Volume 1 — LETTER XXXV. (continued 2)
Reading edition
VI. Organs of Motion. We are next to consider those organs attached to the trunk of insects which are instruments of motion. These are principally those by which they are transported through the air, and those by which they move on the earth or in the water—their wings and their legs. I shall begin with the first, the wings[1776]. These are not formed precisely after any type at present discovered in vertebrate animals: in some respects they have an analogy to those of birds[1777]; in others, to the dorsal fins of fishes: but, perhaps, altogether they approach the nearest to those of the dragon or flying-lizard (Draco volans L.), which do not, as in birds, replace the fore-legs, are kept expanded by diverging bony rays, and are connected with the hind-legs[1778]. As the Divine Creator appears in his works to proceed gradually from one type of structure to another, it has been supposed by a learned physiologist of our own country, that in winged insects, four of the legs of the Decapod Crustacea are represented by the four wings[1779]: this opinion, however, is not yet fully proved; a remark which may also be applied to a more recent one of a celebrated French writer, who seems to think their origin and structure aërostatic, that they are auxiliary to the legs, and borrowed in part from the respiratory organs[1780]. Were I disposed to enter into these subtile speculations, I might here recall your attention to the analogy that, in their metamorphoses, exists between the Saurian Reptiles or lizard tribe and insects, and conjecture that the wings of the Draco are really representatives of the mid-legs of Hexapods, thus preparing to disappear altogether; but I shall content myself with throwing out this hint, which you are welcome to pursue. The organs of flight in general may be considered as to their number, kinds, and composition.
i. Number. The most natural number is four, for this obtains in the majority. In almost every Order, indeed, there occur instances of insects that have solely a single pair or none[1781].
These, however, are only exceptions to the rule; but in the Diptera, unless we consider the alulæ, the representatives of the secondary wings[1782], as a distinct pair, there are never more than two wings, and one instance is known in which an insect of this Order has none[1783]. Certain genera or individuals of the Tetrapterous Orders are also furnished with alulæ: besides Dytiscus, Blatta, Phalæna hexaptera, which have been before noticed[1784], they may be detected in miniature in Ammophila K. and affinities; these all may be regarded in some slight degree as insects with six wings.
ii. Kinds. Under this head we may consider the organs of flight as to their situation and as to their substance. As to their situation, usually the first pair are attached to the mesothorax, and the second to the metathorax; but in one instance, as has been before observed[1785], in the Strepsiptera K., the anterior pair belong to the manitrunk, and the posterior to the mesothorax. As to their substance, they take the several denominations of elytra, tegmina, hemelytra, and wings, for the most part according to its variations, as will be seen more at large hereafter. Under this head I shall only further observe, that in many instances the organs of flight appear to be mere abortions or rudiments, which serve to exemplify what has been more than once stated, that the CREATOR has seen it good to approach to new organs gradually as well as to new forms. Thus elytra are mere rudiments that do not serve to protect the wings in Atractocerus; tegmina in some species of Phasma, Acrydium, &c.; hemelytra in the bed-bug[1786]; wings in many female moths, in Cryptus hemipterus a Hymenopterous insect, &c.
iii. Composition. The structure of wings has been before explained to you[1787], and I shall again have occasion to allude to it; but here I wish to call your attention to a circumstance that has not hitherto, that I recollect, been adverted to; I mean that all kinds of organs of flight, and it may be traced as we shall soon see even in elytra, are divided longitudinally into three areas or folds; the first or external one I call the Costal Area[1788] from its beginning with the costal nervure; the second is the Intermediate Area[1789]; and the third is the Anal Area[1790].
Having made these observations with respect to the organs of flight in general, I shall now proceed to consider more at large the elytra, tegmina, hemelytra, and wings.
i. Elytra. These are the wing-covers of the Coleoptera Order, distinguished from tegmina by the absence of nervures, from hemelytra by the want of the membrane at the apex, and from both by their uniting in almost every instance at the suture. I shall consider them as to their substance; articulation with the trunk; expansion; parts; shape; appendages; sculpture; clothing; colours, and uses.
1. Substance. The firmness of the substance of elytra is usually regulated by that of the crust of the insect to which they belong; in hard insects they are hard, and in soft ones they also are soft. The most impenetrable ones that occur to my recollection are those of Illiger's genus Doryphora, and the softest and most flexile those of Telephorus, Meloe and affinities. With regard to individuals, they are mostly as hard as the prothorax, and harder than the back of the abdomen. Elytra also, as far as my observation goes, are never diaphanous.
2. Articulation with the trunk. This is by means of a process of the base of the elytrum which I call the axis[1791] or pivot, attached by elastic ligaments, and certain little bony pieces (osselets Chabr.) in the socket under the side of the anterior angle of the dorsolum[1792]. You may easily remove the elytra attached to the mesothorax from Geotrupes stercorarius, which will enable you to see the mode of articulation with little trouble[1793].
3. Expansion. It is by means of the bony pieces just mentioned that the organs in question are opened and shut[1794] under the action of the antagonist muscles. In opening for flight the two elytra recede from each other, and are elevated so as not to retain their horizontal position, which would interfere probably with the play of the wings, but form an angle with the body. When they return to a state of rest, the sutures usually meet and coincide longitudinally; but in some cases when closed, as in Necydalis, &c., they diverge from each other at the apex; and in Meloe, like the Orthoptera, to which that genus approaches, one laps over the other.
4. Parts. The parts to be considered in an elytrum are the areas, the axis, the suture, the margin, the epipleura, the base and apex, the angles, and the hypoderma. At first it should seem as if an elytrum was not like other wings divided into areas; but I think upon examination it will be found that, though often nearly obsolete, these are represented in it; for the epipleura[1795] with the recurved part of the external margin seems to me analogous to the Costal Area; the inflexed part adjoining the scutellum and often going beyond it to the Anal, and the rest of the organ to the Intermediate. All this you may see in the dung-chafer, Geotrupes stercorarius. The axis[1796] or pivot by which the elytrum articulates with the trunk is generally placed about the middle of its base, but nearer the scutellar than the humeral angle, and varies in length and shape in the different tribes, but not so as to merit particular notice; it may be regarded as composed of three parallel pieces, one belonging to each area, that of the costal being the longest. In many these pieces are marked by no line of distinction, but in Macropus, &c., they may be readily traced[1797]. The suture[1798] is the internal margin of the elytrum from the point of the scutellum to the end. In many beetles the right hand suture, looking from the anus to the head, has a lower ledge or margin, and the other, one more elevated, which when they are closed lies upon the former; in some Dynastidæ there seems a kind of ginglymous structure in this part, each suture being fitted with a kind of ridge which is received by a channel of the other; in these the suture is generally marked out by an adjacent channel: but the most remarkable structure of this part distinguishes the genuine species of the genus Chlamys, in which both the sutures, except at their base, are armed with little teeth, alternating with each other like the cogs of a mill-wheel. In apterous beetles the elytra are often connate, or have both sutures as it were soldered together. The margin[1799] or external edge of the elytra is generally formed by a bead or ridge, which, except in the case of the truncated ones, in which it is straight, curves more or less from the base to the apex; this ridge is often recurved so as to form a kind of channel between it and the disk of the elytrum, as may be seen in the Dynastidæ; in some there are two parallel ridges, as in Copris; in Silpha the margin is dilated; in Helæus and Cossyphus it is remarkably so and recurved, so that, in conjunction with those of the prothorax which are similarly circumstanced, they give the animal some resemblance to a small model of a barge. Though the margin of elytra is most commonly intire, yet in some beetles, as Gymnopleurus Illig., a sinus is taken out of it; in Cetonia it often projects at the base, and in Cryptocephalus in the middle, into a lobe; in Phoberus MacLeay it is denticulated, and in many Buprestes more or less serrulated; sometimes it terminates before it reaches the apex of the elytrum in a tooth, as in many Carabi Latr. The epipleura[1800] or side-cover is that part of the organ in question, below the margin, with which it usually forms an angle, being more or less inflexed, that covers the sides of the body. It varies in different tribes, being sometimes obsolete, as in the weevils (Curculio L.); in the Capricorn beetles it is very narrow; in Carabus, &c., dilated at the base; in many Heteromerous beetles, as Blaps, Pimelia, &c., it is very wide and conspicuous; in Cossyphus it stands out a little from the abdomen, so as to form a kind of fence round it. Its shape generally approaches that of a scythe, being incurved and growing more slender towards the apex[1801]; but it is sometimes straighter and shorter. In Geotrupes and many other Lamellicorns, the base of the elytrum is nearly vertical, forming a right angle with the rest of it; it is usually transverse and straight; but in Calandra Palmarum and many Cassidæ it slants to the scutellum; in Chlamys it is sinuate, and in Elater it has a deep cavity above the axis which receives the points of the phragma mentioned before[1802]. The apex of elytra is usually acute, the angle being formed by the confluence of a curving and straight line: but there are many exceptions; for instance, in Mylabris it is rounded; in Hister obliquely, and in Necrophorus transversely, truncated; in many Capricorns it is emarginate; in others, as Macropus longimanus, it is bidentate; in some Prioni, P. cinnamomeus, &c., it terminates in a mucro at the internal angle; and in Cerambyx Batus, horridus, &c., at the external; and, to name no more, in some species of Necydalis it ends in a long acumen. The scutellar angle in insects that have a large scutellum, as Macruspis MacLeay, is obliquely truncated to admit it, but where it is small it is generally rectangular, with the angle rounded; in Buprestis vittata it is obtusangular; and in Dytiscus marginalis, &c., it is emarginate. In Cassida spinifex, perforata, &c., the humeral angle is producted into an acute lobe that stretches beyond the head, and in C. bicornis and Taurus it forms a horn at right angles with the elytrum. In general it is either rectangular or rounded, with a prominence of the elytrum within it. The sutural and anal angles exist only where the elytra are truncated at the apex. In this case the sutural is generally rectangular, and the anal rather obtusangular or rounded. The Hypoderma is the fine soft membrane before noticed[1803] that lines the underside of the elytra, the use of which is probably to prevent injury to the wings from friction with their usually hard substance; this membrane is commonly of either a pallid or brownish colour; but in some insects, as Staphylinus hybridus, murinus, &c., Buprestis grandis, it is of a beautiful green or blue; and it exhibits the puncta, striæ, and other modes of sculpture of the elytra very distinctly, the pores of which usually perforate this membrane[1804]. Just under the shoulders of these organs you may observe an oblong and sometimes roundish spot, occasioned by the hypoderma in that part being particularly tense, and covering a cavity or pocket which appears to be connected with the axis by the hollow part, which I regard as representing the Costal Area; this pocket is evidently the analogue of a part in the wings noticed by M. Chabrier[1805], and named by me the phialum: from its connexion with the axis by a channel, this part in elytra should also seem destined to receive a fluid to add to the weight of the margin and its means of resistance.
