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developed in many of the higher social wasps. The mother wasp attends to several young simultaneously, and when their development is completed seals up the cells. S. micans var luzonensis (Fig. 30 B) makes the most remarkable nest of all. It is attached to some pendent plant filament under an overhanging bank or under masses of dead leaves supported by twigs or vines and is made of "moist and well-decayed wood chewed up into a pulp and formed into delicate paper which is not rain proof." The basal portion of the nest (Fig. 30 C) is a single comb of about 20 regular, hexagonal cells, enclosed in a pear-shaped covering which is longitudinally grooved and ribbed on the outside and constricted below to form a filigree-work, funnel-like aperture surrounded on one side by a spear-shaped expansion. This species seems also to have been observed in Ceylon by E. E. Green, who remarks that "the nest seems to be the property of one pair only" of wasps. Two other species, S. nigrifrons of Burma and melleyi of Java, are also recorded as social. They make nests consisting of a few pendent, hexagonal-celled combs attached to one another by slender pedicels. All of the descriptions indicate that the colonies of the social species of Stenogaster must consist of very few individuals, and there is nothing to show that the female offspring

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Nest of Polybioides tabida from the Congo, with the involucre partly removed. (After J. Bequaert from a photograph by H. O. Lang).

differ in any way from their mother or that they assist in caring for the brood. Even in the case of S. varipictus, Williams remarks: "In a small way, it seems to be a social wasp; one to several insects

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A. Nest of Polybioides melaena of the Congo. B. The same partly destroyed, showing the pendent combs, which have cells on both sides.

Bequaert, from a photograph by H. O. Lang).

(After J.

attend to a cell group. It may be, however, that each female has her own lot of cells in this cell group." Future investigations may show that none of the species of Stenogaster is really social. in the same sense as are the four other subfamilies, though they approach the definitively social forms in using paper in the construction of the nest, in sometimes making combs of regular hexagonal cells and in caring for a number of larvæ at the same time.

(2). The Epiponinæ are a large and heterogeneous group, comprising a much greater number of genera (23) than any other subfamily of social wasps, and ranging all the way from very primitive forms like Belonogaster to highly specialized forms like Chartergus and Nectarina. Great differences are also apparent in the architecture of the nest, which in the more primitive genera consists of a single naked comb of hexagonal cells attached to some support by a peduncle (Fig. 31), and in the more advanced forms of a single comb or of several combs superimposed on one another and enclosed in an envelope with an opening for ingress and egress (Figs. 32 and 33). The combs are in some cases pedunculate (stelocyttarous), in others attached directly to the support or to the envelope (phragmocyttarous). In nearly all cases the nest is made entirely of paper, but in a few tropical American species. some clay may be added. It is always above ground and attached to the branches or leaves of trees, to the underside of some shelter (roofs, banks, etc.). In primitive forms like Belonogaster (Fig. 31), as a rule, a single fecundated female starts the nest by building a single pedunculate cell and then gradually adding others in circles concentrically to its periphery as the comb grows, but not infrequently the foundress may be joined by other females before the work has progressed very far. Each larva is fed with pellets of malaxated caterpillars till it is full grown when it spins a convex cap over the orifice of its cell and pupates. The emerging females are all like the mother in possessing well-developed ovaries and in being capable of fecundation. In other words, all the females of the colony are physiologically equal, and even such differences in stature as they may exhibit have no relation to fertility. The colonies are small, the nests having usually only about 50 to 60 cells, rarely as many as 200 to 300. In larger colonies there is a certain rude division of labor since the older females devote themselves to egg-laying, the younger to foraging for food and nest materials and the recently emerged individuals to feeding the larvæ and caring for the nest. The males, too, remain on the comb, but behave like parasites and exact food whenever it is brought in by the foraging females. Belonogaster is described as a polygynous wasp because each of its colonies contains a number

