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 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 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: 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 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.
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 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 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]. 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 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]. 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].