5. Shape. The shape of elytra is various; taken together, in which case, in describing insects, they are denominated coleoptra, their most common form is more or less oblong, or forming more or less a considerable portion of an ellipse; taken separately, it inclines to that of an isosceles triangle, with the exterior side curvilinear: truncated elytra are generally quadrangular, sometimes presenting a trapezium, at others nearly a parallelogram, and at others a square. With regard to their proportions they vary considerably, but the most general law seems to be that the length shall exceed twice the width; in some, as Buprestis grandis, it is more than thrice; in many Staphylinidæ they are as wide as they are long and sometimes wider; they are generally narrower at the apex than at the base, but in some species of Lycus, as L. fasciatus, &c., the reverse takes place; in Telephorus they are nearly of the same width every where: with regard to their surface they are sometimes very convex, as in Moluris; at others very flat, as in Eurychora, Akis, &c.
6. Appendages. These, though not so remarkable as those of the head and prothorax of beetles, ought not to be overlooked. In many Capricorns, as Lamia Tribulus, speculifera, &c., the disk and sides are armed with short sharp spines; in others (Stenocorus, &c.) the sutural and anal angles or one of them terminate in a spine or tooth; sometimes the whole surface, as in Hispa atra, &c., is covered, like a porcupine, with a host of slender spines, or its sides defended by spinose lobes, as in H. erinacea: the humeral prominence is armed with a spine pointing to the head in Macropus longimanus, and forming a right angle with the elytrum in some Curculionidæ, as Rhynchites spinifex; but the most remarkable appendage of this kind is exhibited by Cassida bidens and its affinities,—from the centre of the sutures of the elytrum rise perpendicularly a pair of long, slender, sharp processes internally concave, which both apply exactly to each other, so as together to form a single horn which rises, like a mast from a ship, from the body of the animal[1806]. Besides the appendages here mentioned, the elytra exhibit a variety of tubercles and other elevations of various form and size, which it would be endless to particularize.
7. Sculpture. The sculpture of the organs in question is very various and often very ornamental: but as almost every kind of it will be noticed in the orismological tables, it will not be necessary to enlarge upon it here, especially since I have endeavoured upon a former occasion to explain how it may be useful and important as well as ornamental to the animal[1807]. I shall therefore only notice a few instances, amongst many, in which a particular kind of sculpture distinguishes particular tribes. Amongst those that are Predaceous the Cicindelidæ have elytra without striæ or furrows, while the majority of the subsequent terrestrial tribes of this section are distinguished by them: the Dynastidæ in the Lamellicorn section are remarkable for a single crenated furrow next the suture; in the weevil tribes the numerous species of the genus Apion are ornamented by furrowed elytra with pores in the furrows, which give them the appearance of neat stitching; in many of those beetles that have soft elytra, as the glow-worms (Lampyris), the blister-beetles (Cantharis, Mylabris), and still more in Œdemera, two or three slight ridges generally run longitudinally from the base to the apex, and are visible also on the under-side; as the furrows probably lighten a hard elytrum, these ridges may serve to strengthen a soft one, and it is by these that the first approach is made to the reticular structure of tegmina or the wing-covers of Orthoptera: Lycus palliatus, &c., in its elytra exhibits a direct resemblance of the reticulations of nervures.
8. Clothing. To what I have before said on this subject in general[1808] I shall here add a few remarks, which, though they more properly belong to elytra, may in many cases be extended to the whole body of a beetle. In various instances it happens that the beautiful markings of these organs, as in Macropus longimanus, whose elytra when denuded are black, are produced by short decumbent hairs; in some these variegations are the effect of scales resembling those of Lepidoptera, often of a metallic lustre; from these scales is derived all the brilliancy of the diamond-beetle (Entimus imperialis, Germ.); in some the scales are so minute as to resemble the pollen of flowers, as the white marks observable on the green elytra of the rose-chafer (Cetonia aurata).
9. Colour. The organs of flight in the majority of the Orders with respect to colour are usually the most gaily decorated part of insects; I therefore deferred the notice of that subject till I came to treat of them. In general the colour of insects is either inherent in the substance of their crust, or produced by the hairs or scales that either partially or totally cover it. To confine myself to the Coleoptera, of whose elytra we are treating, it may be observed, I think, in general, that the majority of those that feed upon putrescent substances, the saprophagous tribes of Mr. W. S. MacLeay, are commonly of a more dark and dismal aspect and colour than those which feed upon such as are living and fresh, denominated thalerophagous by the same learned author; this you may see exemplified in his Scarabæidæ and Cetoniadæ. Again, in the Predaceous beetles a similar contrast of colours is often observable. How brilliant and gay are the fierce Cicindelæ! those tigers of insects, as Linné calls them; how black as to colour, how horrible in aspect is their near relation the Manticora: what difference exists in the economy of these animals is not known, except, as I learn from Mr. Burchell, that the latter is subterraneous, whereas the former seek the sunbeam and fly rapidly. I shall now point out a few instances in which the colours of their elytra distinguish tribes or families. Amongst the Predaceous beetles a large family of the Cicindelidæ are distinguished by a middle angular white band, and several white dots on their green or brown elytra, as in C. sylvatica; a family of Brachinus, and the majority of Mylabris, Lamia capensis and fasciatus, &c., by black elytra, with yellow or red bands; Carabus violacea and affinities by the violet margin of these organs; Calliochroma Latreille by their sericeous, and Eumolpus by their metallic, lustre. These instances will be sufficient to turn your attention to this subject, which though not of primary importance in discriminating genera &c., is not without its use in a secondary view.
10. Uses. I must not quit this subject without saying something upon the ends which elytra seem designed to serve. Their first and most obvious use is the protection of the wings when unemployed, that they may not be lacerated or soiled, and rendered unfit for flight in the various retreats to which these animals betake themselves either for food, repose, or to lay their eggs; to promote this purpose more effectually, the wings are usually curiously folded and laid up under them; and where the elytra are very short, as in the Staphylinidæ, these folds are very numerous and complex. In some instances, however, as in Molorchus F., Atractocerus, &c., the wings are only partially protected by the elytra and not folded under them; probably they are less in danger of laceration from their peculiar habits than the generality. Another use is to protect the upper-side of the alitrunk, which for reasons before assigned is usually softer than the under-side, and also of the abdomen, often above nearly membranous, from the injury to which they would otherwise be exposed; in the latter part also the spiracles in Coleoptera are not covered by the inosculations of the segments, as is the case in most other Orders, and therefore probably require some covering when the insect is not flying. In the Apterous beetles this appears to be their principal use; where these organs are connate, or as it were soldered together, the back of the abdomen is a thin membrane; the appearance of two elytra in these cases is given, doubtless, for the sake of symmetry and beauty, a subordinate attention to which may be traced in all the works of creation. If we consider the bulk and weight of many flying beetles, we may imagine that they want some assistance, more than the extent and dimension of their wings seem to promise, to support them in the air, and to enable them to move more readily in it; and although it seems clear from the state of their muscular apparatus that elytra do not move much in flight, yet by giving a broad and concave surface to the air, for then they are usually nearly vertical, they may assist in some measure as sails, and help them in flying traversely and before the wind[1809].
ii. Tegmina[1810]. By this name the learned Illiger has distinguished the upper organs of flight of the Orthoptera and Homopterous Hemiptera[1811]. They may be considered under the same heads nearly as elytra.
1. Substance. Tegmina differ very materially from elytra in their substance, being generally more or less diaphanous, though in Blatta Petiveriana the dark parts are as opaque as elytra, and those of the Mantes that resemble dry leaves are only semidiaphanous. These organs are also of a less dense substance than elytra, something between coriaceous and membranous, which I shall express by the term pergameneous, as somewhat resembling parchment or vellum. Another circumstance relative to this head also distinguishes them,—they are not lined with membrane. In some instances, as in B. Petiveriana just named, they approach nearly to the substance of elytra, and in B. viridis, some Mantes, and Tettigonia, &c., they are little different from wings in their substance; but this does not diminish their right to be considered as tegmina, since their structure is altogether the same.
2. Articulation with the trunk. I observed above that the axis of elytra may be regarded as formed of three parts, one appertaining to each of the areas or their representatives[1812]; in tegmina, and indeed in wings in general, these parts are separate and may be more distinctly traced, the axis of the Costal Area being generally the longest, and that of the Intermediate often the shortest; these axes are suspended in the wing-socket by elastic ligaments, intermixed with hard bony plates, the principal one of which, called by M. Chabrier the humerus[1813], is connected both with the tegmen and the trunk, and in some a little resembles the head and neck of a swan. This structure permits the animal to move the lateral areas in some degree separately, so that each, especially the anal, shall form an angle with the intermediate; as the motion of the latter is not wanted, its axis often falls short of the base, or is obsolete, as in Blatta.
3. Composition. The three areas, traces of which we had discovered in elytra, are particularly visible in tegmina. If you take any cockroach (Blatta), you will at first sight see that in it they are divided into three larger portions by stronger nervures or folds; and if you also take a Mantis, or Locusta Leach, a Fulgora or Tettigonia, the same circumstance will strike you, only you will see that in these the intermediate portion terminates also in an axis; these are what I call the three areas. The external one or Costal is usually the longest and narrowest[1814]; the Intermediate one is commonly triangular, with its inner side curvilinear[1815]; and the interior one, or Anal area, in the Orthoptera is rather oblong; in Fulgora angular, and in Tettigonia it presents an isosceles triangle; with its vertex to the apex of the wing[1816]. The first of these may be defined as that portion of the wing that lies between the costal and postcostal nervures; and perhaps, in some cases, as in Mantis, for there is the fold of the tegmen, the mediastinal may be regarded as its limit; the Intermediate Area is that which lies between the postcostal or mediastinal nervure and the anal fold of the wing; and the Anal Area is the remainder. These areas may perhaps best be made out by tracing each to its axis. To study them carefully in tegmina and hemelytra is of considerable importance; for in them we find the first outline of the general plan upon which the wings of insects are constructed, and which, as we shall see hereafter, more or less enters into the composition of them all.