of fecundated females. When it has reached its full development the females leave in small companies and found new nests either singly or together. This phenomenon is known as "swarming" and occurs only in the wasps of the tropics where it seems to be an adaptation to the favorable climatic conditions. In the higher South American genera of Epiponine, however, the females are not all alike but are differentiated into true females, or queens, i. e., individuals with well-developed ovaries and capable of fecundation, and workers, i. e., females with imperfectly developed ovaries and therefore sterile or capable only of laying unfertilized, male-producing eggs. Many of these wasps, according to H. and R. von Ihering and Ducke, are polygynous and regularly form new colonies and nests by sending off swarms of workers with one or two dozen queens. The colonies often become extremely populous and comprise hundreds or even thousands of individuals. Some of the species (Nectarina, Polybia) have a habit of storing a considerable amount of honey in their combs, while others are known to capture, kill and store within the nest envelope, and even in the combs, quantities of male and female termites or male ants as a supply of food to be drawn on when needed.

(3). The Ropalidiinæ are a small group of only three genera, the best known of which is Ropalidia. These are primitive wasps which build a single naked comb like that of Belonogaster and feed their young with pellets of malaxated insects. The colonies. are small and polygynous, but, according to Roubaud, true workers can be distinguished, though they are few in number compared with the true females. Swarming seems to occur in some species.

(4). The Polistinæ are represented by only two genera. One of these, Polistes, is cosmopolitan and, like Ropalidia and Belonogaster, makes a single, naked comb, suspended by a central or ecentric peduncle to the underside of some shelter. As there are several common species in Europe and the United States, the habits of the genus are well known. The nest is usually established and in its incipient stages constructed by a single female, or queen. A certain number of her offspring are workers though they seem often to lay male-producing eggs. True females are rather numerous in the colonies of some species, which may therefore be regarded as polygynous, and some of the tropical forms may, perhaps, swarm. In temperate regions, however, the Polistes colony is an annual development and usually not very populous. The young females are fecundated in the late summer and pass the winter hidden away under bark or in the crevices of walls, whence they emerge in the spring to found new colonies. Several of the species, even in temperate regions, are known to store small quantities of honey in their combs.

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THE SCIENTIFIC MONTHLY

(5). Like the Polistinæ, the subfamily Vespinæ includes only two genera, Vespa and Provespa. The species of the former, the only genus besides Polistes that occurs in the north temperate zone, are the largest and most typical of social wasps. So far as known the species are strictly monogynous. The nest, founded by a single female, consists at first of a small pendent comb, like that of Polistes, but while there are still only a few cells a more or less spherical envelope is built around it. The eggs first laid produce workers, which are much smaller than the mother and incapable of fecundation. They remain with the parent, enlarge the comb and envelope and, to accommodate the rapidly increasing brood, build additional combs in a series from above downward, each new comb being supported by one or more peduncles attached to the comb above it (stelocyttarous). At first large numbers of workers are produced, but later in the summer males and females appear. Owing to the greater size of the females, the cells in which they are reared are considerably larger than the worker cells. After the mating of the males and females the colony perishes, with the exception of the fecundated females, which hibernate like the females of Polistes and during the following spring found new colonies. In the Vespinæ, therefore, a very distinct worker caste has been developed, though its members occasionally and perhaps regularly lay male-producing eggs. The species of Vespa are usually divided into two groups, one with long, the other with very short cheeks. In Europe and North America the long-cheeked forms as a rule build aerial nests above ground, the short-cheeked forms in cavities which they excavate in the ground. The colonies may often be very populous by the end of the summer (3,000 to 5,000 individuals).

After this hasty sketch of the five subfamilies of the social wasps we may consider a few of their fundamental behavioristic peculiarities, especially the trophic relations between the adults and larvæ, the origin of the worker caste, its ultimate fate in certain parasitic species and the question of monogyny and polygyny. In all these phenomena we are concerned with effects of the foodsupply and therefore of the external environment.

The feeding of the larvæ by Vespa and Polistes queens and workers with pellets made of malaxated portions of caterpillars, flies or other insects has often been described and can be readily witnessed in any colony kept in the laboratory. The hungry larvæ protrude their heads with open mouths from the orifices of the cells, like so many nestling birds, and when very hungry may actually scratch on the walls of the cells to attract the attention of the workers or their nurses. The feeding is not, however, a one-sided

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