their place are five valves, which by their sudden opening and closing force in the water to the rectum, the walls of which are furnished with branchial lamellae. The alternate opening and closing of these valves enables the creature to make quick jerks or rushes (incorrectly termed “leaps”) through the water,[1] and, in conjunction with its mouth parts, to make sudden attacks upon prey from a considerable distance. Well-developed Aeschnid larvae have been observed to take atmospheric air into the rectum. The lateral angles of the terminal abdominal segments are sometimes produced into long curved spines. In colour these larvae are generally muddy, and they frequently have a coating of muddy particles, and hence are less likely to be observed by their victims. If among insects the perfect dragon-fly may be termed the tyrant of the air, so may its larva be styled that of the water. Aquatic insects and larvae form the principal food, but there can be no doubt that worms, the fry of fish, and even younger larvae of their own species, form part of the bill of fare. The “nymph” when arrived at its full growth sallies forth from the water, and often crawls a considerable distance (frequently many feet up the trunks of trees) before it fixes itself for the final change, which is effected by the thorax splitting longitudinally down the back, through which fissure the perfect insect gradually drags itself. The figures indicate this process as observed in Aeschna cyanea.
The Complete Insect.—For a considerable time after its emergence a dragon-fly is without any of its characteristic colours, and is flaccid and weak, the wings (even in those groups in which they are afterwards horizontally extended) being held vertically in a line with the abdomen. By degrees the parts harden, and the insect essays its first flight, but even then the wings have little power and are semi-opaque in appearance, as if dipped in mucilage. In most species of Calopterygina, and in some others, the prevailing colour of the body is a brilliant bronzy green, blue or black, but the colours in the other groups vary much, and often differ in the sexes. Thus in Libellula depressa the abdomen of the fully adult male is covered with a bluish bloom, whereas that of the female is yellow; but several days elapse before this pulverulent appearance is attained, and a comparatively young male is yellow like the female. The wings are typically hyaline and colourless, but in many species (especially Calopterygina and Libellulina) they may be wholly or in part opaque and often black, due apparently to gradual oxidization of a pigment between the two membranes of which the wings are composed; the brilliant iridescence, or metallic lustre, so frequently found is no doubt due to interference—the effect of minute irregularities of the surface—and not produced by a pigment. A beautiful little genus (Chalcopteryx) of Calopterygina from the Amazon is a gem in the world of insects, the posterior wings being of the most brilliant fiery metallic colour, whereas the anterior remain hyaline.
These insects are pre-eminently lovers of the hottest sunshine (a few are somewhat crepuscular), and the most powerful and daring on the wing in fine weather become inert and comparatively lifeless when at rest in dull weather, allowing themselves to be captured by the fingers without making any effort to escape. Many of the larger species (Aeschna, &c.) have a habit of affecting a particular twig or other resting place like a fly-catcher among birds, darting off after prey and making long excursions, but returning to the chosen spot. A. R. Wallace, in his Malay Archipelago, states that the inhabitants of Lombok use the large species for food, and catch them by means of limed twigs.
They are distributed over the whole world excepting the polar regions, but are especially insects of the tropics. At the present day about 2200 species are known, dispersed unequally among the several subfamilies as follows: Agrionina, 700 species; Calopterygina, 280; Gomphina, 320; Aeschnina, 170; Corduliina, 130; Libellulina, 600. In Europe proper only 100 species have been observed, and about 46 of these occur in the British islands. New Zealand is excessively poor, and can only number 8 species, whereas they are very numerous in Australia. Some species are often seen at sea, far from land, in calm weather, in troops which are no doubt migratory; the common Libellula quadrimaculata, which inhabits the cold and temperate regions of the northern hemisphere, has been frequently seen in immense migratory swarms. One species (Pantala flavescens) has about the widest range of any insect, occurring in the Old World from Kamtchatka to Australia, and in the New from the Southern States to Chili, also all over Africa and the Pacific islands, but is not found in Europe. The largest species occur in the Aeschnina and Agrionina; a member of the former subfamily from Borneo expands to nearly 612 in., and with a moderately strong body and powerful form; in the latter the Central American and Brazilian Megaloprepus caerulatus and species of Mecistogaster are very large, the former expanding to nearly 7 in., and the latter to nearly as much, but the abdomen is not thicker than an ordinary grass-stem and of extreme length (fully 5 in. in Mecistogaster).
