this seems analogous to one lately mentioned in the mole cricket. In the other section of this Order it is not easy to decipher the parts of the under side of the alitrunk. In Fulgora, Cicada, and many others of its genera, there appears to be more than one opening into the chest; but whether they are of a pneumatic nature or not, can only be ascertained by an inspection of the living animal. There is a very visible[Pg 46] spiracle over each of the four last legs of the Libellulina[195], but in the remainder of the Neuroptera Order they have eluded my search. In the Hymenoptera and Diptera they are nearly in the same situation, being placed behind the wings on each side of the metathorax; in the latter Order with the poiser near them on the inner side[196]: in this also, the spiracles of the trunk are without lips, except in the larvæ, but are often merely an orifice, sometimes fringed with hairs; this is particularly conspicuous in Syrphus, in which these orifices are very large, and in some species closed by an elegant double fringe of white hairs. This is doubtless to prevent the entrance of any particles of dust or the like.

We are next to consider the situation of the spiracles of the abdomen: these which are supposed to be appropriated exclusively to inspiration, are usually more numerous than those of the trunk, by which it is probable that expiration is performed, and have principally attracted the notice of Entomologists: they are either dorsal, lateral, or ventral. In Dytiscus, Copris, &c. amongst the beetles, all the spiracles are dorsal; in the larvæ of Coleoptera and Lepidoptera they are lateral; and in the Heteropterous Hemiptera they are usually ventral: in Dynastes they are commonly found of all three descriptions;—the three first being dorsal, the two next lateral, and the last pair ventral[197]. In some instances, as in Perga Kirbii, and probably other Hymenoptera, these organs are planted in that portion of the dorsal segment which turns under, as was observed in a former letter[198],[Pg 47] and becomes ventral. Generally there is a pair of spiracles to each segment, and in those insects that have a hypochondriack joint[199] there is often a spiracle in it. The last segment of the abdomen is always without these orifices, as is the basal one in Velia, Ranatra, and some other bugs. A singular anomaly distinguishes the Libellulina: they appear to have no abdominal spiracles[200], yet I have seen the abdomen of Libellula depressa when reposing, contract and dilate alternately, from whence it follows that this part is concerned in respiration. Sprengel says that the larvæ in this tribe have seven or nine on each side[201], and Reaumur speaks of them as discoverable in the pupa[202]. I have carefully examined the pupa-skin of most of the genera of Libellulina, under a powerful magnifier, but have not succeeded in discovering any thing like these organs in the abdomen. The Ephemera and probably the other Neuroptera have abdominal spiracles[203]. M. Latreille observed one on each side of the base of the scale on the footstalk of the abdomen in ants[204]. Generally the abdominal spiracles may be described as planted in the crust of the insect; but in many cases their station is in the membranous folds, which I have therefore named the pulmonarium, that sometimes separate the dorsal from the ventral segments: these folds allow of a considerable distention of the abdomen, which is probably necessary when all the air-vessels are full. In a gravid Ichneumon I once saw it enlarged to more than twice its natural size by means of this membrane, through which the eggs were distinctly[Pg 48] visible.—Before I bid adieu to this subject, I must say a few words upon the situation of the organs in question in the myriapods. In Iulus, in each segment is a pair of orifices which have usually been regarded as spiracles, but M. Savi found that these orifices opened into vesicles containing a fetid fluid, and upon a very close examination he discovered the real spiracles above the base of the legs, in connexion with tracheæ[205]. In some of the larger species of Scolopendræ large open spiracles in the same situation are extremely visible[206]. Cermatia presents a singular anomaly:—a single series of spiracles of the usual form, each planted in a cleft of the posterior margin of the dorsal scuta, runs along the back of the animal[207]: unless we may suppose that, like the seeming spiracles of Iulus just mentioned, these are merely orifices by which it covers itself with some secretion.

6. A few words upon the number of spiracles.—If you examine the common dog-tick (Ixodes Ricinus), you will find only one of these organs on each side of the abdomen[208]; the Libellulina, as we have seen, have only four, all in the trunk; in the Dynastidæ, Melolontha, and the larva of Dytiscus, there are fourteen; sixteen in the Copridæ; eighteen in Dytiscus, and probably the majority of Coleoptera, both larva and imago, and Lepidoptera; and a pair to each segment except the last, in the Myriapods.

ii. Respiratory plates (Respiratoria). The nearest approach[Pg 49] to spiracles is made by those remarkable plates that are found in such larvæ of Diptera, as in that state inhabit substances that might impede or altogether stop the entrance or exit of the air by the ordinary spiracles, such as dead or living flesh, dung, or the like. The Creator therefore, as he has seen it good for wise reasons[209] to commission certain insects to feed on unclean food, has fitted them for the offices that devolve upon them, and has placed their orifices for breathing in plates at each extremity of the body. There are usually two of these plates at the head, and two at the tail. In the grub of the common flesh-fly (Sarcophaga carnaria), at the junction of the first segment of the body with the second, two of these plates are planted, which are concave and circular, with a denticulated margin; in the cavity near the lower side is a round spiracle. These plates the animal can withdraw within the body, so as to prevent this spiracle from being stopped up by any greasy substance[210]. The posterior extremity of this grub is truncated, and has a large and deep cavity surrounded by several fleshy prominences: at the bottom of this are two oval brown plates, in each of which are three oval spiracles, placed obliquely: by the contraction of the fleshy prominences, this cavity also can be closed at the will of the animal[211]. In some cases, several stiff rays or[Pg 50] spines replace the prominences[212]. In Echinomyia grossa and others the anal plates appear not to be perforated, being surmounted only by a central boss[213]; but this, most probably, as in the case of Œstrus Ovis[214], is a valve that closes the respiratory orifices. In the gad-fly of the ox (Œ. Bovis) there are no plates at the anterior extremity of the body; but those planted in the other end are very remarkable, and demand particular attention. Each is separated by a curved line into two unequal portions; the smallest of which is contiguous to the convex belly, and the largest to the concave back of the animal. This last is distinguished by two hard, brown, kidney-shaped pieces, a little elevated with the concave sides turned towards each other: in this sinus is a single, small, white spot, which appears to be a spiracle: in the smallest portion are eight minute circular orifices, arranged in a line[215]. As the only communication which this grub has with the atmosphere is at its anal extremity, it has no occasion for respiratory organs at the other. The gad-fly of the horse (Gasterophilus Equi, &c.) which has no communication at all with the external air, breathing that which is received into the stomach, has these plates at both ends of the body.

iii. Respiratory Appendages[216]. These may be divided into two kinds; those by which the animal has immediate communication with the atmosphere, and those by which it extracts air from water.

[Pg 51]

1. To begin with the first. These are often found in insects which, during their two first states, live in the water. No better example, nor one more easy to be examined, of this structure, can be selected, than the gnat (Culex). You must have occasionally observed in tubs of rain-water, numerous little wriggling worm-like animals, which frequently ascend to the surface; there remain a while, and then bending their head under the body rapidly sink to the bottom again. These are the larvæ of some species of the genus just named; and if you take one out of the water and examine it, you will perceive that it is furnished near the end of its body with a singular organ, which varies in length according to the species, and forms an angle with the last segment but one[217]. The mouth of this organ is tunnel-shaped, and terminates in five points like a star; and by this it is usually suspended at the surface of the water, and preserves its communication with the atmosphere: in its interior is a tube which is connected with the tracheæ, and terminates in several openings, visible under a microscope, at the mouth of the organ. The points or rays of the mouth when the animal is disposed to sink in the water, are used to close it, and cut off its communication with the atmosphere. When the animal is immersed, a globule of air remains attached to the end of the tube, so that it is in fact of less specific gravity than that element, and it is not without some effort that it descends to the bottom; but when it wishes to rise again, it has only to unclose the tube, and it rises without an effort to the surface, and remains suspended for any length of time. Its anal extremity is clothed with bunches of[Pg 52] hairs, which are furnished with some repellent material which prevents their becoming wet[218]: it is this repellent quality that probably causes a dimple or depression of the surface, which if you look narrowly you will discover round the mouth of the tube[219].

When the gnat undergoes its first change and assumes the pupa, instead of a single respiratory appendage it is furnished with a pair, each in shape resembling a cornucopia, and, what is remarkable, placed near the opposite extremity of the body, for they proceed from the upper side of the trunk[220]. By these tubular horns, which Reaumur compares to asses' ears[221], they respire, and are suspended at the surface.

Other respiratory tubes or horns are more complex. The rat-tailed grub of a fly (Helophilus pendulus), like the gnat, breathes by a tube: but as if the Creator willed to show those whose delight it is to investigate his works, by how many varying processes he can accomplish the same end, this respiratory organ is of a construction totally different from that we have been considering. It is not fixed to the side of the tail, but is a continuation of the tail itself, and is composed of two tubes, the inner one, like the tube of a telescope, being retractile within the other[222]. The extremity, which is very slender, and through which the air finds admission by a pair of spiracles, terminates in five diverging hairs or rays, which probably maintain it in equilibrio at its station at the surface[223]. As these larvæ seek their food[Pg 53] amongst the mud at the bottom of shallow pools, in which they are constantly employed, they require an apparatus capable of being lengthened or shortened, to suit the depth of the water, that they may maintain their necessary communication with the atmosphere; and for this purpose a single tube would not have been sufficient: therefore Providence has furnished them with two, and both are extremely elastic, consisting of annular fibres, so as to admit their being stretched to an extraordinary length. Reaumur found that these animals could extend their tails to near twelve times their own length. The mechanism by which the terminal piece is pushed forth or retracted, is very curious, though extremely simple. Two large parallel tracheæ, the direction of which is from the head[224] of the grub to its tail, occupy a considerable portion of its interior: near the origin of the tail, where they are very ample, they suddenly grow very small, so as to form a pair of very slender tubes, but so long that, in order to find room in a very contracted space, they form numerous zigzag folds attached to the terminal tube; when this issues from the outer tube they consequently begin to unfold, and when it is intirely disengaged, they are become quite straight and parallel to each other. Reaumur has figured them as being united at the base of the inner tube[225]; most probably, however, they do not here stop short, but, as in[Pg 54] other instances, proceed to the end, and terminate in the two spiracles mentioned above: he conjectures that when the animal has occasion to push forth its respiratory apparatus, it injects into these vessels part of the air contained in the body of the tracheæ, which of course would cause them to unfold and push forth the tube[226]. When this insect assumes the pupa, instead of its anal respiratory organ it has four respiratory horns in the trunk near the head[227].

The larva of the chamæleon-fly (Stratyomis Chamæleon) is furnished with a respiratory organ of a still different and more elegant structure, exhibiting some resemblance to the tentacula of what are called sea anemones. In this larva the last joint of the body is extremely long, and terminates in an orifice to receive the air, which is surrounded by a circle of about thirty diverging rays, consisting of beautifully feathered hairs or plumes[228]. This apparatus serves the same purpose with that above described of the larva of the gnat. The feathery hairs are so prepared as to repel the water, and thus to suspend the animal by its tail at the surface, and preserve a constant access of air. When it has occasion to sink, it turns these hairs in and shuts the orifice, carrying down with it an air-bubble that shines like quicksilver, and which Swammerdam conjectures enables it again to become buoyant when it wants to breathe[229].

In the red aquatic larva of a small gnat (Chironomus plumosus) there are two anal respiratory subcylindrical horns, with the orifice fringed with hairs[230]; and in another[Pg 55] gnat Reaumur discovered four[231]. The larva of Tanypus maculatus, whose remarkable legs I formerly noticed[232], exhibits in the interior of its trunk two long, oval, opaque bodies, which De Geer conjectures may be air-reservoirs; these, when the animal assumes the pupa, according to every appearance become external, and are placed on the back, precisely where the respiratory horns of aquatic pupæ are usually situated,—they appear to terminate in a transparent point[233]. The pupa of a Tipula observed by Reaumur, instead of two has only one of these respiratory organs, in the form of a very fine hair proceeding from the anterior end of the trunk, and considerably longer than the animal itself[234].

It is observable that aquatic insects that come to the surface of the water for air, receive it at the anus, often carrying it down with them as a brilliant bubble of quicksilver. This is generally done by means of spiracles in perfect insects, but in the water-scorpion tribe in that state respiration is by means of a long hollow tube, consisting of two concavo-convex pieces which apply exactly to each other. This is found in both sexes, and therefore cannot be an ovipositor, as some have thought[235].

These respiratory organs, however, are not invariably confined to aquatic larvæ and pupæ, for those of some aphidivorous flies have anal ones, and the pupa of Dolichopus nobilitatus, or a fly nearly related to it, which is terrestrial, has likewise a pair of long sigmoidal ones on the back of the trunk[236]. The pupa also of the rat-tailed[Pg 56] larva just noticed as having four horns, resides under the earth, the insect being only aquatic in its grub state.

2. I am next to consider those respiratory appendages by which aquatic insects, since they do not come to the surface for that purpose, appear to extract air for respiration from the water; so that they may be looked upon in some degree as analogous to the gills of fishes: there is, however, this difference between them—in fishes, the blood is conveyed in minute ramifications of the arteries to the surface of the branchial laminæ, through the membranes of which they abstract the air combined with the water; but as insects have no circulation, the process in them must be different, and their branchiform appendages may be regarded as presenting some analogy rather than any affinity to those of fishes. The first approach to this structure is exhibited by the pupa of a gnat lately mentioned (Chironomus plumosus); for on each side of the trunk this animal has a pencil consisting of five hairs elegantly feathered, which, when they diverge, form a beautiful star; its anus also is furnished with a fan-shaped pencil of diverging hairs[237].

On most of the abdominal segments of the larvæ and pupæ of the Trichoptera are a number of white membranous floating threads, arranged in bundles, four on each segment, two above and two below, and traversed longitudinally by several air-vessels or bronchiæ, which run in a serpentine direction, growing more slender as they approach the extremity, and in some places sending forth very fine ramifications,—these are their respiratory organs[238]. The caterpillar also of a little aquatic[Pg 57] moth (Hydrocampa stratiotata) at first sight appears to be covered on each side with hairs, but which examined under a microscope are found to be branching flattish filaments, each furnished with tubes from the tracheæ. These caterpillars have also the semblance of spiracles, but apparently found in the usual situation[239]. The larva of a little beetle often mentioned in my letters (Gyrinus Natator), is furnished on each side of every abdominal segment with a long, hairy, slender, acute, conical process, of the substance of the segment, through each of which an air-tube meanders; the last segment but one has four of these processes, longer than the rest[240].

Laminose or foliaceous respiratory appendages distinguish the sides of the abdomen of the larvæ and pupæ of the Ephemeræ, whose history you found so interesting[241]. In them these organs wear much the appearance of gills. In the different species they vary both in their number and structure. With regard to their number, some have only six pair of them, while others have seven. In their structure the variations are more numerous, and sometimes present to the admiring physiologist very beautiful forms[242]. They usually consist of two branches, but occasionally are single, with one part folding over the other, as in one figured by Reaumur, which precisely resembles the leaf of some plant, the air-vessels or bronchiæ in connexion with the tracheæ branching and traversing it in all directions, like the veins of leaves[243]. The double ones[Pg 58] differ in form. In the larva and pupa of Ephemera vulgata there are six of these double false gills on each side of the abdomen, the three last segments being without them; each branch consists of a long fusiform piece, rather tumid and terminating in a point, which is fringed on each side with a number of flattish filaments, blunt at the end. An air-vessel from the trachea enters the gill at its base; is first divided into two larger branches, each of which enters a branch of the false gill. These branches send forth on each side numerous lesser ramifications, one of which enters each of the filaments[244]. In another species (E. vespertina) each false gill presents the appearance of a pair of ovate leaves with a long acumen, and the air-vessels represent the midrib of the leaf, with veins branching from it on each side[245]; and, to name no more, in E. fusco-grisea, one branch represents the leaf of a Begonia, the sides not being symmetrical, with its veins, while the other consists only of numerous branching filaments[246]. In other aquatic larvæ, as in that of the common May-fly (Sialis lutaria), these appendages consist of several joints[247].

By the above apparatus these aquatic animals are enabled to separate the air from the water, as the fish by their gills; but how this separation is made has not been precisely explained. The false gills in many species are kept in continual and intense agitation. When they move briskly to one side, Reaumur conjectures they may receive the air, and when they return back they may[Pg 59] emit it[248]. This brisk motion probably disengages it from the water. In many species, when in repose, they are laid upon the back of the animal[249], but in others they are not[250].

The larvæ of the Agrionidæ appear to respire like those of the Ephemeræ, &c. by means of long foliaceous laminæ or false gills filled with air-vessels; but instead of being ventral, they proceed from the anus. They are three in number, one dorsal and two lateral, perpendicular to the horizon, of a lanceolate shape, beautifully veined, with a longitudinal middle nervure, from which others diverge towards the margin, which are probably bronchiæ. They are used by the animal, which swims like a fish, as fins, but it does not appear to imbibe the water like the other Libellulinæ, nor to propel itself by ejecting it,—a circumstance which furnishes an additional argument for the more received opinion, that this action in them is for the purpose of respiration as much as for motion[251].

The larvæ and pupæ of the Libellulinæ, receive the water and air that they respire by a large anal aperture, which is closed at the will of the animal by five hard, moveable, triangular, concavo-convex pieces, all very acute and fringed with hairs. These pieces are placed so that there is one above, which is the largest of all; one on each side, which are the smallest, and two below; when these are closed they form together a conical point[252]. Sometimes only three of these pieces are conspicuous[253]:[Pg 60] three other cartilaginous pieces, resembling the valves of a bivalve shell, close the passage within the pointed pieces[254]. At this orifice the water is received; and when, by an internal process to be described afterwards, it has parted with its oxygen, is again expelled.

Under this head I shall mention a fact which may be connected with respiration of the insects concerned. In dissecting a moth related to Catocala Pronuba, but I do not recollect the particular species,—at the base of the abdomen of the male I discovered two bunches of long fawn-coloured parallel hairs, planted each in an oval plate, plane above, but below convex and fleshy; while the plates remained attached to the insect, they appeared to have a distinct pulsation. The hairs, which are about half an inch long, diverge a little, and form a tuft not very unlike a shaving-brush[255]. I have not since met with this species, but I have preserved the brush and scale. Somewhere in Bonnet's works, but I do not recollect where, I have since found mention of a similar fact in another moth.

II. Having considered the external respiratory organs of insects, by which the air is received, we are next to consider the internal ones, by which it is distributed. These are gills; tracheæ and bronchiæ; and sacs or pouches[256].

i. Gills (Branchiæ[257]). Having lately described what[Pg 61] may be denominated false gills, or branchiform appendages, I shall now call your attention to what may be denominated true ones, which are peculiar to the Arachnida Class: but what is remarkable, the animals that breathe by them are very rarely inhabitants of the water, so that their functions cannot be perfectly analogous to those of fishes.

In the Scorpion, on each side of the four first ventral segments a spiracle may be discovered, which has no lip as in other insects, but is merely a circular orifice. These orifices do not lead to tracheæ or vesicles, but to true gills, which are situated below a muscular web which clothes the internal surface of the crust. Each gill consists of many semicircular very thin plates, of a dead milky white, which are connected together at the dorsal end like the leaves of a book. There appear to be more than twenty of these leaves, which when strongly magnified look transparent and destitute of any vessels. Each gill is fastened at the back to the spiracle[258]. In the spiders also, gills are discoverable, but differently circumstanced. On the under side of the abdomen, near the base, is a transverse depression, on each side of which is a longitudinal opening leading to a cavity, which is covered from above by a cartilaginous plate. In this cavity is situated a true gill, which is white, triangular, and covered with a fine skin; the leaves of this gill are far more numerous and much finer and softer than those of the gills of the scorpion. On account of their softness they have often the appearance of a slimy skin; but their[Pg 62] laminated structure shows itself very clearly in old specimens, and in such as have been immersed in boiling water[259].

ii. Tracheæ and Bronchiæ[260]. Parallel with each side of the body of most insects and extending its whole length, run two cylindrical tubes[261], which communicate with the spiracles[262], and from which issue, at points opposite to those organs, other tubes which ramify ad infinitum, and are distributed to every part of the body[263]. The first of these tubes are called the tracheæ and the latter the bronchiæ. This structure appears, however, not to be universal: it is to be found in caterpillars and many Dipterous larvæ; but in that of the rhinoceros-beetle and other Lamellicorns, the bronchiæ branch directly from the spiracle, the bottom or interior mouth of which is lined by a membrane from which they proceed[263]: something similar has been observed to take place in many insects in other states, as the common cockchafer[264]; in the pupa of Smerinthus Populi[265]; in the Cicadæ[266]; in the Locust tribe[267]; and many others. In the Cossus, or larva of the great goat-moth, the trachea commences with the first spiracle, and finishes a little beyond the last, after which it diminishes considerably in diameter, and terminates in several branches or bronchiæ, which proceed to the anal extremity of the body[268]. The bronchiæ which originate from the tracheæ in the[Pg 63] vicinity of each spiracle, may be considered as consisting in general of three packets;—dorsal ones, which are distributed to the back and sides of the animal; visceral ones, which enter the cavity of the body, and are lost amongst the viscera and the caul; and ventral ones, which dipping from the tracheæ overrun the lower part of the sides and belly[269].

The tracheæ and bronchiæ consist of three tunics[270]: the first or external one is a thickish membrane, strengthened by a vast number of fibres or vessels, which form round it a number of irregular circles; the second is a membrane more thin and transparent, without a vascular covering[271]; the third is formed of a cartilaginous thread running in a spiral direction, which may be easily unwound[272]. This structure gives a great elasticity to these organs, so that they are capable of considerable tension, after which they return to their usual length[273]. The Bronchiæ are cylindrical or slightly conical, insensibly diminishing in size as they leave the trunk, in which they originate. In larvæ, after losing their spiral fibre, they appear to terminate in membrane, but in perfect insects they pass into vesicles[274]. In the Cossus the trachea is flattened, and in every segment, except the first and two last, is bound by a fleshy cord four or five times as thick as its threads. Where this occurs, there is a slight constriction,—probably here is a sphincter, by the contraction[Pg 64] of which Lyonet supposes the trachea may be shut when it is necessary to stop the passage of the air, and direct it to any particular point[275]. The structure here described is admirably adapted for the purpose it is intended to serve; for had these vessels been composed of membrane, they could not possibly have been prevented from collapsing; but by the intervention of a spiral cartilaginous thread this accident is effectually guarded against, and the necessary tension of the tubes provided for. However violent the contortions of the insect, however small the diameter of these vessels, they are sure to remain constantly open, and pervious to the air. And by this circumstance they may be always distinguished from the other organs of the animal, and likewise by their pearly or silvery hue, for from being constantly filled with air, these tubes, when viewed under a powerful microscope in a recently dissected insect, present a most beautiful and brilliant appearance, resembling a branching tree of highly polished silver or pearl:—though sometimes they are blue, or of a lead colour, and sometimes assume a tint of gold. In the dead insect the larger tubes soon turn brown, but the finer ones preserve their lustre several weeks[276]. The ramifications of the tracheal tree may be seen without dissection through the transparent skin of the common louse[277] and most of the thin skinned larvæ.

You will not expect to view in this way the minuter ramifications of the bronchiæ, when I have mentioned their number and incredible smallness. Nothing but[Pg 65] the scalpel of a Lyonet and the most powerful lenses are adequate to trace the extremities of these vessels; and even with every help, they at last become so inconceivably slender as to elude the most piercing sight. That illustrious anatomist found that the two tracheæ of the larva of the Cossus gave birth to 236 bronchial tubes, and that these ramify into no less than 1336 smaller tubes, to which, if 232, the number of the detached bronchiæ, be added, the whole will amount to 1804 branches[278]. Surprising as this number may appear, it is not greater than we may readily conceive to be necessary for communicating with so many different parts. For, like the arterial and venous trees, which convey and return the blood to and from every part of the body in vertebrate animals, the bronchiæ are not only carried along the intestines and spinal marrow, each ganglion of which they penetrate and fill, but they are distributed also to the skin and every organ of the body, entering and traversing the legs and wings, the eyes, antennæ, and palpi, and accompanying the most minute nerves through their whole course[279]. How essential to the existence of the animal must the element be that is thus anxiously conveyed by a thousand channels, so exquisitely formed, to every minute part and portion of it! Upon considering this wonderful apparatus we may well exclaim, This hath God wrought, and this is the work of his hands.

Though in general there is only a pair of tracheæ, yet in some larvæ a larger number have been discovered.[Pg 66] In those of the Libellulinæ there are six. According to M. Cuvier, Reaumur, who mentions only four, overlooked the two lateral ones that are connected with the spiracles[280]. The reason of this and other parts of their internal structure I shall explain under the next head. In the grub of the gad-flies of the horse (Gasterophili,) Mr. B. Clark discovered eight longitudinal tracheæ,—six arranged in a circle and two minute ones, which appeared to him to terminate in a pair of external nipples (spiracles) in the neck of the animal[281]. This is a singular anomaly, as the other Œstridæ have only a pair of tracheæ[282].

iii. Respiratory Sacs or Pouches. Besides their tracheæ and bronchiæ, many insects are furnished with reservoirs for the air, under the form of sacs, pouches, or vesicles. These are commonly formed by the bronchial tubes being dilated at intervals, especially in the abdomen, into oblong inflated vesicles; from which other bronchial tubes diverge, and again at intervals expand into smaller vesicles, so as to exhibit no unapt resemblance—as Swammerdam has observed with respect to those of the rhinoceros-beetle—to a specimen of Fucus vesiculosus. Cuvier compares them in the Lamellicorn beetles in general to a tree very thickly laden with leaves[283]; and Chabrier observes that they particularly occur in the intestinal canal[284]. This structure of the pulmonary organs may be seen also in the common hive-bee, and other Hymenoptera;[Pg 67] but the vesicles are less numerous, and those at the base of the abdomen much larger than the rest[285]. These vesicles, by a very rough dissection, may be distinctly seen in the abdomen of the cockchafer, which appears to be almost filled with them. Not being composed of cartilaginous rings like the air-tubes, but of mere membrane, if a pin pierces one, the air that inflates it escapes, and it collapses. In the larva of a little gnat (Corethra culiciformis) the tracheæ appear to proceed from a pair of oblong vesicles of considerable size[286] in the trunk, and towards the anus they form two other smaller ones[287],—upon piercing the former, De Geer observed a considerable quantity of air to make its escape[288]. Another species, probably of the same genus, described by Reaumur, exhibits something similar[289].

But one of the most remarkable structures, in this respect, is to be seen in the larva and pupa of the dragon-flies (Libellulina). I have before noticed the number of their tracheæ, but I shall here describe their whole internal respiratory apparatus. I must observe that Reaumur, Cuvier, and most modern writers on the physiological department of Entomology, have affirmed that they respire the water, and that they receive it for that purpose at their anal extremity: but M. Sprengel, from having observed in the larvæ abdominal spiracles, is unwilling to admit this as a fact[290]; and De Geer also seems to hesitate upon it, especially as he discovered that the animal seemed to absorb the water to aid it in its motions[291].[Pg 68] But when we consider that it is by the action of a pneumatic apparatus that the absorption and expulsion of the water takes place, and that the animal when it has been taken out of that element, upon being restored to it, immediately has eager recourse to this action[292], we shall feel inclined rather to adopt the opinion of those great physiologists Reaumur, Lyonet, and Cuvier, and admit that it absorbs water for the purpose of respiration. I shall now explain how this takes place. The pieces both internal and external that close the anal orifice have been before described; the others employed in the admission and expulsion of the water are evidently respiratory organs. When this orifice is opened, the parts that are above it are drawn back in an opposite direction, so that the five last segments of the abdomen become entirely empty, and form a chamber to receive the water that enters by it. When the water is to be expelled, the whole mass of air-vessels which had receded towards the trunk, is pushed forwards, and forms a piston that again expels the water in a jet. It consists of an infinite number of bronchiæ, entangled with each other, which proceed from the middle and posterior end of the tracheæ. M. Cuvier in the interior of the rectum of the larva discovered twelve longitudinal rows of little black spots, in pairs, which exhibited the resemblance of six pinnated leaves. These are minute conical tubes, of the spiral structure of tracheæ, which decompose the water, and absorb the air contained in it. He also discovered that each of these tubes gave birth to another outside the rectum, which connected itself with one of the six great longitudinal tracheæ; two of which are of enormous size,[Pg 69] and appear to serve as reservoirs, since they furnish air by transverse branches to two other tubes; they have each a recurrent branch, which follows the course of the intestinal canal, and furnishes it with an infinity of bronchiæ[293]. These tracheæ are found in the perfect insect. The principal ones in some send forth many branches, terminating in vesicles, which in shape resemble the seed-vessels of some species of Thlaspi, while others appear to form a file of oblong ones[294]. Near each of their spiracles also is a vesicle which appears to be a reservoir[295].

But this kind of structure is not confined to insects strictly aquatic. Even such species of terrestrial ones as live upon aquatic plants, and are, consequently, necessarily or accidentally often a considerable time under water, are furnished with some apparatus by means of which they can exist in this element for a considerable period. For example, most of the Weevils (Rhyncophora) die in a short time if immersed in water; yet the species of the genera Tanysphyrus, Bagous, and Ceutorhynchus which feed on aquatic plants, can exist for days under water, as I have ascertained by experiment. C. leucogaster and another of the same tribe, swims like a Hydrophilus, and will live a long time in a bottle filled with water and corked tight. Other insects also, that are not at all aquatic, have pneumatic pouches. A striated or channeled vesicle I have found under the lateral angles of the collar in the humble-bee, where Chabrier supposes the vocal spiracles are situate; and also at the mouth of the spiracles of the metathorax in Vespa, &c.[296] In Sphinx[Pg 70] Ligustri the bronchiæ terminate in oblong vesiculoso-cellular bodies, almost like lungs[297]; in Smerinthus Tiliæ these are preceded by a simple vesicle bound with spiral fibres[298]. M. Chabrier thinks that these air-bladders of insects, amongst other functions, give more fixity and force to the muscles for flight[299].

Many physiologists have seen an analogy between the spiral vessels of plants and the tracheæ of insects; and some of great name, as Comparetti, Decandolle, and Kieser, have thought that in some instances they terminated in the oscula or cortical pores: but Sprengel contends that they are not accurate in this opinion[300]. In fact, the principal analogy seems to be in the spiral structure of both these vessels.

Having considered the different organs of respiration both external and internal, I shall make a few further observations upon this function. We know little more respecting the mode in which insects respire, except that they breathe out the air by the same kind of organs by which they receive it,—namely, the spiracles, or their representatives. This has been satisfactorily proved by Bonnet, who showed that the experiments by which Reaumur thought it established that insects inspire by their spiracles, but expire through the mouth, anus, or pores of the skin, are founded on an erroneous assumption.[Pg 71] This physiologist, having observed on the surface of submerged insects numerous bubbles of air, concluded that they had passed through the above orifices[301]: but Bonnet found by various experiments carefully conducted, that this appearance was caused by air which adhered to the skin and its hairs, and that when the access of this was precluded by carefully moistening the skin with water previously to immersion, this accumulation of air-bubbles on its surface did not take place[302]. And in a variety of instances he observed large ones issue from all the spiracles, especially the anterior ones. These bubbles sometimes were alternately emitted and absorbed without quitting the spiracle[303], and at others were darted with force to the surface of the water, where they appeared to burst with noise[304]. This author is of opinion that the first and last pair of these organs are of most importance to respiration[305]. Reaumur subsequently owned that Bonnet's arguments had shaken his opinion[306]; and some observations of his own, with respect to the respiration of the bot of the ox, go to prove that expiration and inspiration are not by the same spiracles; for he found that the air in this animal was expired by the eight little lower orifices before mentioned[307], from which he clearly saw the air-bubbles issue—the upper one he conjectures receives the air[308]. As the only communication that this grub has with the atmosphere is by its posterior extremity, it follows, reasoning from analogy, that the anterior respiratory plates of Dipterous larvæ, which may be regarded as representing the spiracles of the trunk in[Pg 72] insects in general, are destined for the escape of the air, after it has parted with its oxygen, received by the anal ones[309]. So that there seems very good ground for M. Chabrier's opinion that inspiration is ordinarily by the abdominal spiracles, and expiration by those of the trunk of insects[310]. He seems to have been led to the adoption of this opinion, not so much by experiments similar to that of Reaumur just stated, but by observing that in many instances these two sets of spiracles differ from each other, the latter having a convex and the former a concave mouth or bed[311]. In some cases, however,—for instance during flight,—he supposes the spiracles of the trunk may receive as well as emit the air[312]: he likewise is of opinion, and it seems not improbable, that by means of these openings in the trunk, from the rush of the superfluous air through them, insects produce those sounds for which they are remarkable,—as the humming of bees and flies. In the former he thinks the sound is produced by the pneumatic apparatus covered by the ends of the collar; while in the latter he attributes it to the spiracles in the metathorax behind the wings attended by a poiser[313]. I incline, however, to M. Dufour's opinion[314],—that the vocal spiracles in the Hymenoptera, as well as in the Diptera, are those behind the wings. Perhaps both theories may be right; for if you take any common humble-bee, you will find that, in the hand, it produces one kind of sound when its wings are motionless,[Pg 73] and another more complex and intense when they vibrate. In numerous instances, however, there is no very striking external difference between the spiracles of the trunk and those of the abdomen: this observation applies more particularly to the caterpillars of Lepidoptera; but whether these receive the air by those of the abdomen, and return it by those of the trunk, has not yet been ascertained; and indeed, too little is at present known upon the subject, and too few facts have been collected, to admit of dogmatizing.

The external signs of respiration in insects are not universally to be discovered. The alternate contraction and expansion of the abdomen is, however, very visible in some beetles, bees, the larger dragon-flies, and grasshoppers. In one of the latter, Acrida viridissima, Vauquelin observed that the inspirations were from fifty to fifty-five times in a minute in atmospheric air, and from sixty to sixty-five when in oxygen gas[315]. But M. Chabrier has given the most satisfactory account of these signs: The abdomen, says he, is the principal organ of inspiration; it can dilate and contract, lengthen and shorten, elevate and depress itself. In flight, in elevating its extremity at the same time with the wings, it contracts itself, pushes the air into the trunk, and diminishes the weight of the body by the centrifugal ascending force[316]. In the majority of insects perhaps the dilatation of the abdomen takes place by the recession of the segments from each other by means of the elastic ligaments that connect them; in others, as the Dynastidæ, Galeodes, &c. by the longitudinal folded membrane that unites the dorsal[Pg 74] and ventral segments—in the Libellulina by similar ventral folds; and in Cimbex by membranous pieces in the first dorsal segment, which De Geer observed was elevated and depressed at the will of the animal[317].

Air is as essential to insects in their pupa as in their larva or perfect states. Lyonet, however, Musschenbroek, Martinet, and some other physiologists, have doubted whether quiescent pupæ breathed[318]; but Reaumur and De Geer seem to have proved that they do[319]: and if thrown into water, the same proof of respiration, by the emission and retraction of a bubble of air takes place, as in the larvæ; and De Geer found that if one be transferred under water from one spiracle to another, it will be absorbed by it[320]. Indeed, unless these pupæ had breathed, where would have been the necessity for the spiracles with which all are furnished? It is remarkable, however, that all these spiracles do not seem of equal importance in this respect. Reaumur found that if the posterior spiracles only were closed with oil, the insect suffered no injury; but that if the anterior ones were similarly treated, it infallibly died[321]. The respiration however of pupæ seems more perfect in those that have recently assumed that state, than in those that are more advanced towards the imago; in which at first, from Reaumur's experiments[322], it appears that the posterior spiracles were stopped; and in others still older, from Musschenbroek's[323], even the anterior ones. Those quiescent pupæ that during that state remain submerged, respire[Pg 75] air. De Geer has given an interesting record of this, in the case of Hydrocampa stratiotata. This insect spins a double cocoon, the outer one thin, and the inner one of a close texture. In the pupa there are three pair of conspicuous spiracles on the second, third, and fourth segments of the abdomen, which are placed on cylindrical tubes, and they appear to have no other air-vessels. The respiratory gills of the larva having vanished, like some others of the same genus, they know how to surround themselves with an atmosphere of air in the midst of the water, so that the interior of their inner cocoon is impervious to the latter element—how they renew the air has not been ascertained. Though they respire air, water is equally necessary, for the animal died when kept out of water[324].

The great majority of insects respire in much the same manner in all their states, particularly as to their external organs; for when the larva breathes by the lateral spiracles, the pupa and imago usually do the same. The converse of this, however, by no means holds; for it not unfrequently happens that the two latter breathe by means of lateral spiracles, though they received the air in their larva state by an apparatus altogether different. Thus the larvæ of many Diptera breathe by an anal tube, while the pupa and imago follow the general system. Sometimes a tribe of insects breathe by an apparatus quite different in all their states, as we have seen to be the case with the common gnat[325], which has an anal respiratory tube in its first state, thoracic respiratory horns in its second, and the ordinary lateral spiracles in its third.

[Pg 76]

Changes also take place in their internal organs. In the larvæ the respiratory apparatus, especially the tracheal tubes, is often much larger and more ramified than in the imago; and as the former is the principal feeding state, there seems good ground for Mr. B. Clark's opinion—that the respiration is intimately connected with the conversion of the food[326]. In the imago, there appears to be more provision for storing up the air in vesicular reservoirs, than in the larva. Wonderful is the mode in which some of the changes in the internal structure, which these variations indicate, must necessarily take place. They are, however, probably not more singular than those which less obviously occur in the air-vessels of all insects in their great change out of the larva into the pupa state. But having before enlarged on this subject, I need not repeat my observations[327].

The access of air is as necessary to insects even in their egg state[328], and in many cases its presence seems provided for with equal care, by means as beautiful as those Sir H. Davy and Sir E. Home have shown to occur in the oxygenation of the eggs and fœtuses of vertebrate animals[329]. It is only necessary to view the admirable net-work of air-vessels which Swammerdam discovered spread over the surface of the eggs of the hive-bee while in the ovaries[330],—a provision which, from analogy, we may conclude obtains generally; from the importance[Pg 77] which nature has attached to the oxygenation of the germ while in the matrix. And judging from analogy, we may infer that the access of this element is as carefully secured after the egg is laid, as before. The eggs of most insects being of a porous texture, often attached to the leaves of plants, and some of them embedded in the very substance of a leaf or twig[331], are in a situation for the abundant absorption of oxygen: and the pouch of silk in which the eggs of spiders and Hydrophili are deposited, may probably, from Count Rumford's experiments, be of utility in the same point of view. In the case of the Trichoptera and other insects[332] whose eggs are dropped into the water enveloped in a mass of jelly, this substance perhaps serves for aërating the included embryo, in the same way with the jelly surrounding the eggs of the frog, dog-fish, &c. It would be desirable to ascertain whether the former jelly be of the same nature as the experiments of Mr. Brande have shown the latter to be[333]. It is not improbable that the singular rays that terminate the eggs of Nepa[334] may in some way be connected with the aëration of the egg.

