|Origin and evolution of social behavior: case study on genus Ceratina (Hymenoptera: Apidae)|
Small carpenter bees of genus Ceratina are very suitable models for research of the origin and evolution of eusociality. Most of Ceratina species are solitary (Michener, 2007) and some are socially polymorphic (Sakagami & Maeta, 1989; Rehan et al., 2009). Socially polymorphic species belong to different unrelated clades of genus Ceratina (Rehan et al., 2009; Rehan et al., 2010). Social behavior therefore either must have arisen several times independently, or it represents original state for the whole genus and solitarity arised secondarily.
Females of small carpenter bees of genus Ceratina nests in twigs with soft pith (Michener, 2007; Sakagami & Laroca, 1971). Bees firstly excavate burrow. Afterwards they provision cell by pollen and nectar. When they complete the provisioning of the cell, they oviposit and close the cell by septum. (Maeta et al., 1997). Bees usually provision about eight cells per life. After provisioning and oviposition females stay upstairs in burrow and guard offsprings against predators and parasites (Sakagami & Maeta, 1977; Rehan & Richards, 2010). This behavior is quite unique, because in most solitary species female provisions most of her life and the nest is unprotected from attack of predators and parasites most of the time. Parazitation can markedly reduce number of bee´s offspring (Vickruck et al., 2010). The failure of the whole nest because of usurpation by other Hymenoptera or Salticidae spiders, or predation of brood by Dermaptera (unpublished results) is very common.
Marked female defending her nest by own abdomen and Salticidae spider usurping nest of Ceratina
If female is in the burrow after the completing of the cells and she blocks the entrance by her abdomen, predators and parasites can´t entry nest. Similarly social nesting significantly decreases probability of parazitation and total failure of nest, because nest is always protected by at least one female (Rehan et al., 2011). After new adults are hatched, mother feed them by pollen and nectar (Sakagami & Maeta, 1977). In some species young adults are fed not by mother, but by sisters (Sakagami et al., 1989). Females of some species are able to reuse nest. This ability is probably very important for social nesting, because most of nests inhabited simultaneously by more than one female are nests reused (Sakagami & Maeta, 1984, 1989; Rehan et al., 2010). Temperate species usually leave the nest and make new burrows for overwintering. (Sakagami & Laroca, 1971), but sometimes overwinter in natal nests (Rehan & Richards, 2010).
Nest of Ceratina cucurbitina - on the left are fresh new adults, in the middle pupae and on the right larva and old adult female
Nest od C. chalybea with provisioned and epmty cells
In some species more individuals are in some nests (Sakagami & Maeta, 1977; Rehan et al., 2010). Wilson (1971) stated three features of eusociality: reproductive division of labor, cooperative brood care and generation overlap. Published studies (Sakagami & Maeta, 1977, 1989; Rehan et al., 2010, 2011) indicate that eusociality occurs in some species of genus Ceratina. Females of social nests have usually different size of ovary (Rehan et al., 2010; Maeta & Sakagami, 1995). There are probably differences in reproductive success between females in some nest. There is probably overlap of generation, because females in some nest have distinctive differences in wing and mandible wear (Sakagami & Maeta, 1989). For quantification of proportion different types of social organization (eusocial, semisocial and quasisocial) are essential high resolution molecular marker. Azuma et. al. (2005) developed microsatellites for species Ceratina flavipes, but they are not sufficiently variable for assess relationship between individuals.
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