Benefits and costs of earwig (Forficula auricularia) family life

The evolution of parental care and family group formation critically depends on offspring survival benefits and parental fecundity costs of care under given ecological conditions. Investigations of the functional significance of care in insect species that exhibit facultative parental care have been relatively rare but may be of particular interest for better understanding of benefit and cost schedules at an early evolutionary stage. In this study, aspects of benefits and costs of care were addressed in the sub-social European earwig (Forficula auricularia; Dermaptera: Forficulidae) by manipulating the presence of tending mothers and brood size in a fully crossed experimental design. Larvae growing in broods tended by their mother or of reduced size showed a higher survival probability than larvae growing in untended or large broods, as predicted if maternal care is beneficial and shaped by a trade-off between number and quality of offspring. Analysis of patterns of food consumption and developmental time further suggested that the benefit of maternal attendance is mediated by the maternal provisioning of food, while the quality–quantity trade-off seemed to be driven by sibling rivalry. Further, tending mothers delayed the production of a second clutch, indicating a potential cost of care in terms of lifetime fecundity. This study experimentally shows benefits and potential costs of maternal care and family group formation in the European earwig. More detailed behavioural experiments will be required to fully understand how behavioural interactions among family members mediate these reproductive outcomes.     

Extra-pair paternity and the evolution of testis size in a behaviorally monogamous tropical mammal,the large treeshrew (<Emphasis Type="Italic">Tupaia tana</Emphasis>)

Monogamy is rare in mammals (<5% spp.) but occurs in greater frequency among primates (15%) and their close relatives, the treeshrews (100%; Order: Scandentia). Two genetic studies of parentage in monogamous primates revealed high rates of extra-pair paternity (EPP), but to date parentage has not been studied in a treeshrew species. I analyzed the genetic parentage of 22 offspring from two populations of large treeshrews in Sabah, Malaysia (NE Borneo), using seven autosomal microsatellite loci and one mitochondrial DNA marker. Half of these offspring were sired by males that were not the presumed partner of the mother (50% EPP), and three litters exhibited evidence of multiple paternity. However, comparative analysis indicated that the high rate of EPP in Tupaia tana is not associated with intense sperm competition. Relative testis size of treeshrews was similar to testis size in 22 primate species with uni-male mating systems but smaller than 44 primates with multi-male mating systems. After factoring out the effects of body size and phylogeny, I also found that the evolution of multi-male mating systems was significantly associated with the evolution of larger testis size. Male–female pairs of T. tana occupy joint territories but forage and sleep alone (“dispersed pair-living”), and I argue that this form of behavioral monogamy renders mate guarding ineffective. The adaptive advantages of behavioral monogamy likely differ from the advantages driving EPP in large treeshrews. However, small testis size suggests that behavioral monogamy is not masking a dispersed multi-male mating system in this species.     

黄泥土微团聚体颗粒组对Cu2+的吸附与解吸研究

采用低能量超声波分散和冷冻机干燥法提取太湖地区黄泥土不同粒径微团聚体颗粒组样品,用平衡液吸附法和CaCl2与HCl溶液的连续解吸法研究原土和不同粒径微团聚体颗粒组对重金属Cu2 的吸附和解吸特征。结果表明,对于Cu2 的吸附,原土用Langmuir方程拟合最佳,而不同粒径微团聚体颗粒组用Freundlich方程拟合最佳。原土和不同粒径微团聚体颗粒组吸附量大小顺序为粘粒级>砂粒级>原土>粉砂级>粗粉砂级,这与其游离氧化铁和有机质含量呈显著正相关。原土和不同粒径微团聚体颗粒组对Cu2 的专性吸附质量分数随吸附量增加而减少,而非专性吸附质量分数则相反。不同粒径微团聚体颗粒组吸附总量小于原土,所以传统风干磨细的研究方法可能高估了土壤对铜固持能力。     

Constraints on foraging success due to resource ecology limit colony productivity in social insects

The benefit of group living is a fundamental question in social evolution. For sociality to evolve, each individual must gain in terms of some fitness component by living in larger groups. However, in social insects, a decrease in per capita success in brood production has been observed in larger groups. While it has been proposed that this decrease could be outweighed by an increase in the predictability of success, a functional basis to this hypothesis has so far never been demonstrated. In this paper, using foraging economics as a functional proxy to colony productivity, we construct a model to explore how number of foragers in the colony interacts with the ecology of resources to influence per capita foraging success and its predictability. The results of the model show that there is no increase in per capita foraging success in larger colonies under most circumstances, though there is an increase in its predictability. We then test the model with empirical data on the foraging behavior of the primitively eusocial wasp, Ropalidia marginata. The consistency between the data and the model suggests that foraging economics could provide a robust functional basis in explaining the relationship between colony size and productivity.     

Selection on social traits in greater spear-nosed bats, Phyllostomus hastatus

Many studies assume that selection molds social traits and have investigated the manner in which this occurs, yet very few studies have measured the strength of selection on social traits in natural populations. In this paper, I report results of phenotypic selection analyses on two social traits – the size of social groups and the frequency of group foraging – in Phyllostomus hastatus, the greater spear-nosed bat. I found significant positive directional selection on individual group foraging frequency, but no directional selection on individuals in different-sized social groups. These results have implications for the structure of social groups, cooperative behavior among group mates, and maternal investment strategies. I argue that combining studies of natural selection on wild populations with experiments to identify the agents and mechanisms of selection can do much to increase our understanding of social evolution.     