To what I have before remarked with regard to the vital heat of insects[335], I may under this head very properly add a few further observations. I there stated, that the temperature of these animals is usually that of the medium they inhabit, but that bees, and perhaps other gregarious ones, furnish an exception to this rule[336]. A confirmation of this remark is afforded by Inch, a German[Pg 78] writer, who, upon putting a thermometer into a bee-hive in winter, found it stand 27° higher than in the open air; in an anthill, he found it 6° or 7° higher; in a vessel containing many blister-beetles (Cantharis vesicatoria,) 4° or 5° higher. A thermometer, standing in the air at 14° R., put into a glass vessel with Acrida viridissima, in nine minutes rose to 17°, and a similar result was observed with respect to other insects[337]. Dr. Martine says that caterpillars have but two degrees of heat above that of the air they live in[338]. Coleopterous insects are said to move slowly and with difficulty when the thermometer sinks to 36°, to become torpid at 34°, and to lose muscular irritability at a lower degree[339]. I have before observed that some insects will bear to be frozen into an icicle, and yet survive[340]: they share this power with reptiles, fishes, and amphibia. But, however small the excess of it in some insects above that of the medium they inhabit, it proves that they possess the power of generating heat. Whether, like the warm-blooded animals, they generally possess that of resisting heat by perspiration, &c. is not so clear. Yet the heat to which some can bear to be exposed, basking at noon, as Dr. Clarke informs us[341], on rocky and sandy places, exposed to the full action of the sun, appears sufficient, if not resisted by some principle of counteraction, to roast them to a cinder. That bees perspire is well known, but probably not singly.

When the respiration of insects is suspended by immersion[Pg 79] in any fluid, it is often resumed, even when it has been long and they are apparently dead, if they be brought into contact with the atmosphere. Reaumur found this to be the case with bees[342]; and Swammerdam tells us that the maggot of the cheese-fly (Tyrophaga Casei) lived six or seven days in rain-water[343]: he found it so difficult to kill the larva of Stratyomis Chamæleon, which he first immersed twenty-four hours in spirits of wine, and then put them several days in water, without killing them,—that he lost his patience, and dissected them alive. He tried to drown them also in vinegar, in which they held out more than two days[344].

That the suspended animation and subsequent death of most terrestrial insects when thrown into water is caused by the want of air, is evident from this,—that the same effect ensues if the spiracles be covered with any oily or fatty matter. In this case too, their vital powers soon become suspended: they revive, if the suffocating matter be soon removed; and if this be not done, infallibly perish. This fact was known to the ancients, for Pliny observes that bees die if dipped in oil or honey[345]. One exception to this law has been before mentioned[346]: a similar contrivance secures the cheese-maggot from having its respiration interrupted by its moist and greasy food; the grub also of Sarcophaga carnaria, and of other Muscidæ probably, has its posterior spiracles placed in a plate at the bottom of a kind of fleshy pouch, which has the shape of a hollow, truncated, and[Pg 80] reversed cone. This pouch the grub can close whenever it pleases, so as to cover its spiracles[347]. And numerous other larvæ, both of Diptera and Coleoptera that devour unclean and oily food, have doubtless some protection of this kind for their spiracles and respiratory plates.

I am, &c.

[Pg 81]

LETTER XXXIX.

INTERNAL ANATOMY AND PHYSIOLOGY OF INSECTS, CONTINUED.

CIRCULATION.

We learn from the highest authority, that the blood is the life of the animal[348]: every object of creation, therefore, that is gifted with animal life, we may conclude, in some sense, has blood, which in this large sense may be defined—The fluid that visits and nourishes every part of a living body[349]. But the Great Author of nature has varied the machinery by which this nutritive fluid is formed and distributed, gradually proceeding from the most simple to the most complex structure; in which he seems to have seen it fit to invert the process observable in the systems of sensation and respiration, where the ascent is from the most complex, to the most simple structure. In the lowest members of the animal creation, the blood seems the portion they imbibe of the fluid medium in which they reside, which when chylified, distributes new molecules to all parts of their frame[350]. In others, as in insects, it is formed by the chyle that transpires[Pg 82] through the intestinal canal into the general cavity of the body, where it receives oxygen from the air-vessels, and is fitted for nutrition[351]. In these animals it is accompanied by a long dorsal vessel, the first step towards a heart, which alternately contracts and dilates with an irregular systole and diastole, but appears to have no vascular system connected with it, though in their preparatory states it has an extra-vascular circulation which ceases in the perfect insect. Again: in others, as the Tubicoles, Annelida, &c., a real circulation has been discovered; that is to say, a system of veins and arteries, but unaccompanied by a muscular heart[352]. In the Arachnida and Branchiopod Crustacea the long dorsal vessel is also found; but in these it is connected with an arterial and venous system, which receives, distributes, and returns the blood[353]. It has therefore now become a true heart, and there is a regular circulation; and in the Decapod Crustacea the dorsal vessel is contracted into an oval form, and placed nearly in the centre of the trunk[354]. In the great majority of invertebrate animals the blood is white, but in the Annelida, to which Class the common dew-worm belongs, a curious anomaly takes place—for it is red[355]. Thus a gradual ascent is made to the circulating system of the vertebrate and red-blooded animals. In all, however, the blood is the principal instrument of nutrition and accretion; and is on that account properly so denominated, though not connected with a circulating system.

Having given you this general outline of the means[Pg 83] by which the blood is distributed in the different Classes of animals, I shall now confine myself to the case of insects and Arachnida, beginning with the former.

I. If you examine attentively the back of any smooth caterpillar with a transparent skin, you will perceive in that part an evident pulsation, as though a fluid were pushed at regular intervals towards the head, along a narrow tube which seems to run the whole length of the body. Accurate dissections have proved that this appearance is real, that there is actually present in the back of most insects, placed immediately under the skin and furnished with numerous air-vessels, a longitudinal vessel[356] originating in the head near the mouth[357], running parallel with the alimentary canal nearly to the anus, containing a fluid which is propelled in regular pulsations of from 20 to 100 per minute, more or less as the weather is colder or warmer[358], causing a sensible alternate systole and diastole from the anal extremity towards the head. In the Cossus these pulses were observed by Lyonet to begin in the eleventh segment, from which they passed from segment to segment, till they arrived at the fourth, where they terminated[359]. This vessel is what Malpighi, who first discovered it, termed a heart, or rather series of hearts[360]; but which Reaumur, who injected it, regarded as a simple artery without striking contractions[361]: but to steer clear of any hypothesis, I shall merely call it the dorsal vessel (Pseudo-cardia). When carefully taken out of the body it is found to be a membranous tube, appearing to be closed[Pg 84] at each end[362], in many larvæ of equal diameter every where, but in perfect insects usually widest at the anal extremity[363], and attenuated into a very slender filament towards the head. In some insects, however, as in the larva of the chamæleon-fly (Stratyomis Chamæleon), it is attenuated at both ends, and in the Ephemera is alternately constricted and dilated as Malpighi describes that of the silkworm[364], a dilated portion belonging to each segment[365]. In the Cossus, and probably others, after the third segment, it is furnished with nine pair, the three posterior pair being the largest, of triangular transverse bundles of muscular fibres, which Lyonet denominates its wings[366], the action of which produces its systole and diastole, and their propagation from the tail towards the head[367]. Under the last pair of these wings it is strengthened by a large number of circular muscular fibres[368]. I have stated it as appearing to be closed at each extremity, because Cuvier and most writers have so regarded it, and probably it is so closed in the perfect insect; but from Lyonet's words it should seem that, in the larva of the Cossus, he considered it as open and expanded at its anterior end[369]. He seems also to suspect, that, by means of what he calls the frontal ganglions, a fluid is derived from the dorsal vessel to the spinal marrow. He likewise describes a large nerve as passing through it and becoming recurrent[370]. Carus, as we shall soon see, has also proved that this tube is not closed in larvæ.

[Pg 85]

The fluid which this vessel contains is very abundant; in the animal it appears colourless and transparent like water, but when collected in drops it becomes more or less yellow, and even orange[371]. Examined under the microscope it appears filled with a prodigious number of transparent globules, of incredible minuteness[372]. When mixed with water, which it does readily, its globules lose all their transparency, and coagulate into small clammy masses. After evaporation it becomes hard, and cracks like gum, as blood does also. This gummy substance is so abundant, that the fluid contained in the dorsal vessel of the caterpillar of the Cossus yields a mass of it of the size of a grey pea[373].

From the situation of this dorsal vessel, which is precisely the same with that of the heart in Arachnida and the Branchiopod Crustacea, and from the systole and diastole which keep its fluid contents in constant motion, who can wonder that the physiologists who first discovered it, reasoning analogically, maintained that it was a true heart? But modern comparative anatomists, and those of the highest name, from the absence of a vascular system for a circulation, have contended that it is not a true heart, but an organ appropriated to other purposes: a third hypothesis, and intermediate between these two, has very recently been promulgated, that the organ in question, namely, is a real heart, and in the preparatory states of insects, the centre of a real circulation, which, in the imago state, ceases with the full development of the wings; but that this circulation is[Pg 86] extravascular, or without peculiar vessels analogous to veins and arteries.

I shall now enlarge a little upon each of these hypotheses, beginning with the first or original one.

No one will deny that the argument from analogy is strongly in favour of this: I need not therefore dwell upon it, but proceed to others. Swammerdam, to whose exactness in observing, and scrupulous accuracy, every reader of his immortal work will bear testimony, expressly asserts that he has seen vessels issuing from the dorsal vessel in the silkworm, and even succeeded in injecting them with a coloured fluid[374]. Now it seems extremely improbable that so practised and expert an anatomist should have been deceived, especially upon a point which would naturally excite his most earnest and undivided attention. Without this recorded experiment, perhaps, it might be thought, though this was very unlikely, that he had mistaken bronchiæ for veins and arteries: but how could they have been injected from the supposed heart? Another great physiologist, Reaumur, in the caterpillar of the saw-fly of the rose (Hylotoma Rosæ) observed, besides the dorsal vessel, a ventral one of similar form, in which also was a pulsation, but slower than that of the other. This he supposes may be the principal trunk of the veins[375]. Bonnet thought he discovered a similar vessel in a large caterpillar, but with[Pg 87] all his attention could perceive no motion in it[376]. Reaumur also fancied he perceived in the grub of Musca vomitoria, in which he in vain looked for the dorsal vessel, a fleshy part which exhibited alternate pulsations; and when with a pair of scissors he made a lateral incision in the insect, amongst other parts that came out, there was one that had movements of contraction and dilatation for several minutes,—this experiment was repeated with the same result upon several grubs[377]. De Geer, whose love of truth and accuracy no one will call in question, saw the appearance of blood-vessels in the leg of the larva of a Phryganea L. (as Lyonet did in those of a flea[378]); and in the transparent thigh of Ornithomia avicularia he discovered a pulse like that of an artery[379]. Baker, whose only object was to record what he saw, speaks of the current of the blood being remarkably visible in the legs of some small bugs[380]: what he meant by that term is uncertain, but they could not be spiders, which he had just distinguished. This author has likewise seen a green fluid passing through the vessels of the wings of grasshoppers[381]; and M. Chabrier is of opinion that insects possess the power of propelling a fluid into the nervures of their wings and withdrawing it at pleasure, as they are elevated or depressed[382]; but this last fact may be independent of a circulation.

But though these arguments, which I have stated in their full force, appear strong, and at first sight conclusive,[Pg 88] those which may be urged for the more modern opinion—that no circulation exists in insects, properly so called,—appear to have still greater weight. Lyonet, whose piercing eye and skilful hand traced the course of so many hundred nerves and bronchiæ long after they became invisible to the unassisted eye, and which were a thousand times smaller than the principal blood-vessels, opening into so large an organ as the supposed heart of insects, might be expected to be, could never discover any thing like them. His most painful researches, and repeated attempts to inject them with coloured liquors, were unable to detect the most minute opening in the dorsal vessel, or the slightest trace of any artery or vein proceeding from or communicating with it[383]. And Cuvier, whose unrivalled skill in Comparative Anatomy peculiarly qualified him for the investigation, repeated these inquiries, and tried all the known modes of injection, with equal want of success; and is thus led to the conclusion, that insects have no circulation, that their dorsal vessel is no heart, and therefore ought not to be called by that name: that it is rather a secretory vessel, like many others of that kind in those animals. As to the nature of the fluid that it secretes, and its use, he thinks it impossible, from our present information on the subject, to form any satisfactory conclusion[384]. Marcel de Serres informs us—which further seems to prove that it can be no real heart—that this vessel may be totally removed without causing the immediate death of the insect[385]. This opinion receives additional confirmation from the mode in which respiration is performed in insects. In[Pg 89] those animals that have a circulation, this takes place by means of lungs or gills;—thus we find, even in the Crustacea and Arachnida so nearly related to insects, that the organs of this function are true gills; whereas in insects, though in some of their states their respiratory tubes are branchiform, yet they are not gills, and the respiration is by tubes and spiracles. And these tubes, as you have seen, are so numerous and so infinitely ramified and dispersed, as to occupy the place of arteries and veins, and to imitate their distribution,—and thus to oxygenate what may be deemed the real analogue of the blood, which bathes every internal part of the body of an insect. Those animals likewise that have a circulation are furnished with a liver, as is the case with the Arachnida and even many aggregate animals that have a heart; but in insects there are only hepatic ducts. M. Cuvier has also proved that the conglomerate glands, which exist in all animals that have a heart and blood-vessels, do not exist in insects, in which they are replaced by long slender secretory tubes, which without being united float in the interior of the body: from this circumstance, he is led to conclude that their nutrition is by imbibition or immediate absorption, as in the Polypi and other zoophytes, the chyle transpiring through the alimentary canal, and running uniformly to all parts of the body[386].

These arguments appear so satisfactory, that Physiologists in general seem to have been convinced by them that no circulation, at any time, takes place in insects, and that their supposed heart is merely a secretory vessel, though of what kind they were at a loss to[Pg 90] conjecture[387]. But, convincing as they seem, they appear to have been founded in error, and on the idea that a circulation, as well as a heart, necessarily implies a vascular[Pg 91] system consisting of veins and arteries; for by the recent discoveries of M. Carus, it has been satisfactorily proved that insects in their preparatory states, have an extravascular circulation, the arterial and venose currents not being confined by parietes. The observations upon which M. Carus' hypothesis is founded, were made in the Autumn of 1826; and an abstract of their results presented to the Union of German Naturalists and Physicians, which then held its meeting at Dresden, many of the members of which, as MM. Oken, Husche, Heyne, Purkinje, Otto, Weber, and Müller, had ocular proofs of the reality of the phenomena.

His first observations were made on the larva of Agrion Puella, which swims by means of three vertical laminæ attached to the tail; which, when the wings first appear as rudiments, begin to be exsiccated and are finally detached. Each of these laminæ, in its natural vertical position, presents an inferior abdominal and a superior dorsal edge, has two tracheæ running along its centre with ramifying bronchiæ, and consists of granular substance contained between two strata of the external integuments. A current of blood-globules enters each[Pg 92] lamina somewhat nearer to its abdominal than to its dorsal edge, and running through the greater part of its length suddenly turns and bends its course back towards the body, somewhat nearer to the dorsal than to the abdominal margin of the lamina. The channel thus formed in the midst of the granular substance is perfectly transparent, except where it is occupied by the blood-globules, or crossed by the bronchiæ. The parietes of the channel are not strictly defined, nor formed by any thing like the coats of a vessel, the blood circulating through the granular Parenchyma; a circumstance however which is not peculiar to this case, but also occurs generally in the first states of the circulation, as it presents itself for instance in the embryo of Fishes, and in the figura venosa of the incubated egg[394]. The blood-globules are elongated like a grain of wheat, considerably larger than those of the human blood, and float in a fluid which is invisible because of its transparency, but the existence of which is proved by the variations in the position of the globules in the current, sometimes following its direction, at others crossing it transversely, or more or less obliquely.

When the animal is vigorous, the current is uninterrupted, although its velocity is accelerated at regular intervals; and that not only in the excurrent (arterial), but also in the recurrent (venous) part of its course through the lamina. When the animal becomes exhausted, or the laminæ exsiccated, the circulation is interrupted, and in the same manner, as under the same circumstances, in the larvæ of frogs and lizards; the disturbance displaying itself not merely by a cessation of the process, but also[Pg 93] by retrograde movements of the currents, or by oscillatory motions of the blood-globules.

In proportion as the wings are developed, the circulation in the laminæ diminishes, and ultimately ceases, preparatory to the detachment of the laminæ themselves. At the same time, however, it presents itself under a new form in the wings. In these the excurrent or arterial stream takes its course along the inner margin of the wing, and the recurrent or venous returning along the outer; whilst, occasionally, other transverse currents take their course through the net-work of the wing from its inner to its outer margin. As the wings are further developed, the circulation in them, like that in the caudal laminæ, gradually becomes weaker and ultimately ceases[395].

The next observations were made on the transparent larva of a neuropterous insect (probably a Semblis or Sialis), in which the pulsations of the dorsal vessel were distinctly seen at its posterior extremity, from which they were propagated towards the anterior; these two divisions of that vessel appearing to bear to each other the relation of a heart and aorta. There were no traces of other vessels, though regular and rapid currents of blood-globules, exterior to the tracheæ, proceeded from the head towards the posterior extremity of the body, where each of these currents entered the heart, which again propelled its contents with accelerated velocity through the anterior part of the dorsal vessel towards the head. The lateral currents also were accelerated upon each contraction of the heart, proving that they must communicate with the dorsal vessel at the anterior part of the body, though the[Pg 94] opacity of the head rendered it impossible to ascertain the mode of anastomosis. An excurrent and returning current were also traced to each of the legs[396]. But the phenomena of the circulation was most distinctly visible in the larva of Ephemera vulgata, even more distinctly than it is possible to trace it in the larvæ of frogs and newts. In this animal the circulation, with the help of the microscope, is at once visible in the three last segments of the body; and with a little attention is discoverable not only in the three terminal caudulæ, and in the upper joints of the legs, but also in the head, and particularly the roots of the antennæ. In the posterior part of the body there are on each side two currents of blood, not bounded by parietes, situate on each side of the intestinal canal, the inner one being the most considerable. The external one communicates with the internal by several intermediate branches; from this probably the streams are detached, which in the form of loops are seen at the upper joints of the legs, though it is not possible precisely to ascertain this, nor even whether these lateral currents continue distinct in the thorax, which probably they do. At the ninth abdominal segment these currents which flow posteriorly from the head, change their direction, and are inflected so as to enter the pulsating heart, from which the current again flows towards the head. Before they enter the heart they give off three streams, one for each of the three caudulæ. The currents in these caudulæ present the phenomena of the circulation with peculiar distinctness, and are particularly remarkable from the circumstance, that the excurrent and recurrent streams, though closely approximated without any visible separation,[Pg 95] flow without disturbing each other. The excurrent stream is accelerated in correspondence with the pulsations of the heart; the recurrent on the contrary being always somewhat more sluggish, and the first to stagnate and cease when the strength of the animal is impaired. In the anterior part of the head currents can be discovered, forming loops like those of the legs, at the roots of the antennæ; each current proceeding from the cranial surface, and in returning taking its course towards the region of the larynx[397].

M. Carus has likewise observed currents of blood in the larvæ of water-beetles (Hydrophilus and Dytiscus)[398]; but at present he appears to have detected it in no terrestrial larva. Whether this is occasioned by their opacity, or it exists only in the ovum, as he seems to suspect[399], must be left for determination to future observers; it is scarcely probable, however, that the larvæ of Dytisci and Hydrophili should differ from other Coleoptera in their circulation.

The endeavours of M. Carus to discover any proofs of a circulation in insects in their last state, except in the wings at their first development, were without success[400]. He observes that the fact of the currents of fluids in larvæ not being defined by vascular parietes, enables us to comprehend the rapidity and facility with which the traces of the circulation are lost in the perfect insect. On the other hand, that the existence of a circulation at one period, and its cessation at another, elucidate many circumstances connected with the physiology of these animals: for instance, the contrast between the[Pg 96] rapid growth and transformations of the larvæ, and the stationary existence of the imago, &c. Lastly he remarks, that the phenomena of this circulation do not throw any light on the obscure subject of the mode of nutrition in perfect insects; which therefore must still be supposed to be effected according to the idea of Cuvier, without the intervention of vessels[401].

Whatever be the functions of the dorsal vessel, this seems the most proper place to state to you what further is known respecting it. Its construction is nearly alike in insects in all their states, except that in the imago it is shorter and narrower. Reaumur has affirmed, and before him Malpighi made a similar observation, that in chrysalises newly disclosed from the larva, and yet transparent, the motion of the included fluid is the reverse of what it has been in that state, it being propelled from the head to the tail, which he found to be the case also in the imago[402]. If this be true, and there is no reason to doubt his accuracy, when they are more advanced, it resumes its old course, as Lyonet observed, from the tail to the head[403]. But probably it is not always uniformly in the same direction, since Malpighi states that a very slight cause will change its course, and that the pulsations differ in quickness in different portions of the heart[404]. If its course were really always the same, and in one direction, without any reflux, it would seem to follow that the fluid must be absorbed at one end, and, if there was no outlet, transpire at the other, which would be a kind of circulation. In Syrphus Pyrastri and other aphidivorous[Pg 97] flies, this dorsal vessel, instead of the usual form which it had in the larva, assumes a very peculiar appearance. If, taking one of these flies by the head and wings and holding it up to the light, you survey under a lens the base of the lower part of its abdomen, you will see through its transparent skin, which exactly forms such a window as physicians have sometimes wished for in order to view the interior of their patients, a flask-shaped vessel having its long end directed towards the trunk, in which there is a manifest pulsation and transmission of some fluid. This vessel extends in length from the junction of the trunk with the abdomen to about the termination of the second segment. The included fluid does not run in the dorsal vessel in a regular course, but is propelled at intervals by drops, as if from a syringe, first from the wide end towards the trunk, and then in the contrary direction, forming a very interesting and agreeable spectacle. One circumstance led Reaumur to conjecture that the neck of this vessel, which he at first regarded as simple, is in fact composed of two or more approximated tubes, and that the blood is conveyed forward by the outward ones, and backward by the intermediate one[405]: he even thinks that he saw a kind of secondary heart, at the extremity next the trunk, for the purpose of causing the reflux. This illustrious author observed the above remarkable structure not only in the Syrphi, but in many of their affinities, and thinks that it is also widely diffused amongst the Muscidæ[406].

I must now say something upon what I conceive to be the real blood of insects; for I think no one will object to that name being given to their nutritive fluid, especially[Pg 98] in the larva, though it does not circulate by means of a vascular system. The chyle that is produced in the intestines of animals from the food, is that fluid substance from which their blood is formed: in insects it is not absorbed by the lacteals, but transpires through the pores of the intestinal canal into the general cavity of the body, where, being exposed to the influence of the oxygen in the air-vessels, it becomes, though retaining its colour, a different fluid from what it was before, and analogous to blood in its use and office[407]; only that in these animals, as Cuvier has observed, at least in their perfect state, the blood, for want of a circulating system, not being able to seek the air, the air goes to seek the blood[408]. The dispersion of this fluid appears to be universal, so that all the parts and organs contain it in a greater or less degree[409]. In many insects, if you break only an antenna or a leg, a drop of fluid flows out at the wound. In larvæ, the fluid which bathes[410], or visits, all the internal parts and organs is not only sufficient for their nutriment, but a large quantity of seemingly superfluous blood remains that is not wanted for this purpose. This is expended in the production of the caul or epiploon (Corps graisseux[Pg 99] Reaum.), which laps over and defends all the viscera of the animal, and goes principally to the formation of the imago[411]. I have said that Cuvier conceives nutrition in insects to take place by imbibition or immediate absorption; that is, I suppose, the different parts and organs thus constantly bathed in the blood, imbibe from it the particles necessary for their constant accretion. M. Chabrier seems to think that it is the compression and dilatation of the trunk that duly distributes the nutritive fluid[412]; Lyonet compares the nutrition of insects by their fibres from this fluid, when formed into the corps graisseux, to that of plants that draw their support by their roots from the earth[413]. Much obscurity, however, at present rests upon this subject—much for future investigation to explore; but in all the works of the Most High there is always something inscrutable, something beyond the reach of our senses and faculties, which teaches us humbly to adore his infinite perfections.

II. The circulation of the Arachnida is next to be considered; and the term applied to these becomes strictly proper. Two great tribes, in our view of the subject, constitute this Class,—the spiders (Araneidea) and scorpions (Scorpionidea): I shall give you some account of the circulating vessels of each.—In spiders, the heart in general is a long dorsal vessel as in insects, but supposed to be confined to the abdomen, growing slenderer towards each extremity, particularly the anal. In some also, as in Aranea domestica, like that of insects, it has lateral muscular appendages; but in others, as in[Pg 100] Clubiona atrox, it is without them[414]. It exhibits a pair of vessels that appear to connect with the gills, by which the oxygenation of the blood takes place, and a number of others that ramify minutely and are lost in the analogue of the epiploon, supposed to be their liver[415]. Whether these last are to be regarded merely as veins, has not been ascertained; they seem rather to convey the blood outwards, than to return it back to the heart: but this question must be left for future investigation. I may observe, however, that though the heart of the spider has been traced only in the abdomen, it may probably extend into the trunk.

The heart of the scorpion has been examined both by Treviranus and Marcel de Serres; but I shall principally confine myself to the description of the latter, as the most clear and intelligible. The heart, then, of these animals is elongated, almost cylindrical, but attenuated at each end; it is extended from the head to the extremity of the tail, and appears to have four pairs of lateral muscles. On each side are four pairs of principal vessels, which go to the pulmonary pouches, and there ramify. These may be assimilated to veins. Besides these, there are four other vessels that cross them, forming with them an acute angle, and which, with four branches of smaller size, receive the blood from the pulmonary pouches, and distribute it to the different parts of the body,—these are the arteries. Before it enters the tail, the heart throws out two vascular branches which do not go to the gills, but distributing the blood[Pg 101] to different parts, ought to be considered as arteries[416]. Treviranus mentions bunches of reticulated vessels, concerning the use and origin of which he seems uncertain[417]; but as they approach the gills they are probably the branching extremities of what M. de Serres considers as the veins.

I am, &c.

[Pg 102]

LETTER XL.

INTERNAL ANATOMY AND PHYSIOLOGY OF INSECTS, CONTINUED.

DIGESTION.

"The immense Class of insects," says the immortal Cuvier, "in the structure of its alimentary canal exhibits as many variations as those of all the vertebrate animals together: there are not only the differences that strike us in going from family to family and from species to species; but one and the same individual has often a canal quite different, according as we examine it in its larva or imago state; and all these variations have relations very exact, often easily estimable, with the temporary or constant mode of life of the animals in which it is observable. Thus the voracious larvæ of the Scarabæi and butterflies have intestines ten times as large as the winged and sober insects—if I may use such an expression—to which they give birth[418]."

In the natural families of these creatures, the same analogy takes place with respect to this part that is observable in the rest of the Animal Kingdom; the length and complication of the intestines are here, as in the[Pg 103] other Classes, often an index of a less substantial kind of nutriment; while their shortness and slenderness indicate that the insect lives by prey[419].

In considering therefore the parts connected with the digestive functions of the insect world, it will not be amiss to have reference to their food, and their mode of taking it; but first it will be proper to state and define the parts of this important organ.

In general the alimentary canal[420] is composed of the same essential tunicks as that of the vertebrate animals, consisting of an interior epidermis, a papillary and cellular tunick, and an exterior muscular one[421]. The first is usually tender, smooth, and transparent; but not always discoverable, perhaps on account of its tender substance[422]. Ramdohr does not notice the papillary and cellular tunicks; they are probably synonymous with what he denominates—the flocky layer (Die flockige lage), and which he describes, when highly magnified, as appearing to consist of very minute globules or dark points, and as being of a cellular structure[423]. The exterior tunick is thicker and stronger than the interior, and composed of muscular fibres, running either longitudinally, or transversely so as to form rings round the canal. This tunick mostly begins at the mouth, and goes to the anus, changing its conformation in different parts of the above intestine. Sometimes however it originates only at the beginning of the stomach[424]. With respect to its general disposition, that canal—in its relative length, in[Pg 104] the size of its different parts, in the number and form of its dilatations, and particularly of its stomachs and its cœcums, and in the folds of its interior—exhibits variations altogether analogous to those of vertebrate animals, and which produce similar effects[425]. As to its parts, it may be considered as consisting of two larger portions, between which the biliary or hepatic vessels form the point of separation. In the first, the most universal parts are the gullet and the stomach; and in the second, the small intestine and the large intestine[426].

1. The gullet (Œsophagus[427]) is that portion of the intestinal canal which, receiving the food from the pharynx, or immediately from the mouth, conveys it to the stomach. Though it often ends just behind the head[428], it is usually continued through the trunk, and sometimes even extends into the middle of the abdomen[429]; it therefore seldom much exceeds in length half the body. It is constantly long when the head is connected with the trunk by a narrow canal—as in the Hymenoptera, Neuroptera, Lepidoptera, &c.; but is frequently short when these parts are more intimately united[430]. It often ends in a kind of sac analogous to the crop of birds. Under this head I must mention a part discovered by Ramdohr, which he calls the food-bag (Speisesack), as he thinks, peculiar to Diptera[431]. From the mouth in these proceeds a narrow tube into the abdomen, where[Pg 105] it expands into a blind sac having no connexion with the stomach; so that the fluid food, as blood, &c. stored in it, must be regurgitated into the mouth before it can pass into that organ[432]. Thus these animals, besides their stomach, have a reservoir in which to store up their food; the product therefore of a single meal will require several days to digest it.

2. The stomach (Ventriculus[433]) is that part of the intestinal canal immediately above the bile-vessels, which receives the food from the gullet for digestion, and transmits it when digested to the lower intestines[434]. By its admixture with the gastric juice, the food acquires in the stomach a quite different colour from what it had in the gullet. In herbivorous insects it contains no acid, but, like the gastric juice of herbivorous quadrupeds, is of an alkaline nature[435]. The chyle is forced through this organ, probably in part by the pressure of the muscular fibres during the peristaltic motion; and being pressed through the inner skin, is first collected in the intermediate cellular part, and ultimately forced through the outer skin[436]. At its posterior end it terminates in the pylorus, a fleshy ring or sphincter formed of annular muscular fibres[437]. The stomach often consists of two or more successive divisions, which are separated from each other, and are often of an entirely different conformation and shape[438]. In the Orthoptera, Predaceous Coleoptera, and several other insects, an organ of this[Pg 106] kind precedes the ordinary stomach, which from its structure Cuvier denominates a second stomach or gizzard[439]; Posselt improperly calls it Cardia[440]; and by Ramdohr it is named the plaited-stomach (Falten-magen[441]). It is a short fleshy part consisting of two skins, placed above the opening of the stomach, and perhaps rather belongs to the gullet. The inner skin is formed into longitudinal folds, and sometimes armed with horns, teeth, or bristles. Its cavity is very small and compressed, so as to admit only small masses of food, and yet present them to a wide surface for the action of the teeth or bristles;—in this stomach therefore, as in the gizzard of birds, to which it seems clearly analogous[442], the food is more effectually comminuted and rendered fit for digestion. The muscles, by which its action upon the food is supported, in some species amount to many thousands[443]. Rudiments of a gizzard are sometimes found concealed in the gullet of many insects[444]. The idea of Swammerdam, Cuvier, &c. that grasshoppers and other insects that have this kind of stomach, chew the cud[445], Ramdohr affirms is entirely erroneous[446]. Besides its divisions, the stomach has other appendages that require notice. In most Orthoptera, a pair or more of blind intestines or cœca may be found at the point of union of the gizzard with the stomach[447], which have been regarded as forming a third stomach: they also begin the stomach in the louse[448]; they form a coronet round[Pg 107] the apex of that organ, in the grub of the cockchafer[449]; and in that of the rose-beetle, there is one at the apex, one in the middle, and a third at the base[450]. Besides these appendages, which are formed of the skin of the stomach, there are others that are not so. In the Predaceous and some other beetles, the whole external surface of this organ is covered with small blind appendages opening into the space between its two skins, which cause it to resemble a shaggy cloth; these Ramdohr calls shags (zotte[451]), and Cuvier, hairs[452] (villi). These appendages the latter author seems to regard as organs that secrete the gastric juice and render it to the stomach[453]; but the former thinks their use uncertain[454].

3. The small intestines (Intestina parva) are the portion of intestines next the stomach, and consist often of three distinct canals;—the first is supposed to be analogous to the duodenum; it is found only in the Coleopterous genera Silpha L. and Lampyris L., and is distinguished from the succeeding intestine by being perfectly smooth[455]. Next follows the thin intestine (Dünndarm), which in the above insects is wrinkled; it most commonly immediately follows the stomach. Sometimes it is wholly wanting, as in Agrion, the Hemiptera[456], &c. Ramdohr conjectures that it is not solely destined for conveying the excrement, but that probably some juices are separated in it from the food especially for the nutrition of the gall-vessels, as their principal convolutions are mostly near this intestine[457]; which perhaps[Pg 108] may in some cases be regarded as analogous to the jejunum in vertebrate animals. The third pair of the small intestines, which perhaps represents the ileum, Ramdohr distinguishes by the name of club-shaped (Keulförmigen Darm[458]). It may generally be regarded as only a continuation of the former thickened at the end so as to resemble a club reversed. It is however sometimes separated from the thin intestine, as in Cerambyx moschatus[459].

4. The large intestines (Intestina magna) consist sometimes of two portions. The thick intestine (Dicken-darm), which may be regarded as a kind of cœcum, is found only in the larvæ of the Lamellicorn beetles, but never in the perfect insect. In shape it is oval and folded; whence it is thicker than the rest of the intestinal canal, and is constantly filled with excrement[460]. The second portion of these intestines is the rectum (Mastdarm), which terminates in the anal passage. This part is scarcely ever wanting, except when the insect evacuates no excrement, which is the case with the grubs of bees, wasps, and the antlion (Myrmeleon). In the imago of Telephorus, at least in T. fuscus, it is also obsolete[461]: in most cases, however, it is very distinct from the preceding intestine. Sometimes it consists of only one tunick composed of muscular fibres[462]. When the gullet is wide, the rectum is usually so likewise; but when it follows a club-shaped or thick intestine, it is narrow[463]. It generally may be termed short[464]. When[Pg 109] wide, it often contains a great quantity of excrement, as the gullet does of undigested food; but when narrow, the excrement seldom remains long in it. This intestine also in a few cases has a lateral enlargement or cœcum (Blind-darm), being a continuation of the same skin; but perhaps this enlargement is really analogous to what Ramdohr calls the thick intestine, though in these cases he regards it as an appendage of the rectum[465].

I must now call your attention to the bile-vessels of insects. These, by Malpighi[466] and the earlier physiologists, who regarded them as a kind of lacteals, were denominated varicose vessels: but Cuvier—and his opinion after some hesitation has been adopted by Ramdohr—considers them as vessels for the secretion of bile, and as analogous to the liver of animals that have a circulation[467]. As the want of blood-vessels prevents insects from having any gland, the bile is produced with them, as all their other secretions, by slender vessels that float in their nutritive fluid, and from thence secrete the elements proper to form that important product, which usually tinges them with its own yellow hue; though in the Lamellicorns and Capricorns they are of an opaque white, and in the Dytisci of a deep brown colour[468]. Their bitter taste further proves that they contain the bile[469]. They are long, slender, filiform, tortuous or convoluted, and mostly simple vessels; sometimes gradually smaller toward the base[470], at others towards the apex[471]. In[Pg 110] some, screw-shaped[472]: in one larva, with hemispherical elevations[473]: in the cockchafer, part of them are fringed on each side with an infinity of short, blind, minute, setiform tubes, while the rest are naked[474]; they are composed of a single, thin, transparent membrane, according to Ramdohr[475]; but Cuvier thinks their texture is spongy[476]. They appear to contain a number of small, irregular, dark granules, which float in a peculiar fluid, with which, however, they are not always filled throughout, nor are they constantly permeable from one end to the other. Thus in the meal-worm beetle (Tenebrio Molitor), the common trunk by which they are attached to the intestinal canal is composed of gelatinous granules[477]. The place of their insertion is generally a little below the pylorus, but in the common cockroach they are inserted into the stomach just above that part[478]. Usually each vessel opens singly into the intestinal canal, which the whole number surround at an equal distance from each other[479]. Sometimes, however, they are connected with it by a common tube in which they all unite, as in the asparagus-beetle (Lema Asparagi[480]), and the mole-cricket (Gryllotalpa vulgaris[481]); in the house-fly (Musca domestica), and other Muscidæ, each pair unites so as to form a single branch on each side of the canal previously to their insertion[482]; in the field-cricket (Gryllus campestris) they are all inserted in one spot[483];[Pg 111] and when numerous, they are generally attached singly though irregularly[484]. These vessels at their base do not open into the cavity of the intestinal canal, but merely into the space between its outer and inner tunicks, the last being constantly imperforated[485].

With regard to their apex, the bile-vessels are sometimes fixed singly or connectedly to the intestine merely by a few muscular fibres; for they do not enter it, their ends having no orifice. This structure is mostly to be met with in the Coleoptera[486]. In caterpillars, the tops of these vessels perforate the outer skin of the rectum, and proceeding in dense convolutions to the anus, become at last so fine that their terminations cannot be discovered[487]. In other cases, the extremities of a pair of these vessels unite so as to form a double one: this may be seen in those of Philonthus politus[488], and probably other rove-beetles: and lastly, in others the bile-vessels are free, hanging down by the intestinal canal, without being attached to it or to each other. This structure is constantly found in the Orthoptera and Hymenoptera Orders, &c.[489].

With regard to their number, the bile-vessels vary from two to upwards of one hundred and fifty, yet so that their whole amount is constantly the product of the number two,—at least as far as they have been counted: and even when those on one side are not alike, a similar variation takes place in the other, as may be seen in Galleruca Vitellinæ, where on each side are two long ones and one shorter[490]; the most usual numbers are, four—six—or many, that is, more than twenty—

[Pg 112]

The bile-vessels vary considerably in length: in many cases where they are free they are short[498]; they are often very long, and perhaps those that are fixed may be generally stated as the longest. In the Lamellicorn beetles they are remarkable for their great length[499].

Having given you this general account of the intestinal canal and its parts and appendages, I shall now state some of the peculiarities that in this respect distinguish particular tribes and families.

