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 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 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 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 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 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 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 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.