4. Position, and folding in repose. With regard to their position when not expanded, tegmina vary somewhat in the different tribes. In the Coleoptera we have seen that, except in a few instances, the elytra unite at their suture. Something like this takes place in Fulgora, Cercopis and affinities, in the Homopterous Hemiptera; in these, though the union is not near so exact, yet the tegmina do not lap over each other; they are usually more or less deflexed, with scarcely any portion in a horizontal position: in Tettigonia F., Chermes, Aphis, &c., the middle part only of these organs meets, from which point they diverge both towards their base and apex[1817]. In the Orthoptera the position is quite different, for one tegmen more or less lies over the other. In Blatta, in which the tegmina are nearly horizontal, the left hand one covers almost half the other[1818]: in the other tribes of the Order, with little variation, the Anal Area of the tegmen is horizontal, and covers the back of the animal, and the Intermediate and Costal are vertical and cover its sides; the former, however, in some cases, only forms the angle between them. Sometimes in these the right-hand one is laid upon the left, as in Acheta; and sometimes the reverse of this takes place, as in Acrida K. With regard to the folding of the tegmina, the most remarkable instance that occurs is that of Acheta monstrosa, in which the ends of both these organs and the wings, in repose, are folded like a fan, and then rolled up like a serpent[1819].
5. Shape. The shape of tegmina is various. In the Blattæ and some Mantes they are more or less oblong; in Mantis precaria, strumaria[1820], and others, they incline to elliptical; in Phasma grandis and Acheta monstrosa they are rather panduriform[1821]; in M. gongyloides they are semi-cordate[1822]; in Pterophylla trapeziformis they are rhomboidal[1823]; in Conocephalus erosus they are sinuated; in Locusta Leach they are usually linear or linear-oblong[1824]; in Pterophylla K. they generally terminate in a short mucro[1825]; and in some of those Mantidæ whose tegmina simulate arid leaves, in a recurved one[1826]. In the Homopterous Hemiptera the shape of these organs is less various. In the Fulgorellæ Latr. they incline to a trapezium, sometimes to a pentagon[1827]; in the Tettigoniæ F. they approach to an obtuse-angled triangle; and in others of the tribe they are nearly wedge-shaped[1828].
6. Neuration. The circumstance that most strikingly distinguishes tegmina from elytra is their neuration or veining; which adds much to their strength, without increasing their weight so much as to render them unapt for flight. To look at these organs in Blatta Petiveriana, you would imagine them at first to be deprived of this distinction; but if you observe them attentively, particularly their white spots, you will soon detect their nervures; and if you further examine their lower surface, you will find them very visible. The gibbous Blattæ also, Blatta picta and affinities, the analogues of Erotylus amongst the Coleoptera, have tegmina which, except at their apex, exhibit but faint traces of the nervures of their tribe, and approach to elytra besides by the innumerable minute impressed points that cover them. In the Orthoptera and some Homopterous Hemiptera the nervures may be divided into longitudinal ones more or less ramified, and traversing ones. In the Blattæ the traversing nervures cut the longitudinal ones nearly at right angles, but not at regular intervals, so as to cover the tegmen with quadrangular areolets; in Mantis precaria and affinities the longitudinal nervures of the Anal Area diverge from the base, and are traversed nearly as in Blatta, while those of the Costal diverge from the mediastinal nervure, but the traversing ones form innumerable irregular reticulations; in Mantis sinuata K.[1829] the whole tegmen has such reticulations but less numerous; in Locusta Leach it is regularly reticulated at the base, but the areolets of the apex are quadrangular; in the Mantes, with oblong wings, all are quadrangular; in Pterophylla K. the longitudinal diverging nervures are not numerous, and the traversing ones cut them into quadrangular and triangular areolets, besides which they are covered by innumerable impressed points, so as altogether to exhibit a most exact resemblance of the leaf of some evergreen: in Gryllotalpa the longitudinal nervures of the Anal Area rather converge towards the apex, are traversed by few transverse nervures, and those of the Costal Area which diverge from the mediastinal nervure by still fewer; the neuration of Acheta F. has been before described[1830]; I shall only observe here, that the constructors of stringed instruments of music might, perhaps, from the tegmina of the male, the nervures of which probably modulate the sounds which it produces, take a hint for giving the strings in them a serpentine or convolute direction, and so might produce something new in that department, corresponding with the serpents and French-horns in wind instruments. Of the Homopterous Hemiptera in the Fulgorellæ Latr., which are most analogous to the Orthoptera of all that tribe, the longitudinal nervures are more numerous and branching, more especially toward the apex of the tegmen, and are traversed as much by transverse ones, sometimes reticulating the wing with roundish areolets, as in F. laternaria, and at others with quadrangular ones, as in F. candelaria; in some of these however, as Otiocerus K., Flata F., &c.[1831], there are no traversing nervures; and these lead to the Cercopidæ and others in which the longitudinal nervures become few, and some are without any[1832], and these terminate those of this section of the Order in which the nervures in question are continued to the margin of the wing. We next come to those, Darnis, Centrotus, Membracis, &c., in which they are circumscribed a little within the apex by a traversing nervure, so that the tegmen ends in a margin of pure membrane, and thus some approach seems to be made to the Hemelytra, from Tettigonia, the most conspicuous genus of this tribe, in which the areolets, few in number, like those of Lepidoptera, are not formed, except the terminal ones, by traversing nervures, but by the ramifications of the longitudinal ones; in Chermes the Intermediate Area, which is connected with the base of the wing by a single nervure, is the only part that has any areolets[1833].
7. Colour. Orthopterous insects are seldom remarkable for tegmina of brilliant colours; there is in them none of that gilding or metallic lustre which so often distinguishes elytra: they are also frequently less ornamented in this respect than the wings, with which they usually form an agreeable contrast. Their reticulations and nervures, which are sometimes of a different colour from the rest of the tegmen, decorate them considerably: a remarkable circumstance belonging to this head attends the black tegmina of Blatta Petiveriana; one has four white spots, and the other only three; but as one laps over the other, the symmetry of the arrangement is preserved: the Homopterous Hemiptera are more distinguished in this respect, and some of the Fulgoridæ imitate the Lepidoptera both by their ocelli and spots: Fulgora laternaria, Candelaria, serrata, and Diadema, sufficiently exemplify this remark, as do several Flatæ likewise[1834].
We may observe here—that tegmina are more calculated for flight than elytra, both from their thinner substance, and from the angle that their Anal Area, and often the Costal, forms with the rest of the tegmen; a circumstance which, in wings, M. Chabrier thinks presents some facilities in that kind of motion.
iii. Hemelytra[1835]. The next species of wing-covers, which though varying in the substance of their base, terminate in a part distinct from the three areas, consisting in almost every case of mere membrane, peculiar to the Heteropterous Hemiptera, are called hemelytra, or half-elytra:—this term was also formerly employed, but certainly incorrectly, to denote tegmina. I shall consider them with respect to such of the particulars noticed under the former heads as apply to them, but without repeating them formally.
1. As to their substance, they must be separately considered with regard to their base and apex. In various instances the base, or part consisting of the three areas, is almost corneous, as in Cydnus Morio and bicolor, bugs not uncommon with us, and many others[1836]; in these cases it is lined with a hypoderma like elytra; and in many the points, which are impressed upon it, also perforate the hemelytrum, and seem to act as pores: but in Lygæus, Reduvius, Capsus, Miris, and the majority of the Heteropterous Hemiptera, the organs in question being soft and flexible, may be stated as rather resembling leather than horn;—on this account this part of a hemelytrum is denominated the corium. In Scutellera the portion covered by the scutellum is membranous; and in Acanthia paradoxa, and the cucullated species of Tingis, the wing-covers are entirely so. The apex of these organs is almost universally either membranous or coriaceo-membranous, on which account it is called the membrana. I say almost, because in Aradus and the Hydrocorisæ Latr., this part, though rather thinner than the rest of the Hemelytrum, is also coriaceous; in the latter tribe usually with a very narrow membranous edge; and in many Reduvii and Zeli there is scarcely any difference in the substance of the base and apex.
2. As to the articulation of Hemelytra with the trunk, it seems not strikingly different from that of tegmina: the point or base of the Intermediate Area, which falls short of that of the lateral areas, seems connected by a slender ligamentous piece, with its axis, which is thick; and I do not discern Chabrier's humerus shaped like a swan's head and neck[1837].
3. The composition of these organs differs from that of tegmina in more respects than one: in the first place, they consist, as was lately observed, of four instead of three areas; in the next, they appear to have, at least several of them, a part, which I suspect to be analogous to that above described in Coleoptera, supposed to represent the phialum of wings[1838]. I shall first speak of the areas. In some apterous species related to the bed-bug, Lygæus brevicollis Latr.[1839], &c., there is no trace of the usual areas, and the membrana is a very narrow strip; in L. apterus the former are very faintly traced out, but they are present in all those that are furnished with wings; whence we may conjecture that they are of the same importance in flight with the folds observable in those organs[1840]. The three basal areas may be said most commonly to present three isosceles triangles, the Costal one being narrow and curvilinear[1841], the Intermediate the most ample[1842], and the Anal one the narrowest and shortest[1843], with its vertex towards the apex of the Hemelytrum, while in the two former it is at its base. In Lygæus compressipes (Rhinuchus K. MS.) the Anal Area is cultriform; and in most of the Hydrocorisæ it has an angle in the middle of its posterior margin. The proportion that the membrana or apical area bears to the rest of the wing varies in the different tribes. In some, as before stated, it is obsolete, in others nearly so; in the majority, perhaps, it occupies about a third of the hemelytrum; in Lygæus compressipes, cruciatus, &c., full half; in Alydus calcaratus, two-thirds; in Reduvius, nearly three-quarters[1844]; and in Aradus depressus the corium,—divided, however, though indistinctly, into the three areas,—is driven to the base of the wing: two ends are answered by this structure—as this insect lives under bark, its thin hemelytra take less room; and as it flies, though it has only rudiments of wings, they are more fit to supply their place: the part we are speaking of usually runs obliquely from the vertex of the Anal Area to the base of the Costal.