The Coleoptera alone, exhibit as many variations in the structure of the alimentary tube as all the other Orders of insects together:—to particularize these would occupy too large a portion of this letter, I shall therefore only notice a few of the most remarkable. In general they may be stated as having universally a stomach, a[Pg 113] small intestine and rectum, and not more than three pairs of fixed or united bile-vessels. In the Predaceous beetles, the gullet mostly widens at the base into a considerable crop, followed by a gizzard, a shaggy stomach, and two pairs of united bile-vessels. The whole alimentary canal in these, is never less than double, and sometimes treble the length of the body[500]. In the carnivorous beetles, at least the Staphylinidæ and Silphidæ, there is little or no crop, and the gizzard is hidden: in the former, the whole length of the intestinal canal is not twice, while in the latter it is more than four times that of the body[501]. In these also the intermediate portion of the large intestine is singularly annulated[502]. In the Petalocera the stomach is usually longer than all the rest of the intestines together, and often convoluted: in the cockchafer the whole intestinal canal is nearly five times the length of the body, four parts of which is occupied by the stomach[503]. In the grub the canal scarcely exceeds the length of the animal[504]. In Lampyris the stomach exhibits a remarkable appearance, having on each side a series of spherical folds or vesicles[505]. Have these any thing to do with the secretion of its phosphoric matter? Tenebrio has a gizzard armed internally with calluses, and a shaggy stomach, and Blaps does not differ materially; their entire canal is more than twice the length of the body[506]. In the vesicatory beetles (Cantharis, Meloe, &c.) there is no gizzard, and the canal is less than twice the length of the body[507]. Little is known with regard to the alimentary canal of the beetles distinguished by a rostrum (Rhyncophora).[Pg 114] In the only two that appear to have been examined, Rhynchites Betuleti and Cryptorhynchus Lapathi, that canal is moderately long, the stomach partially shaggy, and the small intestine inversely claviform; but in other respects they differ materially[508]. In the former there is no crop or gizzard, the stomach is fringed on each side, except at its upper extremity, with a series of small cœca or shags, and there are three pairs of bile-vessels[509]; while in the latter the gullet is dilated into a crop which includes a gizzard in which the skill of a Divine artist is singularly conspicuous:—though so minute as scarcely to exceed a large pin's head in size, it is stated to be armed internally with more than 400 pairs of teeth, moved by an infinitely greater number of muscles[510]. A transverse section of this gizzard represents two concentric stars, with nine rays each[511]: the object of this structure is, the comminution of the timber which this beetle has to perforate and probably devour[512]. The stomach is very slender, but dilates in the middle into a spherical vesicle[513], and there are only two pairs of bile-vessels[514]. In the Capricorn beetles, the part we are considering varies much: in general we may observe that it is more than double the length of the body, that the stomach is long and slender, and usually naked, that the gullet terminates in a crop without a distinct gizzard, and that there are three pairs of bile-vessels[515]. In the Herbivorous beetles (Chrysomela, Cassida, &c.) the canal[Pg 115] is more than double the length of the body, and in some much longer[516], the stomach is long, and commonly naked; but in Chrysomela violacea it is covered with hemispherical prominences[517], and in Chrysomela Populi it is shaggy[518]; in the insect last named and Galleruca Vitellinæ the rectum consists of two pieces[519]. In this tribe the intestines of the larva resemble those of the perfect insect[520].

In the Orthoptera the alimentary canal, which continues the same in every state, is short, or only moderately long; the gullet has one or two lateral pouches or crops[521], and terminates in a gizzard of curious construction, with singular folds and teeth[522]; then follows a short stomach, usually with a pair or more of cœca at its upper extremity[523]; the lower intestines are not distinct, and the bile-vessels numerous, short and free[524].

In the Neuroptera, many of the genera are distinguished by the remarkable length of the gullet, and by the lower intestines forming one short piece[525]. In the Libellulina the bile-vessels are numerous, short, and free, as in the Orthoptera[526]. In Hemerobius and Myrmeleon there is a gizzard[527], and just above it a cœcum, in the former very remarkable, is connected with the gullet[528].

The Hymenoptera appear all to be distinguished by a long slender gullet, terminating in a dilated crop forming the honey-bag; their stomach is variable, their small intestine slender, and the rectum dilated;—their bile-vessels,[Pg 116] like those of the two preceding Orders, are numerous, short, and free[529]. In the ants and ichneumons there is an approach to a gizzard[530]. In the wasp and humble-bee the stomach is very long, with muscular rings surrounding it[531]. In this Order the larvæ at first have no lower intestines and void no excrement[532], but as they approach to the pupa state one begins to appear[533].

The next insects whose alimentary canal we are to consider, are those which, taking their food by suction, have no occasion for masticating organs: this may in part be predicated of the preceding Order, in which most of the tribes in their perfect state imbibe fluid food, and use the ordinary organs of mastication principally in operations connected with their economy; and their crop, in which the honey in many is stored up for regurgitation, may be regarded in some degree as analogous to the food-bag of the Diptera and other suctorious insects.

The two sections of the Hemiptera Order differ widely in the canal we are considering, and I shall therefore give a separate account of each. In the Heteropterous section, appended to the gullet by a long convoluted capillary tube, besides the usual saliva-reservoirs there is often a double vessel, which Ramdohr regards as discharging the same function, but which in many respects seems rather analogous to the food-reservoir of the Diptera[534]. As I have had no opportunity of examining this vessel, I shall content myself with stating this idea, and describe the vessel more fully hereafter. The gullet, in[Pg 117] these, usually terminates in an ample crop consisting of many folds[535], followed by a long, slender, cylindrical tube, dilated at its base into a spherical tumour; these two may be said to form the first stomach: to this succeeds a second[536], which Ramdohr denominates the bug-stomach (Wanzen-magen), which varies in its figure, and in Pentatoma consists of four demi-tubes, so as to form a quadrangular canal[537]. In the Homopterous section of this Order Ramdohr seems to have examined but few; Chermes however and Aphis exhibit one remarkable feature; they have no bile-vessels, at least he could discover no trace of these organs[538]. Their intestinal canal is very simple, their stomach very long, widest above, and somewhat convoluted, with a very slender gullet[539]. In Cercopis spumaria the structure is more complex, and extremely singular. It has two or rather three stomachs; the two first of a horny substance, and the last a slender somewhat convoluted membranous tube, which becoming reversed, is attached by what should be deemed its lower extremity to the first stomach, from the other side of which emerge the lower intestines, terminating in a thick pear-shaped rectum. At the same point of the first stomach the four bile-vessels are attached, they grow gradually thicker for about a third of their length, when they become twisted like a cord, and taper towards the rectum, to which also they are attached[540]. From this structure it should seem that the food has to pass twice through the first stomach, before the process of digestion is complete, and it is rejected at the anus.

[Pg 118]

The next suctorious Order is the Lepidoptera: in these the gullet is long and slender, surrounded at the beginning with a loose transparent skin, and at the base furnished with a pair of lateral sacs, forming the honey-stomach, and probably analogous to the food-reservoirs of the Diptera, which when blown up are of an oval form; the stomach, as in the bugs, consists of two portions, the first being the longest[541]. There are three free bile-vessels on each side, proceeding from a single branch[542]. It will not be uninteresting here to abstract from Herold the progressive changes which take place in the intestinal canal in this Order, during the transition of the animal from the larva to the imago state. In the larva, the gullet, the small intestine, and the rectum, are short and thick[543], there are a pair of silk reservoirs (sericteria), as well as vessels for the secretion of saliva (sialisteria): if you examine it two days after its first change, you will find the gullet and the small intestine much lengthened and become very slender; the stomach contracted both in length and size; the rectum also changed, and the silk vessels contracted[544]. These in a pupa eight days old have wholly disappeared; the gullet is become still longer, its base is dilated into a crop or food-reservoir; the stomach is still more contracted, and instead of a cylinder represents a spindle; the small intestine also is lengthened[545]: at a still more advanced period, when it is near appearing under its last form, the gullet and small intestine are still more drawn out; and the honey-bag, though very minute, has become a lateral[Pg 119] appendage of the gullet[546]; and lastly, in the butterfly it appears as a large vesicle[547]; the small intestine is grown very long[548]; and the rectum has changed its form and acquired a cœcum[549]. When we consider the adaptation of all these changes of form, the loss of old organs and the acquisition of new ones, to the new functions and mode of life of the animal, we see evidently the all-powerful hand of that Almighty Being who created the universe, upholding by his providence, and the law that he has given to every creature, the system that he at first brought into existence.

We now come to the Diptera. These have a very slender gullet, to which is attached on one side a long filiform tube, terminating in the food-reservoir, which in some instances is simple[550], but most generally consists of two or more vessels[551], collapsing when empty, but varying in shape and size when inflated with food: the mouth of the stomach in many cases is dilated into a kind of ring[552]; sometimes there is on each side a blind appendage or cœcum opening into it, in Bombylius covered with shags, which though not connected with the mouth by a tube, Ramdohr regards as saliva-reservoirs[553]; in Musca vomitoria the beginning of this organ below the mouth is covered with hemispherical prominences, and in Tipula it is dilated and marked with transverse folds. There are usually two pairs of bile-vessels; in the Muscidæ pedunculate and free[554]; in Tipula, Bombylius, and Leptis,[Pg 120] sessile and united[555]; and in Tabanus sessile and fixed[556]. It is remarkable that in some of this Order—the reverse of what usually happens—the alimentary canal appears to be much longer in the larva than it is in the imago; in Musca vomitoria, its length in the former is two inches and a quarter, while in the latter it is only one inch and one third[557]. A singular organ distinguishes the imago of this species, the use of which appears not to be discovered. It succeeds the rectum, and has on each side two short club-shaped appendages, open at the end, which receive tracheæ, and terminate in a short piece that opens into the anus[558].

In Hippobosca and its affinities the canal in question differs from that of other Diptera, in having no food-reservoir; in other respects it resembles it[559].

From the above statement it appears that the principal character which distinguishes those that take their food by suction, from those that masticate it, is the faculty with which they are furnished by means of an ample crop, honey-stomach, or food-reservoir, of regurgitating the food they may have stored up. Another distinction still more striking, which will appear more evidently hereafter, is to be seen in the saliva-secretors with which the suctorious tribes are furnished, to be found in very few masticators, by which they are enabled to render the juices more fluid and fit for suction.

The only insect amongst the Aptera whose alimentary[Pg 121] canal I shall notice, is the common harvest-man (Phalangium Opilio): in this, though the stomach and lower intestine are remarkably simple, yet their cœcal appendages are numerous and singular: the former, which has no distinct gullet, is pear-shaped[560]; and the latter, tapering downwards, and truncated at the end[561]; connected with it above are no less than twenty-three cœca or blind appendages, of various forms and dimensions; the last pair but one of which is very remarkable, being bent like a bow, and furnished externally with four short clavate processes[562]. It is probable that some of these organs are analogous to the bile-vessels of other insects.

When the Creator in his wisdom fixed the limits of the various tribes of animals, he united them all into one harmonious system by means of certain intermediate forms, exhibiting characters taken some from those that were to precede, and others from those that were to follow them, and this not only in their external structure, but likewise in their internal organization; so that we are not to wonder if in the same individual we meet with organs that belong to two distinct tribes, or if, remaining nearly the same in their prima facie appearance, they begin to exercise new functions. An instance of this we have seen in the dorsal vessel of insects, which in the Arachnida, though not materially different in situation or general form, by the addition of a small apparatus of arteries and veins becomes the centre and fountain of a regular system of circulation[563]. From the circumstances here alluded to, physiologists have been led to entertain[Pg 122] very different sentiments with regard to the structure of the alimentary organs of the Class we are now to enter upon, the Arachnida: what some regard as a real liver, others look upon as an epiploon or caul; and what the last denominate bile-vessels are by some of the former considered as appropriated to the secretion of chyle[564]. Yet both these opinions have some foundation in nature. When, in the Arachnida, we discover a lobular substance consisting of granules filling the whole cavity of the body and wrapped round the intestines, every one will see in it no small analogy to the epiploon which in insects performs the same function: but when, upon a further examination, we detect certain vessels communicating with this substance and the intestinal canal[565], the idea that these may be hepatic ducts, and this substance analogous to the liver, immediately strikes us as not improbable. Again: when we discover pairs of other capillary and tortuous vessels connecting with the intestinal canal either at the pylorus[566] or below it[567], which in appearance strikingly resemble the bile-vessels which we so constantly find in insects, we seem warranted in concluding that they are of the same nature and use: but when a nearer inspection enables us to detect the hepatic ducts just mentioned in the scorpion, and we find that these capillary vessels in the spider are in a very different situation from those in insects which we suppose them to represent, it occurs to us as not unlikely, that their function may be different.

[Pg 123]

Let us now consider how the intestinal canal is circumstanced in the two sections into which the Class Arachnida is divided; the Scorpionidea, and Araneidea. In the Scorpions, this organ proceeds from the mouth to the anus without any flexure or convolution, so that its length is scarcely equal to that of the body[568]; it is slender, and its diameter, with the exception of an irregular dilatation here and there, is nearly the same in its whole extent; the gullet is short; the stomach long, and nearly cylindrical; the duodenum shorter and thicker than the stomach, from which, as well as from the rectum, it is separated by a valve; the latter is cylindrical, and opens at the anus above the insertion of the vesicle that secretes the poison[569]. With regard to the biliary system and its organs: The liver is of a pulpy granular consistence and of a brownish colour, fills the whole cavity of the trunk and abdomen, and serves as a bed for the other intestines. It is divided longitudinally into two portions, by the channel in which the heart reposes—its anterior part is formed into many irregular lobes, by the sinuosities of the trunk; at the other extremity, it terminates in two acute ends, which enter the first joint of the tail; its surface presents a reticular appearance, the result of the approximation of polygonous lobuli; its interior is a tissue of infinitely minute glands: in Scorpio occitanus there are about forty pyramidal lobuli detached from each other, the summits of which, by their union, form bunches that have their excretory canals, varying in number in different species, which convey the bile to the alimentary[Pg 124] tube; in the above insect there are six pairs, three in the trunk and three in the abdomen, and in S. Europæus a smaller number[570]; these vessels run transversely from the liver, or aggregation of conglomerate glands, to the intestinal canal[571]; the bunches consist of an infinite number of spherical glands, generally filled with a brown thick fluid[572]: besides the transverse vessels, from the base of the stomach there issue two pairs of very slender tortuous ones, seemingly analogous to the common bile-vessels; one pair of which runs upwards, one on each side that organ towards the mouth, forming here and there some ramifications which enter the liver; and the other runs nearly transversely to it[573]. As the fluid contained in these vessels is different from that contained in the glands of the liver, M. Marcel de Serres supposes they may be chyliferous[574].

In the Araneidea also the alimentary canal is nearly straight, and scarcely exceeds the length of the body: the gullet is rather thick and cylindrical[575]; the stomach is distinguished anteriorly by two pairs of sacs, the upper pair being much the largest and nearly triangular, the lower linear[576]; from these sacs a narrow tube runs towards the rectum, but which is so entangled with the liver, muscles, &c., as not to be easily made out[577]; the rectum is rather tumid, and has a lateral cœcum[578]. The disposition of the liver or conglomerate glands is stated to be similar to that of the scorpion[579]; it is usually white,[Pg 125] but in some species it is yellowish, or reddish, and its lower surface has sometimes regular excavations[580]; no transverse hepatic ducts connecting it with the alimentary canal, as in the scorpion, appear to have been at present discovered: two pairs of capillary free vessels are attached to the base of the rectum on one side, which, except in their situation, seem analogous to the bile-vessels of insects[581].

From the above detailed account of the alimentary canal of the animals whose internal anatomy we are considering, it appears that M. Cuvier's observation—that the length and complication of the intestines indicate a less substantial kind of nutriment—does not hold universally: thus, in Necrophorus and Silpha, carnivorous insects, the intestinal canal in its length and convolutions exceeds those of most herbivorous ones, and in Cassida viridis and some others of the latter tribe are not longer than those of the predaceous beetles. In herbivorous larvæ also, in general, the length of the alimentary canal does not exceed that of the body, but in those of some flesh-flies (Musca vomitoria) it very greatly exceeds it[582]. So true is the observation—that there is no general rule without exceptions.

In this letter it may not be out of place to say a few words upon the excrements of insects; which, strange as the observation may seem, but it is no less true than strange, are sometimes pleasing to the eye, from their symmetry, and to the taste, from their sweetness. In those that masticate their food they are solid, and in those that take it by suction, fluid or semi-fluid. In the[Pg 126] caterpillars of Lepidoptera they are of the former description, and every grain wears some resemblance to an insect's egg: as the passage in many of these consists of six fleshy parts separated by channels, so the excrement represents six little prisms separated by six channels[583]. The Aphides all secrete a fluid excrement as sweet as honey, of which the ants are so fond[584], which is ejected not only at the anal passage, but, in many, by two little siphonets also above it[585]. A semi-fluid excrement is produced by some species of Chermes, as that which inhabits the Box, which often comes from the animal in long convoluted strings resembling vermicelli. Reaumur says its taste is agreeable, much more so than that of manna[586]. Under this head should be included the abundant spume with which the larva of Cercopis spumaria envelopes itself[587].

I am, &c.

[Pg 127]

LETTER XLI.

INTERNAL ANATOMY AND PHYSIOLOGY OF INSECTS, CONTINUED.

SECRETION.

Having given you so full an account of the system of digestion in insects, I am now to say something concerning their secretions, and the organs by which they are elaborated. Though no individual amongst them perhaps secretes so many different substances as the warm-blooded animals; yet in general the Class abounds in secretions perhaps as numerous and extraordinary as in the last-mentioned tribes, to some of which a few of them are analogous, while others are altogether peculiar. We know little or nothing of the mode in which the process of secretion in insects is accomplished; in most cases we cannot even discover, except in general, whence the secreted substance originates; and in others, though we are able to trace the vessels that contain it, we are often in the dark as to their structure.—Cuvier, as has been before hinted, from not being able to detect any thing in them like glands, and from their being constantly bathed in the blood or nutritive fluid, conceives that they separate the peculiar substances they contain, by imbibition[Pg 128] or infiltration, through the pores of the skin[588]; a circumstance which seems to indicate a certain conformation of the pores both as to size and figure, so as to enable them to admit only one peculiar product.

In treating on this subject, I shall first consider the organs of secretion, and next their products.

I. Organs of Secretion. In general, these are membranous vessels that float in the blood or nutritive fluid, and secrete from it a peculiar substance. They may be denominated according to their products—Silk-secretors, Saliva-secretors, Varnish-secretor, Jelly or Gluten-secretor, Poison-secretor, and Scent-secretors.

i. Silk-Secretors (Sericteria). These organs are most remarkable in the caterpillars of the nocturnal Lepidoptera or moths, especially in that tribe called Bombyces, to which the silkworm belongs: but this faculty is not confined to these insects, but is shared by many other larvæ in different Orders; and in one instance at least, by the imago. In general, the outlet of the silk-secretors is at the mouth; sometimes, however, as in the larva of Myrmeleon and the imago of Hydrophilus, its exit is at the anus. The first is the organ which in the silk-worm provides for us that beautiful substance from which the animal takes its name. There are always two of these vessels, which are long floating tubes, growing slender towards the head of the insect, where they unite to form the spinneret (fusulus) before described[589], which renders the silk. Their lower extremity also is commonly more slender than the middle, and is closed at the end. These organs are usually very much convoluted and twisted[590].[Pg 129] According to Ramdohr[591], they consist of two transparent membranes, between which is found a yellow or transparent jelly. The greater the quantity of silk employed by the caterpillar in the construction of its cocoon, &c., the longer are the silk-secretors. Those of the silkworm are a foot long[592], while those of the larva of the goat-moth are little more than three inches[593].

Other insects spin silk with the posterior extremity of their body. In the great water-beetle (Hydrophilus piceus) the anus is furnished with two spinnerets, with which it spins its egg-pouch[594]; these are in connexion, probably, with the five long and large vessels containing a green fluid, described by Cuvier[595], which surround the base of each branch of the ovaries. The larva of Myrmeleon, which also spins a cocoon with its anus, differs remarkably in this respect from other insects, since its reservoir for the matter of silk is the rectum; this is connected with a horny tube, which the animal can protrude, and thus agglutinate the silk and grains of sand that compose its cocoon[596].

The web of spiders is also a kind of silk remarkable for its lightness and extreme tenuity. It is spun from four anal spinnerets, which never vary in number; two longer organs peculiar to some species have been mistaken for additional ones, but Treviranus affirms that they are merely a kind of anal feeler. Their structure, as far as known, has been before described[597]. The web is secreted in vessels varying in form. In some (Clubiona[Pg 130] atrox) they consist of two larger and two smaller ones, at the base of which lie many still more minute[598]. The four larger vessels are wide in the middle, branching at top, and below terminating in a narrow canal leading to the spinnerets[599]. Treviranus thinks the fluid contained in the lower minute vessels different from that furnished by the larger ones—but for what purpose it is employed has not been ascertained.

ii. Saliva-secretors (Sialisteria). These are organs, rendering a fluid to the mouth or stomach, that are found in many insects, especially those that take their food by suction, as the Hemiptera, Lepidoptera, and Diptera, though they are not confined to the perfect insect, being also in some cases visible in the larva. Swammerdam was one of the first that discovered them, and he suspects that they may be salival vessels; though he, as well as Ramdohr, thinks they are the same with the silk vessels of the caterpillar[600]; an opinion which Herold has sufficiently disproved, by showing that at one period of the insect's life they co-exist[601], and Lyonet discovered a very conspicuous pair in the caterpillar of the Cossus, co-existent with the silk-secretors[602]. But the physiologist who has given the fullest account of these organs is Ramdohr:—I shall therefore extract chiefly from him what I have further to communicate with respect to them.

They are variously constructed blind vessels, that are present in almost all insects that take their food by suction, but are mostly wanting in those that masticate it. They have been found, however, in Cryptorhynchus Lapathi,[Pg 131] Chrysopa Perla, and Iulus terrestris. The most usual number of the saliva-secretors is two[603]; but sometimes, as in the first of the last-named insects, there is only one[604]; in others (Pentatoma Baccarum) there are three, the exterior one consisting of a pair of reservoirs connecting with the gullet by a single capillary tube[605]; in Pentatoma prasina there appear to be four[606]; in Nepa cinerea, even six—the exterior double pair in this insect, under a powerful lens, is found to consist of spherical vesicles, resembling somewhat a bunch of currants[607]; and in Syrphus arcuatus they are covered with four rows of similar ones[608]. In the flea they consist of two pair of spherical reservoirs, each of which is connected with a short tube, which uniting with that of the other forms a common capillary one connecting with the mouth or gullet[609]; these organs sometimes terminate below in slender vessels;—thus, in Nepa, the inner pair terminates in a single vessel of this description[610], and in Tabanus and Hemerobius apparently in many[611]. It admits of a doubt however, as was lately observed, whether in the Hemiptera, which have usually more than a pair of these organs, some are not rather food-reservoirs as in the Diptera.

The saliva-secretors open either into the instruments of suction themselves (Tabanus, Musca); or into the entrance of the gullet (Pentatoma, &c.); or, lastly, into that of the stomach (Syrphus, Bombylius). Those which lie at the entrance of the stomach consist only of a blind uniform[Pg 132] tube[612]; but there is commonly to be distinguished in those that open into the mouth, a reservoir, varying in shape in different species, and terminating in a capillary tube, or tubes, at one or both extremities[613]. In Bugs, two pair of these vessels are often present, one of which opens into the stomach (Reduvius), or gullet (Pentatoma), but the other into the instruments of suction[614]. In the Diptera they open into the stomach when the insect feeds only upon the nectar of flowers (Syrphus), and into the proboscis when it feeds upon both animal and vegetable juices (Tabanus, Musca). The function of the fluid secreted by these organs is to moisten or dilute the food before it is received by the instruments of suction and passed to the stomach[615]. When a common house-fly applies its proboscis to a piece of sugar, it is easy to see that it moistens and dissolves it by some fluid.

iii. Varnish-secretor (Colleterium). In butterflies, moths, and several other insects, one or more vessels called blind vessels open into the oviduct, concerning the use of which, physiologists are not agreed. In the cabbage butterfly there is a pair of ovate ones, or rather a bilobed one, each lobe of which externally terminates in long perplexed convolutions, not easily traced, filled with a yellow fluid, which Reaumur and Herold think is used for varnishing or gumming the eggs, so that they may adhere to the leaves on which they are deposited: it may probably serve likewise for other uses[616]. Another vessel is also to be found in the above butterfly, which enters[Pg 133] the oviduct above this, filled with a thick white fluid, the function of which is, probably, to lubricate the passage[617]. A similar organ is found in Phryganea grandis[618].

iv. Jelly-secretor (Corysterium). This is a remarkable organ, related to the preceding, which secretes the jelly of Trichoptera, some Diptera, &c.; this organ in the former, at least in Phryganea grandis, is of an irregular shape, with four horns or processes[619].

Poison-secretor (Ioterium). This organ, which is most conspicuous in the Hymenoptera Order, has not received much notice, except in the case of the Hive-bee and the Scolia: in the former, it is an elliptical membranous vesicle or reservoir, furnished at its lower extremity with a tube which renders to the sting, and at the other by a blind, long, filiform, secretory, vessel, which according to Swammerdam divides into two terminal blind branches[620], though Reaumur could detect but one[621]; in this vessel the poison is secreted and stored up. In Scolia there are two secretory vessels, which enter the reservoir in the middle on each side[622]. In the Scorpion, we learn from Marcel de Serres that the poison-secretor is clothed externally with a horny thickish membrane, containing two yellowish glands, composed of an infinity of spherical glandules, terminating in a canal, enlarged towards its base so as to form a reservoir, and leading to the extremity of the sting[623]. Connected by a slender tube with each mandible in spiders is a vessel with spiral folds, which seems properly to belong to this head—though[Pg 134] Treviranus calls it a saliva-vessel[624]—since in the Mygale avicularia and other spiders, the effect of the bite is said to be so venomous as to occasion considerable inflammation, and sometimes death[625].

v. Scent-secretors (Osmateria). Amongst other means with which insects are gifted for the annoyance of their foes and pursuers, are the powerful scents which many of them emit when alarmed and in danger. Concerning the internal organs by which these effluvia are secreted we possess but little information, but more notice has been taken of the external ones by which they are emitted. We may conclude in general, that the secretory organs are membranous sacs or vesicles, perhaps terminating in longer or shorter blind filiform vessels, sometimes secreting a fetid fluid, and at others a fetid gaseous effluvium. The Iulidæ, at least Iulus and Porcellio[626], cover themselves when alarmed, with a fluid of this kind, or emit one, for this faculty is not peculiar to the species noticed by Savi. I observed early in the year, when I handled Iulus terrestris, that it was covered with a slimy secretion, of a powerful scent, which stained my fingers of an orange colour. The spiraculiform pores that mark the sides of the animal are the outlets by which this fluid is emitted, and not spiracles as has been supposed: each of these orifices, as we learn from Savi, terminates internally in a black vesicle, which is the reservoir of the fluid[627]. The most remarkable insect for its powers of annoyance in this way, is one on that account called the bombardier (Brachinus crepitans), which can fire numerous[Pg 135] volleys of stinking vapour at its assailants before its ammunition is exhausted[628]. M. Dufour has given a very particular account of the organ that secretes this vapour;—it consists of a double apparatus, one on each side, in the cavity of the abdomen, both formed of two distinct vessels. The first, which is the innermost, presents itself under two different aspects, according as it is contracted or dilated: in the former case it is a whitish, irregularly rounded, soft body, apparently glandular, placed under the last abdominal segments; communicating at one end with the reservoir, and terminating constantly at the other in a very long and slender filament: in the second case, or when it is dilated, it resembles an oblong, membranous, diaphanous sac, filled with air, then occupying the whole length of the abdomen, and appearing free except where it communicates with the reservoir. The second vessel or reservoir is a small, spherical, brown or reddish body, constant in its form, internally hollow, placed under the last dorsal segment, precisely above the rectum, and opening by a small pore into the anus[629]: so that the tail of this little beetle may be regarded as a battery mounted with two pieces of cannon, which our alert bombardier fires alternately without intermission till all his ammunition is expended. The ground-beetles (Eutrechina) in general have a pair of these anal scent-secretors, which discharge an acrid and caustic fluid, and sometimes a volatile one[630]. The external organ of the scent-secretors in Gyrinus consists of two minute hairy cylindrical retractile tubes, of a red colour[631]. Numerous insects of other tribes and genera[Pg 136] emit scents from their anus, and from various other parts of the body, of which having before given you a very full account[632], I shall proceed to the consideration of the secretions themselves: but first I must observe, that in many cases, as in some of the cottony and powdery Aphides, Chermes, &c., the substance secreted appears to be a transpiration through the pores of the body, a kind of excretion from the superabundance of its fluid contents[633]. In many, however, this secretion transpires through appropriate orifices: thus in Chermes Abietis, which produces those curious galls resembling the cone of a fir[634], the flocoons of seeming cotton that cover it proceed from little oval concavities on its back, four of which are arranged in a transverse line on each dorsal segment of the abdomen: these concavities have minute tubercles probably terminating in a pore[635]. In Aphis Fagi the cottony flocoons are almost an inch long[636].

The secretions of insects may be considered under the following heads—Silk; Saliva; Varnish or Gum; Jelly; Oils; Milk; Honey; Wax; Poisons and Acids; Odorous fluids and Vapours; and Luminous matter.

i. Silk. This valuable product of insects, while in the silk-secretor, assumes in the Lepidoptera the appearance of a viscid gum, but the moment it is exposed to the air it hardens into a silken thread. It is remarkable for the following qualities:—it dries the instant it comes in contact with the air; it is then insoluble not only in water[Pg 137] but in the most active solvents, and even heat has no effect upon it to melt or soften it: indeed, without these qualities it would be of no use to us[637]. As soon as it leaves the spinneret it becomes the thread we call silk, which being drawn through two orifices is necessarily double through its whole length. This thread varies considerably in colour and texture, as has been before stated[638], and sometimes resembles cotton or wool rather than silk. In spiders it is of a much softer and more tender texture than that of other spinning insects; and Mr. Murray seems to have proved that it is imbued, in the case of the gossamer, with negative electricity: in the sericterium the fluid that produces it is sometimes white or grey, and at others yellow[639]. A remarkable gnat (Ceroplatus tipuloides), living on an agaric, carpets its station of repose and its paths with something between silk and varnish, which it spins, not in a thread, but in a broad riband[640].

ii. Saliva. Many insects have the power of discharging from their mouth a fluid which seems in some degree analogous to the saliva of larger animals. Thus many, as Lepidoptera, Hemiptera, Diptera, &c., can dilute their food, and render it fitter for deglutition. I have seen a common fly when not employed in eating, emit a globule of fluid as big as a grain of mustard-seed from its proboscis, and retract it again. On a former occasion[Pg 138] I observed to you that many predaceous, carnivorous, and some herbivorous beetles, when alarmed emit a drop of coloured acrid fluid from the mouth[641]. That this is not secreted in any of the ordinary salival vessels is evident from Ramdohr's dissections of those beetles[642], who, had there been such an organ, would doubtless have discovered it: but as the stomach of all of them is distinguished by those minute cœca or blind vessels, which he denominates shags (zotten)[643], perhaps these may be the secretors of this fluid, probably analogous to the gastric juice[644]; in which case its primary office would be the digestion of the food. We are not however warranted in considering every fluid effused from the mouth as saliva. The glutinous material with which wasps cement the woody fibres for their paper edifices[645]; that with which some sand-wasps moisten the sand which they scrape away, of which they form the singular tubes that lead to their nests[646]; and that with which the aphidivorous larvæ fix themselves previously to their becoming pupæ[647],—may be a secretion distinct from saliva; possibly intermediate between it and gum or the matter of silk, and secreted by peculiar organs. In the wasp, however, Ramdohr discovered nothing of the kind[648]; and in Syrphus, as before observed, the saliva-secretors are very peculiar in their structure, as if appropriated to the secretion of a peculiar fluid[649]. Something[Pg 139] similar has been observed by Reaumur with regard to the larva of Crioceris merdigera, which forms its cocoon with a kind of froth produced from the mouth[650].

iii. Varnish or Gum. The eggs of various insects, when they leave the oviduct, are covered with a kind of varnish or gum by which they adhere to the substances that the young larvæ are to feed upon, or are placed in a proper position for their hatching in an appropriate station. Several instances of this have been already mentioned[651]; I shall therefore not enlarge further upon the subject. With regard to the secretion itself, little has been recorded except its colour, which has been before noticed. Some Lepidoptera also as we learn from Reaumur and Bonnet[652], use a varnish in the construction of their cocoons.

iv. Jelly or Gluten. This secretion is particularly conspicuous in the Trichoptera and some Diptera, serving as a bed or nidus for those eggs that are committed to the water,—upon which I have nothing to add to what has been before said[653]. Under this head also may be noticed the fluid, secreted in peculiar vesicles, that lubricates the oviduct and the passages of the sexual organs[654].

v. Oils. Oily substances are sometimes produced by insects. The common oil-beetle (Meloe Proscarabæus) when touched sends forth a drop of this kind of fluid, of an orange colour, from each joint of its legs[655]: something similar I have observed in Coccinella bipunctata:[Pg 140] Ray mentions a locust taken in Spain which emits a yellow oleaginous fluid from between the claws of its fore legs[656]; but the precise nature of these substances has not been ascertained, nor whether they are secreted by peculiar organs.

vi. Milk. A milky fluid is produced by the larva of Chrysomela Populi. Willughby observed a similar effusion from pores in the upper surface of the body of Acilius sulcatus; and other insects emit it from other parts of their body[657].

vii. Honey. It is certain that honey is not an animal secretion; yet the saccharine matter collected from the nectaries of flowers, from which it is derived, seems to undergo some alteration in the stomach; for the consistence of honey is greater than that of any vegetable nectar, and its taste does not vary greatly, while that of the nectar in different plants is probably not the same. Reaumur also has observed, that each honey-cell in a bee-hive is always covered by a cream-like layer of a thicker consistence than the rest, which apparently serves to prevent the more liquid honey, which from time to time is introduced under it, from running out[658]. Now if honey were the unaltered nectar of plants, it is difficult to conceive how this cream could be collected in proper proportions. The last-mentioned naturalist likewise ascertained, that if bees, in a season in which the fields afford a scarcity of food, be supplied with sugar, they will from this substance fill their cells with honey which differs in no respect from the common sort, except that its flavour is a little heightened[659]:—a similar[Pg 141] argument may be deduced from the circumstance of the bees imbibing the juices of fruits of various kinds as they are well known to do[660]. It seems therefore evident that the honey collected by bees undergoes some modification in their honey-stomach before it is regurgitated into the cells, and therefore may be regarded in some degree as a peculiar secretion.

Huber says that he has ascertained by a great number of observations that electricity is singularly favourable to the secretion of the substance of which honey is formed by flowers; the bees never collect it in greater abundance, nor is the formation of wax ever more active, than when the wind is in the south, the air humid and warm, and a storm gathering[661].

viii. Wax generally transpires through the pores of the skin of those insects that produce it, either partially or generally, and it is secreted from honey or other saccharine substances taken into the stomach. In the hive-bee, as has been before stated, it is produced partially[662], but in many other insects it is a general transudation of the body. This is particularly the case with a large number of the Homopterous Hemiptera; and those flocoons that look like cotton, and cover the body of several Chermes and Aphides, if closely examined will be found of the nature of wax: this I have particularly noticed with respect to Chermes Fagi, in which the cotton-like flocoons are often so long as to cause the insect to look like a feather, and a leaf covered by them exhibits a very singular appearance, as if clothed with the[Pg 142] fine down of a swan[663]. Probably the white powder or threads that appear to transpire through the skin of many other insects is of a waxy nature. In the larva of a beetle described by Reaumur, the flocoons are so arranged as to give the animal some resemblance to a hedgehog, and when rubbed off they are reproduced in twelve hours[664]. Gyllenhal, speaking of Peltis limbata, observes, that when alive it is covered with a white powder resembling mould, which if rubbed off returns again as long as the animal lives[665].

It will not be improper to include under this head what further account I have to give of Lac, which though regarded as a resin, since Cocci sometimes certainly produce wax[666], probably has some analogy with the latter substance. When the females of this Coccus (C. Lacca) have fixed themselves to a part of the branch of the trees on which they feed (Ficus religiosa and indica, Butea frondosa, and Rhamnus Jujuba[667]), a pellucid and glutinous substance begins to exude from the margins of the body, and in the end covers the whole insect with a cell of this substance, which when hardened by exposure to the air becomes lac. So numerous are these insects, and so closely crowded together, that they often entirely cover a branch; and the groups take different shapes, as squares, hexagons, &c., according to the space left round the insect which first began to form its cell. Under these cells the females deposit their eggs, which after a certain period are hatched, and the young ones eat their way out. Though indisputably an animal[Pg 143] secretion, many of the properties of lac are not very different from those of the juices of the trees on which the animal feeds, and which therefore would seem to undergo but little alteration.

Wax seems also to form a constituent part of some insects which are not found to secrete it. The yellow substance deposited in vessels containing spiders in alcohol is said to be a true wax, and may be obtained from these animals by gently heating them[668].

ix. Poisons and Acids. The bite as well as the sting of many insects is followed by inflamed tumours, so that the sialisteria of some bugs, Diptera, Aptera and spiders, may be regarded as producing a poisonous fluid; but we know nothing of the real nature of it, nor of that of other venomous insects, except the ant—whose celebrated acid may be considered under the present head,—the bee, the wasp, and the scorpion.

Contrary to the once received doctrine that no acid was to be found in any animal, except as the effect of disease in the alimentary canal, many insects secrete peculiar and powerful ones. I have on a former occasion related an instance in which an acid of this description, secreted in its sialisteria, is employed by a moth to soften its cocoon[669]; and Lister mentions a species of Iulus which produced one resembling that of ants[670]; but this last is the most powerful of all. The fact that blue flowers when thrown into an ant-hill become tinged with red has been long known; but Mr. Fisher of Sheffield, about 1670, seems to have been the first who ascertained that this effect is caused by an acid with which ants abound, and which may be obtained from them by distillation or[Pg 144] infusion in water[671]. Margraff and other chemists confirmed this discovery[672]; and concluding that this acid was of a peculiar kind, they gave it the name of the Formic acid. This name, however, is now exploded; the subsequent experiments of Deyeux, Fourcroy and Vauquelin having ascertained that the acid of ants is not of a distinct kind, but a mixture of the Acetic and Malic[673]. These acids are in such considerable quantities, and so concentrated in these animals, that, when a number of Formica rufa are bruised in a mortar, the vapour is so sharp that it is scarcely possible to endure it at a short distance. It also transpires from them, for they leave traces of it on the bodies which they traverse: and hence, according to the experiments of Mr. Coleridge, the vulgar notion that ants cannot pass over a line of chalk is correct; the effervescence produced by the contact of the acid and alkaline being so considerable, as in some degree to burn their legs[674]. The circumstance of much of the food of ants being of a saccharine nature may account for this copious secretion of acid, the use of which is probably to defend themselves and their habitations from the attack and intrusion of their enemies: if a frog be put into a nest of Formica rufa that has been deranged, it will be suffocated in five minutes[675]. That which they ejaculate from their anus when attacked, as formerly stated[676], must be secreted in an ioterium; but their very blood seems of an acid nature. It is very probable, as Dr. Thomson has observed[677], that acids may be obtained from many other insects, and that they are various modifications of the acetic.