Emergence of increased division of labor as a function of group size

Empirical evidence suggests that division of labor in insect societies is positively related to group size both within and across taxa. Response threshold models (RTM) have been commonly used to analyze patterns of division of labor. However, these models have been explored empirically and theoretically for only a limited number of tasks, and few studies have examined predictions of the model as colony size and work availability change. We theoretically examine how group size influences division of labor using a fixed response-threshold model. We simultaneously explore how expected by-products of increased colony size, including demand (total work need relative to total work force available) and task number, affect this relationship. Our results indicate that both low demand and high task number positively influence division of labor. We suggest that these changes parallel what is observed within social groups as their size increases, and that, in part, the commonly observed increased division of labor with increasing group size is emergent.     

Size-dependent mating and gender choice in a simultaneous hermaphrodite, Bulla gouldiana

Simultaneous hermaphrodites are predicted to optimally divide resources between male and female function, which can result in both size-dependent mating behaviors and conflict between potential mates. Predicted strategies include size-assortative mating, conditional exchange of gametes, and mating patterns where relative size affects investment in each sexual role. This study investigated the effect of body size on the mating strategies of a hermaphroditic opisthobranch, Bulla gouldiana. Although individuals were spatially aggregated in the field with high levels of movement and size variation, there was little evidence for predictions. Laboratory experiments, however, revealed complicated effects of mass on the probability and duration of mating, as well as gender choice. Pairs were more likely to mate if they included at least one large animal, with the larger animal typically inseminating the smaller. When both individuals were large, they were more likely to each mate in both sexual roles by switching roles once. Although B. gouldiana did not usually alternate between sexual roles multiple times within mating events, paired individuals behaved similarly (neither or both mating as sperm donors) more often than expected by chance. This suggests some level of reciprocity, which is unlikely to be conditional given rates of unilateral mating. When the larger member of the mating pair inseminated the smaller, the duration of insemination increased with the size of the smaller sperm recipient. Copulations lasted longer in pairs that switched sexual roles than in those that did not switch roles. This study suggests that variation in body size can lead to size-dependent mating patterns, but only some of the patterns in B. gouldiana support theoretical predictions. We review other studies that have addressed similar issues, providing inconsistent mating patterns in sperm-storing hermaphrodites.     

Sex-specific energy requirements in nestlings of an extremely sexually size dimorphic bird, the European sparrowhawk (Accipiter nisus)

Allocation of parental investment is predicted to be equal at the population level between both sexes of offspring, and should lead to sex ratio biases in species that exhibit a sex-difference in parental care. Sex-differences in parental care are rarely quantified. We measured daily energy expenditure in free-living nestlings of the extremely sexually size dimorphic European sparrowhawk (Accipiter nisus), using the doubly labelled water method. These data were combined with measured growth characteristics to estimate daily and total metabolised energy intake of male and female young during the nestling stage. Females reached an asymptotic body mass 1.6 times higher than males. This resulted in a total metabolised energy an estimated 1.4 times higher for the nestling stage. Furthermore, we observed a decline in daily metabolised energy with an increase in brood size, which was significantly stronger for females than for males. These results are discussed in the context of Fishers equal allocation theory. Empirical evidence of a sex ratio bias at the end of parental care, with an overall excess of males, is lacking in this species. Consequently, our data do not support the idea of equal allocation between the sexes. The observed sex difference in daily metabolised energy in response to brood size may give scope for sex ratio bias at the level of the individual brood.     

Soil Lead and Children’s Blood Lead Levels in Syracuse, NY, USA

A geo-referenced data set of 12 228 first-time blood lead screening values for Syracuse, NY, children was established for the 4-year period 1992–1996. Soil lead values were measured in a 600 m by 600 m tessellation grid covering the city. The two data sets were merged for evaluation of relationships between them. Strong seasonal variation in blood lead levels suggests the importance of contaminated soils as an exposure source. When the data were aggregated at a large geographic scale (3 km²), a logarithmic model showed an R² of > 0.65 for the regression of geometric mean blood lead on median soil lead values. Results showed a striking similarity to those obtained by Mielke et al. (1999) for a study in New Orleans, LA, USA.     

Fitness costs of an immune response in the house martin (<Emphasis Type="Italic">Delichon urbica</Emphasis>)

Immune responses constitute a major way for hosts to defend themselves against parasites. Because hosts do not habitually produce strong responses all the time, immune responses might be costly to produce or maintain. We tested experimentally if the production of a response to a challenge with a novel antigen resulted in a cost in terms of fitness using the highly colonial house martin Delichon urbica as a model system. We injected adult breeding birds during laying of the first clutch with either Newcastle disease virus (NDV) or a control injection, and the clutch was subsequently removed to induce relaying. NDV stimulates the non-specific immune system, causing production of antibodies during a period of more than 2 weeks. Accordingly, we found an increase in leukocyte counts in experimental birds compared to controls. Experimental treatment reduced the frequency of re-laying, caused a delay in timing of relaying and a reduction in brood size. Quality of nestlings in terms of body size, body mass and T-cell-mediated immune response did not differ significantly between treatments. Overall, seasonal reproductive success differed significantly between treatments, showing that the production of an immune response by adult birds is costly in terms of future fecundity.