4. As to their position and folding in repose, Hemelytra are usually nearly or altogether horizontal; but in Notonecta and Plea they are deflexed and cover the sides of the body; and the apical area of one wing precisely covers that of the other; where the scutellum does not intervene, as in Scutellera, Pentatoma, &c., the vertical angles of the Anal Area meet in the middle of the back, so as to exhibit the appearance of a cross. In Notonecta, in which the hemelytra are deflexed, at the apex of the membrana is a fissure which permits the two sides to form an angle with each other, and to apply exactly to the body. In Plea, in which there is no apical area, the posterior margins of the tegmina, as they ought rather to be termed, unite, but do not lap over each other. With regard to the appearance of something like a phialum, if you examine the hemelytra of most species of bugs on the underside, you will see that the costal nervure at the base is inflexed and covers a kind of channel; if you next take one of Belostoma grandis, where the structure is most conspicuous, or even the common Nepa cinerea, you will find in the same situation, adjacent to the inflexed costal nervure, a hollow tube running from the base of the wing, and terminating, after proceeding about one-fourth of its length, in a hollow cavity, which, as it is covered by a membrane, appears to me to be a collapsed pouch. This circumstance is worthy of further and more general investigation.
5. In their shape, with few exceptions, hemelytra more or less represent a wedge, being wider at their apex, where they are usually obliquely truncated, than at the base; but in Plea Leach they are obtusangular, with the angle in the sutural margin; in Notonecta, on the contrary, an obtusangular sinus distinguishes that part; in Naucoris they are curvilinear and every where of equal width; in Ranatra they are linear and straight; in Aradus they are oblong, usually with an external lobe or dilatation at their base: a remarkable instance of the intention of this is observable in a nondescript Brazilian species, in which the head, prothorax, and abdomen, are edged with a number of broad foliaceous appendages; if the base of the hemelytrum had not been furnished with a similar appendage, the symmetry of the whole body would have been destroyed by the hiatus between the prothorax and abdomen, as may be seen by removing the hemelytra; but by this compensating contrivance of Providence, the gap is filled, the above lobe fitting exactly into it.
6. The neuration of these organs will not occupy us long, since the corium or harder part, though in some species there are traces of nervures, is often without them. Those of the cucullated species of Tingis resemble many tegmina in being ornamented by them with a kind of network, which looks like the finest lace; in several Lygæi, Edessa, and some Reduvii, there are a few diverging longitudinal nervures which occasionally by a ramification here and there form an areolet[1845], but there are seldom any traversing nervures. The Apical Area is usually most distinguished by nervures, in some forming several areolets, as in Aradus, in others running parallel to each other, nearly to the end of this area, as in Belostoma grandis, where they are met by a traversing nervure; the object of this is doubtless to strengthen the membrane.
7. Both tegmina and hemelytra are most commonly naked, yet very short hairs are found on those of some species of Cercopis, and in many more instances in those of the latter description, as in Notonecta, several Lygæi and Reduvii, &c.
8. Colours in hemelytra are very various, and in many instances are peculiar to families; in certain Lygæi (L. Hyoscyami, &c.) black and red; in Lygæus compressipes and affinities a dingy black; in some Reduvii black with a large white spot;—but it is needless to enlarge further on this subject.
9. That hemelytra are used in flight is evident not only from the large space allowed for their muscles[1846], but likewise from a circumstance noticed by M. Chabrier, that in flight, in the Pentatomæ Latr., the corium of the hemelytrum is fixed to the wing[1846]; in which case both must describe the same arc.
iv. Wings. We are next to consider organs which are exclusively appropriated to flight, and therefore are properly denominated wings. These in the Orders that have elytra, tegmina, or hemelytra, are the pair that correspond with the secondary wings of the other Orders. It may be said, indeed, that in several instances both tegmina and hemelytra do not differ at all in substance or use from the wings that they cover. This is true; but as their structure in other respects is the same with that of those that are more solid and less apt for flight, it was convenient to consider them under the same name.
1. To begin with the articulation of these organs with the trunk; in general it may be stated that this, as in tegmina and hemelytra, is usually by the intervention of three axes, formed by the conflux of the nervures of the three areas at the base of the wing, which either immediately or by other pieces are implanted in the trunk, so as to receive from it the aërial and other fluids, necessary for its expansion and motions[1847]. Having given this general statement, I shall next apply it to the wings in some of the different Orders. If you carefully extract one from the stag-beetle (Lucanus Cervus) or any large species of the Dynastidæ, in the Coleoptera; the first thing that will strike you, upon examining the base, will be the plate before mentioned called by Chabrier the humerus, which is a stout transverse corneous piece, with a deep sinus towards the wing, filled with ligament: if you again follow the costal, mediastinal, and postcostal nervures, you will find them unite to form an axis, consisting of three parallel pieces, which connects by its intermediate internal piece with one end of this plate. The nervures of the Intermediate Area terminate also in a corneous axis at a greater distance from the base than the other two, which connects with Chabrier's humerus by means of the ligament of the sinus just named. Those of the Anal Area are received by a ligament attached to a transverse plate, widest at its anterior end, which connects with the posterior part of the said humerus; and at its posterior end is united to the postfrænum[1848], with which it forms a right angle. In the Orthoptera Order the structure is not very different, but the axes and other plates of the base of the wing are less distinct and rather cartilaginous; the nervures of the Anal Area often terminate in a transverse one that there forms the segment of a circle[1849]; the inner base of this circle is ligament connected with the postfrænum[1850]. In the Homopterous Hemiptera the three axes may be readily traced, but the humeral plate, with which they all are connected, is more irregular in shape, and in Fulgora longitudinal, with an angular surface; in this Order the nervure, in some cases consisting of cartilaginous rings[1851], in which the frænum and postfrænum terminate in the tegmina and wings, is attached posteriorly to the ligament of the Anal Area. In the Heteropterous section the three axes are evident, but the humeral plate is not easily made out. In the Libellulina the axes of the Costal and Intermediate Areas are the coloured broad plates at their base, formed by the dilatation of their nervures; that, however, of the Anal is not dilated, but forms one nervure, in the primary wing, with the frænum, and in the secondary with the postfrænum. Having given you this clue to trace the axes in those tribes in which they are most conspicuous, it will assist you in searching for them in the remaining Orders, in all of which they may be traced, except perhaps in those minute Hymenoptera whose wings have solely the costal nervures; probably in these there is only one axis. In the Lepidoptera and Hymenoptera a circumstance connected with the present head is observable, which is not to be discovered in the other Orders: these are the tegulæ or base-covers, which appear intended to defend the base of the anterior wings. They are concavo-convex scales, which in the Lepidoptera are large and of an irregular shape[1852], but in the Hymenoptera are smaller and semicircular[1853].
2. Wings, with regard to their substance, may generally be termed membranous; but they vary in this respect, some being much thicker than others, either partially or totally: in spotted wings, as in those of many Libellulina, Tettigoniæ F., &c., the dark opaque parts are denser than those that are transparent: in several Orthopterous insects, as in Phasma, some Mantes, &c., the Costal Area or covering part of the wing is of a substance equally firm with that of the tegmen. This is a compensating contrivance, that where the latter is shorter and smaller than the former, its membranous part, when folded, may be protected from injury. Another similar contrivance of Divine Wisdom is exhibited by those Pterophyllæ K. (Locusta F.) whose tegmina resemble the leaves of plants (Pt. laurifolia, &c.); in these the tip of the wings when folded being longer, is not covered by the tegmina, and therefore exposed to injury; to prevent which this small piece, while the whole wing, as far as covered by those organs, is membranous, is of the same substance with them[1854]. The wings of most Coleoptera, Orthoptera, Hemiptera, and Thereva coleoptrata, in the Diptera, &c., are of a firmer substance than those of the other Orders; in many Locustæ Leach, Fulgoræ, &c., they are nearly as firm as the tegmina; and in Ascalaphus italicus, except at their base, the secondary wings are less membranous than the primary. M. Chabrier has observed[1855] that the wings of insects in general diminish in thickness from their base to their apex, and from their anterior to their posterior margin.