Fossils.—Among fossil insects dragon-flies hold a conspicuous position. Not only do they belong to what appears to have been a very ancient type, but in addition, the large wings and strong dense reticulation are extremely favourable for preservation in a fossil condition, and in many cases all the intricate details can be as readily followed as in a recent example. From the Carboniferous strata of Commentry, France, C. Brongniart has described several genera of gigantic insects allied to dragon-flies, but with less specialized thoracic segments and simpler wing-neuration. These form a special group—the Protodonata. True Odonata referable to the existing families are plentiful in Mesozoic formations; in England they have been found more especially in the Purbeck beds of Swanage, and the vales of Wardour and Aylesbury, in the Stonesfield Slate series, and in the Lias and Rhaetic series of the west of England. But the richest strata appear to be those of the Upper Miocene at Oeningen, near Schaffhausen in the Rhine valley; the Middle Miocene at Radaboj, near Krapina in Croatia; the Eocene of Aix, in Provence; and more especially the celebrated Secondary rocks furnishing the lithographic stone of Solenhofen, in Bavaria. This latter deposit would appear to have been of marine origin, and it is significant that, although the remains of gigantic dragon-flies discovered in it are very numerous and perfect, no traces of their subaquatic conditions have been found, although these as a rule are numerous in most of the other strata, hence the insects may be regarded as having been drowned in the sea and washed on shore. Many of these Solenhofen species differ considerably in form from those now existing, so that Dr H. A. L. Hagen, who has especially studied them, says that for nearly all it is necessary to make new genera. It is of great interest, however, to find that a living Malayan genus (Euphaea) and another living genus Uropetala, now confined to New Zealand, are represented in the Solenhofen deposits, while a species of Megapodagrion now entirely Neotropical, occurs in the Eocene beds of Wyoming.
A notice of fossil forms should not be concluded without the remark that indications of at least two species have been found in amber, a number disproportionately small if compared with other insects entombed therein; but it must be remembered that a dragon-fly is, as a rule, an insect of great power, and in all probability those then existing were able to extricate themselves if accidentally entangled in the resin.
See E. de Selys-Longchamps, Monographie des Libellulidées d’Europe (Brussels, 1840); Synopses des Agrionines, Caloptérygines, Gomphines, et Cordulines, with Supplements (Brussels, from 1853 to 1877); E. de Selys-Longchamps and H. A. L. Hagen, Revue des Odonates d’Europe (Brussels, 1850); Monographie des Caloptérygines et des Gomphines (Brussels, 1854 and 1858); Charpentier, Libellulinae europeae (Leipzig, 1840). For modern systematic work see various papers by R. M‘Lachlan, P. P. Calvert, J. G. Needham, R. Martin, E. B. Williamson, F. Karsch, &c.; also H. Tumpel, Die Geradflugler Mitteleuropas (Eisenach, 1900); and W. F. Kirby, Catalogue of Neuroptera Odonata (London, 1890). For habits and details of transformation and larval life, see L. C. Miall, Natural History of Aquatic Insects (London, 1895); H. Dewitz, Zool. Anz. xiii. (1891); and J. G. Needham, Bull. New York Museum, lxviii. (1903). For geographical distribution, G. H. Carpenter, Sci. Proc. R. Dublin Soc. viii. (1897). For British species, W. J. Lucas, Handbook of British
- ↑ A similar contrivance was suggested and (if the writer mistakes not) actually tried as a means of propelling steamships.
