|dc.description.abstract||Many arthropods exhibit behaviours precursory to social life, including adult
longevity, parental care, nest loyalty and mutual tolerance, yet there are few examples of
social behaviour in this phylum. The small carpenter bees, genus Ceratina, provide
important insights into the early stages of sociality. I described the biology and social
behaviour of five facultatively social species which exhibit all of the preadaptations for
successful group living, yet present ecological and behavioural characteristics that seemingly
disfavour frequent colony formation. These species are socially polymorphic with both
solitary and social nests collected in sympatry. Social colonies consist of two adult females,
one contributing both foraging and reproductive effort and the second which remains at the
nest as a passive guard. Cooperative nesting provides no overt reproductive benefits over
solitary nesting, although brood survival tends to be greater in social colonies.
Three main theories explain cooperation among conspecifics: mutual benefit, kin
selection and manipulation. Lifetime reproductive success calculations revealed that mutual
benefit does not explain social behaviour in this group as social colonies have lower per
capita life time reproductive success than solitary nests. Genetic pedigrees constructed from
allozyme data indicate that kin selection might contribute to the maintenance of social nesting
as social colonies consist of full sisters and thus some indirect fitness benefits are inherently
bestowed on subordinate females as a result of remaining to help their dominant sister. These
data suggest that the origin of sociality in ceratinines has principal costs and the great
ecological success of highly eusociallineages occurred well after social origins.
Ecological constraints such as resource limitation, unfavourable weather conditions
and parasite pressure have long been considered some of the most important selective
pressures for the evolution of sociality. I assessed the fitness consequences of these three ecological factors for reproductive success of solitary and social colonies and found that nest
sites were not limiting, and the frequency of social nesting was consistent across brood
rearing seasons. Local weather varied between seasons but was not correlated with
reproductive success. Severe parasitism resulted in low reproductive success and total nest
failure in solitary nests. Social colonies had higher reproductive success and were never
extirpated by parasites. I suggest that social nesting represents a form of bet-hedging. The
high frequency of solitary nests suggests that this is the optimal strategy when parasite
pressure is low. However, social colonies have a selective advantage over solitary nesting
females during periods of extreme parasite pressure.
Finally, the small carpenter bees are recorded from all continents except Antarctica. I
constructed the first molecular phylogeny of ceratinine bees based on four gene regions of
selected species covering representatives from all continents and ecological regions.
Maximum parsimony and Bayesian Inference tree topology and fossil dating support an
African origin followed by an Old World invasion and New World radiation. All known Old
World ceratinines form social colonies while New World species are largely solitary; thus
geography and phylogenetic inertia are likely predictors of social evolution in this genus.
This integrative approach not only describes the behaviour of several previously
unknown or little-known Ceratina species, bu~ highlights the fact that this is an important,
though previously unrecognized, model for studying evolutionary transitions from solitary to