3. I should have had, it is probable, but little original matter to communicate under the head of the composition and neuration of wings, had M. Jurine, who has written so ably on those of Hymenoptera, undertaken a survey of the organs of flight in every Order of insects: but as his views were confined to only two of the Linnean Orders, it is not wonderful that his system and set of terms should fail where a generalization is necessary; and I may stand acquitted of presumption and conceit if I attempt to substitute a system and body of terms more universally applicable. Had the plan of this able Entomologist led him to pay attention to tegmina and hemelytra, their division into three longitudinal areas would have immediately struck him; and having acquired this outline of the greater natural divisions, he would have applied it to the Orders that have wings only, and having discovered that it is to be traced in all, the result would have probably superseded my labors. Had his life been longer spared, perhaps something of this kind would have been effected by him; but as he, alas! is gone, and no abler hand seems to have undertaken the task, I will do what I can to give you satisfaction on this subject[1856]. You have already got a tolerably good idea of these areas from what has been said upon the subject under tegmina and hemelytra; but I shall now more particularly state to you how they are circumstanced in wings. I shall first explain the general law as to their limits. The Costal Area[1857] is all that longitudinal portion of the wing that lies between the anterior margin and the postcostal nervure; the Intermediate Area[1858] is all that longitudinal portion of the wing that lies between the postcostal and the anal nervures; and the Anal Area[1859] is all that longitudinal portion of the wing that lies between the anal nervure and the posterior margin. But there are other helps to enable you to distinguish the areas in the different Orders. The Anal Area in all Orders forms the posterior fold of the wing; in Coleoptera turned under when in repose; in Orthoptera folded like a fan; in Lepidoptera, in some Papilionidæ, forming an arch over the abdomen. Again, in Blatta, the Costal Area is distinguished chiefly by longitudinal nervures; the Intermediate by oblique ones; and the Anal by radiating ones; and in both this tribe and the Mantidæ this last Area is marked out from the Intermediate by a marginal notch, which is not present in Phasma, but is found in both sections of the Hemiptera. In Locusta Leach the notch is between the Costal and Intermediate Areas: in Phasma the nervures of the Intermediate Area are branches of the externo-medial, while those of the Anal, as they do in all the Orthoptera, diverge from the base of the wing: in many, as in Pterophylla K., the part of the wing lately alluded to, that is longer than the tegmen, and of the same substance, points out the limit of the Costal Area; and in others this part terminates in a segment of a circle and is differently reticulated at the apex from the Intermediate: in the Homopterous Hemiptera and the Libellulina, in which the areas at first seem indistinct, they may generally be easily traced by following them from the axes. The separation of the Costal from the Intermediate in the remaining Orders seems less easy on account of the branching of the nervures: in the rest of the Neuroptera and the Lepidoptera, if the posterior branches of the postcostal nervure are not included, you will have a narrow Postcostal Area, which in most cases forms an angle more or less prominent, in Corydalis almost a right angle, with the Intermediate: in Hemerobius and affinities this part is distinguished by areolets formed by transverse nervures, while those of the rest of the wing are longitudinal[1860]: but if the posterior branches are included, the Costal Area will be more ample: a similar observation applies to the Hymenoptera and Diptera; in these, in all cases, the areolets adjoining the anterior margin, which follow the stigma, should be regarded as belonging to the Area in question[1861]. In those tribes of the former Order, whose wings are without nervures, the areas are often marked by folds.
M. Chabrier has observed that in Coleoptera the specific weight of the margin of the wing, and its means of resistance, are augmented by a liquid which is introduced, at the will of the animal, into a long pocket under the brachial, here called the costal and mediastinal nervures, covered by a supple membrane, which in a state of repose becomes flaccid[1862]: it is easily detected, being of a paler colour than the nervures between which it lies; this is what I call the Phialum; we have before seen that it exists also in Elytra and some Hemelytra[1863]; but I have not detected it in any other wings.
I have before given you a sufficiently full account of the alulæ or winglets of Diptera[1864]; and shall here only observe that they are not confined to one particular tribe, as has been usually imagined; but though sometimes extremely minute, simple, and not easily detected, are an universal distinction of the Order.
Having thus endeavoured to elucidate the larger Areas into which wings appear to be divided; I shall next say something on the smaller ones produced by the intersection or ramification of the nervures; these had been named areolets (areolæ) several years before M. Jurine's work, in which he calls them, I think improperly, cellules (cellulæ), was published; I therefore retain the prior term. The general structure of the nervures of the wings of insects having been before explained[1865], I shall not here repeat what I then said; but there is a curious circumstance connected with it, particularly visible in the wings of certain Hymenoptera, that I must not pass without notice. If you examine attentively with a microscope against the light the wing of any Nomada or Andrena, you will discover little transparent points in some of the smaller transverse nervures that form the middle areolets, in which the nervure becomes white and looks as if it was interrupted, though in substance it seems continued: these little points, somewhat resembling minute air bubbles detained in the tubes, are what M. Jurine, who first discovered them, has, on that account, named bullæ, which he thus further describes:—"When the tube (of the nervure) arrives at the spot where a bulla is to be formed, it extends itself on all sides in minute threads in the upper membrane of the wing, losing its colour and tubular structure, which it resumes immediately after the formation of the bulla[1866]." But if you look closely at them you will find that there is always a slight fold of the wing that cuts the nervure exactly at the bullæ, and if the fold changes its direction they accompany it; their object, therefore, is clearly to relax the tension so as to admit a little motion where the fold is; consequently, rather than bullæ (bubbles), they should be denominated articulations. A similar construction, but on a larger scale, may be observed in the wings of Coleoptera[1867] and some others, as Psocus, where the folds traverse the nervures. I shall next make a few observations on the principal nervures; and first a word upon their names. M. Jurine, being of opinion that a striking analogy exists between the wings of insects and those of birds, in which M. Chabrier seems to agree with him, has named the nervures in the anterior margin of the wings of the former, radius and cubitus, as corresponding with the bones so named in the fore-arm of the latter, and the plate which often terminates these nervures in Hymenoptera, he names the carpus; it may look like presumption to differ from two such weighty authorities, but as their observations seem to have been too limited, in one case to the Hymenoptera and Diptera only; and in various Orders there is nothing analogous to the stigma or carpus, and all the other nervures of an insect's wing have no analogue in that of a bird, but more especially as M. Latreille seems to think with me on this subject[1868], I have retained Linné's term for the marginal nervure, and for most of the others have adopted those of the great French Entomologist just mentioned. I shall here only further observe,—and it seems to me an observation of prime importance, in the determination of the question of the analogy of the wings of insects,—that they are not, as in birds, the fore-leg converted into an organ of flight, but, like the wing of the Draco, an organ superadded to the legs; and, further, that the connection is not with the fore-legs, but, as has been before observed[1869], with the two posterior pairs.
The Costa[1870] is usually the strongest of the nervures, and that upon which the wing seems to be built; but in some cases, as in Blatta, Scutellera, Cynips, &c., it is represented by the mere membrane of the anterior margin; in some Coleoptera, as in Geotrupes, Dytiscus, &c., its structure, except at the base, appears to be annular or nearly so, at least a vast number of corrugations, running transversely, are observable on its upper and lower surfaces; it is thus capable of greater tension and relaxation, and more flexile. The stigma or carpus[1871], though most conspicuous in the Hymenoptera Order, may be traced in some Coleoptera, Heteropterous Hemiptera, the Libellulina, &c.; but it has no representative in the Orthoptera, Lepidoptera, Trichoptera, &c. The mediastinal is usually a very slender nervure, placed between the costa and postcosta, sometimes terminating in the former[1872], and at others in the latter[1873]: in the Orthoptera, Lepidoptera, &c., however, and some others, it is a very conspicuous and principal one[1873]; in the Hymenoptera it is obsolete, merging in those nervures[1874]. The Postcosta is the principal nervure of the wing in Scutellera, but in Staphylinus it is wanting; in Chalcis sispes it is the only true nervure of that organ, the others being represented by spurious ones[1875]. The externomedial and internomedial are sometimes distinct at their origin, but more frequently are branches from a common stem.
Having made these general remarks, I shall now consider particularly the neuration of the wings in the different Orders, beginning with the Coleoptera. The first thing that strikes the physiologist in surveying a wing belonging to an insect of this Order, is the general arrangement of the nervures[1876]; which are so placed that the required degree of tension may be given to every part of this organ: thus some are nearly straight[1877]; others run in a serpentine direction[1878]; others are forked with one branch recurrent and another proceeding onwards[1879]; others again are insulated, or do not originate from the base of the wing, or from other nervures, but are merely placed to strengthen an open space of it[1880]: these nervures are also usually broader and more substantial than those of the wings of the subsequent Orders. Another striking circumstance with regard to them is that the nervures form few or no closed areolets, except in the Costal Area, where they are inconspicuous; in Dytiscus marginalis, indeed, and Tenebrio Molitor one or two may be found, but in general there are none. In many of this tribe the postcosta, which terminates at the joint of the wing, becomes recurrent, so as to form a hook, which perhaps represents the stigma, as in Dynastes[1881]; in Creophilus K., a rove-beetle, there is no hook but a broad plate adjacent to the costa. In the Strepsiptera Order the neuration is extremely simple, the nervures, except one insulated one, diverging from the base of the wing[1882]: in this respect, as well as in the form of that organ, an approach is made to the Orthoptera. In the Dermaptera this approach is still more evident; in the common earwig[1883], the diverging nervures become numerous; between each is an insulated one, taking its origin in the middle of the wing, and running to the margin; a little nearer to the latter all the nervures are dilated into a plate; those of the anal area are angular[1884], and the exposed part of the costal is as hard as the elytra. The neuration in the Orthoptera Order may be called radiate, the longitudinal nervures for the most part diverging from the base of the wing like rays: in some few instances[1885], but not often, I believe, an insulated nervure intervenes between each; traversing or connecting nervures, cutting the longitudinal ones in various directions, ornament these wings with an infinity of areolets, causing them to resemble fine gauze or beautiful lace or net-work; very often these areolets are quadrangular, sometimes rhomboidal, frequently nearly circular, and differing occasionally, as has been before observed[1886], in the different areas: it sometimes occurs that there are no traversing nervures[1887], when the wing of course is without areolets. In the Heteropterous Hemiptera the type of neuration, as to the wing, seems borrowed from the Coleoptera, a further proof that these are the analogues of that Order amongst the Haustellata Clairv. In these the nervures usually are few and dispersed, and seldom form any closed areolets. If you examine any Scutellera, Pentatoma, or Lygæus, you may trace the uncinated, forked, serpentine, and insulated nervures of Coleopterous insects; in Gerris and Velia there is an approach to the neuration of some Homopterous species, and in Belostoma &c. the wing is reticulated by spurious nervures. In the Homopterous section there are several types of neuration; thus the Fulgoræ resemble the Orthoptera in this respect; while the Tettigoniæ F., &c., approach nearer to the Hymenoptera and Diptera, and have their apical areolets circumscribed within the margin by a traversing nervure; in Flata, &c., the areolets are mostly formed, not by traversing nervures, but by the branching of the longitudinal ones; in this respect they are not unlike the Lepidoptera. In this last-named Order there are some variations with regard to their neuration—thus, amongst the butterflies in Urania, &c., there is no closed areolet in any of the wings, and almost all the nervures diverge from the base[1888]; in Morpho, &c., there is only one in the primary wing[1889]; in Heliconia, &c., there is one in both wings; amongst the moths, in the Bombyces L., this is divided into two, and in Cossus labyrinthicus Don. into three areolets: in some butterflies (Lycæna) there is one insulated nervure[1890], and in others (Hesperia) there are two[1891]; in these two last, and Heliconia, Urania, &c., the end of the Costal Area is divided into several areolets by oblique nervures[1892], which gives them some analogy to the wings of many Neuroptera; and at the base of this Area, in Morpho, is a roundish areolet[1893]. In this Order the externo-medial and interno-medial nervures coalesce into one, and are only represented separately by their first and third branches[1894]. In the Neuroptera Order the general type of neuration is borrowed from the Orthoptera; but in Osmylus, Termes, &c., there is an approach to that of Flata in the Homopterous Hemiptera, and in Psocus to others of that section; in the second of these genera the nervures, except those of the costal margin, are spurious.