[Pg 145]

From the circumstance that water is absorbed by greasy moths, that crystals of a salt are occasionally found adhering to them, that they change blue litmus paper red,—it has been inferred that their supposed oiliness is in fact an acid or acid salt, having the property of attracting moisture from the air, the infected moths being in fact not greasy, but wet; hence the application of chalk and clay, usually recommended in this case, can have only a temporary and superficial effect. The only effectual remedy, is steeping the body in spirits of wine till all the acid is extracted[678]. This acid is probably the same as Chaussier obtained from silkworms, since called Bombic Acid[679].

The poison of bees and wasps, as to its chemical qualities, is a transparent fluid, at first sweet to the taste, but immediately afterwards hot and acrid like the milky juice of the spurge[680]; soluble in water, but not in alcohol; and separable from the former in the state of white powder, when the latter is added giving a slight red tinge to paper stained with vegetable blue, and when dry and chewed appearing tenacious, gummy and elastic. This last property, as well as solubility in water and not in alcohol, is common also to the poison of the viper, which however differs in being tasteless, and not affecting vegetable blues. From hence Fontana concludes that this fluid is united with an acid, but in a very small proportion, and not with an alkali[681]. The venom of bees is extremely active; a grain in weight, it is conjectured, would kill a pigeon in a few seconds[682]. It is[Pg 146] remarkable, however, that while in some constitutions the sting of a single bee or wasp is sufficient sometimes to induce alarming symptoms, in others numerous punctures will produce little or no pain or inflammation. That this fluid, and not the puncture of the sting, is the sole cause of the inflammation that usually follows the wound inflicted by one of these animals, is proved by the facts, that if it be introduced into one made by a needle, the same effect ensues, and that when the whole contents of the poison-bag have been exhausted by the insect's stinging three or four times in succession, its weapon then becomes harmless[683].

The venom of scorpions, though much more potent, probably resembles that of bees, &c., in many of its chemical qualities: it issues from two pores in the sting before described[684], where, when the animal is irritated, it accumulates under the form of two little drops of a whitish colour: spread upon paper this fluid produces a spot like what would be caused by oil or grease, and this part of the paper becomes by desiccation firmer and transparent[685].

x. Odorous fluids and Vapours[686]. The powerful scents which different insects emit are extremely numerous, much more so indeed than the generality of Entomologists have been aware, for there is scarcely a scent odious or agreeable that may not be met with in the insect world. This you will be convinced of, by following a practice which I would recommend to you—that of smelling[Pg 147] the insects you take. Some of these scents are peculiar to particular parts or organs, and some are exhaled generally by the whole body; some are emitted by a fluid secretion, and others are gaseous effluvia. On a former occasion I gave you a rather full account of these scents and their organs[687]; I shall relate here only what I there omitted. To begin with sweet odours. Many beetles emit an agreeable scent. The rose-scented Capricorn or musk-beetle (Cerambyx moschatus) has long been noted for the delicious scent of roses which it exhales; this is so powerful as to fill a whole apartment, and the insect retains it long after its death. Captain Hancock also informed me that another species of the same genus, C. sericeus, has in a high degree a scent resembling that of the cedar[688] on which they feed. Though most of the micropterous tribes (Brachyptera) have a fetid smell, yet there are some exceptions to this amongst them. One species (Philonthus suaveolens K. MS.) related to P. micans, which I once took, smelt precisely like a fine high-scented ripe pear; another, Oxytelus morsitans, like the water-lily; a third, O. rugosus, like water-cresses; and lastly, a fourth (P. fuscipes) like saffron[689]: Trichius Eremita, one of the Petalocerous beetles, is stated to have the scent of Russia leather; Geotrupes vernalis, in spite of its stercorarious food, of lavender-water[690]. Mr. Sheppard has observed that Dytiscus marginalis when recently taken smells not unlike liquorice: Bonnet mentions a caterpillar that had the scent of new hay. A little gall-fly (Cynips Quercus Ramuli) has[Pg 148] the remarkable odour of Fraxinella: the larva of another species of this genus (C. Rosæ) has an odour which seemed to Reaumur as attractive to cats as that of Nepeta cataria or Teucrium Marum[691]: some Phalangia smell like walnut leaves[692]; and the various species of the genus Prosopis (Melitta * b. K.) have a very agreeable scent of Dracocephalum moldavicum[692].

We next come to fetid odours. These in numerous cases are known to be secreted and emitted by appropriate vessels and organs; they are often exhaled from a fluid secretion, of which, in the letter lately referred to, I gave you almost all the known instances. Savi, in his history of Iulus fœtidissimus, informs us that it emits a yellow fetid fluid from its supposed spiracles, which if applied in sufficient quantity imparts a red colour to the skin, to be removed neither by friction nor washing, but only disappearing by time; when removed from the black vesicles in which it is stored, it shoots into very transparent octahedral crystals[693].

I have before mentioned the coloured fluid which some insects emit when they are disclosed from the pupa, and that it probably exhales some powerful odour which attracts the males[694].

The great Hydrophilus, in its larva state, when first taken into the hand remains without motion; in a minute afterwards it renders itself so flaccid as to appear like a cast skin. Taken by the tail it contracts itself considerably, it then agitates itself briskly, and ejaculates with a slight noise a fetid and blackish fluid[695].

[Pg 149]

In other cases these odours are produced by gaseous vapours. That of the Bombardiers (Brachinus) is the most celebrated and remarkable. It is whitish, of a powerful and stimulating odour, very like that exhaled by nitrous acid. It is caustic, producing upon the skin the sensation of burning, and forming instantly upon it red spots which soon turn brown, and which, in spite of frequent lotions, remain several days. It turns blue paper red[696]. That amiable, intelligent, and unfortunate traveller Mr. Ritchie,—whose premature death, when attempting to penetrate to the interior of Africa, all lovers of Natural History so deeply lamented, and whose ardour in the pursuit of that science I had an opportunity of witnessing, when, in company with him, Messrs. Savigny, Du Fresne, and W. S. MacLeay in 1817, I visited the forest of Fontainebleau,—in a letter to the last-mentioned gentleman[697], relates that his companion M. Dupont, near Tripoli took a nest consisting of more than a thousand of a species of this genus. "I am making a few experiments," says he, "on the substance which they emit when they crepitate, but do not know whether I can collect enough to arrive at any conclusion. It made Dupont's fingers entirely black when he took them. It is neither alkaline nor acid, and it is soluble in water and in alcohol." From this we may conjecture that it formed crystals.

xi. Phosphorus. On this remarkable secretion I have so fully enlarged on a former occasion[698], that here I shall merely add a few observations which Mr. Murray obligingly[Pg 150] communicated to me. He remarks that in a box in which glow-worms were kept—five luminous specks were found secreted by the animal, which seemed to glow and were of a different tinge of light. One put into olive oil at eleven p. m. continued to yield a steady and uninterrupted light until five o'clock the following morning, and then seemed, like the stars, to be only absorbed by superior effulgence. The luminous spherical matter of the glow-worm is evidently enveloped in a sac or capsule perfectly diaphanous, which when ruptured discloses it in a liquid form, of the consistency of cream. M. Macaire, he observes, in the Bibliothèque Universelle, draws the following conclusions from experiments made on the luminous matter of this animal;—that a certain degree of heat is necessary to their voluntary phosphorescence—that it is excited by a degree of heat superior to the first, and inevitably destroyed by a higher—that bodies which coagulate albumen take away the power—that phosphorescence cannot take place but in a gas containing no oxygen—that it is not excited by common electricity, but is so by the Voltaic pile—and lastly, that the matter is chiefly composed of albumen.

xii. Fat. There is one product found in the body of insects most copiously in their larva state, but more or less also in the imago, which may be called their fat. In the former it is a many-lobed mass, occupying the whole of the interior, except the space that is required for the muscles and the internal organs, which it wraps round and protects. It is contained in floating membranes, very numerous, which fill all the interstices, and assume the appearance sometimes of small globules, and sometimes of a thickish mucilage, which easily melts and inflames;[Pg 151] in colour it is most commonly white, but sometimes yellow or green. It is imagined to be a kind of epiploon or caul, and is accumulated in the larva as a store of nutriment for the growth and development of the organs of the perfect insect while in the pupa state[699]. The blood in which the different organs float that is not required for their nutriment, is supposed to be expended in the formation of this substance. Marcel de Serres is of opinion that it is secreted from the chyle by passing through the pores of the dorsal vessel, formerly called the heart of insects[700].

Under this head I may mention what little is known with regard to the perspiration of these animals[701]. That a considerable quantity of fluid passes off from them when in the pupa state, is sufficiently proved by the loss of weight which they undergo, and by the experiments of Reaumur, who collected the fluid in closed glass tubes; and that in their perfect state they are constantly passing off perspirable matter by the pores of their skin or crust, is not only rendered probable by the succulent nature of their food and the absence of any urinary discharge, but is proved by what takes place in a swarm of bees. These insects, when crowded together in hot weather in a large mass, become heated to such a degree, and perspire so copiously, that those near the bottom are quite drenched with the moisture it produces, which so relaxes their wings that they are unable to fly[702].

I am, &c.

[Pg 152]

LETTER XLII.

INTERNAL ANATOMY AND PHYSIOLOGY OF INSECTS, CONTINUED.

REPRODUCTION.

The reproductive organs of insects in their general denominations and functions correspond with those of the higher classes of animals; but as to number, proportions, and other particular details of their structure, they differ from them very considerably. I shall not now, however, enter at large upon this subject, but confine myself principally to the consideration of those organs in the female which are appropriated to the formation, fecundation, maturation, exclusion and deposition of their eggs, and other circumstances relating to that subject. The organs connected with this function are the Sperm-reservoir; the Oviduct; the Ovaries; and the Ovipositor.

I. The Sperm-reservoir (Spermatheca) is an organ connecting the vagina with the oviduct, which, according to Herold, receives the male sperm as into a reservoir[703], and fecundates the eggs in their transit through that passage. This vessel, which consists of a double tunic, in the cabbage-butterfly terminates the vagina, and is connected with the oviduct by a lateral undulating tube: in shape it is a rather irregular oblong, and is surmounted by a[Pg 153] small orbicular vesicle, connected by a short tubular footstalk with the main reservoir[704]. A similar organ was discovered by Malpighi in the imago of the silkworm, who denominates it the uterus; to which indeed it seems analogous, and which he also regards as a reservoir for the sperm for the gradual fecundation of the eggs[705]. But in that fly the organ is of a rather different shape, and the interior vessel terminates in several spherical vesicles[706]. John Hunter by the most decisive experiments, such as covering the eggs of the unimpregnated moth, after exclusion, with the liquor taken from the spermatheca in those which had been impregnated, and rendering them fertile, he demonstrated that this organ was a reservoir for the spermatic fluid, to impregnate the eggs as they were ready for exclusion, and that coition and impregnation were not simultaneous[707]. It is not improbable that in all insects whose eggs are gradually laid, this provision for their gradual fecundation, if carefully sought for, might be detected[708]. Rifferschweils is of opinion, that[Pg 154] in these cases the eggs are fertilized in their transit through the oviduct by sperm adhering to the folds of the cloacæ[709]: but this opinion seems less analogous to what takes place in other cases, with regard to the due preparation of the eggs for a safe and effectual transit[710].

II. The Oviduct (Oviductus) is the canal, always separate from the vagina, which receives the eggs from the ovary, transmitting them, often by a peculiar and complex instrument in which it terminates, to their proper station. This canal sometimes opens into the anal passage or cloaca, and at others, as in the cabbage-butterfly[711], is distinct, and lies between the sexual organ and the anus. In the Arachnida there are two oviducts[712].

III. The Ovaries (Ovaria) in insects are the viscera in which the eggs are generated and grow till they arrive at maturity, when they pass through the oviduct, and are extruded or deposited in their appropriate station. They vary considerably in their structure. In all however, except the Iulidæ, in which there is only a single ovary[713], the oviduct at its upper or inner extremity terminates in two branches, usually further subdivided into a number of smaller conical ones, which several ramifications constitute the ovaries, or egg-tubes as they are sometimes called: these tubes generally consist of a[Pg 155] single membrane, and are joined to the oviduct by membranous rugose cloacæ[714]: in the Phalangia, however, there are two tunics; the outer one of a cellular substance, and the inner one consisting of spiral fibres like tracheæ—a kind of structure which renders them capable of great extension[715]. Rifferschweils considers the ovaries as formed upon two primary types.—First, flagelliform ovaries, consisting of conical tubes equal in length, and inserted at the same place at the end of the primary branches as in the Lepidoptera, the Bee, &c. Secondly, racemose ovaries, consisting of short conical tubes, so proceeding from the primary branches as to render the ovary racemose or pinnated, as in certain Neuroptera, Coleoptera, and Diptera[716]: but perhaps their structure will be better understood if they are divided into agglomerate ovaries and branching ovaries: in the first the egg-tubes form two bundles, in which the branches are not discernible, as in the Ephemera, the chamæleon-fly, and spiders[717]: and in the second the branches are distinct, as in the Lepidoptera and the majority of insects.

The number of branches varies in different genera and species. In Echinomyia grossa, a large fly, there are only the two primary branches[718]; in the common dung-beetle (Geotrupes stercorarius) these appear divided at their apex into fingers[719]: in Scolia, a Hymenopterous genus, and the butterfly of the nettle, there are three secondary branches on each side[720]: in many other[Pg 156] Lepidoptera and the humble-bee there are four[721]; in the common louse there are five[722]; in the rhinoceros-beetle and the cockchafer, six[723]; in the wasp, seven[724]; eight in the cockroach[725]; twelve in the Carabi and the mealworm-beetle[726]; thirty in the large green grasshopper (Acrida viridissima[727]); thirty-two in the cheese-maggot-fly[728]; and in the hive-bee more than a hundred and fifty[729].

The number of eggs also contained in the ovaries varies. In Echinomyia grossa there is only one egg in each, and only two at once in the matrix[730]: in another fly produced by the cheese-maggot there are four[731]; in the louse there are five; in the cockchafer six[732]; in the hive-bee sixteen or seventeen are visible at the same time[733]; and in the silkworm-moth sixty or seventy[734]. Besides the eggs, the tubes contain a pellucid mucus, and at their upper extremity the eggs are lost in a granular mucous mass, in which, however, they may still be discovered with a microscope[735]. With regard to the termination of the ovaries or egg-tubes internally,—in those that have agglomerated ones it is not to be traced, the whole appearing like an oblong obtuse or acute body[736]: but in the branching ones it is more easily traced; at first they converge in most cases to a point; this is seen to advantage in the caterpillar of some butterflies, when near assuming the pupa, in which they are readily discovered, and represent with great truth and elegance the[Pg 157] bud of some blossom[737]; but in time they diverge, and sometimes become convoluted[738]; they generally terminate in a slender simple filament, but in the louse in a fork[739]; they are sometimes extremely long, as in the wasp and Lepidoptera[740]; in the hive-bee they appear to be shorter[741].

IV. We are next to consider the Ovipositor, or instrument by which numerous insects are enabled to introduce their eggs into their appropriate situations, and where the new-born larva may immediately meet with its destined food. As this instrument is one of the most striking peculiarities with which the wisdom of the Creator has gifted these little animals, and in many cases is extremely curious and wonderful, both in its structure and modes of operation—though on a former occasion I gave you a brief account of several kinds of them[742], I shall now enter more at large into the subject, and describe these often complex machines, as they are exhibited in most of the different Orders of insects.

With regard to the Coleoptera Order, there are doubtless numerous variations in the structure of this organ; but very few have been noticed, and those chiefly belong to insects whose grubs feed on timber. In these it is usually retractile one part within another, like the pieces of a telescope: in Buprestis it consists of three long and sharp laminæ, the two lateral ones forming a sheath to the intermediate one, which probably conveys the eggs[743]: in Elater it is a cylindrical organ, terminating in a pair of conical joints, which seem to form a forceps, and including a tube probably conveying the egg to the forceps,[Pg 158] which perhaps introduces it[744]. The Ovipositor of Prionus coriarius differs from that of Callidium violaceum, and many Capricorns before described[745]: it consists merely of a long bivalve piece ending in a kind of forceps, and hollowed above into a channel for the passage of the eggs[746].

In the Orthoptera the instrument of oviposition is more simple; in Locusta consisting merely of four robust three-sided pieces, two above and two below, the former pair at the end curving upwards and the latter downwards[747], these pieces seem calculated when they have entered the earth to enlarge the burrow, and the animal appears able to separate them very widely from each other[748]. The ovipositor of Acrida viridissima, which like that of many Hymenopterous insects forms a kind of appendage or tail to the body, has been described both by De Geer and Latreille as consisting of two valves only[749]; but in reality it consists of six, two upper and four lower, as you may ascertain by means of a pin or the point of a penknife, which will readily separate them. This is confirmed by a figure of Stoll's of a species which seems to connect Conocephalus with Gryllus. In this the ovipositor is considerably longer than the body of the animal, and is composed of six distinct pieces; viz. two external ones stouter than the rest, and within these four others finer than a hair and convolute at the apex[750]. There is a considerable variety in the shape of the ovipositors of the Acridæ and the cognate genera:—thus in[Pg 159] A. viridissima this organ is straight, in A. verrucivora bent like a sabre, and in Pterophylla citrifolia and some others, the whole machine is short and boat-shaped; in Scaphura Vigorsii it is also rough with sharp little tubercles[751]. I had an opportunity of observing, with respect to the first of these insects, that in boring, as is the case with the Cicadæ and saw-flies, the motion of the valves was alternately backwards and forwards. It appeared also to me that the two outer pieces of each of the apparent valves were fixed in a groove in the margin of the intermediate one. I saw this clearly with respect to the upper pieces, and it is most probable that the lower are similarly circumstanced. In the cricket tribe (Gryllus) the ovipositor is as long as the abdomen, very slender, terminating in a knob[752]. It is apparently bivalve like that of Acrida, but I believe is resolvable into the same number of pieces.

In the Homopterous Hemiptera there seems to be more than one type on which the ovipositor is constructed. In an insect very common with us, the froth froghopper (Cercopis spumaria), some approach is made to the ovipositors last described, at least the number of pieces is the same—for it has a pair of external valves forming a sheath, which includes three sharp laminæ resembling the blades of a lancet, the middle one of which can be separated into two; this instrument De Geer had reason to think was scored transversely like a file[753]. In the insects of this Order so noted for their song[754] (Cicada), there are only five pieces; namely, two valves forming the[Pg 160] sheath, two augers or borers, and an intermediate piece upon which they slide, each being furnished with an internal groove for that purpose, and the middle piece with a ridge to fit; a contrivance of Divine Wisdom, to prevent their dislocation when employed in boring; the augers terminate in a knob which is externally toothed[755]. This structure approaches that of the Hymenoptera, especially the saw-flies. With regard to the Heteropterous section of this Order—as they usually do not introduce their eggs into any substance, they have no call for any remarkable ovipositor, and therefore are not so furnished. A remark which will also apply to the Lepidoptera Order.

In the Libellulina amongst the Neuroptera, an organ of this kind is sometimes discoverable. In Agrion, Reaumur noticed a part which he conjectured to be an ovipositor; it consists of four laminæ or lancets, the interior pair slender, the exterior wider, and all externally serrated[756].

The insects of the Hymenoptera Order have long been celebrated for the organs we are describing, whether used as saws, augers, or darts. I formerly gave you a very general account of the saws,—I shall now give you a very interesting one in detail copied from an admirable little essay of Professor Peck. "This instrument," says he, "is a very curious object; and in order to describe it it will be proper to compare it with the tenon-saw used by cabinet-makers, which being made of a very thin plate of steel, is fitted with a back to prevent its bending. The back is a piece of iron, in which a narrow and deep groove is cut to receive the plate,[Pg 161] which is fixed: the saw of the Tenthredo is also furnished with a back, but the groove is in the plate, and receives a prominent ridge of the back, which is not fixed, but permits the saw to slide forward and backward as it is thrown out or retracted. The saw of artificers is single, but that of the Tenthredo is double, and consists of two distinct saws with their backs: the insect in using them, first throws out one, and while it is returning pushes forward the other; and this alternate motion is continued till the incision is effected, when the two saws receding from each other, conduct the egg between them into its place. In the artificial saw the teeth are alternately bent toward the sides, or out of the right line, in order that the fissure or kerf may be made sufficiently wide for the blade to move easily. To answer this purpose in some measure, in that of the Tenthredo the teeth are a little twisted, so as to stand obliquely with respect to the right line, and their point of course projects a little beyond the plane of the blade, without being laterally bent; and all those in each blade thus project a little outwards: but the kerf is more effectually made, and a free range procured for the saws, by small teeth placed on the outer side of each; so that while their vertical effect is that of a saw, their lateral effect is that of a rasp. In the artificial saw the teeth all point outward (towards the end) and are simple; but in the saw of the Tenthredo they point inward, or toward the handle, and their outer edge is beset with smaller teeth which point outwards (towards the end)[757]." Valisnieri, Reaumur, and De Geer describe the groove as being in the back; but in Mr. Peck's insect, if there is no error in his account, it is, as[Pg 162] in the Cicadæ, in the saw itself[758]. In the genus Cimbex, belonging to the same tribe, the saw differs in shape, being somewhat sigmoidal or resembling the letter S, while in that of other saw-flies it is cultriform with a concave edge: other minor differences distinguish them, which need not be particularized.

A similar structure, with regard to the organ in question, obtains in the rest of the Hymenoptera, even those that use it as a weapon of offence; but the backs of the saws in them, composed of a single piece, become a sheath for the darts. The valves, however, vary. In most of those with an exerted sting, as Pimpla, they are linear, exerted, and as long as the aculeus itself[759]. In Proctotrupes they appear to be united so as to form a tube for the ovipositor, and are produced by a prolongation of the last abdominal segment. The darts usually run in two grooves of the sheath, and at their apex are retroserrulate[760]. In some cases the sheath itself is serrated[761]. The shanks of the darts are connected with the valves; so that when these open they are pushed out: sometimes on their outer side they have a triangular plate towards the base, which prevents their being pushed out too far[762].

In Sirex and many ichneumons, in which the ovipositor is too long to be withdrawn within the abdomen, it remains always exerted; but in general it is retracted within that part when unemployed. In the gall-fly (Cynips) this instrument is really as long as in Pimpla, &c.;[Pg 163] but as it is infinitely more slender, when in repose it is rolled up spirally and concealed within the abdomen. It is the puncture of this minute organ that produces the curious galls formerly described to you[763]. But the most anomalous ovipositor in this Order appears to be that of Chrysis (C. ignita, &c.), which is covered by several demi-tubes or scales enveloping and sliding over each other: when these scales are removed, the true ovipositor appears, which is of a structure similar to that of the rest of the Order, but the valves are long and slender with their summit generally visible without the anus[764].

Though the ovipositor of the majority of Dipterous insects is a tube with retractile joints[765], in the crane-flies this organ is different, and, like that of Acrida above described, consists of what at first sight appear two valves, but each of which is formed of two pieces, the upper ones sharp and longer, and the lower pair blunt. The upper pair forms the auger that bores a hole in the ground, and the lower conducts the eggs into it after it is bored[766].

In the Aptera and Arachnida in general there seems no remarkable instrument of this kind; but Treviranus has described one in spiders for extruding the eggs of a singular construction. It is an oval plate lying between the external genitals and spinning organs, and is composed of a number of small screw-shaped cartilages, connected together in the most wonderful manner. There are few organs, he observes, in the animal kingdom which[Pg 164] for their artificial mechanism can be compared with this. Each cartilage inosculates very closely in the adjoining one, and all are besides bound together by a strong skin[767].

The manner in which the eggs of insects are fecundated by the male sperm is one of those mysteries of Nature that are not yet fully elucidated and understood. We can readily conceive that all the eggs may be fertilized by a single intercourse in the case of insects which, like the Ephemeræ and Trichoptera, exclude the whole mass at once; or like many moths and butterflies, in a very short time afterwards; but the subject becomes much more difficult to explain when we advert to the female of the hive-bee, the whole number of whose eggs, deposited in two years, are, as Huber has demonstrated, in like manner fertilized by a single act[768]:—if you bear in mind, however, what I have lately observed with regard to Malpighi's discovery of a sperm-reservoir in insects, you will more readily comprehend how in this case a gradual fecundation may take place. The principal objection to this solution of the difficulty in the case before us, is derived from the very small size of the organ supposed to be destined for this purpose—it being scarcely bigger than the head of a pin[769]: it seems therefore incredible that it should retain any portion of an extraneous fluid at the end of twelve or eighteen months, and still more unlikely that the fluid should in the interval have sufficed for the slightest moistening of not fewer than 30,000 or 40,000 eggs. The only hypothesis that seems at all to square with this fact, is that of Dr. Haighton,—that impregnation is the result not of any[Pg 165] actual contact of the sperm with the eggs, but of some unknown sympathetic influence[770], or rather perhaps of some penetrating effluvia or aura seminalis, which, though small in quantity, it may retain the power of emitting for a long period.

Certain female moths, of the species of that family which, from the remarkable cases or sacs the larvæ inhabit, the Germans call sack—träger, before noticed[771], have been supposed to have the faculty of producing fertile eggs without any sexual intercourse; and various observers, after taking great pains, appeared to have satisfactorily proved the fact; so that some doubted whether these insects produced any males at all[772]. The enigma was at length explained by the accurate Von Scheven. At first his experiments were attended with the same result as those of his predecessors; but upon making them more carefully, and separating what he conceived to be the female from the male pupæ, he ascertained not only the existence of a female in the species he examined (Psyche vestita), but that when thus secluded she laid barren eggs; evidently proving that in the contrary instances above alluded to, an unperceived sexual intercourse must have taken place[773]. Though he thus ascertained that these insects do not in this respect deviate from the general rule, he remarked or confirmed several facts in their economy sufficiently anomalous and striking;—as that the female is not only without wings, but with scarcely any feature of a moth, much more closely resembling a caterpillar; and that in ordinary circumstances[Pg 166] she never attempts to leave the pupa-case in which she has been disclosed, but that being there impregnated by the male, she there also, apparently after the manner of the female Cocci, deposits her eggs, which hatching produce young larvæ that make their way out of the case, and thus seem to originate without maternal interference[774].

But the most remarkable fact bearing upon this head, though as relating to a viviparous insect it does not strictly belong to it, is the impregnation of the female Aphides, or plant-lice, before alluded to[775]. If you take a young female Aphis at the moment of its birth, and rigorously seclude it from all intercourse with its kind, only providing it with proper food, it will produce a brood of young ones: and not only this; but if one of these be treated in the same way, a similar result will ensue, and so on, at least to the fifth generation!! to which period Bonnet, who first made an accurate series of observations on this almost miraculous fact, successfully carried his experiments, till the approach of winter and the want of proper food forced him to desist[776]; and Lyonet extended it still further[777]. It is now generally admitted as an incontestible fact, that female Aphides[Pg 167] have the faculty of giving birth to young ones without having had any intercourse with the other sex. How are we to explain this most extraordinary fact? Are we to suppose with Bonnet that these insects are truly androgynous, as strictly uniting both sexes in one? This supposition, however, is completely overturned by the circumstance, that there are actually male as well as female Aphides, and that these, as was first observed by Lyonet, are united towards the close of the summer in the usual manner[778]. The most likely supposition therefore is, that one conjunction of the sexes suffices for the impregnation of all the females that in a succession of generations spring from that union. It is true that at the first view this supposition appears incredible, contradicting the general laws and course of nature in the production of animals. But the case of the hive-bee, stated above, in which a single intercourse with the male fertilizes all the eggs that are laid for the space of two years, and in the case of a common spider mentioned by Audebert[779], for many years, shows that the sperm preserves its vivifying powers unimpaired for a long period, indeed a longer period than is requisite for the impregnation of all the broods that a female Aphis can produce; and if immediate contact with the fluid be not necessary, who can say that this is impossible? It is, however, one of those mysteries of the Creator that human intellect cannot fully penetrate. But this anomaly in nature is not wholly confined to the Aphides; since Jurine has ascertained that the same thing takes place with Daphnia pennata Müll (Monoculus Pulex L.), one[Pg 168] of Branchiopod Crustacea[780]. It is worth observing whether the female Aphides in their natural state, I mean those of the summer or viviparous broods, have intercourse with the male. I think I have noticed males amongst them; but they seem to become most numerous in the autumn, preparatory to the impregnation of the oviparous females. The object of this law of the Creator is probably the more ready multiplication of the species[781].

As to the period of gestation, most insects begin to lay their eggs soon after fecundation has taken place: but in some Arachnida, as the Scorpion, which seems to be both oviparous and ovo-viviparous, nearly a year intervenes, and the eggs increase to four times the size which they had attained at that period, before they are extruded[782]. The time that is required to lay the whole they are to produce, varies also in insects. In this respect they may be divided into two great classes:—those namely which deposit the whole at once, as Ephemerina, Trichoptera, &c., and those which deposit them in succession, occupying in this operation a longer or shorter period. Many in the first class, as the Trichoptera or caseworm-flies, envelope their eggs in a gelatinous substance[783], which renders their extrusion in a mass more easy. Of the second class, which includes by far the[Pg 169] greater proportion of insects, some exclude the whole number in a very short period, others require two or three days or a week, as the cockroach[784]; and others, as the queen-bee, not less than two years. The eggs in the ovaries of the last vary infinitely in size; those that have entered the oviduct have arrived at maturity, while the rest grow gradually smaller as they approach the capillary extremity of the tubes, where they become at length invisible to the highest magnifier[785]. In many insects the eggs seem nearly to have reached their full growth previously to the exclusion of the female from the pupa; and this exclusion and the impregnation and laying of the eggs rapidly succeed each other. One moth (Hypogymna dispar), which is remarkable for the number of eggs she contains, sometimes deposits them, even before they are fecundated, in the pupa-case[786]. But in other cases the sexual union is not so immediate, and some time, longer or shorter, is requisite for the due expansion of the eggs; and the ovaries of the animal swell so much, as often to enlarge the abdomen to an extraordinary bulk: this is seen in a very common beetle (Chrysomela Polygoni) that feeds upon the knot-grass; but in no insect is it so striking as in the female of the white ants, whose wonderful increase of size after impregnation I have related to you on a former occasion[787].

I shall conclude this subject with a few observations upon ovo-viviparous insects; supposed neuters, and hybrids, which, though they do not fall in regularly under any of the foregoing heads, may very well have a place in this letter.

[Pg 170]

1. It has already been observed that there are a few ovo-viviparous insects[788], the young of which exist in the ovaries at first as eggs, but are hatched within the body of the mother, and come forth in the living form of a larva and sometimes even of a pupa. Of the first description are certain Diptera, the Aphides, and the Scorpion.

Reaumur has described two modes in which the larvæ of the first are arranged in the matrix of the mother. In some they are heaped together without much appearance of order, being placed merely parallel to each other[789]; but in others they are arranged in a kind of riband—the length of the little animals, which are also parallel, forming its thickness—rolled up like the mainspring of a watch[790]. These larvæ in general are not divided into two masses corresponding with the pair of ovaries in other insects, but form only a single one[791]. You must not suppose that these little fetuses lie naked in the womb of the mother; each has its own envelope formed of the finest membrane, which, however, is not entirely divided from that of those adjoining to it, but appears to be one tube, which becomes extremely slender between each individual, so as when drawn out to look like a chain[792]. Reaumur seems to have thought that in these flies the larvæ were never confined in any other case or egg[793]; but De Geer sometimes found eggs in the body of Sarcophaga carnaria, though most generally larvæ, from which he conjectures that it is really ovo-viviparous, the eggs being hatched in the body of[Pg 171] the mother[794]. As these flies are all carnivorous, and their office is to remove putrescent flesh, you may see at one glance the object of Providence in this law of nature—that no time may be lost, and the animal exercise its function as soon as it is disclosed from the matrix.

The Aphides, so fruitful in singular anomalies, are ovo-viviparous, as I have before hinted[795], at one period of the year, that is during the summer, but strictly oviparous at its close. From the experiments of De Geer, however, upon Aphis Rosæ, it would appear that this faculty is not conferred upon the same individuals, but only upon those of different generations of the same species; all the generations being ovo-viviparous except the last, which is oviparous[796]: nor does it appear, as has been sometimes imagined, that it is common to the whole genus. De Geer observed a species in the fir, which makes curious galls resembling a fir cone (Aphis Abietis), which appeared never to be ovo-viviparous[797].

With regard to scorpions, it does not seem clear that they are always ovo-viviparous: M. Dufour twice found in the midst of the eggs nearly mature, a young scorpion which appeared to him at large in the cavity of the abdomen; it was so large that it was difficult to comprehend how it could possibly be excluded from the animal, without an extraordinary operation[798]. The pupiparous insects (Hippobosca, &c.) have been sufficiently noticed before[799].

2. I have already in several of my former letters stated[Pg 172] to you what the modern doctrine of physiologists is with respect to certain individuals, usually forming the most numerous part of the community with insects living in society, that were formerly supposed to be neuters, or as to their sex neither male nor female—that they are in almost every instance a kind of abortive females, fed with a different and less stimulating food than that appropriated to those whose ovaries are to be developed, and in consequence in most instances incapable of conception[800]. Upon these sterile females, you also heard, devolve in general the principal labours of their respective colonies, showing the beneficent design of Providence in exempting them from sexual cares and desires, and meriting for them the more appropriate name, now generally used, of workers. The differences in the structure of the female bee and the workers were also then accounted for; and similar reasoning may be had recourse to with regard to those of ants, in which the worker and the female differ still more materially. My reason for introducing this subject here, is to observe to you that I have some grounds for thinking that this system extends further than is usually supposed, and that to each species in some Coleopterous and other genera there are certain individuals intermediate between the male and female; this I seem to have observed more especially in Copris and Onthophagus. For in almost every British species in my cabinet of these genera I possess such an individual, distinguished particularly by having a horn on the head longer than that of the female, but much shorter[Pg 173] than that of the male. I once observed a pair of Pentatoma oleracea, a very pretty bug, in coitu, both sexes being ornamented with white spots, and by them stood a third distinguished from them by red ones. I do not, however, build on this circumstance, though singular; but mention it merely that you may keep it in your eye. It would be curious should it turn up, that, to answer some particular end of Providence, in some tribes of insects there are two kinds of males, as in the gregarious ones two descriptions of females.

I am, &c.

[Pg 174]

LETTER XLIII.

INTERNAL ANATOMY AND PHYSIOLOGY OF INSECTS, CONCLUDED.

MOTION.

We have seen upon a former occasion the great variety of movements that insects can perform, and of the external organs with which they perform them[801]: but we are now to consider the internal apparatus, by the immediate action of which they take place—their system of muscles. When we reflect upon the wonderful velocity, their size considered, with which many insects move, and the unparalleled degree of muscular force that many exert[802], we feel no small degree of curiosity to know something of that part of their internal structure that produces these almost incredible effects. I shall in the present letter endeavour in some degree to gratify that curiosity, and give you an account of the muscles of these little animals,—first considering them in general; and then, as far as my information goes, adverting to those in particular that move the different parts and organs of an insect's body.

[Pg 175]

I. The muscles of insects may be considered in general as to their Origin; Substance and Parts; Shape; Colour; Kinds; Attachment; and Motions.

i. Origin. The origin of the muscular fibre in the higher animals is from the blood, the globules of which, by their coagulation in a series, appear to form it[803]; and in insects it is derived from the same universal source of nutrition and accretion, but not till it has been concreted into the adipose tissue or epiploon before noticed[804]. In the pupa of the cabbage-butterfly, Herold observed that this substance first assumed a fine flocky appearance and a blue-green colour, and that from it so changed were produced tender bundles of muscular fibres, extending in various directions, the epiploon itself decreasing in proportion as they were formed[805].

ii. Substance and Parts. The muscular fibre in vertebrate animals appears to consist of globules arranged in a series, and of no larger diameter than those of the blood,—the mean diameter of which in the human subject, when measured under the microscope by a micrometer, is found to be about ¹⁄₅₀₀₀th part of an inch[806]. When Cuvier published his immortal work in 1805, the powers of any magnifier then constructed were not sufficient to enable this great physiologist to arrive at the simple fibre[807]; but Mr. Bauer, by the use of improved glasses, amongst other discoveries that will immortalize his name, was the first to detect, under the directions of Sir E. Home, the ultimate thread of which the muscular bundles are composed[808]. Chemists distinguish the substance[Pg 176] of which we are speaking, by the name of fibrine. By the abundance of azote or nitrogen that enters into its composition, it possesses a character of animalization more marked than any other animal substance; and its elements are so approximated in the blood, that the slightest stagnation causes them to coagulate: and the muscles are without doubt, in the living subject, the only organs that can separate this matter from the mass of blood, and appropriate it to themselves[809]. The primary bundles of muscles are formed of the simple fibres, and the secondary are the result of an aggregation of the primary. The smaller bundles are not always exactly parallel to each other, but must in many cases diverge more or less, to produce those variations in shape observable in the muscles themselves: there are intervals therefore between the bundles, which in some animals are filled by a cellular substance[810]. Probably much of this statement will apply in most instances to the muscles of insects, but we may conclude that the globules that form them are infinitely smaller[811]. Lyonet has given some interesting observations with regard to those of the caterpillar of the Cossus: he describes them as of a soft transparent substance, capable of great extension, covered and filled by silver tubes of the bronchiæ, penetrated by the nerves, and containing oily particles. Each muscle was enveloped in membrane, and was composed of many parallel bands, consisting of bundles of fibres enveloped likewise in separate membranes. The fibres themselves, (but it is doubtful whether he arrived at the ultimate term of muscular fibre,) in a favourable light and under a good magnifier,[Pg 177] appeared to be twisted spirally[812]. In spiders the muscles seemed to him to consist of two substances, the one soft and the other hard, the last forming a kind of stiff twisted filament[813]. A muscle thus composed of different bundles of fibres may be stated as to its parts, in insects, to consist of base, middle, and apex: the base is that part by which they are fixed to any given point of the internal surface of the crust, or of one of its processes, which serves as their fulcrum; the apex is that part by which they are fixed, either mediately or immediately, to the organ to be moved; and the middle is the remainder of the muscle. We usually discover in them no inflation of the middle corresponding with the belly of the muscles in vertebrate animals; they occasionally, however, terminate in a tendon, as those of the thighs and legs; but these tendons are of a different nature from the fibrous ones of warm-blooded animals; for they are hard, elastic, and without apparent fibres: the fleshy ones of the muscle envelope them, and are inserted in their surface[814].

iii. Shape. The muscles of insects are usually linear, with parallel sides; some are cylindrical, as those of the wings of the Libellulina[815]; and others, as those that move the legs in the caterpillar of the Cossus, are triangular[816]. In the suctorious mandibles of the grub of a common water-beetle[817] they are penniform, or shaped like a feather; and some in the Cossus are forked[818]. Under[Pg 178] this head I may also observe, that the muscles are sometimes extremely slender threads, crossing each other, and often curiously interwoven in various directions, so as to resemble lace or fine gauze, as may be seen in the alimentary canal of some caterpillars[819]; sometimes also they surround part of this organ, like a series of minute rings[820].

iv. Colour. The most usual colour of the muscles of insects is white: those for flight however, according to Chabrier, differ from the rest, by being of a deeper and reddish colour[821]; and I have observed likewise that those in the head of the stag-beetle, when dried at least, are red, and look something like the flesh of warm-blooded animals.

v. Kinds and Denomination. In general, muscles may be regarded as divided into primary and secondary—the primary being the muscles by which the principal movements of any organs are effected, and the secondary their auxiliaries which are the cause of subordinate movements[822]. Every muscle almost has its antagonist, the action of which is in an opposite direction; so that when it is equal, the organ to which they are attached remains without motion; but when that of one preponderates, a movement in proportion takes place[823]. The principal antagonist muscles that may be found in insects are the following. 1. Levator muscles that raise an organ, and Depressors that depress it. 2. Flexors that bend an organ, and Extensors that unbend or extend it.[Pg 179] 3. Abductors that draw an organ back, and Adductors that draw it forwards. 4. Constrictors that contract an opening, and Laxators that relax it. 5. Supinators that turn the underside of an organ upwards, and Pronators that return it to its natural situation. Some of these muscles in insects, like some of their articulations and their spinal chord[824], seem to exercise a double function,—thus the levators and depressors of the wings are constrictors and laxators of the trunk[825]. At first it may seem that insects, not having the power of turning up the hand, cannot have the Supinator and Pronator muscles; but some muscle of this kind must be in the Gryllotalpa, and in those that have a versatile head[826].

v. Attachment and Insertion. The attachment and insertion of the muscles in insects in general is to the interior of the crust, or to some of its internal processes as a fulcrum, and to the organ to be moved. In some cases, however, the muscles act upon the organ by the intervention of other bodies. Thus, those that move the wings are often attached to little bones, as Chabrier calls them[827], which are connected with the base of the wings by ligaments. In the Dynastidæ and other Lamellicorns, and the Libellulina, &c., a remarkable provision is made for giving a vast increment of force to the muscles of the wings, by means of caps or cupules surmounted by a tendon, which receive their extremity; the tendon terminating in a fine point attached to the wing, and thus more muscles are brought to bear upon it[828].[Pg 180] Chabrier seems to think that, in some cases, the back that intervenes between each pair of wings is the medium by which the muscles act upon it[829].

vi. Motions. Irritability is the universal distinction of the muscular fibre,—when put in action by the will or involuntarily, it causes it to contract or become shorter; and the intermediate agents of the will and other causes are the nerves, which, as galvanic experiments seem in some degree to prove, are the conductors of an invisible fluid or power which immediately causes that action. If a nerve is divided, the muscles to which it renders obey it no longer, evidently proving that the nerves cause muscular irritability[830]. How this contraction is immediately effected,—whether the fibre, as some suppose, undergoes any crispation, or becomes zigzag[831], or whether there is any sudden change in their chemical composition that rapidly and strongly augments their cohesion, as Cuvier hints[832], cannot be clearly ascertained, unless a Bauer could submit the living fibre to his glasses. All that we know certainly on the subject is, that muscles alternately contract and relax at the bidding of the will or involuntarily, and so occasion all the movements of animal bodies.

