already mentioned. The second (so-called “first”) abdominal segment is often very constricted, forming the “waist” so characteristic of wasps and ants for example. The constriction of this segment and its very perfect articulation with the propodeum give great mobility to the abdomen, so that the ovipositor or sting can be used with the greatest possible accuracy and effect.
Mention has already been made of the series of curved hooks along the costa of the hind-wing; by means of this arrangement the two wings of a side are firmly joined together during flight, which thus becomes particularly accurate. The wings in the Hymenoptera show a marked reduction in the number of nervures as compared with more primitive insects. The main median nervure, and usually also the sub-costal become united with the radial, while the branches of radial, median and cubital nervures pursuing a transverse or recurrent course across the wing, divide its area into a number of areolets or “cells,” that are of importance in classification. Among many of the smaller Hymenoptera we find that the wings are almost destitute of nervures. In the hind-wings—on account of their reduced size—the nervures are even more reduced than in the fore-wings.
The legs of Hymenoptera are of the typical insectan form, and the foot is usually composed of five segments. In many families the trochanter appears to be represented by two small segments, there being thus an extra joint in the leg. It is almost certain that the distal of these two segments really belongs to the thigh, but the ordinary nomenclature will be used in the present article, as this character is of great importance in discriminating families, and the two segments in question are referred to the trochanter by most systematic writers.
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| After C. Janet, Aiguillon de la Myrmica rubra (Paris, 1898). |
| Fig. 5.—Ovipositor or Sting of Red Ant (Myrmica rubra) Queen. Magnified. The right sheath C (outer process of the ninth abdominal segment—9) is shown in connexion with the guide B formed by the inner processes of the 9th segment. The stylet A (process of the 8th abdominal segment—8) is turned over to show its groove a, which works along the tongue or rail b. |
The typical insectan ovipositor, so well developed among the Hymenoptera, consists of three pairs of processes (gonapophyses) two of which belong to the ninth abdominal segment and one to the eighth. The latter are the cutting or piercing stylets (fig. 5, A) of the ovipositor, while the two outer processes of the ninth segment are modified into sheaths or feelers (fig. 5, C) and the two inner processes form a guide (fig. 5, B) on which the stylets work, tongues or rails on the “guide” fitting accurately into longitudinal grooves on the stylet. In the different families of the Hymenoptera, there are various modifications of the ovipositor, in accord with the habits of the insects and the purposes to which the organ is put. The sting of wasps, ants and bees is a modified ovipositor and is used for egg-laying by the fertile females, as well as for defence. Most male Hymenoptera have processes which form claspers or genital armature. These processes are not altogether homologous with those of the ovipositor, being formed by inner and outer lobes of a pair of structures on the ninth abdominal segment.
Many points of interest are to be noted in the internal structure of the Hymenoptera. The gullet leads into a moderate-sized crop, and several pairs of salivary glands open into the mouth. The crop is followed by a proventriculus which, in the higher Hymenoptera, forms the so-called “honey stomach,” by the contraction of whose wails the solid and liquid food can be separated, passed on into the digestive stomach, or held in the crop ready for regurgitation into the mouth. Behind the digestive stomach are situated, as usual, intestine and rectum, and the number of kidney (Malpighian) tubes varies from only six to over a hundred, being usually great.
In the female, each ovary consists of a large number of ovarian tubes, in which swollen chambers containing the egg-cells alternate with smaller chambers enclosing nutrient material. In connexion with the ovipositor are two poison-glands, one acid and the other alkaline in its secretion. The acid gland consists of one, two or more tubes, with a cellular coat of several layers, opening into a reservoir whence the duct leads to the exterior. The alkaline gland is an irregular tube with a single cellular layer, its duct opening alongside that of the acid reservoir. These glands are most strongly developed when the ovipositor is modified into a sting.
Development.—Parthenogenesis is of normal occurrence in the life-cycle of many Hymenoptera. There are species of gall-fly in which males are unknown, the unfertilized eggs always developing into females. On the other hand, in certain saw-flies and among the higher families, the unfertilized eggs, capable of development, usually give rise to male insects (see Bee). The larvae of most saw-flies feeding on the leaves of plants are caterpillars (fig. 6, b) with numerous abdominal pro-legs, but in most families of Hymenoptera the egg is laid in such a situation that an abundant food-supply is assured without exertion on the part of the larva, which is consequently a legless grub, usually white in colour, and with soft flexible cuticle (fig. 7, a). The organs and instincts for egg-laying and food-providing are perhaps the most remarkable features in the economy of the Hymenoptera. Gall-fly grubs are provided with vegetable food through the eggs being laid by the mother insect within plant tissues. The ichneumon pierces the body of a caterpillar and lays her eggs where the grubs will find abundant animal food. A digging-wasp hunts for insect prey and buries it with the egg, while a true wasp feeds her brood with captured insects, as a bird her fledglings. Bees store honey and pollen to serve as food for their young. Thus we find throughout the order a degree of care for offspring unreached by other insects, and this family-life has, in the best known of the Hymenoptera—ants, wasps and bees—developed into an elaborate social organization.
Social Life.—The development of a true insect society among the Hymenoptera is dependent on a differentiation among the females between individuals with well-developed ovaries (“queens”) whose special function is reproduction; and individuals with reduced or aborted ovaries (“workers”) whose duty is to build the nest, to gather food and to tend and feed the larvae. Among the wasps the workers may only differ from the queens in size, and individuals intermediate between the two forms of female may be met with. Further, the queen wasp, and also the queen humble-bee, commences unaided the work of building and founding a new nest, being afterwards helped by her daughters (the workers) when these have been developed. In the hive-bee and among ants, on the other hand, there are constant structural distinctions between queen and worker, and the function of the queen bee in a hive is confined to egg-laying, the labour of the community being entirely done by the workers. Many ants possess several different forms of worker, adapted for special duties. Details of this fascinating subject are given in the special articles Ant, Bee and Wasp (q.v.).
Habits and Distribution.—Reference has been already made to the various methods of feeding practised by Hymenoptera in the larval stage, and the care taken of or for the young throughout the order leads in many cases to the gathering of such food by the mother or nurse. Thus, wasps catch flies; worker ants make raids and carry off weak insects of many kinds; bees gather nectar from flowers and transform it into honey within their stomachs—largely for the sake of feeding the larvae in the nest. The feeding habits of the adult may agree with that of the larva, or differ, as in the ease of wasps which feed their grubs on flies, but eat principally vegetable food themselves. The nest-building habit is similarly variable. Digging wasps make simple holes in the ground; many burrowing bees form branching tunnels; other bees excavate timber or make their brood-chambers in hollow plant-stems; wasps work up with their saliva vegetable fibres bitten off tree-bark to make paper; social bees produce from glands in their own bodies the wax whence their nest-chambers are built. The inquiline habit (“cuckoo-parasitism”), when one species makes use of the labour of another by invading the nest and laying her eggs there, is of frequent occurrence among Hymenoptera; and in some cases the larva of the intruder is not content with taking the store of food provided, but attacks and devours the larva of the host.
Most Hymenoptera are of moderate or small size, the giants of the order—certain saw-flies and tropical digging-wasps—never reach the bulk attained by the largest beetles, while the wing-spread is narrow compared with that of many dragon-flies and moths. On the other hand, there are thousands of very small species, and the tiny “fairy-flies” (Mymaridae), whose larvae live as parasites in the eggs of various insects, are