I now come to the Order in which M. Jurine has laboured with so much success, I mean the Hymenoptera; and I only regret that his labours were directed to so small a portion of the Class Insecta, and in that portion only to a part of the upper wing; I say only a part, because all those areolets of the posterior part of the wing, in some cases amounting to five[1895], that lie behind his cubital cellules, are not employed by him as diagnostics, and are left without a name. By dividing the areolets of the Intermediate Area of these wings into three portions, the basal, medial, and apical[1896], I have endeavoured to remedy this defect, and by naming each set of areolets in the middle portion, as you will see in the Orismological Definitions, under the term Areolets, you will find it easy to describe any given areolet and its place in the wing; those of the base may be called the anterior, intermediate, and posterior, where three occur; and the first and last of these terms will suffice where there are only two; the apical areolets, or those that are open to the margin, may be called, first, second, and third in the order of their occurrence, reckoning from the anterior or costal margin.
In this Order it is curious to trace the progress of neuration in the wings of different genera. Thus in Psilus only the costal nervure and the stigma are to be traced[1897]; in Chalcis the postcostal and stigma[1898]; in Codrus and Leucospis the costal, postcostal, stigma, and a nervure representing the externo-medial and interno-medial coalescing into one[1899]; in Omalus the basilar areolets appear[1900]; in Crabro both basilar and medial[1901]; in Cynips basilar, medial, and apical[1902]; and in Hylotoma the wing is filled with its greatest complement of areolets[1903]. The medial areolets of the Intermediate Area, as you will see in the definitions, form three distinct series; these may be called the protomesal, deuteromesal, and tritomesal, reckoning from the postcostal areolets; the first of these corresponds with the cubital cellules of Jurine. These series may be expressed, according to the number of their areolets; by figures, the protomesal standing first. They vary much in this respect in the different genera. Thus in Cyclostoma K.[1904], reckoning the didymous areolet as two, the numbers will stand 4:2:1; in Hylotoma, &c., 3:2:1[1905]; in Aulacus, &c., 2:2:1[1906]; in Bracon, &c., 2:1:1[1907]; in Chelonus, 2:0:1[1908]; in Cynips erythrocephalus Jur., 2:0:0[1909]; in Formica, 1:1:1[1910]; in Oxybelus, 1:0:1[1911]; in Chrysis, 0:1:1[1912]; and in Cynips Rubi K., 1:0:0[1913]. The most natural number is 3:2:1. The next in importance to the medial areolets of the Intermediate Area are the apical, or those open to the margin; the most usual number of them, excluding the postcostal areolets which belong to the Costal Area, is three; but in Sirex there is an approach to four[1914]; in Evania there are only two[1915]; and in Philanthus there are none[1916]; in many, as Prosopis, Nomada, Andrena[1917], though there is the usual number, they are incomplete and do not reach the margin. The basal areas are of little importance in assisting to determine genera; they are most commonly two in number, but in Cynips, &c., there is only one[1918]. The shape and other circumstances of the areolets vary considerably in different genera and species: upon these however I shall not enlarge further, but proceed in the next place to consider very briefly the wings of the Diptera Order as to their neuration. These are not so easily made subservient to a general plan. The basilar areolets are now reduced considerably in length, occupying merely the base of the wing[1919]; the medial are become less numerous and important[1920]; and the apical, in a variety of instances, are the most conspicuous[1921]; in some wings, as in those of Penthetria, the Intermediate Area has no nervures or areolets, or only spurious ones; in Psychoda the nervures diverge from the base almost without branching, so as to form no closed areolets[1922]; in many, the lower medial areolets are very long, resembling the basilar in Hymenoptera[1923]; these are often crowned by a single small one, as in the Stratyomidæ, Tipula, &c., from which numerous branches proceed to the margin[1924]; but in Musca two large ones approach the margin, the anterior one having an angle open to it[1925]; in the Hippoboscidæ almost the whole of the wing is occupied by the apical areolets[1926]; though in some cases they are incomplete[1927].
4. I am next to consider the position of wings in repose and their folding. The most important object of this is that when unemployed they may occupy less space, be less in the way of the insect, and be most effectually protected from injury. Another end is also served by this structure,—that wings can thus be very ample, and present a large surface to the action of the atmosphere without incommoding the insect when it has not occasion to use them.
With respect to this head, insects may be divided into two classes—namely, those whose wings in repose are covered by wing-cases harder than the wings themselves, and those that have no such protection. In the former the wings, though the rule admits several exceptions, have more folds than in the latter. As the different mode of folding the wings has been assumed for a characteristic of the earlier Orders, I shall explain to you with as much brevity as possible how each is circumstanced in this respect, beginning as usual with the Coleoptera.
There are two principal folds of the wing in this Order, which may be named the anal and the apical: the former is when the Anal Area or part of it is folded on the under surface of the base of the wing; this fold is always more or less longitudinal: the latter, the apical fold, is by means of the commissura or joint of the postcosta lately mentioned: which in Hister, Staphylinus, &c., for obvious reasons[1928] is nearer the base of the wing; in Necrophorus in the middle; in Dynastes Aloeus beyond the middle[1929]; in Tenebrio Molitor near the apex; and in Dytiscus marginalis there appears to be no joint at all; but the fact is, that in this insect the postcosta,—the termination of which really forms the joint, the costa itself being only flexible at that point,—stands at a greater distance from the latter at its end. Well, at this joint the above fold is made, the apex of the wing, being first folded longitudinally, turning under and inwards, and forming an angle, more or less acute, with the joint or costal margin, so that the fold is not quite but nearly transverse: this at least is the case in Geotrupes stercorarius and other Lamellicorns: in Staphylinus, &c., there are several transverse and longitudinal folds, and thus the wing is more easily packed under the short elytra; in Molorchus, Necydalis, &c., in which it is left uncovered, except at its base, the anal fold takes place, and the apical in some degree; a short portion near the apex forming an obtuse angle with the margin; in Atractocerus the wing appears to be only longitudinally folded; and in Buprestis vittata only the anal fold is to be detected. Besides these transverse and longitudinal folds these organs, in many beetles, have an infinity of fine corrugations, which ramify like the nervures of the tegmina of Flata[1930], &c., proceeding from the Costal Area or the disk of the wing to the posterior margin; the object of these plicatures is doubtless to present a more ample surface to the action of the atmosphere in flight[1931]. When all these folds have been made in a Coleopterous wing, the apex of the one at its posterior margin crosses or rests upon that of the other[1932].
In the Dermaptera[1933], at least the common earwig, there is a triple transverse fold of the wing, and besides this it has numerous longitudinal ones like those of a fan, each of the diverging nervures representing one of the sticks. In the Strepsiptera the folds are only longitudinal; a circumstance which, besides the form and neuration of the wing, sufficiently attests that its station is more near the Orthoptera and Coleoptera than the Diptera. We next come to the Orthoptera[1934]; in these the folds in general are longitudinal; and those of the Anal Area in particular, either in whole or in part, exact counterparts of a fan: wherever there is a straight nervure, there is usually a fold or a tendency to it; this is the case even with the short oblique ones observable in the Intermediate Area of Blatta: in this tribe the Anal Area, or a considerable portion of it, is folded under the rest of the wing, and the whole lies on the back of the animal, so that in this wing there are only two primary folds; but in those with a narrower body, as Phasma, &c., there are more, and the Anal Area, folded like a fan, lies horizontally on the back; the Costal is vertically applied to the sides, and the Intermediate is between both, as in the tegmina[1935]. In Gryllus Latr., Gryllotalpa, &c., when the wings are folded, the end of the Anal Area projects so as to present the appearance of two tails[1936]; and in that remarkable Chinese animal Gryllus monstrosus, in which these tails are very long, they are convolute like those of some quadrupeds[1937]. It is to be observed that in the secondary folds of these wings the angles of the folds are surmounted by a nervure.
In both sections of the Hemiptera Order, as in the Coleoptera, the Anal Area is turned under the wing and lies over the back of the insect; this is the only primary fold, but besides there are several longitudinal semifolds or secondary ones, in which one part of the surface forms an obtuse angle with another; and in Tettigonia, &c., these folds ramify in the wings as well as in the tegmina at the margin: a number of semifolds also, sometimes transverse and sometimes oblique, run in pairs from each side of every nervure of the disk of both tegmina and wings in the genus last named, the use of which has been before mentioned[1938].
We now come to those Orders that have four membranous wings: first, I shall consider the Lepidoptera. With respect to the position of their wings in repose some variations take place. In the majority of the day-fliers (Papilio L.), when the animal reposes the wings are applied to each other by their upper surface so as to be vertical; but in the skippers (Hesperia), the secondary wings assume a horizontal position, while the primary are vertical but applied to each other. In the Crepuscular tribes (Sphinx L.) the upper wings are incumbent on the lower, and deflexed. In the night-fliers (Phalæna L.) the types of position are various. In some Attacus, Saturnia, Noctua, &c., the wings cover each other, and are a little inclined from a horizontal position; in Gastropacha, Odenesis, and some other Bombycidæ, they are deflexed, and the anterior margin of the under wing projects beyond that of the upper: in some of the Tineæ L., as Crambus, the wings are convoluted, and in others, Galleria, they are applied close to the sides of the body, and being elevated at the apex, terminate, to use a French term—en queue de coq: in Noctua, Geometra, &c., the wings usually cover the abdomen, and are nearly horizontal. With regard to the folds of their wings, the Anal Area of the secondary is the only part that has any striking one; in Papilio Hector and affinities it turns up so as to defend the sides and part of the back of the abdomen; in Morpho Teucer it turns down, and meeting that of the opposite wing, forms a semitube which receives and shelters that part below. In the Crepuscular and Nocturnal Lepidoptera this fold, especially in the former, is very slight. With respect to semifolds in the Diurnal, there is one originating in the disk, between each of the nervures, that goes to the margin of the wing; likewise the under wings, particularly of many Noctuæ, Arctiæ, &c., have many longitudinal semifolds.