II. Having considered the muscles of insects in general, I must next make a few observations, as far as my means of information will enable me, upon those that move their different parts and organs—at least the principal ones; since to descend to minutiæ would be an[Pg 181] endless and unprofitable labour. As larvæ, except those whose metamorphosis is semicomplete[833], differ widely in their system of muscles from perfect insects, I shall begin my observations with them.

We owe by far the most accurate and detailed account of the muscles of larvæ to the illustrious Lyonet, who, with incredible labour and patience without example, dissected the caterpillar of the Cossus, and has described every air-vessel, every nerve, and every muscle that could be detected by the microscope. Cuvier also has given a description of the muscles not only of caterpillars, but of the larvæ of the Lamellicorn beetles, the Hydrophili, and the Capricorn beetles[834]. From these sources are derived what I have now to lay before you. If you look at one of Lyonet's plates[835], the layers of longitudinal muscles look like so many parallel ribands, others run in an oblique, and others again in a transverse direction[836]. He divides them into dorsal, ventral, and lateral muscles[837], terms which sufficiently explain themselves. Of the longitudinal muscles there are four principal rows[838], the others are more numerous. The principal object of these muscles, which are flexors and extensors, is to shorten or lengthen the body, or to act on any particular segment as the circumstances of the animal may require. I shall not here notice the muscles of the head and legs, as they are not remarkably different from those of perfect insects. The prolegs are moved by two muscles—the anterior one covering in part the posterior—of a remarkable structure: one of their points[Pg 182] of attachment is by many branches or tails to the sole of the foot, and by several heads to the skin of the animal; so that they can draw the proleg within the body or push it out, and perform other necessary movements[839].

I shall now call your attention to the muscles of the perfect insect, as they move the head and its organs; the Trunk; the Abdomen; and the Viscera.

i. The Head. This part in insects moves upwards, downwards, inwards, to right and left, is pushed forth or drawn in, is often capable in part of a rotatory movement, and is sometimes versatile, turning as it were upon a pivot. All these movements are of course produced by an appropriate apparatus of muscles, which have their attachment in the anterior part of the trunk, mostly in the manitrunk, while their insertion is in the posterior part of the head, in the margin of the occipital cavity. To enumerate and describe them all would be tedious and uninteresting—I shall only mention some of the principal ones. The levators of the head are usually a pair of muscles situated in the manitrunk, to the upper side of which they are attached, and perhaps in Coleoptera and some others to the phragma, which probably Cuvier means by the anterior part of the scutellum[840]; they are inserted in the posterior margin of the upper part of the head, in Coleoptera in a pair of notches (Myoglyphides[841]), or a single one[842]. In Cordylia Palmarum these muscles as they approach the head, to judge from the dead animal, divide into two branches or a fork:[Pg 183] thus, as the muscle-notches are wide in this insect, the muscle acts upon each extremity of the sinus—these branches appear to be tendinous[843]. The depressors of the head are the antagonist muscles to the above, and have their attachment to the antepectus and its antefurca[844]. A circumstance distinguishes these muscles in many Coleoptera, that seems hitherto to have been overlooked. If you take the common dung-beetle (Geotrupes stercorarius), and carefully extract the head with its muscles from the trunk, you will see on each side of the depressors a subovate corneous scale, of a pitch colour[845], which is attached only to the muscle, and designed to strengthen it: if you then examine the anterior cavity of the manitrunk, you will perceive on each side, just within the lower margin, a minute triangular scale, of a similar substance; these ligaments, like the pax-wax, or ligamenta nuchæ, in mammalia, though in a lower situation, are doubtless intended to sustain the action of the muscles.

With regard to the moveable organs of the head—the antennæ, maxillæ, palpi, tongue, mandibulæ, &c., have each their appropriate apparatus of muscles: but I shall only notice those of the last, the mandibulæ. These are principally abductors and adductors to open and shut them: from the work that the jaws of some insects have to do, you may conjecture that they must be furnished with powerful muscles. In caterpillars and other larvæ, in which state the action of the mandibles is most in requisition, the muscles are what Cuvier calls penniform[846], and are attached on each side to a tendinous lamina or[Pg 184] cartilage. In the grub of Dytiscus the power and magnitude of the adductor muscle is wonderful[847]. In the Orthoptera this structure of the mandibular muscles takes place also in the imago[848]; but in the Coleoptera, at least in the stag-beetle and some others that I have examined, these muscles in this state have no cartilage or tendon. Their attachment is always to the parietes of the head, of the cavity of which the adductors, in some cases, occupy a considerable portion[849]. As to their insertion—these last, in some Orthoptera, enter more or less the interior of the mandible[850]; but commonly they are inserted at or near the interior angle of the mandibular basal cavity, and the abductors at the exterior.

ii. The Trunk. We have little information with regard to the muscles of the parts of the trunk itself, by which, in some insects, the manitrunk is enabled to move independently of the alitrunk: it is more probable that the levators have in part at least their attachment to the anterior surface of the prophragm[851], than that the levators of the head should be there fixed, as Cuvier seems to think; since both the phragma and the ligament that appears in many cases to close the cavity of the manitrunk round the viscera[852], would prevent all communication between those muscles and any part connected with the scutellum: probably the depressors have their attachment partly on the anterior face of the medifurca[853]. These points, however, must be left to future investigators.

[Pg 185]

With regard to the organs of the trunk, we have more certain and satisfactory information;—the muscles of the legs having been described by Lyonet and Cuvier, and those of the wings most particularly by Chabrier. In caterpillars, the muscles are situated in the interior of the articulations that form the legs: they consist of several bundles appropriated to each, which have their attachment in the parietes of the preceding joint, near the margin, and are inserted in the margin of that they move[854]. Lyonet counted twenty-one muscles in the leg of the caterpillar of the Cossus; but eight of these were appropriated to the claw, or rather formed a pair of semipenniform muscles, having their insertion at the inner angle of its base[855]. In perfect insects, according to Cuvier, each joint of the legs is furnished with a pair of antagonist muscles—a flexor and extensor, the former being the lower, and the latter the upper muscle; and this pair has its insertion in the joint it moves, and its attachment usually in the preceding one: but those of the coxæ—which are rotators, causing it to turn backwards or forwards—and the extensor of the thigh, have their attachment in the parietes of the trunk, and to the endosternum; one of the rotators of the anterior coxa, and the extensor of the anterior thigh to the antefurca; of the intermediate pairs to the medifurca, and of the posterior to the postfurca[856]. Every joint of the tarsus has also its flexor and extensor. In the ground- and water-beetles (Eutrechina and Eunechina), &c., whose posterior coxæ are immoveable, the thigh includes two pair of[Pg 186] antagonist muscles[857]. In extracting the posterior leg of Necrophorus Vespillo I observed more than a single pair of muscles that had their attachment in the coxa; and probably many other variations in this respect exist.

Little was known with respect to the most interesting part of the muscular apparatus of insects, that by which such wonderfully rapid and varied motions are imparted to their organs of flight, till Chabrier undertook to elucidate it; which he has done in a manner that will confer a lasting honour upon his name, as one of the most able successors to Swammerdam and Lyonet in their peculiar department. He has given a most admirable account of the internal anatomy of the trunk of insects in general, as far as it relates to their flight; particularly of that of the cockchafer (Melolontha vulgaris), of one of the Libellulina (Æshna grandis), and of a bumble-bee (Bombus); and I believe he has thus illustrated insects of some of the other Orders, but his memoirs on these I have not had an opportunity of consulting. What I have to say on this subject, therefore, will be principally derived from what he has communicated with respect to the above insects.

A considerable difference in the volume of the muscles of the wings takes place in insects according to the force of their flight. Where it is rapid and powerful, the alitrunk is nearly filled by them, and the alimentary canal is much attenuated; but in those whose flight is feeble, they occupy less space, and the alimentary canal is proportionally enlarged[858]. In the Lepidoptera, Hymenoptera and Diptera, the principal muscles of both[Pg 187] wings have their attachment in the anterior portion of the alitrunk[859]; in the Coleoptera, in the posterior[860]; and in the Libellulina, those of the anterior wings are confined to the anterior portion, and those of the posterior pair to the posterior[861]. The muscles for flight in general differ from others by their mass, length, and colour; the bundles of fibres are very distinct, strong, and parallel; their direction is uniform, according to the motion they are to produce; their fibres are either attached to the solid parts to be moved, or to cupules, but they never terminate in a tendon; the muscles are perfectly independent of each other, and the wings can be moved by them separately[862]. As to their denomination and kind—the principal ones are the levators and depressors, which with respect to the trunk, as was before observed, are constrictors and laxators. The levator muscles form several distinct bundles in Coleoptera, Lepidoptera, &c.; in the Diptera there are three[863]; in the Libellulina they seem to be single, are all environed with a blackish pellicle, with numerous aërial vesicles, symmetrically arranged, filling the interstices[864]. The most common number is a levator to each wing; there are often, however, as in the cockchafer and the dragon-fly, two depressors[865]: but in the Hemiptera, Lepidoptera, and saw-flies (Serrifera) amongst the Hymenoptera, the secondary wings have distinct levators, but not depressors[866]; the other insects of that Order have only a pair of each[867].[Pg 188] The other wing-muscles are of a secondary description, and auxiliary to the above. Their office is to extend and close the wings: so that though the denomination of extensor will suit the former, that of flexor is not so proper for their antagonists; their office being not so much to bend, as to bring back the wing to its station of repose. The folding of certain wings, as those of Coleoptera, Dermaptera, the Vespidæ, &c., seems more the function of the abdomen than of the wing-muscles; this you may easily see, as I have often done, if you attend to any Staphylinus, when after alighting from flight it proceeds to fold up its wings under the elytra. Perhaps the term retractor might not be inapplicable to the muscles in question. Both these and the extensors are usually small slender muscles, but sometimes numerous[868]. They are larger in the Coleoptera, Lepidoptera, and saw-flies[869]. The muscles that open and shut the elytra of Coleoptera, and probably of Heteropterous Hemiptera, and which also aid their movements during flight, are very slender[870]. With regard to the attachment and insertion of the wing-muscles, it is according to two very distinct types, one of which appertains to insects in general, and the other is peculiar to the Libellulina. In insects in general, the principal muscles for flight have not their insertion in the wings, but act upon their bases by the intervention of small long pieces. The depressors occupy the middle and upper region of the alitrunk, and are inserted anteriorly and posteriorly upon the concave surfaces of two transverse horny semi-partitions, adapted by their elasticity to dilate the trunk—and thus acting[Pg 189] the part of both diaphragm and ribs[871]: but in the Libellulina, as in birds, these muscles are placed on each side of the point of support of the humerus[872]; the depressors being attached immediately to the wings without it, and the levators within it, with this sole difference, that they are connected to the internal extremity of the base of the wing by the intervention of a cupule terminating in a tendon; all are disposed perpendicularly to the arms of the levers on which they act, and all incline more or less outwards, the one to dilate, and the other to contract the trunk[873]. It may be observed in general, that in insects formed upon the first type, the great action of these muscles is the dilatation and contraction of the alitrunk, the main tendency of which is to depress and raise the wings[874]. I shall add here a few words upon the attachment of the wing-muscles in the different Orders: but first I must request you to read what I have said on the partitions and chambers of the alitrunk in a former letter[875]. In most insects of the first type, the depressors are longitudinal dorsal muscles that have their posterior point of attachment in the metaphragm (costale Chabr.); but the anterior varies:—in those that have elytra, tegmina, or hemelytra, the muscles for them seem to be contained in the chamber, varying in size, that lies between the prophragm and mesophragm; and the anterior point of attachment of their depressor muscles is the mesophragm: they are also attached in some to the metathorax or back of the posterior portion of the alitrunk[876].[Pg 190] The levator muscles in Coleoptera, at least in the cockchafer, by a long tendon have their posterior attachment in the lower part of the posterior coxæ[877], their anterior attachment to the solid parts to be moved. In the Cockchafer and the Dynastidæ, but not in Geotrupes, on each side of the cavity of the metathorax under the base of the wing is a large and small cupule, which from their lateral situation one would think must receive the levator muscles—apparently unnoticed by M. Chabrier; but as there is a pair of these cupules on each side, there must have been also a pair of muscles attached to them, which does not agree with his statement[878]. In the Hymenoptera and Diptera the anterior attachment of the depressors is to the back of the alitrunk and to the prophragm, and the levators to the breast, and the sides of the back of the trunk[879]. In the Libellulina the depressors and levators that terminate, by a tendon surmounting a cupule, in the base of the wings, have their posterior attachment in the breast. These cylindrical muscles with their cupule and tendon look like so many syringes[880].

Having thus described to you the powerful muscular apparatus by which, either mediately or immediately, the wings of insects are moved, it will not be out of place if I add a few words upon their flight itself. The great object in this is to generate a centrifugal force which may counteract the weight of the body. Its wings are the external organs by which the insect as it were takes hold of the air when they fall, and is impelled by it when[Pg 191] they rise; its head makes way for it; its abdomen, as a rudder, steers it; and by alternately increasing and diminishing in volume, and rising and falling, enables it to win an easy way through the fluctuations of the atmospheric sea. The trunk by its elasticity admits the internal action of antagonist muscles, which by turns compress and dilate it; an action promoting the elevation and depression of the wings, and keeping up the elasticity of the internal air, which is thus now rarified and now condensed: in the former state flowing like a tide, accompanied by the blood, into the nervures of the wings[881], and thus increasing their tension and centrifugal force;—in the latter ebbing and receding to the trunk, thus relaxing the one and diminishing the other. The spiracles by which the air enters or is expelled, open and shut at the animal's pleasure[882]; and besides, many insects are furnished, as we have seen[883], with numerous vesicles or reservoirs, which can give out a supply of internal air when wanted: and thus they can vary their aërial motions, diminish or increase the counteracting centrifugal force; rise and fall, and move onwards and in different directions, as their occasions demand.

iii. The Abdomen is perhaps capable of the greatest variety of motions of the three primary sections of the body. Even when the insect is reposing, a constant dilatation and contraction usually takes place in it[884]; and from its annular structure, its parts capable of separate motion are numerous:—it expands and contracts; it[Pg 192] rises and falls; it bends in various directions; and its segments can often be lengthened or retracted. Besides all this, its spiracles open and shut, and its reproductive and other anal organs have their appropriate motions. In numerous Coleoptera, however, and some Hemiptera, the upper-side of the abdomen is almost the only part that is moveable, especially near the trunk; the under-side, having its first segments soldered together, is only capable of motion near the tail[885]. The muscles that produce the various motions of this part must be entitled to all the denominations stated above[886]. I have on a former occasion explained to you how, in insects that have a petiolate abdomen, that part is elevated and depressed[887]. In those with a sessile one the base is attached to the metaphragm by strong ligaments[888], and the muscles that move the first piece act from one segment to another. The partial movements of the segments of this part, where they have place, are produced by muscular fibres which extend from the whole anterior margin of one to the whole posterior one of that which precedes it. If those, for example, of the back contract, the abdomen becoming shorter above, bends upwards; and if those of the sides or belly, it bends sideways or downwards[889]: this is a beautiful as well as simple contrivance.

The alternate rush of air from the abdomen into the alitrunk, and from the atmosphere into the abdomen, is attended by the constriction or expansion of that part[Pg 193] as it rises or falls in flight[890], which seems to require the action of constrictor and laxator muscles.

iv. The Viscera. Having before had occasion sufficiently to notice the muscles by which the systole and diastole of the dorsal vessel of insects is maintained[891], I shall now only mention those that are woven round their alimentary canal, by which the peristaltic motion of that organ, causing its contractions and the propulsion of its contents, takes place. One would at first think that a view of the intestines of any animal could under no circumstances afford any very pleasing spectacle to the eye of any but a scientific spectator; but any lady who is fond of going to Disons to be tempted with an exhibition of fine lace, would experience an unexpected gratification could she be brought to examine those of a caterpillar under a microscope: with wonder and delight she would survey the innumerable muscular threads that in various directions envelope the gullet, stomach, and lower intestines of one of these little animals; some running longitudinally, others transversely, others crossing each other obliquely, so as to form a pattern of rhomboids or squares; others again, surrounding the intestine like so many rings, and almost all exhibiting the appearance of being woven, and resembling fine lace,—one pattern ornamenting one organ; another, a second; and another, a third. This will suffice to give some idea of this part of the muscular structure of these little animals[892].

Lyonet counted the muscles contained in the body of the caterpillar of the Cossus. In the head he found 228; in the body, 1647; and enveloping the intestines,[Pg 194] no less than 2186; which, after deducting 20 that are common to the gullet and the head, gives a total of 4061[893]. In the human subject only 529 have been counted[894]: so that this minute animal has 3532 muscles more than the Lord of the creation!

The muscles of the Arachnida seem less numerous than those of insects. In the Scorpionidea they appear to be robust, formed of simple straight fibres, of a whitish gray colour: a muscular web, rather strong, clothes the parietes, but rarely adheres to them, of the abdomen, and envelopes the viscera, with the exception of the lungs, and probably of the heart. The dorsal part of this web gives birth to seven pairs of filiform muscles, which traverse the liver, and are attached to a muscular riband which, passing above the lungs, runs the whole length of the ventral parietes. These muscles when exposed to view resemble extended cords. The abdominal segment preceding the tail is filled with a powerful muscular mass which moves that organ[895]. Treviranus discovered two longitudinal muscles in Scorpio europæus, running from the breast to the tail, which above and below each gill were connected by another running transversely across the heart, thus forming a quadrangular area in which the gills are situate[896]. The heart appears to be moved by muscles not very dissimilar to those of the Cossus[897], as is likewise that of the Araneidea; in Clubiona atrox the wider part of this organ is muscular, and incloses a considerable cavity[898]. In this tribe the muscles of the[Pg 195] abdomen, the skin of which is soft and unfit to act as a lever to them, are attached to a cartilage, and thus their action is better sustained[899].

Having thus laid before you all of importance that I can collect with regard to the apparatus of muscles discoverable in insects, I shall next say something upon a few other points connected with that subject. When I enlarged upon their motions, I related a few instances of the extraordinary power of that apparatus[900] in leaping ones; but this power is not confined to that circumstance. The flea, not more remarkable for its compressed form, enabling it to glide between the hairs of animals, and its elastic coat of mail, by which it can resist the ordinary pressure of the fingers, than for its muscular strength, has attracted notice on this account from ancient times. Mouffet relates that an ingenious English mechanic, named Mark, made a golden chain of the length of a finger, with a lock and key, which was dragged by a flea;—he had heard of another that was harnessed to a golden chariot, which it drew with the greatest ease[901]. Another English workman made an ivory coach with six horses, a coachman on the seat with a dog between his legs, a postillion, four persons in the coach, and four lacqueys behind—which also was dragged by a single flea. At such a spectacle one would hardly know which most to admire, the strength and agility of the insect, or the patience of the workman. Latreille mentions a flea of a moderate size dragging a silver cannon on wheels, that was twenty-four times its own weight, which being charged with powder, was fired[Pg 196] without the flea appearing alarmed[902]. Many caterpillars are accustomed to extend their bodies from a twig, supported merely by the four hind feet, in one fixed attitude, either in an oblique, horizontal, or vertical direction, either upwards or downwards, and that for hours together. We may conceive what prodigious muscular force must be exerted upon this occasion, by reflecting that the most expert rope-dancer, though endued with the power of grasping with his feet like a bird with its claws, could not maintain himself in a horizontal position even for an instant. Bradley asserts that he has seen a stag-beetle carry a wand half a yard long and half an inch thick, and fly with it several yards[903]. Some insects have the faculty of resisting pressure in a wonderful degree. If you take a common dung-chafer (Geotrupes) in your hand and press it with all your strength, you will find with what wonderful force it resists you; and that you can scarcely overcome the counteraction, and retain the insect in your hand: was it not for this quality, the grub of the gad-fly must be crushed probably in passing through the anal sphincter of the horse[904]. But that of Eristalis tenax affords a more surprising instance of this power of counteraction:—an inhabitant of muddy pools, it has occasionally been taken up with the water used in paper-making, and strange to say, according to Linné, has resisted without injury the immense pressure given to the surrounding pulp[905]; like leather-coat Jack mentioned by Mr. Bell[906], who, from a similar force of[Pg 197] muscle, could suffer carriages to drive over him without receiving any injury. Almost as remarkable is the state of extreme relaxation into which the muscles of some larvæ fall, when their animation is suspended; and the revived tension to which a subsequent resumption of the vital powers restores them. Bonnet having suspended the animation of the caterpillar of Sphinx Ligustri by keeping it submerged, squeezed it between his fingers, until it had wholly lost its cylindrical form and was as flat and supple as the empty finger of a glove; yet in less than an hour the very same caterpillar became as firm, as compact, as cylindrical, and in short, as well, as though it had never been submitted to treatment so rough[907].

It is fortunate that animals of a large size, as has been well remarked, especially noxious ones, have not been endowed with a muscular power proportionable to that of insects. A cockchafer, respect being had to their size, would be six times stronger than a horse; and if the elephant, as Linné has observed, was strong in proportion to the stag-beetle, it would be able to pull up rocks by the root, and to level mountains[908]. Were the lion and the tiger as strong and as swift for their magnitude as the Cicindela and the Carabus, nothing could have escaped them by precaution, or withstood them by strength. Could the viper and the rattlesnake move with a rapidity and force equivalent to that of the Iulus and Scolopendra, who could have avoided their venemous bite? But the Creator in these little creatures has manifested his Almighty power, in showing what he could have done had he so willed; and his goodness in not creating the[Pg 198] higher animals endued with powers and velocity upon the same scale with that of insects, which would probably have caused the early desolation of the world that he has made. From this instance we may conjecture, that after the resurrection, our bodies by a change in the structure and composition of their muscular fibre—for we know that their locomotive powers and organs, as far as the muscle is concerned, will then be of a very different nature[909]—may become fitted for motions and a potent agency of which we have now no conception.

This wonderful strength of insects is doubtless the result of something peculiar in the structure and arrangement of their muscles, and principally their extraordinary power of contraction, excited by the extent of their respiration: for animals that respire but little, as the fœtus in the womb and the pullet in the egg, have very little contractile muscular power[910]. To get some idea from facts of this extraordinary contractile power in insects,—extract the sting of a bee or a wasp, with its muscles, which appear to be attached to powerful cartilaginous plates[911], and you will find it continue for a long time to dart forth its spicula, almost as powerfully as when moved by the will of the animal. A still more extraordinary instance of irritability is exhibited by the antlia, or instrument of suction of the butterfly. If this organ, which the insect can roll up spirally like a watchspring or extend in a straight direction, be cut off as soon as the animal is disclosed from the chrysalis, it will continue to roll up and unroll itself as if still attached[Pg 199] to its head: and if after having apparently ceased to move for three or four hours it be merely touched, it will again begin to move and resume the same action. This surprising irritability and contractility of muscle doubtless depends upon the peculiar structure of the antlia, which is composed of an infinite number of horny rings, acted upon by muscles, more numerous probably than those which move the trunk of the elephant. The motion only ceases when the muscles become dry and rigid.

I have already, under another head[912], considered the annual sleep, or winter state of torpidity of insects, during which an intermission for the most part of muscular motion and action takes place. I shall now make a few observations with respect to their diurnal sleep, which may very properly have its place in the present letter. That insects, usually so incessantly busy and moving in every direction, require their intervals of repose, seems to call for no proof. We see some that appear only in the day, and others only in the night, others again only at certain hours; which leads to the conclusion, that when they withdraw from action and observation, it is to devote themselves to rest and sleep. The cockchafer flies only in the evening; but if you chance to meet with it roosting in a tree in the earlier part of the day, you will find it perfectly still and motionless, with its antennæ folded and applied to the breast:—we cannot indeed say that its eyes are shut; for as insects have no eyelids, that sign of sleep can never be found in them. Again, if a Lepidopterist goes into the wood to capture moths in[Pg 200] the day-time, he finds them often perched on the lichens that cover the north side of the trunk of a tree, with their wings and antennæ folded, and themselves without motion, and insensible of his approach and their own danger. Thus it was that I captured that rare insect the lobster-moth (Stauropus Fagi) in the New Forest. Some, however, have asserted that the caterpillar of the silkworm, except when they moult, never intermits feeding day or night, and consequently does not sleep: but the accuracy of this statement, both from analogy and observation, admits of great doubt. Malpighi informs us that these caterpillars for an hour and more, twice a day, remain immoveable with their heads bent down as though asleep, and even if disturbed, resume again the same inactive posture[913]; and other larvæ in great numbers certainly seem to have regular intermissions from eating of considerable duration: those called Geometers, for hours together remain motionless projected from a twig, to which they adhere by their posterior prolegs alone; and the processionary caterpillars make only nightly sorties from their nests, passing the day in inaction and repose[914]. Bees have been often seen by Huber, when apparently wearied with exertion, even in the middle of the day, to insert the half of their bodies into an empty cell, and remain there, as if taking a nap, without motion for half an hour or longer[915]; and at night they regularly muster in a state of sleep-like silence. Mr. Brightwell once observed an individual living specimen of Haltica concinna, which appeared to remain motionless on the same spot of a wall for three successive days.

[Pg 201]

Before concluding these remarks on the Internal Anatomy and Physiology of Insects, I shall explain to you, as you will probably feel inclined occasionally to pursue the subject, the best mode of dissecting them.—By far the most useful dissecting instruments for this purpose are very fine-pointed and sharp scissors, as these will enable you to divide the integument and separate other parts with much less risk of injuring their delicate structure than any knife. These scissors are what Swammerdam chiefly used; and he had some so extremely small and fine, that he was necessitated to employ a lens when he sharpened them. If to these be added a sharp and fine-pointed knife or two, some needles fixed in handles, also fine-pointed—(you will find them more convenient than any other instrument for detaching minute parts and fibres,) a pair of fine and accurately adjusted pliers, and an assortment of camel's-hair brushes,—you will be nearly set up as an Entomological dissector. You will still, however, require a small dissecting table, with a projecting and moveable arm for lenses of various descriptions, so as to admit both the hands to be employed upon the subject under examination; and for this purpose probably no contrivance can be better adapted than that of Lyonet, of which the figure in Adams On the Microscope will convey a better idea than any description[916].

Previously to dissecting any insect, it must be killed by plunging it into boiling water, which is recommended by Lyonet, or spirits of wine or of turpentine; and it is often useful to let larvæ remain a few days in the latter,[Pg 202] by which means the vessels become firmer and stronger. The parts of pupæ become much more distinct if they are boiled for a few minutes: and the same mode may be adopted in the examination of spiders.

The most convenient mode of proceeding, which was that also of Lyonet, is to dissect the insect in water, or, to avoid putridity, in diluted spirits,—if small, upon a concave glass, to which it should be fastened by means of a little melted wax; if larger, in the bottom of a common chip box, surrounded with a border of wax to retain the fluid. The integuments of the insect, being carefully divided longitudinally with scissors, should if flexible be turned back, and fixed by small pins stuck in by a fine pair of pliers, while the skin at the same time is stretched by another. After making such observations as present themselves without further dissection, the viscera must be cautiously extracted, washing away the fat which surrounds them with spirits of turpentine, in which it is soluble, applied by camel's-hair pencils. After separation they may conveniently be examined by putting them into water, and gently shaking them so as to cause the parts to unfold. If endowed with the patience of Swammerdam, you may even arrive at injecting these minute parts with wax or coloured fluids, conveyed by delicate glass tubes having one end as fine as a hair, which he also employed to fill the viscera with air; and afterwards drying them in the shade, and anointing them with oil of spike in which a little resin had been dissolved, he succeeded in preserving them. If it is not convenient to finish the dissection of an insect at once, it should be covered with spirits of wine. Swammerdam found a mixture of spirits and distilled vinegar[Pg 203] very useful for keeping caterpillars previously to dissecting them, as it consolidated the parts[917].

And now having brought to a close my long wanderings in this ample and intricate field, and having threaded, as well as my slender powers and limited knowledge enabled me, the infinite turnings and convolutions of this Dædalean labyrinth—the Anatomy and Physiology of insects,—will you not own that the volume of wonders I have laid before you proves irrefragably that, though these minims of nature apparently rank so low in the scale of being, yet in their structure, instead of being, as might be expected, more simple, they are infinitely more complex and highly wrought than those animals that are placed the nearest to ourselves? the Creator in the latter doing every thing by a beautiful simplicity; while in the former, the more to magnify his power and skill, because they afford no apparent space for it, by a wonderfully curious and intricate multiplicity: and whether we study the one or the other, we shall in both trace the footsteps of that adorable Love which has shown attention to the comfort and well-being of the lowest insect, as well as of the highest of his creatures.

I am, &c.

[Pg 204]

LETTER XLIV.

DISEASES OF INSECTS.

Having laid before you what observations I thought might sufficiently explain all the principal features of the Anatomy of insects both external and internal, you will next expect to be informed whether, like the higher animals, they are subject to have the admirable order observable in their frame interrupted by Disease; and you will perhaps imagine, from the multiplicity of their organs and vessels, that they must be peculiarly exposed to derangements of the vital and other functions. That they have their diseases is certain; but, except in the case of their appropriate parasitic assailants, which is a part of their economy, it does not appear that their maladies are more numerous and frequent than those of other animals. The same Almighty Power which endowed them with so complex a structure, generally upholds them in health during their destined career, until they have fulfilled the purpose of their creation, when they die and return again to their dust[918].

But perhaps I may seem to you as making too great a parade about these little insignificant creatures if I assign a separate letter to the consideration of their diseases: but when you recollect that Aristotle has a chapter on[Pg 205] this subject[919], and that the learned Willdenow has devoted a distinct portion of his excellent introductory work on Botany to the diseases of Plants[920],—you will perhaps be of a different mind: indeed, some facts I shall have to communicate are so remarkable and interesting, that I am sure, when you have read this letter, you will not think the subject one that deserves to be slighted.

Insect diseases may, I think, be divided into two great classes; those resulting, namely, from some accidental external injury or internal derangement, and those produced by parasitic assailants.

I. Under the first head we may begin with wounds, fractures, mutilations, and other extraneous causes of disease. To these—insects are peculiarly subject; and though they are not, like the Crustacea and Arachnida[921] and some other invertebrate animals, endowed with the power of reproducing a mutilated limb, yet their wounds appear to heal very rapidly, and at the time they are inflicted to produce little pain[922]. But if those important members, their antennæ, are mutilated, insects seem to suffer a kind of derangement; the great organ of their communication with each other, and in various respects with the external world, being removed, all their instincts at once fail them. I formerly related how the amputation of these affects the queen-bee[923]. A similar result, as Huber[Pg 206] tells us[924], follows, when the same experiment is repeated on the workers or drones: they immediately become unable to take any further part in the labours of the hive; they can no longer guide themselves except in the light; if they petition one of their fellow-citizens for honey, they are unable to direct their tongue to its mouth to receive it; they remain near the entrance of the hive, and when the light is intercepted they rush out of it to return no more.

Insects occasionally are subject to tumours or a preternatural enlargement of their parts and organs. The antennæ of bees sometimes swell at their extremity so as to resemble the bud of a flower ready to open, becoming at the same time very yellow, as does the fore part of the head[925]. I once saw a specimen of a Hydrobius—agreeing with H. fuscipes in every other respect even to the most minute punctum—which had a large tumour on each side of the prothorax, evidently accidental, occasioned probably by the stoppage of the pores by which the superfluous moisture and air escape when it undergoes its last change. The converse of this I have observed to take place sometimes in the same part of Geotrupes foveatus, the ordinary lateral foveæ becoming very considerably enlarged;—this was the case with the specimen from which Mr. Marsham made his description of that insect. The species is, however, very distinct in other respects, and may always be known by its small size. It happens now and then also, that these tumours represent blisters. I saw one once on one elytrum of a beetle and not on the other. Those of Serropalpus (as[Pg 207] Mr. MacLeay, on the authority of M. Clairville, informs me) are particularly subject to this disease. But, of all the organs, the wings are most exposed to derangements of this kind. De Geer, in a specimen of Pieris Cratægi just excluded from the chrysalis, observed that one of these was distended by a considerable quantity of extravasated green fluid—two or three large drops following an incision. This disease appeared to arise from the lower membrane not adhering to the upper; so that the nervures—which are rather longitudinal channels, being open below, than tubes—were not closed to confine the fluid to its proper course. The malady, which might be called a dropsy of the wing, carried off the insect the day after its exclusion[926]. Reaumur observed that the wings of some flies were affected by an air-dropsy, as he calls it, which appeared to arise from the air escaping from its natural channels, and thus separating, the two membranes that form the wing, and filling the cavity produced by their separation[927].

Sometimes also monstrosities are to be met with in these animals, or variations from a symmetrical structure in organs that are pairs. I have a beetle in which the terminal joint of one of the maxillary palpi is short, ovate, and acute; and that of the other, long, semiovate, and rather obtuse. A specimen of Blaps mortisaga in my cabinet, taken by Mr. Denny, besides the terminal mucro of the elytra, has a long diverging lateral one. Goeze had the larva of a Semblis brought to him in which one of the two fore-legs, though perfect in all its parts, was only half the length of the other[928]; which he regarded[Pg 208] as a reproduction, but it seems rather a malformation. Müller mentions a most extraordinary fact of one of the Noctuidæ, which when disclosed from the pupa retained the head of the larva[929]. One of the most remarkable instances of this kind that have fallen under my own observation, may be seen in a specimen of Chrysomela hæmoptera in the cabinet of our friend Curtis; in which one of the thighs produces a double tibia, but only one of these is furnished with a tarsus.

The diseases of insects which arise from some internal cause are not very numerous. The first that I shall mention is a kind of vertigo. "Ants have also their maladies," says M. P. Huber: "I have noticed one extremely singular; the individuals attacked by it lose their power of guiding themselves in a straight line, they can walk only by turning round in a circle of small diameter and always in the same direction. A virgin female shut up in one of my glasses was seized on a sudden with this distemper; she described a circle of an inch in diameter, and made about a thousand turns in an hour, or not quite seventeen in a minute. She continued constantly turning round for seven days, and when I visited her in the night I found her still in motion. I gave her honey—and I think that she ate some of it." He observed that some workers were attacked by a similar disease: one of these, however, had the power of walking from time to time in a straight line; when placed upon its head it continued its gyrations[930]. Similar motions of a little moth, mentioned on a former occasion[931], may perhaps have been produced by the same cause. Bees are also[Pg 209] subject to vertigo, which has been attributed to their eating poisonous honey[932]—but may not this disease in all these cases arise from some derangement of the nervous system? One of the ants which was so affected had lost one of its antennæ; but as this was not the case with the others, no great stress is to be laid upon the circumstance. Huber does not inform us whether those attacked by this disease recovered or not.

I have observed more than once, that the flesh-fly and some others of the same tribe are subject in particular seasons to a kind of convulsions. When thus attacked, they kick and struggle, and seem unable to fly. Sometimes they lie upon their backs without motion, but if a finger be placed near them their convulsive motions are renewed. When thrown into the air, instead of flying, they fall to the ground. Had this distemper occurred earlier or later in the year I should have attributed it to the benumbing effects of cold; but as my observations were made one year (1816) in May, and in another (1811) in the latter end of June, this could scarcely be the case. In the year last mentioned I observed that many flies died under its influence. In wet seasons this tribe is subject to another disease, which proves fatal to many of them, and indeed to other Diptera. A white crust appears to be formed upon the abdomen both above and below, of a granular appearance, much resembling fine moist sugar. On the back of that part this crust does not cover the margins of the segments, which gives it the appearance of white bands; so that deceived by it, I have often at first flattered myself that I had met with[Pg 210] some new species. The under-side of the abdomen is wholly covered by it, divided in the middle into two longitudinal masses, the anal segment being bare. De Geer has noticed this or a similar disease, which, when flies are attacked by it, causes the abdomen to swell so as even to burst, and the segments become dislocated. Upon opening the abdomen it is found filled with a white unctuous substance, which often accumulates (as above described) on its external surface[933]. Dr. Host says that in this disease when the animal is dead, the wings, which were before incumbent, become extended, and its almost invisible pubescence grows into long hairs[934]. De Geer seems to think that these flies are thus affected in consequence of having eaten some poisonous food[935]; but I rather suspect, as I have observed it become prevalent chiefly in wet seasons, that it arises from a superabundance of the nutritive fluid, or of the fat, so that it seems to be a kind of plethora. I once observed a fly fixed to a pane of glass, round which was a semicircle of what appeared to be merely vapour, whose radius was nearly three-fourths of an inch. Taking it for an aqueous fluid that had transpired from the dead animal, I paid no further attention to it at that time: but observing from day to day that the moisture did not evaporate, after two or three months had elapsed, I had the curiosity to examine it more closely, and, upon scraping some of it off with a penknife, I found it was a white substance of a fatty nature. In this case, then, the fat must have exploded on all sides with considerable violence from half the body or the abdomen. Probably this was[Pg 211] a more intense degree of plethora. When I examined this appearance the fly had fallen off, and I could not find it.

