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Natural History and Behavior of a Primitively Social Wasp, Auplopus Semialatus, and Its Parasite, Irenangelus Eberhardi (Hymenoptera: Pompilidae)

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Natural History and Behavior of a Primitively Social Wasp, Auplopus Semialatus, and Its Parasite, Irenangelus Eberhardi (Hymenoptera: Pompilidae) Journal of Insect Behavior, Vol. 1, No. 3, 1988 Natural History and Behavior of a Primitively Social Wasp, Auplopus semialatus, and Its Parasite, Irenangelus eberhardi (Hymenoptera: Pompilidae) William T. Wcislo,1 Mary Jane West-Eberhard,2 and William G. Eberhard2'3 XccgpW 74 Xpnf 7987, Mvu«f 74 Ju/y 7987 Two to eight females of a neotropical, primitively social wasp, Auplopus sem- ialatus (Pompilidae), cooperatively build and maintain mud nests. Females cap- ture non-web-building spiders as provisions for their offspring. Cohabiting females are usually tolerant of one another and defend the nest against natural enemies, including the cleptoparasitic wasp, Irenangelus eberhardi (Pompili- dae). They often become intensely competitive, however, when a spider is brought to the nest. Auplopus females steal spiders from both uncapped and newly capped cells and eat the previous owner's egg. Many observations high- light the primitive level of sociality in this species, and the discussion relates these observations to those based on other primitively social wasps. KEY WORDS: Auplopus semialatus; Irenangelus eberhardi; Pompilidae, social behavior; clep- toparasitism. INTRODUCTION Spider wasps (Pompilidae) usually lead solitary lives, although some species are obligate cleptoparasites that lay their eggs in the nests of other pompilids (e.g., see Olberg, 1959; Wcislo, 1987). Occasionally two or more conspecific females are found on the same nest (Williams, 1919, Iwata, 1976; Kimsey, 1980). In only one species, however, have the interactions of cohabiting females been recorded. In his study of Philippine wasps Williams (1919) found that 'Department of Entomology, Snow Hall, University of Kansas, Lawrence, Kansas 66045. ^Smithsonian Tropical Research Institute, Balboa, Panama. 3Escuela de Biologia, Universidad de Costa Rica, Ciudad Universitaria, Costa Rica. 247 0892-7553/88/0700-0247$06.0O/0 © 1988 Plenum Publishing Corporation 248 Wcislo, West-Eberhard, and Eberhard nest-sharing females of Paragenia argentifrons were ' 'never aggressive'' to one another during their brief social interactions. This paper reports on another trop- ical nest-sharing ("primitively social") pompilid, Auplopus semialatus Dreis- bach (Pepsinae, Auplopodini). In this species cohabiting females vigorously contested both the use of empty brood cells and captured prey via egg canni- balism and overt fighting. These observations, although brief, provide dramatic illustrations of both the disadvantages and the benefits of social nesting. Notes on a cleptoparasite, Irenangelus eberhardi Evans [Pompilidae, Ceropalinae (Evans, 1987)], are also given. METHODS Field observations were made from 10 to 12 March 1984 on nest A (17 h of observation), from 2 to 8 April 1986 on nest B (42 h), and from 6 to 8 June 1986 on nest C (3 h) at Finca La Selva (elevation, 50 m), near Puerto Viejo, Heredia Province, Costa Rica. Observations on nest B commenced each day before first light (5:00 AM) and continued until after sunset (6:30 PM). Nest A had 16 cells and 4 adult females, nest B had 19 cells with 8 adult females and 1 male, and nest C had approximately 20 cells with at least 5 adult females. All observed females from nests A and B were dissected (M.J.W.E.), but some of the unusually small spermathecae (about 0.02 mm in diameter) were not found. Wasps and parasites were allowed to emerge in captivity near San Jose (elevation, 1300 m), where temperatures are somewhat cooler than at La Selva. Means are given with standard deviations. RESULTS Nest Architecture. Nests were groups of cojoined cylindrical mud cells usu- ally attached to the long spines on the trunks of pejibaye palm trees (Bactris gasipaes) that grew in an otherwise open clearing. Nests were separated from the substrate (trunk or leaf) by about 1 cm and were always on the undersides of the trunks, which were not perfectly vertical. Active nests found in 1986 had 2, 3, 8, 15, and 21 complete or partially complete cells (the nest with three cells was attached to a leaf and disappeared after a night of heavy rain). The number of cells per nest for 24 abandoned nests found in April 1986 ranged from less than 1 completed cell to 20 cells (X = 4.2 + 2.99) (Fig. 1). These cell counts are underestimates since some cells had probably weathered away. A large fraction of nests probably fails early, in the one- to five-cell stage. Cooperative Cell Construction, Renovation, and Reuse. Construction of new brood cells, and the reconditioning of old cells for reuse, was usually coop- erative in that more than