In the Neuroptera Order several variations take place with regard to the position of these organs in repose: thus, in Æshna, Libellula, &c., they continue expanded; in Argion they are applied to the body; in Myrmeleon the upper are horizontally incumbent on the lower; in Hemerobius they incline to the horizon. With regard to their folds in Æshna, &c., the longitudinal nervures alternately form the summit or the bottom of a semifold, as do those branches that terminate in the posterior margin; this kind of plicature may be observed, but in a less degree, in Ascalaphus, Myrmeleon, &c.; in Panorpa every nervure is the ridge of a slight fold; in Termes, on the contrary, it forms its bottom. In the Trichoptera, the under wing being much more ample than the upper, the Anal Area forms a fold under the wing, and there seem longitudinal secondary folds besides.
We now come to the Hymenoptera. In this Order the wings, as to their position in repose, are usually incumbent upon each other, and cover the abdomen; in the Vespidæ, however, they are placed parallel to the body, but do not cover it. Before I notice the plicature of these wings, I must recall your attention to what I lately observed[1939] with regard to Jurine's bullæ (bubbles), but which are really the joints of the nervures, as they are to be found only where the folds pass; and where they exist they are an index by which the folds, or rather semifolds, may be traced. I counted eleven of these little joints in the upper wing of Andrena cineraria; sometimes, however, instead of a bulla, a nervure stops short to admit the fold. Wings in this Order have often three longitudinal semifolds more or less conspicuous; these you may trace in the saw-flies (Tenthredo L.), whose wings Linné terms tumidæ, by which term he would indicate the elevation of the whole surface produced by this structure; in the under wings of these, and Scolia, Bembex, &c., the Anal Area is turned under the wing, as in many preceding tribes[1940]: in Sirex, &c., that Area of the upper wing turns upwards, forming an acute angle with the rest of the organ; the same circumstance distinguishes the under wing in the Ichneumonidæ. Several apical semifolds, marked by a pellucid streak, distinguish Tiphia F., and in Bombus, Bembex, &c., an infinity of branching ones, like those before described in Coleoptera, corrugate the apical margin. In the Vespidæ the upper wings are folded longitudinally into three nearly equal portions, but in the under ones the Anal Area only forms the fold.
In the Diptera Order, as to their position when at rest, the wings are mostly incumbent one on the other; but in Psychoda they are deflexed, so as to form a kind of penthouse. With regard to their plication, in some, Tipula oleracea, &c., a slight oblique semifold runs from the stigma to the apical margin, and the Anal Area has two, as it has in many Muscidæ, itself forming nearly a right angle with the rest of the wing; besides these it is corrugated with minute transverse semifolds, which are observable also in several other Dipterous insects; in many Stratyomidæ they are oblique, and run from the disk to the posterior margin; and in Asilus, Bombylius, &c., they are wavy.
5. We are next to say something upon the shape of wings: this, though apparently extremely various in the different Orders and tribes, may I think be traced in every wing to one original prototype, a triangle with the largest angle rounded and subtended by the anterior or costal margin: in some, as the Coleoptera, Orthoptera, &c., this type of formation is a right-angled triangle[1941]; and in others, as in the Hymenoptera, Diptera, &c., the majority of the Neuroptera, &c., it is an obtusangled one[1942]; it may be further observed, that in receding from these forms wings very often assume that of the half or quadrant of some regular figure, as we shall see when we consider those of the different Orders. Another general observation I shall first mention,—that these organs are universally narrowest at their base and widest at the apex, provided we consider as the apex the termination outwards of the three Areas; otherwise we might say that wings in the Coleoptera, Orthoptera, &c., were wider at the base than at the apex[1943]. The wings in the former Order, and in several of the Heteropterous Hemiptera, as Gerris, Velia, &c., may in general, as to their shape, be termed semicordate or semiovate[1944]; in the Dermaptera they incline to an oval figure[1945]: in the Strepsiptera, Orthoptera, most Homopterous and many Heteropterous Hemiptera, they approach to the quadrant of a circle; in a considerable portion of the Lepidoptera the two under wings, if united at their posterior margin, approach a circular form; the upper ones vary a little from the prototype of the under ones, forming an obtusangled triangle[1946]; in many Neuroptera the primary wings may be called oblong or linear-oblong, while the secondary betray more evidently the right-angled or obtusangled triangle; in the Hymenoptera this latter form is every where conspicuous, with little deviation, except in the rounding of the angles[1947]; and, finally, in the Diptera this form shades off again into an oblong, ovate, or linear shape, the wing being most commonly attenuated at the base into a kind of footstalk[1948]. Some singular variations with respect to the termination or marginal processes of the wings are exhibited by many Lepidoptera; thus in Attacus Atlas, &c., the primary wings are falcated or hooked at their apex[1949]; and in great numbers both wings are there scolloped into alternate bays and capes, if I may so speak, varying in depth and length[1950]. There is usually a sinus between every pair of nervures, each of which terminates in the adjoining prominence, as a fold does in the sinus[1951]. Where present, in the primary wings there are eight of these sinuses, and in the secondary, where they are most usual, seven; some are remarkable for the long tails which distinguish their secondary wings; those in Papilio are usually an elongation of the fifth, from the anterior margin, of the prominences before mentioned, into a spathula-shaped diverging process, varying in length and width[1952]: but in P. Ulysses it does not diverge; and in P. Podalirius it is linear. They are found also in other subgenera; thus in Urania Patroclus there are two; in U. Riphæus three; in Erycina Cupido five; and in E. Endymion six of these tails; in some, as in E. Dorylas, the whole wing seems to form the tail; in others again, as in Hesperia Proteus and Bombyx Luna, it is an elongation of the anal angle. Other wings in this Order are divided into lobes resembling feathers, as you may see in Pterophorus hexadactylus, &c.[1953]
6. We are next to consider the clothing of wings: these, in the Orders in which they are covered by elytra, tegmina, or hemelytra, are generally naked, except that the spots in those of Fulgora laternaria, serrata, &c., and the whole wing in Flata, Aleyrodes, and others, are covered with a kind of farinaceous powder; but in all the remaining Orders, hairs or scales are more or less implanted in these organs: as the Lepidoptera are the most remarkable for the clothing of their wings, I shall leave them till the last, and begin with the Neuroptera. If you lightly pass your finger over the wing of any dragon-fly (Libellula F., Æshna F.), from the apex towards the base, you will find that the longitudinal nervures are, as it were, serrulated with very minute bristles, which point towards the extremity; if you next move the finger across the wing, from the posterior to the anterior margin, a similar circumstance will strike you. M. Chabrier conjectures that, amongst other uses[1954], these hairs may contribute to fix the atmospheric fluid when the wings are depressed in flight, while it glides over them as they rise[1955]; in Ascalaphus, Myrmeleon, Nemoptera, Hemerobius, &c., the nervures are more visibly bristled; the bristles diverging on each side from the longitudinal ones, but all pointing towards the apex from the connecting or transverse ones; in Panorpa, besides these bristles, short hairs, pointing the same way, are thickly planted in the membrane of the wing; and in Hemerobius the margins of the wing are fringed; in the Ephemerina, Corydalis, &c., the wings are naked. In the Trichoptera Order, as their name imports, they are covered with minute decumbent hairs, less easily seen but still existing in the secondary pair. In the Hymenoptera in general the wings are covered with minute hairs or bristles; but in Tiphia, Scolia—with the exception of S. Radula and affinities in which they are hairy—and others, the wings are nearly naked; in Pompilus, Pepsis, &c., the hairs are infinitely numerous and very short; in the Sphecidæ, Mutilla, &c., they are more distinct, longer, and less numerous; in the humble-bee (Bombus) and many others the apex of the wing is darkened by a large number of more conspicuous hairs, each of which seems to spring from a minute tubercle: as these tubercles are in a part of the wing that is strengthened by few nervures, they may probably be intended to supply their place, in giving firmness and tension to this part. The wings of Diptera, under the present head, may be viewed with regard to the hairs that are implanted in the membrane of the wing, in its nervures, and in its margin. In the first view, in Stratyomis and immediate affinities the wing is nearly naked; but in Xylophagus, Beris, and the great majority of the Order, the membrane of the wings is thickly planted with innumerable very minute bristles, not to be seen but under a powerful lens, often black, and seemingly crowning a little prominence, and giving the wing an appearance of the finest net-work. As to the clothing of the nervures, the costal, in Anthrax, Bombylius, &c., is often remarkably bristly at the base, with hairs intermixed; in Œstrus Ovis, in the inner margin or edge of this nervure, is a single series of bristles, or rather short spines, like so many black points; in Œ. Equi the whole costa is covered with short decumbent hairs or bristles; in Musca pagana F., just at the apex of the costal areolet, that nervure is armed with a spur or diverging bristle larger than the rest, which is also to be found in many others of the Muscidæ, some of which have two and others more of these spurs. The little moth-like midges (Psychoda Latr., Hirtæa F.) at first appear to have the whole surface of their wings covered with hairs; but upon a closer examination it will be seen that they are planted in the nervures, from each of which they diverge, so as under a lens to give it a very elegant appearance[1956]. This fly has its wings beautifully fringed with fine hairs, the third circumstance to be attended to under this head; in the Tipulidans, and many others of this Order, the apex and posterior margin are also finely fringed with short hairs. Some Dipterous insects make a near approach to the Lepidoptera in the covering of their wings: in the common gnat, when the wings are not rubbed, the nervures are adorned by a double series of scales, and the marginal fringe also consists of them[1957]; and in a Georgian genus, which appears in some degree to connect Culex with Anthrax &c., there are scales scattered upon the membrane as well as upon the nervures; besides, its antennæ[1958] and abdomen are also covered with them.