Mr. Sheppard once brought me a panicle of grass, the glumes of which were rough with hairs, or small bristles, to which several specimens of a fly related to Xylota pipiens adhered by their proboscis. At first I thought that having been entrapped by the bristles, and unable to extricate themselves, they had perished from want of food; but since when touched they readily dropped from the glumes, some other cause, perhaps disease, probably occasioned this singular suspension of themselves.

The maladies to which bees and silkworms are subject are more interesting to us than those of flies, on account of their utility as cultivated insects. One of the worst distempers which attacks the first of these animals is a kind of looseness or dysentery: this happens early in the year, when they are fed with too much honey without any portion of bee-bread[936], and sometimes destroys whole hives. Their excrements, instead of a yellowish red, then become black, and the odour they emit is insupportable; the bees no longer observe their usual neatness, inducing them to leave the hive when they void their excrements, but they defile it, their cells, and each other. Several remedies have been prescribed for this disease. To prevent it, a syrup made by an equal mixture of[Pg 212] good wine and honey is recommended; and as a cure, to place in the hive combs containing cells filled with bee-bread[937]. But one of the worst maladies to which these useful animals are subject, is that called by Schirach Faux Couvain. It originates with the larvæ; and is caused either by their being fed with unwholesome food, or when the queen, as sometimes happens, lays her eggs so that the head of the grub is not in a proper position for emerging from the cell when the period for its disclosure is arrived:—the consequence is, that in both cases it dies and becomes putrid, which sometimes produces a real pestilence in a hive. The remedy for this evil is to cut away the infected combs, and to make the bees undergo a fast of two days[938]. The hive should be cleaned and fumigated, by burning under it aromatic plants.

The cultivators of the silkworm in France have given names to several diseases to which that animal is subject. One is called La Rouge, and is supposed to be occasioned either by too great heat, or by too sudden a transition from cold to heat. It takes place when the caterpillar is first hatched; which lives perhaps, but in a very sickly state, till it should spin its cocoon and assume the pupa, when it expires. Another degree of the same disease is called Les Harpions or Passis. A second distemper of this animal is Des Vaches, Le Gras or La Saune: this is a mortal disease, supposed to be of a putrid nature, and produced by mephitic air; it shows itself after the second moult, but rarely after the subsequent ones. When a caterpillar is first attacked, changing the air may prove a remedy; but when the disease has made[Pg 213] progress, it is best to burn or bury them, since if the poultry pick them up they might be poisoned by them. A third disease of silkworms is called Les Morts Blancs, or Tripes, which is also occasioned by impure air, when the leaves the animal feeds upon are heaped so as to produce fermentation. The caterpillars attacked by it die suddenly, and preserve after their death the semblance of life and health. Too great heat, whether artificial or natural, occasions La Touffe, a fourth, which, when the heat continues long, destroys all those that are arrived at their last stage of existence in their larva state. Black points scattered over different parts of the body, or livid and blackish spots in the vicinity of the spiracles, followed by a yellowish or reddish tint, are symptoms of a fifth malady, called La Muscardine. After this the animal soon dies, and becomes mouldy, but does not stink. This disease is not contagious, and is thought to be caused by a moist heat, attended by pernicious exhalations. La Luzette, Luisette, or Clairène, is another malady, which shows itself most commonly after the fourth moult. It seems to arise from some original defect in the egg. The caterpillars attacked by it may be known by their clear red and afterwards dirty white colour; their body becomes transparent, and the matter of silk exudes in drops from their spinnerets; consequently, though as voracious as the rest, they are never able to construct a cocoon, and should be destroyed. Les Dragées is the name given to cocoons which include a larva that never becomes a pupa. The cause of this disorder has not been ascertained, and whole broods are sometimes subject to it, which, as in the last, seems to imply some defect in the eggs. But as the caterpillar spins its cocoon, and the silk is as good as usual,[Pg 214] it is a malady of no great importance. Lastly, sometimes the mulberry leaves have a gummy rather acrid secretion, which purges the silkworms; their excrement is no longer solid; they become weak and languid; and if the secretion is abundant, their transpiration is impeded, and at the time of moulting they are become so feeble as to be unable to cast their skin[939].

In the case of many caterpillars of Lepidoptera that died, Bonnet found by dissection that the disease was remotely occasioned by a diarrhea, which taking place immediately before they became pupæ, prevented the inner membrane of their intestines from being rejected, as it would have been if no extraordinary cause had prevented it, attached to the hard excrement. He found this membrane converted into a jelly occupying great part of the stomach, which he conjectured was the proximate cause of their death[940].

To conclude this head—spiders are reputed to be subject to the stone: I do not say Calculus in Vesica; but we are informed by Lesser that Dr. John Franck having shut up fourteen spiders in a glass with some valerian root, one of them voided an ash-coloured calculus with small black dots[941].

II. I now come to that class of diseases which appears to prevail almost universally amongst insects—I mean those resulting from the attack of parasitic enemies. Thus millions and millions annually perish before they have arrived at their perfect state. Diseases of this kind proceed either from vegetable or animal parasites. I shall begin with the first, which will not occupy us long.

i. As insects pass often no small portion of their life[Pg 215] in a state of torpidity, in which they remain chiefly without motion, it will not seem wonderful, should any partial moisture accidentally accumulate upon them, that it affords a seed plot for certain minute fungi to come up and grow in. Persoon observes with regard to his genus Isaria, that one species grows upon the larvæ of insects (I. truncata), and another upon pupæ (I. crassa[942]):—as he does not say upon dead larvæ and pupæ, as upon a former occasion[943], perhaps in these cases these plants may constitute an insect disease; but I lay no stress upon it, and only mention the circumstance here as connected with the history of these animals. Mr. Dickson has described a Sphæria under the name of entomorhiza that grows upon dead larvæ; it has a slender long stipes and spherical granulated head: on the pupa of a species of Cicada in my cabinet, another kind of Sphæria, with a twisted thickish stipes and oblong head, springs up in the space between the eyes. I observed something similar but longer, in the grub of some large beetle in M. Du Fresne's museum at Paris; and I have a memorandum of having noticed something of the kind on the rostrum of a Calandra. Bees and humble-bees have been sometimes thought to have some species of mucor or other Fungilli occasionally growing upon them; but Mr. Brown is of opinion that stamina which they have filched from flowers have been mistaken for these Fungilli, since he has detected those of Orchideæ in some of this tribe, and upon a beetle shown to him by Mr. MacLeay, one which he knew to be the stamen of an Aristolochia. I once observed a bunch of what I mistook for a singular mucor that adorned the vertex of a humble-bee, between the antennæ, which doubtless were[Pg 216] of the same description; and I even saw one upon its wing. Upon a former occasion I mentioned a parallel circumstance with respect to a species of Xylocopa[944].

ii. The animal parasites that infest insects are either themselves insects; or worms.

1. Their insect infesters, as far as we know at present, are confined to the Orders Strepsiptera, Hymenoptera, Diptera, and Aptera: they attack them sometimes in their egg state, most frequently when they are larvæ, occasionally when pupæ, and very rarely in their perfect state. Upon many of these I have formerly enlarged[945], and I shall now add such further circumstances as I then omitted. The Strepsiptera Order, as at present known, consists only of two genera, Stylops and Xenos; the first being appropriated to the imago of Andrena, a kind of bee, and the latter to that of the wasps. Their eggs appear to be deposited in the abdomen of these insects in which they feed, till having attained their full growth they perforate the membrane that connects its segments; and at the proper time their pupa-case bursts, they emerge, and take their flight. Sometimes four or five infest a single bee. Whether the latter dies upon their quitting it I have not been able to ascertain, but from their flying, when the little parasite is very near leaving them, with their usual activity, it should seem that this disease is not mortal; but it probably prevents their breeding: I do not recollect observing the exuviæ of one in a male bee[946].

The great body of insect parasites, however, belong to the Hymenoptera Order, and chiefly to the Linnean genus Ichneumon. The insects of this order have been[Pg 217] denominated Principes, because of the wonderful instincts of ants, wasps, bees, and other gregarious tribes that belong to it; and they merit a name of honour not less for the benefits that they confer upon mankind, by keeping within their proper limits the various insect-destroyers of the produce of the globe. It deserves notice that when these latter increase to a degree to occasion alarm, their parasites are observed to increase in a much greater, so as to prevent the great majority of them from breeding[947]. Though these benefactors of the human race constitute numerous genera, at present not well ascertained, I shall speak of most of them under the common name of Ichneumon.

The appearance of these little four-winged flies puzzled much the earlier naturalists:—that a caterpillar usually turning to a moth or butterfly should give birth to myriads of flies, was one of those deep mysteries of nature which they knew not how to fathom[948]: even the penetrating genius of our great Ray, though he ultimately ascertained the real fact[949], was at one time here quite at fault; for he seems at first to have thought, when from any defect or weakness nature could not bring a caterpillar to a butterfly, in order that her aim might not be entirely defeated, that she stopped short, and formed them into more imperfect animals[950].

Before I detail more particularly the proceedings of Ichneumons, I shall make a few general remarks upon them. The structure of the instrument by which they are enabled to deposit their eggs in their appropriate station has been before sufficiently described[951]; it is long[Pg 218] or short according to the situation and circumstances of the larva which receives them: if this lives in the open air, and the access to it is easy, it is usually short and retracted within the body; but if it lies concealed in deep holes or cavities, or shuns all approach, it is often very long. Thus in Pimpla Manifestator, which commits its eggs to the grub of a wild bee inhabiting the bottom of deep holes bored in posts and rails, the ovipositor is nearly an inch and half in length, and in some extra-European species three inches. How the egg is propelled so as to pass in safety from the oviduct, along this extended and very slender instrument to the grub for which it is destined, has not been certainly ascertained: but from an observation of Reaumur's[952] it should seem that it is aided in its passage by some fluid ejected at the same time with it, or is so lubricated as to slide easily without being displaced. The flies we are speaking of, by some authors are called Muscæ vibrantes, because when searching for the destined nidus of their eggs their antennæ vibrate incessantly, and it is by the use of these wonderful organs that they discover it wherever it lurks. Bergman observed that Fœnus Jaculator searches for the latent grub of certain bees and other Hymenoptera with its antennæ[953]: and from Mr. Marsham we learn that Pimpla Manifestator, before it inserts its ovipositor in the nest of the grub of Chelostoma maxillosa, explores it first with one antenna and then with the other, plunging them all the while intensely quivering up to the very root[954]. With respect to their size, Ichneumons vary greatly; some being so extremely minute as to be invisible to the naked eye, unless moving upon glass; while others, as to[Pg 219] their length, emulate the giants amongst insects. The former, unless appropriated to the eggs themselves, usually commit many eggs to a single larva, while the latter are directed by their instinct to introduce into them only one. Some of the former description are endowed with the faculty of leaping[955]. The food of Ichneumons, and indeed of other internal parasites, is chiefly the epiploon or fat of the larva, but they never touch any vital organ; so that it continues to feed, and probably more voraciously, grow, cast its skin, and often it changes to a chrysalis, although at the same time inhabited by an army of these little devourers.

Ichneumons, as far as has been at present ascertained, are parasitic upon other insects chiefly in their three first states, a solitary instance only having been observed of their inhabiting an imago; but from their first exclusion as eggs from the ovary till their assumption of that state they give them no rest. I shall therefore first treat of those that infest the eggs; next those appropriated to larvæ; and lastly those that devour pupæ.

Vallisnieri appears to have been the first naturalist who discovered that Ichneumons were appropriated to the eggs of other insects. He observed one proceed from those of the emperor-moth (Saturnia Spini): finding two holes in each egg, one larger than the other, he conjectured that one was made when it entered, and the other when it emerged. In this case the egg of the Ichneumon must be fixed on the outside of the egg it[Pg 220] was to feed upon; though some appear to pierce it with their ovipositor, and consequently introduce their egg within: for he says afterwards; "I have seen with my own eyes a certain kind of wild flies deposit their eggs upon other eggs, and bore and pierce others with an aculeus—by which they have introduced the egg[956]." Count Zinanni, a correspondent of Reaumur's, saw an Ichneumon pierce the eggs with her ovipositor repeatedly; which in about fifteen days were filled with the pupa, and in six more produced the imago[957]. I. Ovulorum L. is the only known species of egg-devourers; but most likely there are many, varying in size, according to the size of the egg they inhabit. Probably I. Atomus L., and I. Punctum Shaw, are of this description[958]. It is wonderful what a number these little flies destroy:—out of a mass of more than sixty eggs which was brought to De Geer, not one had escaped the Ichneumon[959]. But the most extraordinary thing is, that even these little creatures we are told are destroyed by another still more minute[960].

Though the animals we are speaking of usually destroy only a single egg, yet some appear not so to confine themselves. Geoffrey informs us that the larva of one of the Ichneumons whose females are without wings (Cryptus) devours the eggs of the nests of spiders, and from its size—it is nearly a quarter of an inch long—it must require several of them to bring it to maturity[961]. One of those also which destroys the gnat infesting the wheat (I. inserens) appears to devour them in[Pg 221] their egg state, and could not be brought to perfection by the food that a single one would furnish[962].

The Ichneumons that are parasitic upon larvæ are the most numerous of all. Some of them are deposited by the parent fly on the outside of their prey, and others introduced into its interior. Ophion luteum is one of the former tribe; it plants its eggs in the skin of the caterpillar of the puss-moth (Cerura Vinula). Each egg is furnished with a footstalk terminating in a bulb[963], which is so deeply and firmly fixed that it is impossible to extract it without detaching a portion of the animal with it, and even when the caterpillar changes its skin it is not displaced. After it is hatched, the grub, while feeding, keeps its posterior extremity in the egg-shell, to which it adheres so pertinaciously, that it is scarcely possible to disengage it without crushing it. It fixes itself by its mandibles to the skin of the caterpillar, and keeps constantly sucking the contents of its body till it dies: sometimes nine or ten of these larvæ inhabit a single caterpillar[964]. Reaumur has given an account of other external Ichneumons. Upon one caterpillar that he examined, they were so numerous as to render the poor animal quite a spectacle, and they underwent their metamorphosis attached to it[965]. One species of this description avenges the cause of insects upon their most pitiless foes, the all-devouring spider—for in the midst of her toils and lines of circumvallation it makes her its prey. De Geer, meeting one day with a young spider of a common kind, observed with surprise, engaged in sucking it, a small white grub, which was firmly attached to the abdomen near the trunk. Putting it by in a[Pg 222] glass, after some days he examined it again; when he observed that it had spun the outline of a vertical web, had stretched threads from the top to the bottom of the glass and from one side to the other, and had also spun the radii that meet in the centre, and this was all;—but what was remarkable, the larva that had fed upon it was suspended in the centre of this web, where it was engaged in spinning its own cocoon, while the spider, exhausted by this last effort, had fallen dead to the bottom of the glass. It cannot be asserted positively that this suspension of the larva of the Ichneumon in the centre of the web always takes place; but if it does, as seems most probable, it shows that this little parasite is endowed with an instinct which causes it so to act upon the spider as may induce it to spin a web so nicely timed as to be sufficiently complete at the period of its death and of the change of the Ichneumon, for the latter to cast it down and assume its station[966].

But the great bulk of the parasitic Hymenopterous devourers of larvæ have their assigned station within the body. As Entomologists in breeding insects have paid their principal attention to Lepidoptera, it necessarily follows that their Ichneumon infesters must be most generally known; but doubtless the larvæ of the other Orders are not wholly liberated from this scourge: they also require to be kept within due limits, and have their appropriate parasites. Some, however, in most of them have been detected: of which I shall now proceed to state to you the most interesting examples, beginning with the Coleoptera.

Alysia Manducator[967], remarkable for having mandibulæ[Pg 223] that do not close, and toothed at the end, usually attends masses of dung, both of man and cattle, probably for the purpose of depositing its eggs in some of the Coleopterous larvæ that inhabit it. Mr. Stephens, one of the most accurate observers as well as one of the best Entomologists of the present day, informs me that he once captured three specimens of Timarcha tenebricosa, from each of which forty or fifty minute Ichneumons emerged. An insect also of this Order, that is a great benefactor to mankind, as a destroyer of the plant-lice,—I mean the lady-bird (Coccinella), in its larva state is itself subject to the attack, as we learn from De Geer, of one of these small parasites[968]. He detected them also in that of two species of weevils: and in the pupa of some large grub of a beetle inhabiting the wood of the elm, perhaps that of the stag-beetle, he found the pupa of one of those Ichneumons that have an exserted ovipositor[969]. Doubtless, did we know their history, we should find that numberless species have their internal assailants belonging to this tribe.

Orthopterous larvæ seem not to have been yet announced as affording a pabulum to these animals: but the late Dr. Arnold, whose tact for observation with regard to the manners and economy of insects has rendered his loss irreparable, discovered that the remarkable parasitic genus Evania was appropriated to the all-devouring Blatta. Whether it attacked it in its egg or larva state I have not been informed. This little benefactor is here extremely rare, at least in the country; perhaps in towns, where the cock-roach abounds, it may be more common.

[Pg 224]

The observations of naturalists have chiefly been confined to the Hemipterous genus Aphis; but these early attracted their notice. Leeuwenhoek has given a particular and entertaining account of the proceedings of I. Aphidum. As soon as the little flies approached their prey, they bent their abdomen, which is rather long, between their legs, so that the anus projected beyond the head; then with their ovipositor they pierced the body of the Aphis, at the same time carefully avoiding all contact with it in every other part: whenever they succeeded in their attempt, a tremulous motion of the abdomen succeeded. Only a single egg is committed to one Aphis: when hatched, the latter becomes very smooth and appears swelled; it is, however, full of life, and moves when touched. Those that are thus pricked separate themselves from their sound companions, and take their station on the underside of a leaf. After some days the inclosed grub pierces the belly of the Aphis, and attaches the margin of the orifice to the leaf by silken threads; upon this it dies, becomes white, and resembles a brilliant bead or pearl[970]. De Geer observed also an Ichneumon on the Coccus of the elm, I. Coccorum[971].

Amongst the Neuropterous tribes likewise, probably the Ichneumonidæ commit their usual ravages; but their exploits, as far as I recollect, have met with no historian. I have a small species related to Chelonus, which a memorandum made when I took it tells me was obtained from Æshna viatica; yet I do not remember ever tracing that species to its final change, so that I must have taken this Ichneumon from the perfect insect. It suffices, however, to prove that this tribe is also exposed to the attack[Pg 225] of these parasites. Where larvæ and pupæ are aquatic, it seems probable, if any attack is made upon them, that it must take place after they have quitted the water.

In the Hymenoptera Order itself, almost every genus has been ascertained to have its Ichneumon parasites. Not even the fortified habitations of the gall-flies (Cynips) can escape them, almost every species becoming their prey; a circumstance which puzzled not a little some of the older naturalists, when they at one time saw a fly not remarkable for its colours or brilliancy emerge from the curious moss-like Bedeguar of the wild rose, and at another were struck by the appearance of one of those splendid minims of nature which almost dazzle the sight of the beholder[972]. Immunity, however, from this pest seems to have been granted to the gregarious Hymenoptera; at least none has yet been discovered to attack the ant, the wasp, the humble-bee, or the hive-bee; in which last, had there been one appropriated to it, it could never have escaped the notice of the Reaumurs and the Hubers. The solitary bees, however, as we have seen above[973], do not escape; and Epipona spinipes, a solitary wasp which feeds its own young with a number of green caterpillars[974], is itself, when a larva, though concealed in a deep burrow, the prey of the grub of an Ichneumon, which by means of a long ovipositor introduces its egg into its body[975]. Even these parasites, whose universal office it is in their first state to prey upon insects, are themselves subject to the same malady. Ichneumonidan devourers are kept in check by other[Pg 226] Ichneumonidan devourers. These in some cases are so numerous as to destroy the tithe of the kinds they attack[976]. Thus an ever-watchful Providence prevents these parasites from becoming so numerous as to annihilate in any place the species necessary for the maintenance of the general economy and proportion of animal and vegetable productions. Amongst the assailants of the Hymenoptera, none seem to have a more laborious task assigned them than those that pierce the various galls in which the larvæ of the Cynips tribe are inclosed. To look at an oak-apple, we should think it a work of difficulty, requiring much sagacity and address, for one of our little flies to discover the several chambers lurking in its womb, and to direct their ovipositor to each of them. Its Creator, however, has enabled it instinctively to discover this, and furnished it with an appropriate elongated instrument, which will open a way to the deep and hidden cells in which the grubs reside, penetrate their bodies, and to each commit an egg. When it prepares to perforate the gall, the Ichneumon begins by depressing this organ, that it may extricate it from its sheath; it next elevates its body as high as possible, and bending the instrument till it becomes perpendicular to the body and to the gall, so as to touch the latter with its point, it then gradually plunges it in, till it is quite buried[977]. A very remarkable Hymenopterous parasite (Leucospis), which when unemployed turns its ovipositor over the back of its abdomen, so that its end points to its head, is said to deposit its eggs in the nest of the mason-bee, most probably in the larva: but the curious observations that are stated to have been[Pg 227] made by M. Amédée Lepelletier upon its history have not yet been given to the public[978].

Dipterous insects, likewise, do not escape from these pests of their Class: but few observations, however, have been recorded as to the species assailed by them. We learn from De Geer, that a gnat (Cecidomyia Juniperi), which forms galls upon the juniper is devoured by an external Ichneumon[979]; that which injures the wheat in the ear, whose ravages I formerly mentioned to you[980], affords food to three of these parasites,—one I lately mentioned as probably devouring its eggs; another pierces the glumes of the floret, where its destined prey is concealed; and the third enters it. I once placed a number of the larvæ of the gnat upon a sheet of paper, at no great distance from each other, and then set down one of these last Ichneumons in the midst of them. She began immediately to pace about, vibrating her antennæ very briskly: a larva was soon discovered, upon which she fixed herself, the motion of her antennæ increasing intensely; then bending her abdomen obliquely under her breast, she inserted her ovipositor, and while the egg was depositing these organs became perfectly motionless. The larva when pricked gave a violent wriggle. This operation was repeated with all that had not already received an egg, for only one is committed to each larva. I have often seen it mount one that was already pricked, but it soon discovered its mistake, and quitted it untouched[981]. The Hessian Fly also (Cecidomyia Destructor) related to the preceding, whose alarming ravages I formerly described to you[982], has a peculiar parasite[Pg 228] attached to it, which keeps it in check. The only other Dipterous insects that I have seen mentioned as affording pabulum to an Ichneumon, are—one of the aphidivorous flies mentioned by De Geer, who does not note the species, to the larva of which the Ichneumon commits only a single egg, producing a grub that intirely devours its interior[983];—and two described by Scopoli, one, the larva of a fly frequenting hemp; and the other, which feeds on a Boletus, that of a gnat[984].

The Lepidoptera, however, is the Order over the larvæ of which the Ichneumons reign with undisputed sway; attacking all indiscriminately, from the minute one that forms its labyrinth within the thickness of a leaf, to the giant caterpillar of the hawk-moth. The most useful of all, however, the silkworm, appears at least with us, exempted from this scourge. De Geer, out of fifteen larvæ that were mining between the two cuticles of a rose-leaf, belonging to the first tribe here alluded to, found that fourteen were destroyed by one of these parasites, only one coming forth to display itself in all its brilliancy and miniature magnificence[985]. One of the most useful to us is that which destroys the clothes-moth, which the same writer also traced[986]. Another, equally serviceable, takes up its abode in the caterpillar that ravages our cabbages[Pg 229] and brocoli (Pontia Brassicæ) which perish by hundreds from its attacks. As this falls frequently under our notice, it will not be uninteresting to give a fuller account of it. Reaumur has traced and related its whole history. One of these little flies that he observed, was so intent upon the business in which she was engaged, that she suffered him to watch her motions under a lens, without being discomposed. She pursued nearly the same plan of proceeding with that of the Ichneumon of the wheat-gnat just described; except that she repeated her operations frequently on the same caterpillar in different parts, alternately plunging in and extracting her ovipositor. She seemed to prefer the spot where the segments of the body are united, particularly where the eighth meets the ninth, and the ninth the tenth. When the fly had completed its work and quitted the caterpillar, Reaumur gave it food, and it did not seem less lively and vigorous than others of its kind; in less than a fortnight it assumed the pupa; and in four days the whole of its interior being devoured, it died: but its parasites, perhaps not finding a sufficient supply of nutriment in it, never came to perfection[987]. Sometimes, however, these little grubs arrive at maturity before the caterpillar has become a chrysalis, when they pierce the skin and begin to emerge. First appears a little white tubercle, which gradually elevates itself in a direction perpendicular to the body; while this is doing, a second appears in another place; and so on, till fifteen or sixteen are seen on each side, giving the caterpillar a very grotesque appearance. By the alternate contraction and relaxation of their bodies the grubs effect their complete liberation, which takes[Pg 230] place with respect to the whole in less than half an hour. When entirely disengaged, they place themselves close to the sides of the caterpillar: even before this they begin spinning, and draw unequal threads in different directions, of which they form a cottony bed, which serves as the base of the separate cocoon of each individual, which they next construct of a beautiful silk thread of a lovely yellow, which, if it could be unwound and in sufficient quantity, would yield a silk unrivalled in lustre and fineness[988].

De Geer has recorded a very singular fact which deserves your notice. An Ichneumon, appropriated to one of the Tortrices, had deposited its eggs in two of their caterpillars; each produced a considerable number; but those that emerged from one were all females, and those from the other, males[989]. He observed a similar fact take place with Misocampus Puparum[990]. One might conjecture from this circumstance, that as in the queen-bee[991], so in these Ichneumons, the eggs producing the two sexes were arranged separately in the ovaries. Reaumur has related, that in one instance three or four males were produced to one female; and in another four or five females to one male[992].

But though the great majority of insects are subject to this Scolechiasis[993] in their larva state, yet sometimes they are not attacked by the Ichneumon till they have become pupæ. Of this kind is one just mentioned (M. Puparum), which commits its eggs to the chrysalis of the butterfly of the nettle, Vanessa Urticæ: the moment this caterpillar[Pg 231] quits its skin to assume that state, while it is yet soft they pierce it and confide to it their eggs[994]. De Geer and others have supposed that this same Ichneumon attacks the Cocci and Coccinellæ[995]; but this probably is an erroneous supposition. Cryptus Compunctor also attacks the pupæ of butterflies[996].

If we consider the great purpose of Providence in giving being to this tribe of destroyers—the keeping of insects within their proper limits,—we may readily conceive that this purpose is more effectually answered by destroying them in their preparatory than in their ultimate state, since at that time the laying of their eggs and a future progeny could not so effectually be prevented;—this will account for there being few or no Ichneumons appropriated to them in their latter state.

The next tribe of insect parasites are to be found in the Diptera Order. The species that has been particularly noticed as such is the Tachina Larvarum; its larva is polyphagous, laying its eggs upon the bodies of caterpillars of different kinds. Sometimes a pair is placed on the first segment, sometimes on the head itself, and sometimes near the anus. These eggs are very hard, convex, of an oval figure, polished and shining like a mirror. They are fixed so firmly that if you attempt to remove them with a penknife the skin comes off with them. When hatched, they enter the body and feed on the interior, and, undergoing their metamorphosis within it, do not emerge till they enter their perfect state. The caterpillar thus attacked lives long enough to spin its cocoon, when it dies[997]. Sometimes, however, these animals[Pg 232] quit their prey sooner. Reaumur saw a grub of one of the Muscidæ come out of a caterpillar, and then become a pupa, which was so large that he wondered how it could have been contained in the animal it had quitted[998].

We have now done with those parasites that produce in insects the disease I have called Scolechiasis[999]: the rest, which belong to the Aptera Order, will afford us examples both of Phthiriasis and Acariasis[1000].

I begin with the first. Mr. Sheppard once brought me a specimen of a bird-louse (Nirmus) which he took upon a butterfly (Vanessa Io): and should such a capture be more than once repeated, it would afford a certain instance of the first of these diseases amongst insects;—but most probably the specimen in question had dropped from some bird upon the butterfly. The only remaining animal belonging to the apterous hexapods that is parasitic on insects, is by many supposed to be the larva of a giant-beetle (Meloe Proscarabæus). I have before alluded to this animal[1001], and shall now resume the subject. Gœdart, Frisch, and De Geer, observed that it deposited in the earth one or two considerable masses, containing an infinite number of very minute orange-coloured eggs adhering to each other, which in about a month were hatched, and produced a number of small hexapods distinguished by two pairs of anal setæ and a proleg, by means of which they could move readily upon glass, as I have myself seen: these little animals precisely corresponded with one found by the latter author upon[Pg 233] Eristalis intricarius; and when that fly was placed amongst them, they immediately attached themselves to it, so as to leave no doubt of their identity[1002]. A congenerous species had been detected upon wild bees, and described by Linné under the name of Pediculus Apis. De Geer is so thoroughly to be depended upon for his veracity and accuracy of observation, that we cannot suppose there is any incorrectness in his statement. If the mass of eggs be, as he represents it, of the size of a hazel-nut, it must have been the product of a very large insect: in confirmation of this opinion it may be further observed, that the larva of the kindred genus Cantharis agrees with it in having anal setæ, though it appears to differ in having only two conspicuous segments in the trunk[1003]. Those which infest wild bees make their first appearance upon acrid plants, which the Meloe likewise feeds upon; from whence with wonderful agility they leap upon the Andrenæ, &c. that visit these flowers. Strong, however, as all these facts appear, still we cannot help exclaiming with the illustrious Swede last named, Who could ever have imagined that the larva of this great beetle would be found upon the body of flies,—and we may add, or bees? Who could ever imagine that it would feed like a bird-louse and resemble it so closely? that in the insertion of its palpi it should exhibit a character exclusively belonging to that tribe[1004]? Another circumstance seems to indicate that these hexapods at the time that they take their station in bees or flies are perfect insects—they do not vary in size, at least not materially. Where, we may also ask, if they are to become[Pg 234] large beetles, where do they take their principal growth? It cannot be as parasites on the little bees or flies that they are usually found upon; they must soon desert them, and like their kindred blister-beetles, as is most probable, have recourse to vegetable food. What an anomaly in rerum natura! It is much to be wished that some skilful insect-anatomist would carefully dissect the Meloe; or perhaps by digging round the roots of the ranunculuses and other acrid plants the larva of that beetle might be discovered in a later stage of growth, and so this mystery be cleared up. I should observe here, that Scopoli has described three parasites as Pediculi; viz. P. rostratus, coccineus, and Cerambycinus; the first of which Fabricius has adopted under the name of P. Gryllotalpæ, but which are all evidently hexapod Acarina[1005].

Acariasis seems a disease almost as universal amongst insects as Scolechiasis; with this difference however, that Acari most commonly take their station upon them in their perfect state. You have doubtless often observed the common dung-beetles (Geotrupes) covered on the underside of their body with small mites (Gamasus Coleoptratorum) which look as if they were engaged in suction—they are often so numerous that no part is uncovered; they also attack other beetles[1006], and are sometimes found on humble-bees. They are easily disturbed, run with great swiftness, and may often be seen in hotbeds and fermenting dung prowling in search of the stercorarious beetles. But the most remarkable insect of this kind is the Uropoda vegetans: it derives its nutriment from the insects it assails not by its mouth, but[Pg 235] by means of a long anal pedicle by which it is attached to them. De Geer found these in such numbers upon a species of Leptura, that its whole body was almost covered with them; they hung from the legs and antennæ in bunches, and gave the animal a most hideous and disgusting appearance. Under this load of vermin it could scarcely walk or move, and all its efforts to get rid of them were in vain: many were attached to its body and to each other by their anal pedicles, but others had cast them off and were walking about. When put into a glass with earth, they began to abandon their prey, so that in a few days it was quite freed from its plagues. He found that these parasites lived long in alcohol[1007].

If you inquire—How are these mites originally fixed by their pedicles? it seems most probable, that as the Hemerobii, when they lay their eggs, know how to place them upon a kind of footstalk, so the parent Uropoda has the same power; and this pedicle appears to act the part of an umbilical chord, conveying nutriment to the fœtus not from a placenta, but from the body of the insect to which it is attached; till having thus attained a certain maturity of growth and structure, it disengages itself and becomes locomotive. Many eggs of the aquatic Acarina (Hydrachna, &c.) are also furnished with a short pedicle by which they are fixed to Dytisci and other water insects. De Geer found some of this description on the underside of the water-scorpion, so thickly set as to leave no void space: they were oval, of a very bright red, and of different sizes on different individuals; whence it was evident that they grow when thus[Pg 236] fixed: when hatched or released—for perhaps they may be regarded as fœtuses in their amnios rather than eggs—they cease to be parasitical. Let us admire on this occasion, (piously observes this great Entomologist,) the different and infinitely varied means by which the Author of Nature has endowed animals, particularly insects, for their propagation and preservation: for it is a most extraordinary sight to see eggs grow, and pump as it were their nutriment from the body of another living animal[1008]. As these mites are fixed to the crust as well as its inosculations, they must have some means of forcing their nutriment through its pores.

Another insect, remarkable for its resemblance in some respects to the scorpion—called in this country the book-crab (Chelifer cancroides), from its being sometimes found in books—occasionally is parasitic upon flies, especially the common blue-bottle-fly (Musca vomitoria). They adhere to it very pertinaciously under the wings; and if you attempt to disturb them, they run backwards, forwards, or sideways, with equal facility.

Spiders also are infested by mites. Mr. Briggs once found a very small Theridion, to the thorax of which were attached four oblong bright scarlet mites, each of which was as large as the thorax itself. He afterwards met with another spider still smaller, attacked by two of these swoln parasites, one of which appeared to him nearly equal to the spider in size. This mite was probably either Leptus Phalangii, or Astoma parasiticum.

2. We now come to a perfectly distinct tribe of insect parasites, which belong to that section or order of[Pg 237] intestinal worms which Rudolph has denominated Entozoa nematoidea, and Lamarck Vers rigidules[1009]. To this tribe belong the Gordius of Linné and the Filaria of modern zoologists, which from the experiments and observations of De Geer, Dr. Matthey, &c. appear to have been too hastily separated, being really congenerous, and living indifferently in water and in the intestines of insects and other animals[1010]. To this genus belong the guinea-worm (Gordius medinensis[1011]), the Furia infernalis, and several others that are found in various vertebrate animals. These little worms have been discovered in insects of almost every Order; and their attack generally produces the death of the animal, though they appear not to devour those parts that are essential to life[1012]. I once took a specimen of Pœcilus azureus, and upon immersing it in boiling water I was surprised to see what at first I mistook for an intestine, thrust itself forth; but upon a nearer inspection, to my great surprise I found it was one of these worms, thicker than a horse-hair and of a brown colour. Mr. W. S. MacLeay also once found one in Abax Striola. It still remains in my specimen, making it appear as if it had a long tail. De Geer long ago found these worms in grasshoppers[1012]; but Dr. Matthey has given the fullest account of one which infested Acrida viridissima. A friend of his noticing one of these insects which had not strength enough to leap and could scarcely even walk,[Pg 238] being struck with the circumstance, caught the animal, upon which its hind legs were immediately detached from it. His surprise was greatly increased when he saw issue from its body a cylindrical worm about two feet and a half in length. Upon being called, Dr. M. soon recognised it for a Gordius or Filaria; and on his putting it into water, it moved in it with great velocity, twisting its long and slender body in all directions. Upon opening the body of the grasshopper, nothing appeared within it but the intestine shrunk up to a thread. A few days after, another was brought, which appeared in full vigour, but its abdomen was enormously distended, and from it another worm was extracted, which remained without motion rolled in a spiral direction: intending to preserve this in spirits of wine—as it had become flat he first immersed it in water, that it might recover if possible its cylindrical form. Upon immersion a movement took place in the animal, and it gradually recovered its plumpness; but it still remained without motion, as if dead, for nearly five days, when another living specimen being brought and placed with it, as soon as water was poured on them, the seemingly dead one began to show by a slight oscillation in its extremities that life was not extinct in it. Fresh water being poured upon it, at the end of the day it had recovered all its strength and agility. He afterwards often repeated the same experiment with a similar result[1013]. From this account it appears that the Gordius or Filaria has a property resembling that of the Vibrio Tritici, so well described and so beautifully figured by M. Bauer[1014], of apparently dying and being resuscitated by immersion in water.[Pg 239] How long it can retain this property remains to be ascertained.

De Geer states that he had seen them of the length of two feet[1015]; but they vary considerably in this respect. In ants, in which Gould detected them, he states their length to be not more than half an inch[1016]. In caterpillars, which they sometimes infest, they are longer; in that of Notodonta Ziczac, De Geer found one three inches and a half long[1017]; and Rösel three, of six inches, in that of Deilephila Euphorbiæ[1018]; and in Phalangium cornutum, according to Latreille, they extend to more than seven inches[1019]. In the larva of a Phryganea L. the author first named found one which was more than a foot long, corresponding exactly with the Gordius aquaticus of Linné; being forked at one extremity, brown above, gray below, and black at each end[1020]. These animals appear to die as soon as they leave the body[1021] they have preyed upon; except this happens in water, when their activity has no repose. In this element they give their bodies every possible inflexion, often tying themselves in knots in various places, interlacing and twisting themselves in a hundred different ways; so that when confined in the body of an insect, from their extreme suppleness and power of contortion they find sufficient space wherein to pack their often enormous length[1022]. Linné makes one of their habitats clay; and Mr. W. S. MacLeay finds them very common at Putney in clay at the bottom of pools.

[Pg 240]

Dr. Matthey asks—How does the Gordius get into Acrida viridissima[1023]? And De Geer—Why do they die after having quitted a caterpillar? and where do they perpetuate their species[1024]? These questions, without further observations, cannot easily be answered. However, it may be supposed that carnivorous insects, such as Harpali, &c. may swallow them when found apparently dead in clay, where the water has been evaporated, or when they have been ejected by other insects; and they may revive in their bodies, as Dr. Matthey found them to do in water. It is not difficult to conjecture that the larvæ of Phryganeæ may meet with them when young in the water, and sometimes unluckily swallow them with their food. Why they become as dead when they emerge from their prey we cannot at present conjecture; but no doubt to answer some wise purpose;—in rainy seasons they probably revive and get into little hollows full of rain-water. Upon De Geer's last question—How they perpetuate their species—at present I can offer no conjecture.

I am, &c.

[Pg 241]

LETTER XLV.