one individual participated. Construction of a new cell was seen only on nests A and C. The cell on Behavior of Auplopus semialatus 249 in <v Z5 i_ <u X! E z UJLLLLJLIJL 5 10 15 20 Number of Cells Fig. 1. Histogram of the number of cells per nest for aban- doned nests of Auplopus semialatus (N = 24 nests). nest A required about 2 wasp-days of work for completion, while the cell on nest C took just over 0.5 wasp-day to build. Each load of mud used in cell construction was visible as a small bump on the outside of the cell [as in illus- trations of other Auplopus nests; see, e.g., Fig. 44 of Williams (1919)], and each cell had about 100-150 such bumps. These bumps were covered by an overlay of smooth mud, which in older nests obscured the outlines of the cells themselves. Females on nests A and C often collected mud away from the nest on trips lasting from 1 to 3 min, while females on nest B used mud only from the nest itself. Collecting trips for water were briefer than for mud, usually lasting less than 30 s. On all three nests females frequently quarried mud from one area of the nest and applied it to another. The wasp first gathered water and then scraped dried mud from the nest surface with her mandibles and worked it into a soft ball by adding fluid (presumably water). The mud ball was spread by bending the gaster forward beneath the thorax and troweling the mud rapidly with the terminal gastral tergite, as described by Williams (1919) and Rau (1928) for other Auplopodini. Females shifted mud on the nest more frequently when a cell was being closed, but sometimes the mud shifting seemed to be function- less. For example, female 2A once shifted mud 14 times in sequence: she removed mud from a newly capped cell 5 times, while 7 times she applied mud to the same cap. At least two wasps emerged from cells on nest B during field observations (see below), and in both cases these cells, as well as others vacated on the nest, were reused after being cleaned and reconditioned. As noted above, the recon- ditioning of these cells involved cooperative work by more than one wasp. There were no vacated cells on nests A and C during our field observations. A behavior possibly associated with nest construction but one whose func- tion we do not understand, involved females biting or rasping with their man- dibles at the spines on the tree trunk or at lichens or debris on the trunk. Females frequently did this, but we could rarely discern that any materials had been gathered, and it did not appear that they applied the materials to the nest. Rau 250 Wcislo, West-Eberhard, and Eberhard (1928), however, observed similar behavior in Pseuadegenia [= Auplopus (see Evans, 1973)] mellipes and was able to see that bits of debris (spiders' webs, bits of wood, etc.) were gathered and applied to the nest. Provisioning, Oviposition, and Cell Closure. Eleven spiders from at least five non-web-building families (Salticidae, Clubionidae, Heteropidae, Any- phaenidae, and Ctenidae) were brought to the nest by wasps. Australian auplo- podines observed by Evans and Matthews (1973) also preyed only on non-orb- weaving spiders. In our observations, the earliest prey was brought to the nest before sunrise at 5:30 AM and the latest after sunset at 6:10 PM. Wasps usually amputated most or all of the spiders' legs before returning to the nest. Wasps carried their prey as described for Auplopus and other auplopodine genera (Wil- liams, 1919; Evans and West-Eberhard, 1970). A provisioning wasp usually did not return directly to the nest but, instead, landed on the tree trunk or on a nearby leaf of a vine (within 30 cm of the nest). Here it paused to suck hemolymph from the stubs of the spider's amputated legs, or imbibed fluids from its mouth, and then waited motionless for several seconds to 5 min. Wasps also often groomed the surface of the spider's abdo- men with chewing movements of the mandibles but never broke the exoskele- ton. Such grooming may serve to remove external parasites or conspecific eggs, as speculated by Richards and Hamm (1939). The wasp then walked onto the nest with her prey, where she repeated the behaviors just described, interacted aggressively with nestmates, or moved directly to an open cell and deposited the spider inside. To deposit a spider, the wasp backed into the cell until most of her abdomen and metathoracic legs were inside the cell. She then passed the spider under her body into the cell as she climbed out and turned to stuff the spider in abdomen-first by pushing it with her head. Unless another wasp stole her spider (see below), the female even- tually oviposited (after waiting from several seconds to 12 h). A female ovi- posited by inserting her abdomen deep into the cell so that only her head, her prothoracic legs, and the anterior part of the thorax were out of the cell. She remained still for about half a minute (X = 25.8 + 4.5 s; N = 12).
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