The Order, the clothing of whose organs of flight excites the admiration of the most incurious beholder, is that to which the excursive butterfly belongs, the Lepidoptera. The gorgeous wings of these universal favourites, as well as those of the hawk-moths and moths, owe all their beauty, not to the substance of which they are composed, but to an infinite number of little plumes or scales so thickly planted in their upper and under surface, as in the great majority entirely to conceal that substance. Whether these are really most analogous to plumes or scales has been thought doubtful. De Geer is inclined to think, from their terminating at their lower end in little quills and other circumstances, that they resemble feathers as much as scales[1959]; Reaumur on the contrary suspects that they come nearer to scales[1960]. Their substance, approaching to membrane, seems to make further for the former opinion, and their shape and the indentations that often occur in their extremity, furnish an additional argument for the latter. Their numbers are infinite; Leeuwenhoek found more than 400,000 on the wings of the silk-worm moth (Bombyx Mori)[1961]; and in those of some of the larger moths and butterflies the number must greatly exceed this. You will observe however that in many Lepidoptera the wings are partially, and in some instances generally, transparent: thus in Hesperia Proteus, a butterfly before noticed for the long tail that distinguishes its secondary wings, there are many transparent spots; in Attacus Atlas, one of the largest of moths, and its affinities, there is as it were a window in each wing formed by a transparent triangular space; in A. Polyphemus, Paphia, &c., the pupil of the ocellus is transparent, which in the former is divided by a nervure. In several of the Heliconian butterflies, and in Zygæna F., &c., the greater part of both wings is transparent, with scales only upon their nervures, round their margin, or forming certain bands or spots upon them; in Parnassius Apollo, Mnemosyne, &c., the scales are so arranged as not wholly to cover the wings, which renders them semidiaphanous; and in some (Nudaria) the wings are intirely denuded. With regard to size, the scales vary often considerably in different tribes; in Heliconia they appear to be more minute than in the rest; and in Castnia they are the largest and coarsest; the extremity of the wings of Lepidopterous insects in general is fringed with longer scales than their surfaces, and even those of the last in the same wing; sometimes vary in magnitude. The little seeming tooth that projects from the middle of the posterior margin in the upper wings of Notodonta, a subgenus of Bombyx L., is merely produced by some longer diverging hairs. The shape and figure also of scales are very various—some being long and slender; others short and broad; some nearly round; others oval, ovate, or oblong; others spathulate; others panduriform or parabolical; some again almost square or rhomboidal; many triangular; some representing an isosceles triangle, and others an equilateral one; lastly, some are lanceolate and others linear; again, some have a very short pedicle and others a very long one: with regard to their extremity; some are intire, without projecting points or incisions, while others are furnished with them: of these some terminate in a single long mucro, others have several shorter ones; some are armed with teeth, varying in number from two to thirteen in different species[1962]. Many other forms might be enumerated, but these are sufficient to give you a general notion of the infinite variety of this part of the works of the Creator. I must next say a word or two upon their arrangement on the wing. In most instances this is in transverse lines, which sometimes vary a little from a rectilinear course, and the extremity of the scales of one row reposes on the base of those of the succeeding one, so that in this respect their arrangement is like that of tiles in a roof: in some cases it is not so regular: thus the minute scales on the wings of Parnassius Apollo, and others with subdiaphanous wings, are arranged without order; in Pieris and other Diurnal Lepidoptera, and many of the Crepuscular and Nocturnal, there appears to be a double layer of scales on both sides of the wing; the under layer usually consisting of white ones. If you denude the wings of any butterfly, which you may easily do by scraping it lightly on both sides with a penknife, you will be amused to trace the lines in which the scales were planted, consisting of innumerable minute dots: the lines of the under side, in some cases, so cut those of the upper side, as by their intersection to form lozenges. With regard to the position of the scales on the wing, they usually lie flat, but sometimes their extremity is incurved: in the beautiful Argynnis Vanillæ a very singular appearance of numerous transverse ridges is produced by the extremity of those scales that cover the longitudinal nervures of the primary wings, except at the base, being recurved.
But though the general clothing of the wings of Lepidoptera consists of these little scales, yet in some cases they are either replaced by hairs or mixed with them. Thus, in the clear parts of the wings of Heliconians, Attaci, &c., short inconspicuous hairs are planted; in a large number of the Orders the upper side of the Anal Area of the secondary wings is hairy; in several Crepusculars (Sphinx Phœnix, &c.), where there is a double layer as before mentioned, the upper one consists of dense hairs, except at the apex, and the lower one of scales; and in most of them the scales of the primary wings are piliform, and the secondary are covered by what approach very near to real hairs; many of the Attaci are similarly circumstanced: the four wings of A. Cytherea are also covered externally with hair.
7. Before I conclude this long diatribe on the organs of flight of insects, I must not omit some notice of the infinite diversity of colours with which their wings are often variegated and adorned by the Creator, who loves to delight us by the beauty, as well as to astonish and awe us by the immensity and grandeur of his works. Though the wings in every Order exhibit instances of brilliant and beautiful colouring, yet those of the Lepidoptera in this respect infinitely excel them all, and to these, under this head, after noticing a few in the less privileged Orders, my observations will be confined. Although in the Coleoptera the wings are seldom distinguished by their splendour; yet those of some Cetoniadæ, as Cetonia africana, are extremely brilliant, and resemble those of many Xylocopæ in the lovely violet hue that adorns them: amongst the Orthoptera some Pterophyllæ, and in the Homopterous Hemiptera some Fulgoræ, emulate the Lepidoptera in the ocelli that give a kind of life to these organs[1963]; and a vast number of the destructive tribe of locusts (Locusta Leach) are remarkable for the fine colours and gaiety of their wings[1964]; in the Neuroptera numerous Libellulinæ emulate the Heliconian butterflies by their maculation; and in the genus Ascalaphus, which represents the Lepidoptera by its clubbed antennæ[1965], many also have the resemblance increased by the painting of their wings, so that some Entomologists have actually considered some of them as belonging to that Order[1966]; the wings of the Xylocopæ, before alluded to, sometimes add to the deep tints of the violet—which also prevail in the wings of several Diptera—towards their extremity the most brilliant metallic green or copper varying,
"As the site varies in the gazer's hand,"
and even those wings that consist of clear colourless membrane are often rendered extremely beautiful from the reflection of the prismatic colours. I should undertake an endless task did I attempt to specify all the modes of marking, clouding, and spotting, that variegate a wing, and all the shades of colour that paint it, amongst the Lepidopterous tribes; I shall therefore confine myself to a few of the principal, especially those that distinguish particular tribes and families. Of whole coloured wings—I know none that dazzle the eye of the beholder so much as the upper surface of those of Morpho Menelaus and Telemachus: Linné justly observes that there is scarcely any thing in nature that for brightness and splendour can be paralleled with this colour; it is a kind of rich ultramarine that vies with the deepest and purest azure of the sky; and what must cause a striking contrast in flight, the prone surface of the wings is as dull and dark as the supine is brilliant, so that one can conceive this animal to appear like a planet in full radiance, and under eclipse, as its wings open and shut in the blaze of a tropical sun: another butterfly, Papilio Ulysses, by its radiating cerulean disk, surrounded on every side by a margin intensely black, gives the idea of light first emerging from primeval obscurity; it was probably this idea of light shining in darkness that induced Linné to give it the name of the wisest of the Greeks in a dark and barbarous age. I know no insect upon which the sight rests with such untired pleasure, as upon the lovely butterfly that bears the name of the unhappy Trojan king (P. Priamus); the contrast of the rich green and black of the velvet of its wings with each other, and with the orange of its abdomen, is beyond expression regal and magnificent. But peculiar beauties of colour sometimes distinguish whole tribes as well as individuals. What can be more lovely than that tribe of little butterflies that flit around us every where in our summer rambles, which are called blues, and which exhibit the various tints of the sky? Lycæna Adonis of this tribe scarcely yields to any exotic butterfly in the celestial purity of its azure wings: our native coppers also, Lycæna dispar[1967], Virgaureæ, &c., are remarkable for the fulgid colour of these organs; in Argynnis the upper side of their wings is tawny, spotted with black, while the under side of the secondary ones is very often adorned by the appearance of silver spots. How this remarkable effect of metallic lustre, so often reflected by spots in the wings of butterflies, is produced, seems not to have occupied the attention of Entomologists. M. Audebert is of opinion that the similar lustre of the plumes of the humming birds (Trochilus) is owing to their density, to the polish of their surface, and to the great number of little minute concave mirrors which are observable on their little beards[1968]. But these observations will not apply to the scales of the wings of butterflies, which are always very thin and generally very flat: in some instances, as in Morpho Menelaus, there appears more than one very slight channel upon a scale; but this takes place also in others that reflect no lustre. Their metallic hues must therefore principally be occasioned by the high polish of their surface and the richness of their tints. It is the purity of the white, in conjunction with their shining surface, contrasted with the dull opaque colour of the under side of the secondary wings, that causes the spots that decorate those of the Fritillaries (Argynnis) to emulate the lustre of silver. In Papilio the Trojans are distinguished by the black wings with sanguine spots, and the Greeks by the same with yellow spots; but these have proved in some instances only sexual distinctions[1969]. In the Danai candidi L. the colour of the tribe may be described as sacred to the day, since every shade, from white or the palest yellow to full orange, is exhibited by them. The yellows prevail also in those Noctuæ, the trivial names of which Linné made to end in ago, as N. Fulvago, Citrago, &c. I must not conclude this part of my subject without noticing one of the most striking ornaments of the wings of Lepidoptera, the many-coloured eyes which decorate so large a number of them. Some few birds, as the Peacock and Argus Pheasant, have been decked by their Creator very conspicuously with this almost dazzling glory; but in the insects just named it meets us every where. Some, as one of our most beautiful butterflies, Vanessa Io[1970], have them both on the primary and secondary wings; others, as Noctua Bubo[1971], only on the primary; others again, as Smerinthus ocellata[1972], only on the secondary: in some also they are on both sides of the wing, as in Hipparchia Ægeria[1973], and in others only on the upper side, as in Vanessa Io; in others again only on the under side, as in Morpho Teucer[1974]: in some likewise they are very large, as in the secondary wings of the same butterfly: and in others very small, as in those in the wings of the blues (Lycæna). Once more, in some they consist only of iris and pupil, as in Hipparchia Semele, and in others of many concentric circles besides, as in Morpho Teucer, &c.