SENSES OF INSECTS.

At first one would think that the senses of insects might be described in very few words, and scarcely afford matter for a separate letter; but when we find that physiologists are scarcely yet agreed upon this subject, and that the use of some of their organs, which appear to be organs of sensation, has not yet been satisfactorily ascertained—we shall not wonder that it requires more discussion than at the first blush we were aware of. In treating on this head I shall first say something on the senses in general, and then confine myself to those of insects.

Touch, taste, smell, hearing, and sight, I need not tell you, is the usual enumeration of the senses: but as the term includes every means of communication with the external world, the list perhaps might be increased; and there is ground for thinking that the number seven, so signalized as a sacred number[1025], may also here have place. Dr. Virey, an eminent physiologist, whose sentiments on various subjects I have before noticed with approbation[1026], appears to be of opinion that there are[Pg 242] really seven senses; which he divides into those that are altogether physical, and those that are more connected with the intellect. The first of these divisions contains four senses,—touch, love, taste, and smell;—the second three,—hearing, sight, and the internal sense of thought, or the brain[1027]. That he is right in adding love to the list seems to me evident, because it is as distinct from touch, as smelling and taste are. With regard to the other, though it may be expected that there should be a transitive sense connecting the intellect (if I may so speak) with the external organ of sense, and as a medium by which the former can receive the notices of the external world furnished by the latter; yet it seems improper to make the entire brain itself a sense. We know that the agent between the common sensory and the sense is the consciousness or perception of the impression. "Seeing we may see and not perceive, and hearing we may hear and not understand." The picture may be painted upon the retina of the eye, the sound may strike upon the tympanum of the ear; but neither the one nor the other be received by the intellect, unless the internal power or faculty of perception be in action and mediate between them. This is what I mean by the internal sense, which, to use a term of Mr. W. S. MacLeay's[1028], is osculant between intellect and sense, or forms the transit from one group of powers to the other.

Of the ordinary senses, sight holds the first rank: it[Pg 243] can dart to the region of the stars, and convey by the perceiving sense, to the sensory, ideas of innumerable objects. Next in rank is hearing, which can receive sounds from a great distance; but the ideas it remits are confined only to one object, the variations of tones. In the other organs the sensitive power is much more confined. There is another difference between the intellectual and physical senses:—the former are the only ones that receive and convey sensations of the beautiful and sublime; of harmony and discord,—the latter, though they minister more to our sensual enjoyments, add little to our intellectual; and therefore too devoted an indulgence in them debases our nature, and levels us with the brutes, which use their eyes and ears only for information, not for pleasure[1029].

In man the ordinary five senses are usually in their greatest perfection, although in some animals particular senses have a greater range. The Vertebrates in general are also gifted with the same number, though there are some exceptions. But in the Invertebrates they are seldom to be met with all together in the same object. The Cephalopods have no smell. Several Gasteropods can neither hear nor see. The animals of bivalve shells have neither eyes, nor ears, nor smell; and the zoophytes and the races below them have, it is affirmed, only the single sense of touch, which in them is so extremely delicate as to be acted upon even by light[1030].

Not so our insects. These, there is good reason to[Pg 244] believe, possess all the ordinary senses. That they can see, touch, taste, and smell, no one denies. Linné and Bonnet, however, thought them deprived of hearing[1031]; but numerous observations prove the contrary. That they hear in their larva state, is evident from facts stated by the latter physiologist. He found that the sound of his voice evidently affected some caterpillars; which he attributes, but surely without reason, to the delicacy of their sense of touch: at another time, when some caterpillars of a different species were moving swiftly, he rang a small bell; upon which they instantly stopped and moved the anterior part of their body very briskly[1032]. That they possess this faculty in their imago state is confirmed still more strongly by facts. I once was observing the motions of an Apion under a pocket microscope: on seeing me it receded. Upon my making a slight but distinct noise, its antennæ started: I repeated the noise several times, and invariably with the same effect. A Harpalus, which I was holding in my hand, answered the sound in the same manner repeatedly. Flies, I have observed, at brisk and distinct sounds move all their legs; and spiders will quit their prey and retire to their hiding places. Insects that live in society give notice of intended movements, or assemble their citizens for emigration by a certain hum[1033]. But the most satisfactory proof of the hearing of these animals is to be had from those Orthoptera and Hemiptera whose males are vocal. Brunelli kept and fed several males of Acrida viridissima (a grasshopper with us not uncommon) in a closet, which[Pg 245] were very merry, and continued singing all the day; but a rap at the door would stop them instantly. By practice he learned to imitate their chirping: when he did this at the door, at first a few would answer him in a low note, and then the whole party would take up the tune and sing with all their might. He once shut up a male in his garden, and gave the female her liberty; but as soon as she heard the male chirp, she flew to him immediately[1034].

But although physiologists are for the most part agreed that insects have the ordinary five senses of vertebrate animals, yet a great variety of opinions has obtained as to their external organs; so that it has been matter of doubt, for instance, whether the antennæ are for smell, touch, or hearing; and the palpi for smell, taste, or touch. Nor has the question, as it appears to me, been satisfactorily decided: for though it is now the most general opinion that the primary use of antennæ is to explore as tactors, yet by the most strenuous advocates of this opinion they are owned not to be universally so employed; so that granting this to be one of their principal functions, yet it seems to follow that there may be another common to them all, which of course would be their primary function. We are warned, however, not to lay any stress upon the argument to be drawn from analogy; and told that we might as well dispute about the identity of the nose of a man, the proboscis of the elephant, the horn of the rhinoceros, the crest of the cock, or the beak of the toucan[1035]. But this is merely[Pg 246] casting dust in our eyes: for though three of these are nasal organs, bearing nostrils; the two others have no relation to the question, the horn of the rhinoceros and the crest of the cock being merely appendages, and have no more analogy to the nose and nostrils, which co-exist with them, than they have to the eyes or ears. I have on a former occasion observed, that a gradual change sometimes takes place in the functions of particular organs; but still, generally speaking, this observation regards secondary functions—the primary usually remaining untouched. We may say, for instance, with regard to the primary use of the legs of animals, that it is locomotion; while the secondary is either walking, running, jumping, flying, or swimming, according to the circumstances and nature of the animal. Thus the fore-legs of the Mammalia, in birds become wings, and both pair in fish are changed to fins. Observe, I do not say always and invariably, but in most cases,—that analogous parts have analogous uses, at least as far as primary uses are concerned. When, therefore, we cannot have demonstrative evidence concerning the function of an organ discoverable in any animal, we may often derive satisfactory probable arguments from the analogies observable in their structure compared with that of other animals, concerning the nature of whose organs we have no doubt. In fact, the chief evidence we have with regard to the office of the organs of sense in the animals immediately below ourselves, is that of analogy;—because we see with our eyes, hear with our ears, &c., we conclude, with reason, that they do the same.

In inquiring therefore into what may be the most general[Pg 247] use of the antennæ of insects, I shall endeavour to discover whether there is any part in the higher animals to which they may be deemed to exhibit any analogy. And here I must refer you to what I have said on a former occasion upon the present subject; where I made it evident, I hope, that the great bulk of the parts and organs of insects, in this particular differing from the majority of Invertebrates, are, some in one respect, some in another, and some in many, really analogous to those of the higher animals[1036]; and that a great many of them, though varying in their structure, have the same functions. Thus the analogues of the eyes of Vertebrates are for seeing; of the jaws for masticating; of the lips for closing the mouth; of the legs for walking, &c. We have seen also very recently, that a similar analogy, more or less strongly marked, holds also in their internal organs[1037]; so that it may be safely affirmed, that if all the invertebrate insects, though gifted with numerous peculiarities, present the most striking picture of those animals that have an internal skeleton, and more particularly of the Mammalia,—we may assume it as a probability, the above circumstances being allowed their due weight, that where facts do not prove the contrary, the function of analogous organs is more or less synonymous, though perhaps the structure and modus operandi may be different.

In the letter lately referred to, I observed that the antennæ of insects are analogous to ears in Vertebrates[1038]. Their number corresponds; they also stand out from the[Pg 248] head; and what has weighed most with me, unless they are allowed as such, no other organ can have any pretension to be considered as representing the ear. If we reflect, that in every other part and organ, the head of insects has an analogy to that of Mammalia, we must regard it as improbable that these prominent organs should not also have their representative. Admitting then that they are the analogues of ears, it will follow, not as demonstratively certain, but as probable, that their primary function may be something related to hearing. I do not say direct hearing, or that the vibrations of sound are communicated to the sensorium by a complex structure analogous to that of the internal ear in Mammalia—but something related to hearing. I conceive that antennæ, by a peculiar structure, may collect notices from the atmosphere, receive pulses or vibrations, and communicate them to the sensorium, which, though not precisely to be called hearing, may answer the same purpose. From the compound eyes that most of them have, the sense of seeing in insects must be very different from what it is in vertebrate animals; and yet we do not hesitate to call it sight: but since antennæ, as we shall see, apparently convey a mixed sensation, I shall have no objection, admitting it as their primary function, to call it after Lehmann Aëroscepsy[1039]. I lately related some instances of sound producing an effect on the antennæ of insects: I will now mention another that I observed, still more remarkable. A little moth was reposing upon my window; I made a quiet, not loud, but distinct noise: the antenna nearest to me immediately[Pg 249] moved towards me. I repeated the noise at least a dozen times, and it was followed every time by the same motion of that organ; till at length the insect being alarmed became more agitated and violent in its motions. In this instance it could not be touch; since the antenna was not applied to a surface, but directed towards the quarter from which the sound came, as if to listen. Bonsdorf made similar observations, to which Lehmann seems not disposed to allow their proper weight[1040]. It has been used as an argument to prove that antennæ are primarily tactors, or instruments of touch, that Fœnus Jaculator, before it inserts its ovipositor, plunges its antennæ into the hole forming the nidus of the bee, to the grub of which it commits its egg[1041]. But had those who used this argument measured the antennæ and the ovipositor of this ichneumon, they would have discovered that the latter is thrice the length of the former: and as these insects generally insert it so that even part of the abdomen enters the hole, it is clear that the antenna cannot touch the larva; its object therefore cannot be to explore by that sense. Others suppose that by these organs it scents out the destined nidus for its eggs; but Lehmann has satisfactorily proved that they are not olfactory organs. We can therefore only suppose, either that by means of its antennæ it hears a slight noise produced by the latent grub, perhaps by the action of its mandibles; or else that by its motions it generates a motion in the atmosphere of its habitation, which striking upon the antennæ of the Fœnus, are by them communicated[Pg 250] to its sensory. A similar disproportion is observable between the antennæ and ovipositor of Pimpla Manifestator, before signalized[1042]. Bees, when collecting honey and pollen, first insert the organs in question into the flowers which they visit; but, as I have more than once observed, they merely insert the tip of them. If anthers are bursting, or the nectar is exuding, these processes probably are attended by a slight noise, or motion of the air within the blossom, which, as in the last case, affects, without immediate contact, the exploring organs.

If the structure of antennæ be taken into consideration, it will furnish us with additional reasons in favour of the above hypothesis, with regard to their primary function. We shall find that these organs, in most of those insects which take their food by suction, are usually less gifted with powers of motion, than they are in the mandibulate tribes; so that in the majority of the Homopterous Hemiptera and Diptera, as is generally acknowledged, they cannot be used for touch. Under this view, they may be divided into active antennæ and passive antennæ: of the former, the most active and versatile are those of the Hymenoptera. By means of them, as was before observed[1043], their gregarious tribes hold converse, and make inquiry—frequently without contact—in the pursuit and discharge, if I may so speak, of the various duties devolved upon them by Providence. Amongst active antennæ, some are much more complex in their structure than others—a circumstance which is often characteristic[Pg 251] of the male insect[1044]: but if we examine such antennæ, we shall find that their most sensitive parts cannot come in contact with the earth or other bodies for exploring their way; but having thus a greater surface exposed to the action of the atmosphere, they have more points to receive vibrations, or any pulses or other notices communicated to it. It is thus, probably, that in their flights, when they approach within a certain distance, they discover the station of the other sex. Even the plumose antennæ of male gnats may in some respects thus be acted upon. In the Lamellicorn beetles, the knob of these organs in both sexes consists of laminæ, the external ones on their outside, of a corneous substance; while their internal surface, and the inner laminæ—which are included between them, as an oyster between the valves of its shell—are covered with nervous papillæ. If you examine the proceedings of one of these little animals, you will find before it moves from a state of repose that its antennæ emerge, and the laminæ diverge from each other; but that it does not apply them to surfaces to explore its way, but merely keeps them open to receive notices from the atmosphere. Even simple antennæ are often employed in this way, as well as for touch. I once noticed a species of Leptocerus, a trichopterous genus, in which these organs are very long, that was perched upon a blade of grass; its antennæ vibrated, and it kept moving them from side to side in the air, as if thus by aëroscepsy it was inquiring what was passing around it. Dr. Wollaston has an observation bearing so[Pg 252] precisely upon this question, and in general so extremely similar to what is here advanced, that I must copy it for your consideration. "Since there is nothing in the constitution of the atmosphere," says he, "to prevent vibrations much more frequent than any of which we are conscious, we may imagine that animals like the Grylli, whose powers appear to commence nearly where ours terminate, may have the faculty of hearing still sharper sounds, which at present we do not know to exist; and that there may be other insects, hearing nothing in common with us, but endued with a power of exciting, and a sense that perceives, vibrations indeed of the same nature as those which constitute our ordinary sounds, but so remote, that the animals who perceive them may be said to possess another sense, agreeing with our own solely in the medium by which it is excited, and possibly wholly unaffected by these slower vibrations of which we are sensible[1045]." That insects, however, hear nothing in common with us, is contrary to fact; at least with respect to numbers of them. They hear our sounds, and we theirs; but their hearing or analogous sense is much nicer than ours, collecting the slightest vibratiuncle imparted by other insects, &c. to the air. In inquiring how this is done, it may be asked—How know we that every joint of some antennæ is not an acoustic organ, in a certain sense distinct from the rest? We see that the eyes of insects are usually compound, and consist of numerous distinct lenses;—why may not their external ears or their analogues be also multiplied, so as to[Pg 253] enable them with more certainty to collect those fine vibrations that we know reach their sensory, though they produce no effect upon our grosser organs? I propose this merely as conjecture, that you may think it over, and reject or adopt it, in proportion as it appears to you reasonable or the contrary; and in the hope that some anatomist of insects, who, to the sagacity and depth of a Cuvier and a Savigny adds the hand and eye of a Lyonet, may give to the world the results of a more minute dissection and fuller investigation of the antennæ of these animals, than has yet been undertaken.

But besides receiving notices from the atmosphere, of sounds, and of the approach or proximity of other insects, &c., the antennæ are probably the organs by which insects can discover alterations in its state, and foretel by certain prognostics when a change of weather is approaching. Bees possess this faculty to an admirable degree. When engaged in their daily labours, if a shower is approaching, though we can discern no signs of it, they foresee it, and return suddenly to their hives. If they wander far from home, and do not return till late in the evening, it is a prognostic to be depended upon, that the following day will be fine: but if they remain near their habitations, and are seen frequently going and returning, although no other indication of wet should be discoverable, clouds will soon arise and rain come on. Ants also are observed to be excellently gifted in this respect: though they daily bring out their larvæ to sun them, they are never overtaken by sudden showers[1046]. Previously to rain, as you well know,[Pg 254] numberless insects seek the house; then the Stomoxys calcitrans, leaving more ignoble prey, attacks us in our apartments, and interrupts our studies and meditations[1047]. The insects of prey also foresee the approach of wet weather, and the access of flies, &c. to places of shelter. Then the spiders issue from their lurking-places, and the ground-beetles in the evening run about our houses. Passive antennæ, which are usually furnished with a terminal or lateral bristle, and plumose and pectinated ones, seem calculated for the action of the electric and other fluids dispersed in the atmosphere, which in certain states and proportions may certainly indicate the approach of a tempest, or of showers, or a rainy season, and may so affect these organs as to enable the insect to make a sure prognostic of any approaching change: and we know of no other organ that is so likely to have this power. I say electric fluid, because when the atmosphere is in a highly electrified state, and a tempest is approaching, is the time when insects are usually most abundant in the air, especially towards the evening; and many species may then be taken, which are not at other times to be met with: but before the storm comes on, all disappear, and you will scarcely see a single individual upon the wing. This seems to indicate that insects are particularly excited by electricity[1048].—But upon this head I wish to make no positive assertion, I only suggest the probability of the opinion[1049].

From all that has been said, I think you will be disposed[Pg 255] to admit that the primary and most universal function of the antennæ is to be the organs of a sense, if not the same, at least analogous to hearing, and answering the same end; something perhaps between it and touch. In some, however, as has been found in the Crustacea, an organ of hearing, in the ordinary sense, may exist at the base of the antennæ, which may act the part in some measure of the external ear, and collect and transmit the sound to such organ[1050].

That numerous antennæ, as a secondary function, explore by touch, is admitted on all hands, and therefore I need not enlarge further upon this point; but shall proceed to inquire whether insects do not possess some other peculiar organs that are particularly appropriated to this sense. First, however, I must make some general observations upon it. Of all our senses, touch is the only one that is not confined to particular organs, but dispersed over the whole body: insects, however, from the indurated crust with which they are often covered, feel sensibly, it is probable, only in those parts where the nerves are exposed, by being covered with a thinner epidermis, to external action. Not that they cannot feel at all in their covered parts; for as we feel sufficiently for walking, though our feet are covered by the thick sole[Pg 256] of a boot or shoe, so insects feel sufficiently through the crust of their legs for all purposes of motion. Besides, the points that are covered by a thinner cuticle are often numerous; so that touch, at least in a passive sense, may be pretty generally dispersed over their bodies; but active or exploring touch is confined to a few organs, as the antennæ, the palpi, and the arms. The two last I shall now discuss.

Various opinions have been started concerning the use of the palpi. Bonsdorf thought that they were organs of smell; Knoch, that this sense was confined to the maxillary ones, and that the labial ones were appropriated to taste[1051]: but the most early idea, and that from which they derive their present name of palpi (feelers), is, that they are organs of active touch; and this seems to me the most correct and likely opinion. Cuvier, himself a host, has embraced this side of the question[1052], and Lehmann also admits it[1053]. The following observations tend to confirm this opinion. The palpi of numerous insects when they walk, are frequently, or rather without intermission, applied to the surface on which they are moving—this you may easily see by placing one upon your hand; which seems to indicate that they are feelers. In the Araneidæ they are used as legs; and by the males at least, as exciting if they be not really genital organs[1054]. In the Scorpionidæ they answer the purpose of hands: besides being usually much shorter than antennæ, they[Pg 257] are better calculated to assist an insect in threading the dark and tortuous labyrinths through which it has often to grope its way, and where antennæ cannot be employed. I have noticed that Hydrophili—in which genus the palpi are longer than the antennæ—when they swim, have their antennæ folded; while the former are stretched out in front, as exploring before them. As these are attached to the under-jaws and under-lip, we may suppose they are particularly useful to insects in taking their food; and upon this occasion I have often observed that they are remarkably active. I have seen Byturus tomentosus, a beetle which feeds upon pollen, employ them in opening anthers; and the maxillary pair appear to me to assist the maxillæ in holding the food, while the mandibles are at work upon it.

The arms or fore-legs of some insects are also organs of active touch, being used, as we have seen, for cleaning the head, digging, repairing their dwellings, and the like[1055]. By the Ephemeræ, which have very short antennæ, the fore-legs, when they fly, are extended before the head, parallel with each other and quite united—probably to assist in cutting the air. The Trichoptera use their antennæ for the same purpose.

Another sense of which the organ seems uncertain is that of smelling, and various and conflicting opinions have been circulated concerning it. Christian thought that insects smell distant objects with their antennæ, and near ones with their palpi[1056]. Comparetti has a most singular[Pg 258] opinion. He supposes in different tribes of insects that different parts are organs of smell: in the Lamellicorns he conjectures the seat of this sense to reside in the knob of the antennæ; in the Lepidoptera in the antlia; and in some Diptera and Orthoptera in certain frontal cells[1057]. At first sight, one of the most reasonable opinions seems to be that of Baster, adopted by Lehmann, and which has received the sanction of Cuvier[1058],—that the spiracles are organs of smell as well as of respiration. Lehmann has adduced several arguments in support of this opinion. Because we both respire and smell with our nostrils, he concludes that neither the antennæ nor any other part of the head of insects can serve for smell, since they are not the seat also of respiration; and that there can be no smell where the air is not inspired[1059]. Again, because nerves from the ganglions of the spinal chord terminate in bronchiæ near the spiracles, they must be for receiving scents from those openings. Though it was necessary, in the higher animals, that the organ of scent should be near the mouth, because they are larger than their food; yet the reverse of this being the case with insects, which often even reside in what they eat, it is to them of no importance where their sense of smelling resides[1060]. By exposing antennæ, by means of an orifice in a glass vessel, to the action of stimulant odours, they appeared quite insensible to it: but he does not name the result of any experiment in which he exposed the mouth to this action; nor at all distinctly how the insect was affected when the spiracles were exposed to it[1061].

[Pg 259]

But though some of these arguments appear weighty, there are others, I think, that will more than counterbalance them, making it probable that the seat of this sense is in the head, either in its ordinary station at the extremity of what I call the nose, between it and the upper-lip, or under those parts. That the nose corresponds with the so-named part in Mammalia, both from its situation and often from its form, must be evident to every one who looks at an insect[1062]; and when we further consider the connexion that obtains between the senses of smell and taste, how necessary it is that the seat of the one should be near that of the other, and that it really is so in all animals in which we certainly know its organ[1063]; we shall feel convinced that the argument from analogy is wholly in favour of the nose, and may thence consider it as probable that the sense in question does reside there. Lehmann seems to be of opinion, because an insect is usually smaller than what it feeds upon, that it makes no difference whether it smells with its head or with its tail: but one would think that a flying insect would be more readily directed to its object by smelling with the anterior part of the body than with the posterior; and that a feeding one would also find it more convenient in selecting its food. As to the argument,—that smell must be the necessary concomitant of the respiratory openings, and that there can be no smell where the air is not inspired,—this seems asserting more than our knowledge of these animals will warrant: for the organs of the other senses, though the senses themselves seem analogous, are so different in their structure, and often in the mode in[Pg 260] which they receive the impressions from external objects, that analogy would lead us to expect a difference of this kind also in the sense of smell. Besides, smell does not invariably accompany respiratory organs even in the higher animals,—for we breathe with our mouths, but do not smell with them. Cuvier says that the internal membrane of the tracheæ being soft and moist, appears calculated to receive scents[1064]. But here his memory failed him; for it is the external membrane alone that answers this description; the internal consisting of a spiral elastic thread, and seeming not at all fitted to receive impressions, but merely to convey the air[1065]. That nerves penetrate to the bronchiæ, does not necessarily imply that they are connected with the sense in question, since this may be to act upon the muscles which are every where distributed.

I shall now state some facts that seem to prove that scents are received by some organ in the vicinity of the mouth, and probably connected with the nose. M. P. Huber, desirous of ascertaining the seat of smell in bees, tried the following experiments with that view. These animals, of all ill scents, abominate most that of the oil of turpentine. He presented successively to all the points of a bee's body, a hair-pencil saturated with it: but whether he presented it to the abdomen, the trunk, or the head, the animal equally disregarded it. Next, using a very fine hair-pencil, while the bee had extended its proboscis, he presented the pencil to it, to the eyes and antennæ, without producing any effect; but when he pointed it near the cavity of the mouth, above the insertion[Pg 261] of the proboscis, the creature started back in an instant, quitted its food, clapped its wings, and walked about in great agitation, and would have taken flight if the pencil had not been removed. On this, it began to eat again; but on the experiment being repeated, showed similar signs of discomposure: oil of marjoram produced the same effect, but more promptly and certainly. Bees not engaged in feeding appeared more sensible of the impression of this odour, and at a greater distance; but those engaged in absorbing honey might be touched in every other part without being disturbed. He seized several of them, forced them to unfold their proboscis, and then stopped their mouth with paste. When this was become sufficiently dry to prevent their getting rid of it, he restored to them their liberty: they appeared not incommoded by being thus gagged, but moved and respired as readily as their companions. He then tempted them with honey, and presented to them near the mouth, oil of turpentine, and other odours that they usually have an aversion to; but all produced no sensible effect upon them, and they even walked upon the pencils saturated with them[1066].

These experiments incontestibly prove that the organ of scent in bees—and there is no reason to think that other insects do not follow the same law—is in or near the mouth, and above the proboscis. It remains, therefore, that we endeavour to discover its precise situation: and as insects cannot tell us, nor can we perceive by their actions, in what precise part the sense in question[Pg 262] resides, the only modes to which we can have recourse to form any probable conjecture, are analogy and dissection. At first, the opinion noticed above, that the palpi are its organs, seems not altogether unreasonable; but as the argument from analogy, except as to their situation near the mouth, is not in favour of them, and there seems no call, were smell their function, for the numerous variations observable in their structure, I think we must consider them, as I have endeavoured to prove, rather as instruments of touch. Let us now inquire, whether there be not discoverable upon dissection, in the interior of the head of any insects, some organ that may be deemed, from its situation, under what we have called the nose and nostrils, the seat of the sense we are treating of. The common burying-beetle (Necrophorus Vespillo) is an insect remarkable for its acuteness of smell, which enables it to scent out and bury, as was formerly related to you[1067], the carcases of small animals. Take one of these insects, and kill it as formerly directed,—examine first its nose: in the middle of the anterior part you will see a subtrapezoidal space, as it were cut out and filled with a paler piece of a softer and more membranous texture. Next divide the head horizontally; and under the nose, and partly under this space, which I call the rhinarium or nostril-piece[1068], you will find a pair of circular pulpy cushions, covered by a membrane transversely striated with beautifully fine striæ. These are what I take to be the organs of smell, and they still remain distinctly visible in a specimen I have had by me more than fifteen years. A similar organ may be discovered[Pg 263] in the common water-beetle (Dytiscus marginalis), but with this peculiarity, that it is furnished with a pair of nipples. I have before described an analogous part covered with papillæ, in Æshna viatica, and you will find it in other insects[1069]. Perhaps at first this part may seem merely a continuation of the palate; but if you consider the peculiarities in its structure just noticed, it is evidently a sensiferous organ; and as the sense of smell appears to reside in the head, this is its most probable seat. But by what channel scents act upon it,—whether they are transmitted through the pores of the part representing the nostrils, or received by the mouth,—I will not venture to assert positively: but from the circumstance of their being membranous in some insects remarkable for acute scent, as in Necrophorus, Staphylinus, &c., there seems some ground for the former opinion, which receives further confirmation from an observation of an eminent Comparative Anatomist, M. Carus, with respect to Acrida verrucivora, in which under the nose and rhinarium, as appears from his description, he found some tracheæ, and two lobes of the cerebral ganglion, which caused him to regard this as the seat of the sense of smell[1070]. He also tells us that Rosenthal, in the blue-bottle-fly (Musca vomitoria) places the sense of smell partly in a delicately folded membrane observable in its head[1071]. As the sense of smell in these little beings is extremely acute, as well as their hearing, the perception of odours may reach their sensory through the above pores;[Pg 264] and even those in the hard rhinarium of an Anoplognathus may receive and transmit them; and besides the upper-lip and nose are often united by membrane, perhaps representing the rhinarium, as in Goerius, &c.[1072] which may facilitate such transmission.

That insects taste, no one hesitates to believe, though some have supposed the palpi to be the organ of that sense; but as they have a tongue, as we have shown, we may with Cuvier conclude, that one of its primary functions is to taste their food[1073]. I shall not therefore launch out further upon this head.

I have now placed before you a picture, or rather sketch, of the insect world. And whether we regard their general history and economy, their singular metamorphoses, the infinite varieties and multiplicity of their structure both external and internal, and their diversified organs both of sense and motion—I think you will be disposed to own, that in no part of his works is the hand of an Almighty and All-wise Creator more visibly displayed, than in these minutiæ of creation; that they are equally worthy of the attention and study of the Christian Philosopher with any of the higher departments of the animal kingdom; and that all praise is due to Him, for placing before our eyes, for our entertainment and instruction, such a beautiful moving picture of little symbols and agents, perpetually reflecting his glory and working his will.

I am, &c.

[Pg 265]

LETTER XLVI.

ORISMOLOGY, OR EXPLANATION OF TERMS.

It was by the language of terms that he invented and employed, as well as by his system and methods of arrangement, that Linné smoothed the way to the study of Natural History;—having therefore led you through a large portion of the flowery fields of the Science of Entomology, I must now conduct you into that arid but not barren or unprofitable region. To enable you to understand descriptions of insects, or to describe them yourself, you must have a knowledge of the technical language by which their parts and characters are expressed. Much of this you already know from the definitions of external parts, furnished in a former letter[1074]: I shall now give you a more full and general explanation of terms, adding many new ones for unnoticed characters, that may be conveniently employed.

The science of terms, which I shall call Orismology[1075], may be divided into two branches—General Orismology, and Partial Orismology; the first containing general definitions, and the last those relating to particular parts and organs.

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A. GENERAL ORISMOLOGY.

I. SUBSTANCE.

1. Membranous (Membranacea). A fine, thin, transparent substance. A Membrane.—Ex. Wings of Hymenoptera and Diptera.

2. Pergameneous (Pergamenea). A thin, tough, and less transparent substance, somewhat resembling parchment.—Ex. The Tegmina of the Orthoptera[1076].

3. Coriaceous (Coriacea). A thicker, flexible substance, resembling leather.—Ex. Elytra of Telephorus and the Malacodermi.

4. Corneous (Cornea). A hard inflexible substance resembling horn.—Ex. Elytra of Lucanus Cervus and many other Coleoptera.

5. Crustaceous (Crustacea). A rigid calcareous substance.—Ex. The Shell of a Lobster or Crab.

6. Callous (Callosa). A substance without pores, harder than the surrounding matter, and usually elevated above it.—Ex. Elevated parts of the Collar in Nomada. (Mon. Ap. Angl. Apis * b.)[1077] Spots on the elytra of Stenocorus (Tylostagmus K. MS.) bimaculatus and affinities.

7. Cartilagineous (Cartilaginea). A gristly substance between bone and ligament.—Ex. The Tongue of many Hymenoptera.

8. Subereous (Suberea). A soft elastic substance[Pg 267] somewhat resembling cork[1078]. The galls of some species of Cynips when mature approach to this substance.

9. Spongiose (Spongiosa). A soft elastic substance resembling sponge.—Ex. The Pulvilli of Thanasimus, Buprestis, &c.

10. Ligneous (Lignosa). A hard unelastic substance like wood.—Ex. Galls of some species of Cynips.

11. Carnose (Carnosa). A soft, fleshy substance.—Ex. Caterpillars and Grubs.

12. Tubulose (Tubulosa). When the interior is hollow or empty.

13. Solid (Solida). When the interior is full.

II. RESISTANCE.

1. Rigid (Rigida). Hard, which does not bend or yield to pressure.—Ex. The weevils (Rhyncophora).

2. Flexile (Flexilis). Which easily bends, or yields to pressure without breaking.—Ex. Elytra of Telephorus.

3. Soft (Mollis). Flexile and retaining the marks of pressure.—Ex. Elytra of Meloe.

III. DENSITY.

1. Foliaceous (Foliacea). Very thin and depressed, scarcely thicker than a leaf.—Ex. Aradus corticalis and Coreus paradoxus.

2. Depressed (Depressa). When the vertical section is shorter than the transverse.—Ex. Trogosita mauritanica.

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3. Compressed (Compressa). When the transverse section is shorter than the vertical.—Ex. Centrotus cornutus: Abdomen in Cynips.

4. Plump (Pinguis). Naturally and proportionably plump.—Ex. Thylacites, &c. Most of the Cicadæ.

5. Obese (Obesa). Unnaturally enlarged and distended, as if from disease or too much food. Ex. Chrysomela Polygoni ♀, Galeruca Tanaceti ♀, Brachycerus.

6. Ventricose (Ventricosa). Bellying out as if filled with air.—Ex. Pneumora.

IV. PROPORTION.

1. Thick (Crassa). Disproportionably thick throughout.—Ex. Copris.

2. Incrassate (Incrassata). Disproportionably thick in part.—Ex. Base of the Abdomen of Æshna and many Libellulina. Plate IX. Fig. 9.

3. Slender (Tenuis). Disproportionably slender throughout.—Ex. Lixus paraplecticus.

4. Attenuate (Attenuata). Disproportionably slender in part.—Ex. Tail of Scorpio, Raphidia ♂, &c.

5. Broad (Lata). Disproportionably broad throughout.

6. Dilatate (Dilatata). Disproportionably broad in part.—Ex. Elytra of Lycus fasciatus, &c. Plate XIII. Fig. 20.

7. Narrow (Angusta). Disproportionably narrow throughout.—Ex. Abdomen of Agrion.

[Pg 269]

8. Angustate (Angustata). Disproportionably narrow in part.—Ex. Elytra of Sitaris humeralis. Plate XIII. Fig. 19.

9. Long (Longa). Disproportionably long throughout.—Ex. Scolopendra.

10. Elongate (Elongata). Disproportionably long in part.—Ex. Abdomen of Libellulina.

11. Short (Brevis). Disproportionably short throughout.—Ex. Copris.

12. Abbreviate (Abbreviata). Disproportionably short in part.—Ex. Elytra of Staphylinidæ, Atractocerus, &c.

V. FIGURE[1079].

1. Circular (Circularis). Having the diameter every way equal. Plate XXIX. Fig. 16, 17.

2. Rotundate (Rotundata). Rounded at the angles or sides. Plate XXIX. Fig. 19.

3. Oval (Ovalis). Having the longitudinal diameter twice the length of the transverse, and the ends circumscribed by equal segments of a circle. Plate XX. Fig. 6.

4. Elliptic (Elliptica). Oval, but having the longitudinal diameter more than twice the length of the transverse. Plate XX. Fig. 19.

5. Oblong (Oblonga). Having the longitudinal diameter more than twice the length of the transverse, and the ends varying, or rounded. Plate XX. Fig. 3, 9.

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6. Ovate (Ovata). Oval, but having the ends circumscribed by unequal segments of circles. Plate XX. Fig. 12, 13.

7. Cordate (Cordata). Heart-shaped. Ovate or subovate and hollowed out at the base, without posterior angles. Plate IX. Fig. 22.

8. Sagittate (Sagittata). Arrow-shaped. Triangular, hollowed out at the base with posterior angles. Plate XXVII. Fig. 41. w´´´.

9. Hastate (Hastata). Halberd-shaped. Triangular, hollowed out at the base and sides with the posterior angles spreading.—Ex. Horn of the prothorax of Dynastes hastatus. Postfurca in many Coleoptera. Plate XXII. Fig. 5. b †.

10. Triangular; Quadrangular; Quinquangular; Sexangular (Triangula; Quadrangula; Quinquangula; Sexangula). Having three, four, five, or six angles.

11. Turbinate (Turbinata). Top-shaped, triangular with curved sides. Plate XXV. Fig. 18.

12. Ensate (Ensata). Gradually tapering till it ends in a point.—Ex. Ovipositor of Acrida viridissima. Plate XV. Fig. 19.

13. Lanceolate (Lanceolata). Oblong and gradually tapering towards each extremity.—Ex. The Cerci in Blatta. Plate XV. Fig. 23. Q´´.

14. Sigmoidal (Sigmoidea). S-shaped. Lanceolate and concave on one side at the base, and on the other at the apex.—Ex. Ovipositor of Cimbex, Plate XV. Fig. 21. H´´.

15. Cuneate (Cuneata). Wedge-shaped. Having the longitudinal diameter exceeding the transverse,[Pg 271] and narrowing gradually downwards. Plate X. Fig. 11.

16. Acinacicate (Acinacicata). Falchion-shaped. Curved with the apex truncate, and growing gradually wider towards the end.—Ex. Abdomen of Ophion, Fœnus, and other Ichneumonidæ[1080].

17. Lunulate (Lunulata). Crescent-shaped. Curved with both ends acute, like the moon in her first quarter.—Ex. Last joint of the labial palpi of Oxyporus. Plate XIII. Fig. 4. a.

18. Falcate (Falcata). Sickle-shaped. Curved with the apex acute.—Ex. Ovipositor of Acrida varia. Antennæ of Atractocerus. Plate XI. Fig. 8.

19. Linear (Linearis). Narrow and of the same width throughout.—Ex. Wings of Pterophorus monodactylus.

20. Arcuate (Arcuata). Linear and bent like a bow.—Ex. Rostrum of Balaninus Nucum. Plate XIII. Fig. 12.

21. Cultrate (Cultrata). Coulter-shaped. Straight on one side and curved on the other.—Ex. Ovipositor of some Saw-flies. Under-wing of many Ichneumonidæ.

22. Spatulate (Spatulata). Spatula-shaped. Broader and rounded at the apex, linear and narrow at the base.—Ex. Abdomen of Ichneumon amictorius Panz.

23. Clepsydrate (Clepsydrata). Hour-glass-shaped.[Pg 272] Broader at the base and apex—Ex. The Prosternum of many Capricorn beetles.

24. Clavate (Clavata). Club-shaped. Linear at the base, but towards the apex growing gradually broader. Plate XI. Fig. 4.

25. Quadrate (Quadrata). Square. Quadrilateral with the sides equal and the angles right angles.

26. Rhomboid (Rhomboidea). Quadrilateral with the sides equal, but with two opposite angles acute, and two obtuse. Plate XXVII. Fig. 62. t´´.

27. Trapezate (Trapezata). Quadrilateral with the four sides unequal, and none of them perfectly parallel. Plate XIV. Fig. 4.

28. Trapezoid (Trapezoidea). Quadrilateral, with two sides unequal and parallel[1081]. Plate XXVI. Fig. 34. b´.

29. Parallelogramical (Parallelogramica). Quadrilateral, with all the angles right angles, and all the sides parallel, but two longer than the others.

VI. FORM[1082].

1. Spherical (Sphærica). The shape of a globe. A body whose diameter every way is equal. Plate XX. Fig. 5.

2. Orbiculate (Orbiculata). A depressed globe, whose horizontal section is circular, and vertical oval. Plate XX. Fig. 10, 11.

3. Lenticular (Lenticularis). Lens-shaped. Whose[Pg 273] horizontal section is circular, and vertical lanceolate.—Ex. Abdomen of Cynips aptera.

4. Ovaliform (Ovaliformis). Whose longitudinal section is oval, and transverse circular. Plate XX. Fig. 6.

5. Ellipsoid (Ellipsoidea). Whose longitudinal section is elliptical, and transverse circular. Plate XX. Fig. 19.

6. Oviform (Oviformis). Whose longitudinal section is ovate, and transverse circular. Plate XX. Fig. 12, 13.

7. Cucumiform (Cucumiformis). Cucumber-shaped. Whose longitudinal section is oblong, and transverse circular. Plate XX. Fig. 18, excluding the neck.

