Evolutionary Origin and Maintenance of Sociality in the Small Carpenter Bees

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 social behaviour.

Soil-Transmitted Helminths, Poverty, and Malnutrition in Honduran Children Living in Remote Rural Communities

Soil-transmitted helminth (STH) infections are endemic in Honduras, but their prevalence according to the levels of poverty in the population has not been examined. The present cross-sectional study is aimed to determine the role of different levels of poverty in STH prevalence and infection intensity as well as the potential associations of STH infections with malnutrition and anemia. Research participants were children attending a medical brigade serving remote communities in Northern Honduras in June 2014. Demographic data were obtained, and poverty levels were determined using the unsatisfied basic needs method. STH infections were investigated by the Kato-Katz method; hemoglobin concentrations were determined with the HemoCue system; and stunting, thinness, and underweight were determined by anthropometry. Data were analyzed using descriptive statistics and univariate and multivariable logistic regression models. Among 130 children who participated in this study, a high prevalence (69.2%) of parasitism was found and the poorest children were significantly more infected than those living in less poor communities (79.6% vs. 61.8%; P = 0.030). Prevalence rates of Trichuris trichiura, Ascaris lumbricoides, and hookworms were 69.2%, 12.3%, and 3.85%, respectively. In total, 69% of children had anemia and 30% were stunted. Households’ earthen floor and lack of latrines were associated with infection. Greater efforts should be made to reduce STH prevalence and improve overall childhood health, in particular, among the poorest children lacking the basic necessities of life.