8. Cordiform (Cordiformis). Oviform and hollowed out at the base without posterior angles. Plate IX. Fig. 22.

9. Conical (Conica). Whose vertical section is triangular, and horizontal circular.—Ex. Abdomen of Cœlioxys conica (Apis * * b. K.). Plate XX. Fig. 7.

10. Turbiniform (Turbiniformis). Whose vertical section is turbinate, and horizontal circular.—Ex. Joints of antennæ of Aleochara socialis, and many others of that genus.

11. Pyramidal (Pyramidalis). Whose vertical section is triangular, and horizontal quadrangular.

12. Cuneiform (Cuneiformis). Whose vertical section is cuneate, and horizontal parallelogramical.

13. Triquetrous (Triquetra). Whose horizontal sections are equilateral triangles. Plate XI. Fig. 6.

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14. Ensiform (Ensiformis). Whose horizontal sections are acute-angled triangles gradually diminishing in diameter from the base to the apex, and propagated in a straight line. Plate XI. Fig. 7.

15. Acinaciform (Acinaciformis). Whose horizontal sections are acute-angled triangles gradually increasing in diameter from the base to the apex, and propagated in a curved line.

16. Cultriform (Cultriformis). Whose horizontal sections are equal acute-angled triangles, or a three-sided body with two equal sides large and the third small.

17. Deltoid (Deltoidea). Short with the horizontal section triangular and decreasing in diameter towards the base.—Ex. Apex of the posterior tibia in Copris lunaris.

18. Trigonal; Tetragonal; Pentagonal; Hexagonal; Polygonal (Trigona; Tetragona; Pentagona; Hexagona; Polygona). Whose horizontal section is triangular; quadrangular; quinquangular; sexangular; multiangular.

19. Triedral; Tetraedral; Pentaedral; Hexaedral; Polyedral (Triedra; Tetraedra; Pentaedra; Hexaedra; Polyedra). That hath three sides; four sides; five sides; six sides; many sides.

20. Prismoidal (Prismoidalis). Having more than four sides and whose horizontal section is a polygon[1083]. Plate VI. Fig. 13. a, b, d´.

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21. Trapeziform (Trapeziformis). Whose horizontal section is a Trapezium.

22. Trapezoidiform (Trapezoidiformis). Whose horizontal section is trapezoid.

23. Rhombiform (Rhombiformis). Whose horizontal section is rhomboidal. Plate VIII. Fig. 11.

24. Two-edged (Anceps). Whose horizontal section is lanceolate.

25. Cylindrical (Cylindrica). Whose horizontal sections are all equal circles. Plate XXI. Fig. 4.

26. Fusiform (Fusiformis). Spindle-shaped. Whose vertical section is lanceolate or lineari-lanceolate, and horizontal circular. Plate XXIII. Fig. 12.

27. Columnar (Teres). Whose vertical section is cuneate, and horizontal circular. Plate XVI. Fig. 2, 3.

28. Claviform (Claviformis). Whose vertical section is clavate, and horizontal circular. Plate XI. XII. Fig. 4.

29. Cubical (Cubica). Six-sided, with sides quadrate.

30. Parallelopipedous (Parallelopipeda). Six-sided, with four parallelogramical and two quadrate sides.

31. Pyriform (Pyriformis). Pear-shaped. Whose vertical section is spatulate, and horizontal circular.—Ex. Apion, &c.

32. Infundibuliform (Infundibuliformis). Funnel-shaped. Whose horizontal sections are circular, at first equal and then progressively larger and larger. Plate XXII. Fig. 12. c.

33. Fornicate (Fornicata). Convex above and concave beneath. Plate XIII. Fig. 18. a.

34. Coarctate (Coarctata). When the diameter of[Pg 276] the middle is less than that of the ends.—Ex. Posterior thigh of Locusta. Plate XIV. Fig. 5.

35. Calceoliform (Calceoliformis). Oblong, and somewhat coarctate in the middle.—Ex. Abdomen of Chelonus.

36. Lageniform (Lageniformis). Bellying out and then ending in a narrow neck, something like a bottle.—Ex. Sperm-reservoir attached to the oviduct in Pontia. Plate XXX. Fig. 12. d.

37. Constrict (Constricta). Suddenly and disproportionably smaller at one end. Plate XXII. Fig. 15.

38. Luniform (Luniformis). Whose longitudinal section is lunate. Plate XIII. Fig. 4.

39. Nodose (Nodosa). Having one or more knobs or swellings. Plate XII. Fig. 5.

40. Geniculate (Geniculata). Bent so as to form a knee or angle. Plate XII. Fig. 7.

VII. SUPERFICIES.

i. PARTS.

1. Disk (Discus). The middle of a surface.

2. Limb (Limbus). The circumference.

3. Margin (Margo). The extreme sides.

4. Apex (Apex). The summit.

5. Base (Basis). The bottom.

6. Supine Surface (Pagina superior). The upper surface.

7. Prone Surface (Pagina inferior). The under surface.

ii. ELEVATION and DEPRESSION.

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1. Navicular (Navicularis). When two sides meet and form an angle like the outer bottom of a boat.—Ex. Notonecta glauca.

2. Convex (Convexa). An elevation the arc of which is the segment of a circle.—Ex. Upper Surface of the body of most Coleoptera.

3. Gibbous (Gibba). An elevation the arc of which is not the segment of a circle[1084].—Ex. Shoulders of the elytra of Prionus coriarius, and of many other Coleoptera.

4. Plane (Plana). Flat. When the disk is not higher than the limb, nor the limb than the disk.

5. Concave (Concava). A depression the arc of which is the segment of a circle.

6. Excavate (Excavata). A depression the arc of which is not the segment of a circle.—Ex. Prothorax of Sinodendrum cylindricum.

iii. SCULPTURE.

1. Equate[1085] (Æquata). Without larger partial elevations or depressions.

2. Smooth (Lævis). Without smaller partial elevations or depressions.

3. Levigate (Lævigata). Without any partial elevations or depressions.

[Pg 278]

4. Pore (Porus). A minute impression that perforates the substance.

5. Porose (Porosa). Beset with many pores.—Ex. Elytra of most Apions.

6. A Point (Punctum). A minute impression upon the surface, but not perforating it.

7. Punctate (Punctata). Beset with many points.—Ex. Impression on the Head and Prothorax of Phyllopertha Horticola, &c.

8. Variole (Variola). A shallow impression like a mark of the small-pox.

9. Variolous (Variolosa). Beset with many varioles.—Ex. Scarabæus variolosus.

10. Umbilicate (Umbilicata). When a variole, tubercle, granule, &c. has a depression in its centre.—Ex. Thorax of Pachygaster scabrosus.

11. Foveolet (Foveola). A roundish and rather deep depression, larger than a variole.

12. Foveolate (Foveolata). Having one or more foveolets.—Ex. Prothorax of Geotrupes stercorarius.

13. Fossulet (Fossula). A somewhat long and narrow depression.

14. Fossulate (Fossulata). Having one or more fossulets.—Ex. Oxytelus rugosus, &c.

15. Unequal (Inæqualis). Having very slight and indeterminate excavations.—Ex. Prothorax of Silpha thoracica, Cerambyx moschatus, &c.

16. Lacunose (Lacunosa). Having a few scattered, irregular, broadish but shallow excavations.—Ex. Elytra of Donacia vittata, Sagittariæ, &c.

17. Rimose (Rimosa). Chinky, resembling the bark of a tree. Having numerous minute, narrow and[Pg 279] nearly parallel excavations, which run into each other.—Ex. Elytra of Colymbetes adspersus ♀, and Cybister Rœselii.

18. Undose (Undosa). Having undulating nearly parallel broader depressions which run into each other, and resemble the sand of the sea-shore when left by the tide.—Ex. Cymatodes[1086] undosus K. MS.

19. Vermiculate (Vermiculata). Having tortuous excavations as if eaten by worms.—Ex. Prothorax of Colymbetes Hybneri and transversalis.

20. Reticulose (Reticulosa). Having a number of minute impressed lines which intersect each other in various directions like the meshes of a net.—Ex. Prothorax of Cybister Rœselii.

21. Acuducted (Acuducta). Scratched across very finely as if with the point of a needle or pin.—Ex. Colymbetes acuductus.

22. Striate (Striata). Having rather slightly impressed longitudinal parallel lines.—Ex. Amara communis, &c.

23. Sulcate (Sulcata). Having deeper impressed longitudinal parallel lines.—Ex. Dytiscus marginalis ♀.

24. Clathrose (Clathrosa). When strias or furrows cross each other at right angles.—Ex. Abdomen of Micropeplus porcatus.

[Pg 280]

25. Rivose (Rivosa). When furrows do not run in a parallel direction and are rather sinuate.—Ex. Prothorax of Elophorus stagnalis, &c.

26. Interstice (Interstitium). The space between elevations and depressions running in lines.

27. Interval (Intervallum). The space between irregular and scattered elevations and depressions.

28. Complanate (Complanata). A convex or irregular surface having a plane slight depression.—Ex. Sides of the Prothorax of Prionus cervicornis.

29. Canaliculate (Canaliculata). Having a longitudinal impressed line or channel.—Ex. Prothorax of Geotrupes, Broscus cephalotes, &c.

30. Carinate (Carinata). Having a longitudinal elevated line.—Ex. Rostrum of Curculio nebulosus. Bicarinate, Tricarinate, &c., having two or three such lines.—Ex. Elytra of Silpha recta.

31. Cristate (Cristata). Having one or two very elevated lines usually crenate.—Ex. Prothorax of Pterophylla laurifolia.

32. Porcate (Porcata). Having several parallel elevated longitudinal ridges.—Ex. Onthophilus striatus.

33. Costate (Costata). Having several broad elevated lines.—Ex. Brachinus bimaculatus, &c.

34. Clathrate (Clathrata). Having several elevated lines which cross each other at right angles.—Ex. Abdomen of Micropeplus porcatus.

35. Reticulate (Reticulata). Having many small elevated lines which intersect each other in various directions like the meshes of a net.—Ex. Lycus reticulatus. Wings of the Libellulina.

[Pg 281]

36. Rugose (Rugosa). Wrinkled. Intricate with approximating elevations and depressions whose direction is indeterminate.—Ex. Elytra of Prionus coriarius.

37. Cicatricose (Cicatricosa). Having elevated spots of a different colour from the rest of the surface, resembling scars.—Ex. Elytra of Ptomaphila lachrymosa K. MS[1087].

38. Embossed (Cælata). Having several plane tracts of a different shape higher than the rest of the surface.—Ex. Prothorax of Prionus damicornis, maxillosus, &c.

39. Gibbose (Gibbosa). Having one or more large elevations.—Ex. Sides of the Prothorax of Brachycerus barbarus.

40. Tubercle (Tuberculum). A pimple-like knob.

41. Tuberculate (Tuberculata). Having several tubercles.—Ex. Apoderus gemmatus. Base of Prothorax of Cerambyx moschatus.

42. Verruca. A small flattish wart-like prominence.

43. Verrucose (Verrucosa). Having several verrucæ.—Ex. Pimelia muricata.

44. Muricate (Muricata). Armed with sharp thick, but not close, elevated points like a Murex.—Ex. Bronchus Tribulus, quadridens[1088], &c.

45. Echinate (Echinata). Armed with sharp spines like a hedgehog or Echinus.—Ex. Hispa atra.

46. Rugged (Salebrosa). When a surface is rough[Pg 282] with mucros, spines and tubercles intermixed.—Ex. Numerous species of Bronchus.

47. Granule (Granulum). A very minute elevation.

48. Granulate (Granulata). Beset with many granules like shagreen.—Ex. Otiorhynchus sulcatus. Prothorax of Copris Molossus.

49. Scabrous (Scabra). Rough to the touch from granules scarcely visible.—Ex. Elytra of Otiorhynchus Ligustici.

50. Papillule (Papillula). A tubercle or variole with an elevation in its centre.

51. Papillulate (Papillulata). Beset with many papillules.—Ex. Elytra of Dynastes Hercules ♀.

52. Catenulate (Catenulata). Having a series of elevated oblong tubercles resembling a chain.—Ex. Carabus catenulatus.

53. Spherulate (Sphærulata). Having one or more rows of minute tubercles.—Ex. Trox lutosus, Limnius tuberculatus.

54. Consute (Consuta). Having very minute elevations in a series at some distance from each other, of a different colour from the rest of the surface, and somewhat resembling stitching.—Ex. Elytra of Oryctes Sylvanus (Cœlosis K. MS.).

55. Intricate (Intricata). When depressions or elevations so run into each other as to be difficult to trace.—Ex. Elytra of Carabus intricatus.

56. Corrugate (Corrugata). When a surface rises and falls in parallel angles more or less acute.—Ex. Front of Nothiophilus aquaticus.

57. Obliterate (Obliterata). Applied to impressions and elevations when almost effaced.

[Pg 283]

iv. CLOTHING.

a. general.

1. Scutate (Scutata). Covered with large flat scales.—Ex. Machilis polypoda.

2. Squamose (Squamosa). Covered with minute scales.—Ex. Lepidoptera.

3. Pulverulent (Pulverulenta). Covered with very minute powder-like scales.—Ex. Ceutorhynchus Sisymbrii.

4. Pollinose (Pollinosa). Covered with a loose mealy and often yellow powder resembling the pollen of flowers.—Ex. Lixus paraplecticus.

5. Farinose (Farinosa). Covered with a fixed mealy powder resembling flour.—Ex. Spots on the Elytra of Cetonia aurata, variegata, &c.

6. Lutose (Lutosa). Covered with a powdery substance resembling mud or dirt, which easily rubs off.—Ex. Trox lutosus.

7. Rorulent (Rorulenta). Covered like a plum with a bloom which may be rubbed off.—Ex. Peltis limbata.

8. Stupeous (Stupea). Covered with long loose scales resembling tow.—Ex. The Palpi of Lepidoptera. Antennæ of some Diptera. Plate XII. Fig. 23.

9. Pilose (Pilosa). Covered with long distinct flexible hairs.—Ex. Thorax of Vespa Crabro.

10. Villose (Villosa). Covered with soft flexible hairs thickly set.—Ex. Prothorax of Amphimalla solstitialis.

11. Lanate (Lanata). Covered with fine, very long,[Pg 284] flexible and rather curling hairs like wool.—Ex. Melolontha lanigera F.

12. Lanuginose (Lanuginosa). Covered with longish very soft fine down.—Ex. Prothorax of Trichius fasciatus. Thorax and base of the Abdomen of Megachile circumcincta (Apis **. c. 2. α K.).

13. Hirsute (Hirsuta). Covered with long stiffish hairs very thickly set.—Ex. Bombus.

14. Plumulose (Plumulosa). When the hairs branch out laterally like feathers.—Ex. Hair on the base of the Maxilla of Eucera (Apis ** d. 1. K.).

15. Hairy (Hirta). Covered with short stiffish sub-distinct hairs.—Ex. Genus Lagria.

16. Tomentose (Tomentosa). Covered with short interwoven inconspicuous hairs.—Ex. Acanthocinus Ædilis.

17. Pubescent (Pubescens). Covered with very fine decumbent short hairs.—Ex. Harpalus ruficornis, &c.

18. Stupulose (Stupulosa). Covered with coarse decumbent hairs.—Ex. Elytra of Melolontha vulgaris.

19. Velutinous (Velutina). Covered with very thick-set upright short hairs or pile, resembling velvet.—Ex. Trombidium holosericeum. Scutellum of Staphylinus hybridus.

20. Holosericeous (Holosericea). Covered with thick-set shining short decumbent hairs, resembling satin[1089].—Ex. Under side of the body of Elophorus stagnalis, Argyroneta aquatica, &c.

[Pg 285]

21. Setose (Setosa). Bristly. Sprinkled with stiff scattered hairs like bristles.—Ex. Echinomyia grossa.

22. Setulose (Setulosa). Setose with the bristles truncated.—Ex. Thylacites setosus.

23. Hispid (Hispida). Rough from minute spines, or very stiff rigid bristles.—Ex. Hispa atra. Phoberus horridus, &c.

24. Rough (Aspera). Rough from pubescence in general.

25. Bald (Calva). A part of a surface with little or no hair, when the rest of it is very hairy.—Ex. Vertex of Melitta and Apis Kirby.

26. Glabrous (Glabra). Without any hair or pubescence.

27. Lubricous (Lubrica). Slippery as if lubricated.—Ex. Dynastes Centaurus.

b. partial.

1. Cirrus (Cirrus). A lock of curling hair.

2. Cirrose (Cirrosa). Having one or more cirri.—Ex. Antennæ of Acanthocinus araneiformis.

3. Fascicule (Fasciculus). A bundle of thick-set hairs often converging at the apex. Plate XIX. Fig. 6. c.

4. Fasciculate (Fasciculata). Having one or more fascicules.—Ex. Catenulated lines in the Elytra of Trox arenosus. Buprestis fascicularis.

5. Penicil (Penicillus). A small bundle of diverging hairs. Plate XIX. Fig. 6. a.

[Pg 286]

6. Penicillate (Penicillata). Having one or more penicils.—Ex. Larva of Orgyia antiqua.

7. Verricule (Verriculum). A thick-set tuft of parallel hairs. Plate XIX. Fig. 6. b.

8. Verriculate (Verriculata). Having one or more verricules.—Ex. Larva of Dasychira pudibunda. Under side of Abdomen of Megachile ♀. (Apis **. c. 2. α. K.).

9. Barbate (Barbata). When any part is clothed with longer hairs, resembling a beard.—Ex. Anus of Macroglossa stellatarum. Antennæ of Cerambyx Ammiralis L. Plate XII. Fig. 26.

10. Ciliate (Ciliata). When the margin is fringed with a row of parallel hairs.—Ex. The base and apex of the Prothorax of Lucanus Cervus.

11. Fimbriate (Fimbriata). When a part is terminated by hairs or bristles that are not parallel.—Ex. Anus of many Andrenæ[1090]. (Melitta **. c. K.).

12. Comate (Comata). When very long flexible hairs thickly cover a space in the upper surface.

13. Crinite (Crinita). When very long hairs thinly cover any space.

14. Jubate (Jubata). Having long pendent hairs in a continued series.—Ex. Intermediate Legs of Anthophora pilipes (Apis **. d. 2. α. K.).

15. Furred (Pellita). When shorter decumbent hairs thickly cover any space, as in the Bombyces dorso cristato L.

v. COLOUR.

1. Niveous (Niveus). The pure unblended white of snow.—Ex. Arctia chrysorhea.

[Pg 287]

2. White (Albus). White less intense than niveous. The colour of chalk.—Ex. Spilosoma mendica ♀.

3. Lacteous (Lacteus). White with a slight tint of blue. The colour of milk.—Ex. Minoa lactearia.

4. Cream-coloured (Lactifloreus). White with a proportion of yellow.—Ex. Pale part of the Primary wings of Euprepia Caja.

5. Flesh-coloured (Carneus). White tinted with red. The colour of young and healthy flesh.—Ex. Secondary wings of Sphinx Ligustri.

6. Hoary (Incanus). White with a small proportion of black. The colour of a gray head. N.B. This term is usually confined to pubescence.—Ex. Curculio sulcirostris.

7. Cinereous (Cinereus). White with a shade of brown.—Ex. Sitona diffinis, Dasychira pudibunda.

8. Griseous (Griseus). White mottled with black or brown.—Ex. Curculio nebulosus.

9. Yellow (Flavus). Pure yellow.—Ex. Bands on the Abdomen of Nomada (Apis *. b. K.), Crabro, &c.

10. Straw-coloured (Stramineus). Pale yellow with a very faint tint of blue.—Ex. Ennomos cratægata.

11. Sulphureous (Sulphureus). Yellow with a tint of green. The colour of brimstone.—Ex. Gonepteryx Rhamni ♂.

12. Luteous (Luteus). Deep yellow with a tint of red. The colour of the yolk of an egg.—Ex. Primary wings of Colias Edusa.

13. Orange (Aurantius). Equal parts of red and yellow.—Ex. Apex of Wings of Pontia Cardamines.

14. Saffron-coloured (Croceus). The colour of saffron.—Ex. Yellow in the Elytra of Trichius fasciatus.

[Pg 288]

15. Miniatous (Miniatus). The colour of red lead.—Ex. Secondary wings of Euprepia Caja.

16. Fulgid (Fulgidus). A bright fiery red.—Ex. Lycæna Virgaureæ and Hippothoe.

17. Rufous (Rufus). A pale red.—Ex. Apion frumentarium.

18. Testaceous (Testaceus). The colour of a tile, a dull red.—Ex. Chrysomela Populi.

19. Scarlet (Coccineus). A bright pale red.—Ex. Elytra of Pyrochroa coccinea.

20. Red (Ruber). Pure red.—Ex. Under Wings of Hypercampa Dominula.

21. Sanguineous (Sanguineus). Red with a tint of black. The colour of blood.—Ex. Spots in Chilocorus Cacti, and Prothorax of Locusta morbillosa.

22. Rose-coloured (Roseus). Colour of the rose.—Ex. Parts of the Wings and Body of Deilephila Elpenor.

23. Crimson (Puniceus). A bright red with a tint of blue.—Ex. Base of the Under Wings of Catocala Sponsa.

24. Purple (Purpureus). Equal parts of blue and red.—Ex. Sagra purpurea. Vitta on the Elytra of Donacia fasciata.

25. Violet (Violaceus). Blue with some red. The colour of Viola odorata.—Ex. Chrysomela Goettingensis, Abdomen of Geotrupes vernalis.

26. Lilac (Lilacinus). Colour of the flowers of the lilac.—Ex. Part of the Iris of the Ocellus, in the Wings of Vanessa Io.

27. Blue (Cyaneus). Pure blue. Colour of Centaurea Cyanus.—Ex. Disk of the Wings of Papilio Ulysses. Callidium violaceum.

[Pg 289]

28. Azure (Azureus). A paler and more brilliant blue.—Ex. Wings of Morpho Menelaus, Telemachus, &c.

29. Sky-Blue (Cæruleus). A paler blue. The colour of the sky.—Ex. Polyommatus Adonis.

30. Cæsious (Cæsius). Very pale blue with a little black. The colour of blue eyes.—Ex. Under side of the Wings of Polyommatus Argiolus.

31. Green (Viridis). Equal parts of blue and yellow.—Ex. Cicindela campestris.

32. Æruginous (Æruginosus). Green with a blue tint. The colour of the rust of copper, verdigris.—Ex. Polydrosus Cnides.

33. Prasinous (Prasinus). Green with a mixture of yellow. The colour of the leaves of leeks or onions.—Ex. Pentatoma prasina. Under side of Wings of Thecla Rubi.

34. Glaucous (Glaucus). Pale blueish green. Sea green.—Ex. Elytra of Dynastes Hercules, Alcides, Tityus, &c.

35. Mouse-coloured (Murinus). Black with a small proportion of yellow. The colour of the common mouse.—Ex. Base of the abdominal segments of Cossus ligniperda.

36. Lurid (Luridus). Yellow with some mixture of brown. Dirty yellow.—Ex. Elytra of Aphodius luridus and nigrosulcatus.

37. Livid (Lividus). A pale purplish brown. The colour of a bruise.—Ex. Berosus luridus.

38. Tawny (Fulvus). A pale dirty orange.—Ex. The pale parts of the Wings of Hipparchia Pamphilus.

39. Fawn-coloured (Cervinus). A reddish brown.—Ex. Lasiocampa Rubi.

[Pg 290]

40. Olive (Olivaceus). A brownish green. The colour of olives.—Ex. Dytiscus marginalis.

41. Fuscous (Fuscus). A dull brown.—Ex. Hipparchia Semele. Prionus scabricornis.

42. Ferruginous (Ferrugineus). A yellowish brown with some red. The colour of the rust of iron.—Ex. Base of Under Wings of Smerinthus Populi. Gastropacha quercifolia.

43. Cinnamon-coloured (Cinnamomeus). A yellowish brown. The colour of cinnamon.—Ex. Prionus cinnamomeus.

44. Brown (Brunneus). Pure brown.—Ex. Dark parts in the Primary Wings of Euprepia Caja.

45. Bay (Badius). Bright red brown of the chestnut.—Ex. Elytra of Melolontha vulgaris when the hairs are rubbed off.

46. Chestnut (Castaneus). Colour of the dark part of the chestnut.—Ex. Elytra of Lucanus Cervus.

47. Piceous (Piceus). Shining reddish black. The colour of pitch.—Ex. Prionus coriarius.

48. Fuliginous (Fuliginosus). The opaque black of soot.—Ex. Wings of Lithosia rubricollis.

49. Black (Niger). A dull black with some brown.—Ex. Pachygaster niger.

50. Atrous (Ater). Pure black of the deepest tint.—Ex. Liparus anglicanus.

vi. SPLENDOUR.

a. gemmeous.

1. Margaritaceous (Margaritaceus). Glossy, white with changeable tints of purple, green, and blue.[Pg 291] The splendour of pearls.—Ex. The drums in Cicada capensis.

2. Opaline (Opalinus). A blueish white reflecting the prismatic colours. The splendour of the opal.—Ex. Wings of Notonecta glauca and some Nepæ.

3. Crystalline (Crystallinus). The white splendour of crystal or glass.—Ex. Stemmata of many Hymenoptera, &c.

4. Topazine (Topazinus). The yellow splendour of the topaz.—Ex. Many Stemmata of Hymenoptera, and Eyes of Spiders.

5. Rubineous (Rubineus). The red splendour of the ruby.

6. Smaragdine (Smaragdinus). The green splendour of the emerald.

7. Amethystine (Amethystinus). The purple splendour of the amethyst.

b. metallic.

1. Argent (Argenteus). The splendour of silver.—Ex. The spots on the under side of the Wings in Argynnis Lathonia, &c.

2. Golden (Aureus). The splendour of gold.—Ex. Entimus imperialis. Spot in the Wings of Plusia Festucæ.

3. Orichalceous (Orichalceus). A splendour intermediate between that of gold and brass.—Ex. Upper Wings of Plusia Chrysitis.

4. Æneous (Æneus). The splendour of brass.—Ex. Elytra of Carabus clathratus.

5. Cupreous (Cupreus). The reddening splendour of copper.—Ex. Carabus nitens.

[Pg 292]

6. Chalybeous (Chalybeus). The blue splendour of steel case-hardened, or of the mainspring of a watch.—Ex. Helops chalybeus. Legs of Lithosia Quadra.

7. Plumbeous (Plumbeus). The colour of lead.—Ex. Prothorax of Clytra dentata?

8. Inaurate (Inauratus). When striæ or other impressed parts have a metallic splendour.—Ex. Margin of Prothorax and Elytra of Carabus violaceus. Striæ of Elytra &c. of Phanæus Mimas.

9. Deaurate (Deauratus). A metallic hue which looks as if the gilding was worn off.—Ex. Donacia ænea, &c.

c. bombycine.

1. Sericeous (Sericeus). The splendour of silk.—Ex. Cryptocephalus sericeus.

2. Tramosericeous (Tramosericeus). The splendour of satin.—Ex. Chlamys Bacca, monstrosa, &c.

d. reflected.

1. Resplendent (Splendens). Reflecting the light intensely.—Ex. The Head and Thorax of Philonthus splendens, æneus, politus, &c.

2. Shining (Nitidus). Reflecting the light, but less intensely.—Ex. Dytiscus marginalis.

3. Pruinose (Pruinosus). When the splendour of the surface is somewhat obscured by the appearance of a bloom upon it like that of a plum, but which cannot be rubbed off[1091].—Ex. Elytra of Serica ruricola and brunnea.

[Pg 293]

4. Obscure (Obscurus). A surface which reflects the light but little.—Ex. Pælobius Hermanni.

5, Opaque (Opacus). A surface which does not reflect the light at all.—Ex. Trox sabulosus, arenarius. Silpha opaca.

vii. TRANSPARENCE.

1. Hyaline (Hyalina). The clear transparency of glass.—Ex. The Wings of many Neuroptera, Hymenoptera, and Diptera.

2. Diaphanous (Diaphana). Transparent, but less purely than hyaline. Semitransparent.—Ex. The Wings of many Coleoptera.

3. Adiaphanous (Adiaphana). Which does not transmit the light at all.—Ex. Elytra of Coleoptera.

viii. PAINTING.

1. Atom (Atomus). A very minute dot.

2. Irrorate (Irrorata). Sprinkled with atoms, as the earth with dew.—Ex. Onthophagus Vacca. Papilio Paris.

3. Gutta (Gutta[1092]). A roundish dot intermediate in size between an atom and a macula.

4. Guttate (Guttata). Sprinkled with guttæ.—Ex. Coccinella.

5. Macula (Macula). A larger indeterminately shaped spot.

[Pg 294]

6. Maculate (Maculata). Painted with such spots.—Ex. Abraxas grossulariata.

7. Litura (Litura). An indeterminate spot growing paler at one end, as if daubed or blotted.

8. Liturate (Liturata). A surface painted with one or more such spots.—Ex. Aphodius conflagratus.

9. Plaga (Plaga). A long and large spot.—Ex. Aphodius plagiatus.

10. Islet (Insula). A spot of a different colour, included in a plaga or macula.—Ex. The Ocelli in the Primary Wings of Hipparchia Semele. A spot in the middle fascia of the under side of the Primary Wing in Papilio Podalirius.

11. Crepera (Crepera). A gleam of paler colour upon a dark ground.—Ex. Elytra of Dytiscus marginalis.

12. Shadow (Umbra). A slight shade, not easily perceptible upon a paler ground.—Ex. Elytra of Acanthocinus Ædilis. Wings of Plutella asperella.

13. Signatures (Signaturæ). Markings upon a surface resembling in some degree letters and characters.

14. Signate (Signatus). Marked with signatures.—Ex. Elytra of Acrocinus longimanus.

15. Inscribed (Inscriptus). When the surface is marked with the resemblance of a letter of any language.—Ex. Plusia Gamma. Vanessa C. album.

16. Hieroglyphic (Hieroglyphicus). Painted with characters somewhat resembling hieroglyphics.—Ex. Acrocinus longimanus. Schizorhina Australasiæ.

17. Annulet (Annulus). A ring-shaped spot.—Ex. Cyclophora omicronaria, &c. Plate XIV. Fig. 1. o.

[Pg 295]

18. Lunulet (Lunula). A small crescent-shaped spot.—Ex. Marginal spots above and below the Secondary Wings in Melitæa Artemis, &c.

19. Reniculus (Reniculus). A small kidney-shaped spot.—Ex. Upper Wings of Mamestra Persicaria.

20. Ocellus (Ocellus). An eye-like spot in the Wings of many Lepidoptera, consisting of annuli of different colours, inclosing a central spot or pupil.

a. Pupil (Pupilla). The central spot of the ocellus. Plate XIV. Fig. 1. t. An ocellus is called bipupillate, tripupillate, &c., when there are two, three, &c. of these spots.—Ex. Primary Wing of Hipparchia Tithonus, &c. Plate XIV. Fig. 1. p.

a. Hastate Pupil (Pupilla hastata). When the pupil is a halberd-shaped spot.—Ex. Pupil of Ocellus of Aglia Tau. Plate XIV. Fig. 1. k.

b. Suffulted Pupil (Pupilla suffulta). When the pupil shades into another colour.—Ex. Primary Wing of Vanessa Io.

b. Iris (Iris). The circle which incloses the pupil. Plate XIV. Fig. 1. u.

c. Atmosphere (Atmosphæra). The exterior circle of the ocellus. Plate XIV. Fig. 1. v.

21. Blind Ocellus (Ocellus cæcus). An ocellus without the pupil.—Ex. Hipparchia Davus.

22. Spurious Ocellus (Ocellus spurius). A circular spot without any defined iris or pupil.—Ex. Spot in the Disk of the Primary Wings of Colias Helice.

23. Simple Ocellus (Ocellus simplex). When the ocellus consists only of iris and pupil.—Ex. Ocelli on the under side of Primary Wings of Hipparchia Semele. Plate XIV. Fig. 1. t, n, u.

[Pg 296]

24. Compound Ocellus (Ocellus complexus). When the ocellus consists of three or more circles.—Ex. Saturnia Spini. Plate XIV. Fig. 1. l.

25. Nictitant Ocellus (Ocellus nictitans). When the ocellus includes a lunular spot of a different colour.—Ex. Under side of Wings of Morpho Perseus. Plate XIV. Fig. 1. m.

26. Fenestrate Ocellus (Ocellus fenestratus). When an ocellus has a transparent spot.—Ex. Attacus Paphia and Cytherea.

27. Dioptrate Ocellus (Ocellus dioptratus). A fenestrate ocellus divided by a transverse line.—Ex. Attacus Polyphemus.

28. Double Ocellus (Ocellus geminatus). When two ocelli are included in the same circle or spot.—Ex. Under side of Secondary Wing of Morpho Perseus. Plate XIV. Fig. 1. r, v.

29. Twin Ocellus (Ocellus didymus). When such ocelli join each other.—Ex. Under side of Secondary Wing of Hipparchia Hyperanthus.

30. Sesquialterous Ocellus (Ocellus sesquialterus). An ocellus with a smaller near it, called also Sesquiocellus.—Ex. Under side of Secondary Wing of Colias Edusa. Plate XIV. Fig. 1. q.

31. Supercilium (Supercilium). An arched line resembling an eyebrow, which sometimes surmounts an eyelet.—Ex. Under side of Secondary Wing of Morpho Achilles. Plate XIV. Fig. 1. i.

32. Nebulose (Nebulosus). Painted with colour irregularly darker and lighter, so as to exhibit some resemblance of clouds.—Ex. Curculio sulcirostris, nebulosus; Catocala nupta.

[Pg 297]

33. Testudinarious (Testudinarius). Painted with red, black and yellow, like tortoise-shell.—Ex. Elytra of Aphodius testudinarius.

34. Consperse (Conspersus). Thickly sprinkled with minute irregular dots often confluent.—Ex. Biston Betularia.

35. Achatine (Achatinus). Painted with various concentric, curved, or parallel lines, resembling the veining of an agate.—Ex. Cossus labyrinthicus. Cerura vinula.

36. Ustulate (Ustulata). So marked with brown as to have the appearance of being scorched.—Ex. Wings of Ennomos dolabraria.

37. Marmorate (Marmorata). So painted with streaks, veins, and clouds, as to resemble marble.—Ex. Under side of the Wings of Vanessa Io. Marmarina marmorata.

38. Tessellate (Tessellata). Painted in checquerwork.—Ex. Abdomen of Sarcophaga carnaria and Musca maculata.

39. Fascia (Fascia). A broad transverse band.

a. Pyramidate Fascia (Fascia pyramidata). A band which juts out into an angle on one side.—Ex. Wing of Apatura Iris. Argynnis Paphia. Plate XIV. Fig. 1. h.

b. Macular Fascia (Fascia macularis). A band consisting of distinct spots.—Ex. Wings of Abraxas grossulariata. Plate XIV. Fig. 1. b.

c. Articulate Fascia (Fascia articulata). A band consisting of contiguous spots.—Ex. Under side of Wings of Melitæa Dictynna. Upper side of[Pg 298] Primary Wing of Morpho Menelaus. Plate XIV. Fig. 1. a.

d. Dimidiate Fascia (Fascia dimidiata). A band traversing only half the wing.—Ex. Primary Wing of Papilio Turnus. Plate XIV. Fig. 1. f.

e. Abbreviate Fascia (Fascia abbreviata). A band traversing less than half the wing.—Ex. Primary Wing of Papilio Podalirius, Ajax, &c. Plate XIV. Fig. 1. g.

f. Sesquialterous Fascia (Fascia sesquialtera). When both wings are traversed by a continued band, and either the primary or secondary by another.—Ex. Endromis versicolor. Plate XIV. Fig. 1. d, c.

g. Sesquitertious Fascia (Fascia sesquitertia). When both wings are traversed by a continued band, and more than half of either the primary or secondary by another; or, when a wing or elytrum contains a band and the third of a band.—Ex. Tortrix Avellana. Plate XIV. Fig. 1. d, e.

40. Striga (Striga). A narrow transverse streak.

41. Strigose (Strigosa). Painted with several such streaks.—Ex. Ennomos prunaria.

42. Line (Linea). A narrow longitudinal stripe.

43. Lineate (Lineata). Painted with several such stripes. N.B. If with two, we say bilineata, with three, trilineata, &c.—Ex. Elater lineatus.

44. Vitta (Vitta). A broad longitudinal stripe.

45. Vittate (Vittata). Painted with several such stripes.—Ex. Chrysomela fastuosa, cerealis, &c.

46. Undulate (Undulata). When fasciæ, strigæ, lines,[Pg 299] &c. curve into alternate sinuses resembling the rise and fall of waves.

47. Sinuato-Undulate (Sinuato-Undulata). When the sinuses are obtuse.—Ex. Boarmia repandaria.

48. Anguloso-Undulate (Anguloso-Undulata). When they go in a zigzag direction, or with alternate acute sinuses.—Ex. Acidalia undulata.

49. Radiate (Radiata). When a dot, spot, &c. appear to send forth rays.—Ex. The large blue area common to all the Wings of Papilio Ulysses.

50. Venose (Venosa). Painted with lines that branch like veins.—Ex. Under side of Wings of Pontia Napi.

51. Cancellate (Cancellata). Painted with transverse lines crossing longitudinal ones at right angles.—Ex. Macaria clathrata.

52. Areolate (Areolata). Painted with lines which intersect each other in various directions, so as to exhibit the appearance of net-work.—Ex. Wings of Tetanocera marginata and Cossus ligniperda.

53. Limbate (Limbata). When the disk is surrounded by a margin of a different colour.—Ex. Dytiscus marginalis.

54. Armillate (Armillata). When a leg, antenna, &c. is surrounded by a broad ring of a different colour.—Ex. Posterior Tibia of Prosopis annulata (Melitta * b. K.).

55. Annulate (Annulata). When a leg, antenna, &c. is surrounded by a narrow ring of a different colour.—Ex. Antennæ of many Ichneumons.

56. Cingulate (Cingulata). When the abdomen or the trunk is wholly surrounded by one or more[Pg 300] belts of a different colour.—Ex. Abdomen of many Nomadæ (Apis *. b. K.).

57. Decolorate (Decolor). When the colour appears to be discharged from any part.—Ex. Margin of the Abdominal segments in Stelis punctulatissima Latr. (Apis ** c. 1. β. K.).

58. Unicolorate (Unicolor). When a surface is of one colour.

59. Concolorate (Concolor). Of the same colour with another part. If speaking of Lepidoptera, when the upper and under sides of the wings are of the same colour.—Ex. Hesperia Linea, Paniscus.

60. Discolorate (Discolor). Of a different colour from another part. When the upper and under sides of the wings of Lepidoptera are of a different colour.—Ex. Polyommatus Corydon, Argiolus, &c.

61. Versicolorate (Versicolor). When a surface changes its colour as the light varies.—Ex. Apatura Iris ♂.

62. Iridescent (Iricolor). When a surface reflects the colours of the rainbow.—Ex. Mesothorax of Xylocopa iricolor. Wings of Hymenoptera, &c.

63. Infuscate (Infuscata). When a colour is darkened by the superinduction of a brownish shade or cloud.—Ex. Apex of the Upper Wings of Cossus ligniperda.