Support for maternal manipulation of developmental nutrition in a facultatively eusocial bee, <Emphasis Type="Italic">Megalopta genalis</Emphasis> (Halictidae)

Developmental maternal effects are a potentially important source of phenotypic variation, but they can be difficult to distinguish from other environmental factors. This is an important distinction within the context of social evolution, because if variation in offspring helping behavior is due to maternal manipulation, social selection may act on maternal phenotypes, as well as those of offspring. Factors correlated with social castes have been linked to variation in developmental nutrition, which might provide opportunity for females to manipulate the social behavior of their offspring. Megalopta genalis is a mass-provisioning facultatively eusocial sweat bee for which production of males and females in social and solitary nests is concurrent and asynchronous. Female offspring may become either gynes (reproductive dispersers) or workers (non-reproductive helpers). We predicted that if maternal manipulation plays a role in M. genalis caste determination, investment in daughters should vary more than for sons. The mass and protein content of pollen stores provided to female offspring varied significantly more than those of males, but volume and sugar content did not. Sugar content varied more among female eggs in social nests than in solitary nests. Provisions were larger, with higher nutrient content, for female eggs and in social nests. Adult females and males show different patterns of allometry, and their investment ratio ranged from 1.23 to 1.69. Adult body weight varied more for females than males, possibly reflecting increased variation in maternal investment in female offspring. These differences are consistent with a role for maternal manipulation in the social plasticity observed in M. genalis.     

Survival and productivity benefits to social nesting in the sweat bee <Emphasis Type="Italic">Megalopta genalis</Emphasis> (Hymenoptera: Halictidae)

Facultatively solitary and eusocial species allow for direct tests of the benefits of group living. We used the facultatively social sweat bee Megalopta genalis to test several benefits of group living. We surveyed natural nests modified for observation in the field weekly for 5 weeks in 2003. First, we demonstrate that social and solitary nesting are alternative behaviors, rather than different points on one developmental trajectory. Next, we show that solitary nests suffered significantly higher rates of nest failure than did social nests. Nest failure apparently resulted from solitary foundress mortality and subsequent brood orphanage. Social nests had significantly higher productivity, measured as new brood cells provisioned during the study, than did solitary nests. After accounting for nest failures, per capita productivity did not change with group size. Our results support key predictions of Assured Fitness Return models, suggesting such indirect fitness benefits favor eusocial nesting in M. genalis. We compared field collections of natural nests to our observation nest data to show that without accounting for nest failures, M. genalis appear to suffer a per capita productivity decrease with increasing group size. Calculating per capita productivity from collected nests without accounting for the differential probabilities of survival across group sizes leads to an overestimate of solitary nest productivity.     

Solitary behavior in a high-altitude population of the social sweat bee Halictus rubicundus (Hymenoptera: Halictidae)

In the subalpine region of the Rocky Mountains of Colorado, United States, Halictus rubicundus has a solitary life cycle, but it is social in other parts of its known range. The brood is protandrous, with a nearly equal investment in the sexes. Productivity averages 6.5 offspring per foundress female, similar to the second brood of social nests in New York, but less than the combined productivity of both New York broods. Leucophora sp. (Diptera: Anthomyiidae) is the principal cause of brood mortality in Colorado. Foundress females in about half the nests survive until brood emerge as adults. Retention of these foundresses decreases offspring mortality by 68%. Comparable abilities to express solitary behavior with a single brood may characterize other eusocial halictine lineages that have successfully invaded high altitudes in the Rocky Mountains. The apparent inability to do this may help explain the absence of other eusocial halictine bees and polistine wasps at high altitudes, despite their success at lower elevations in the same mountains. Presence or absence of this ability may help explain latitudinal distributions of these lineages in North America. Holarctic distributions of lineages with eusocial behavior can be explained by migration as solitary populations from Eurasia to North America across Pleistocene Bering land bridges, with re-expression of double-brooded, eusocial behavior when the species then extended their ranges southward in North America. Received: 4 November 1994/Accepted after revision: 23 October 1995     

Foundress polyphenism and the origins of eusociality in a facultatively eusocial sweat bee, Megalopta genalis (Halictidae)

The reproductive (queen) and nonreproductive (worker) castes of eusocial insect colonies are a classic example of insect polyphenism. A complementary polyphenism may also exist entirely among females in the reproductive caste. Although less studied, reproductive females may vary in behavior based on size-associated attributes leading to the production of daughter workers. We studied a bee with flexible social behavior, Megalopta genalis, to better understand the potential of this polyphenism to shape the social organization of bee colonies and, by extension, its role in the evolution of eusociality. Our experimental design reduced variation among nest foundresses in life history variables that could influence reproductive decisions, such as nesting quality and early adulthood experience. Within our study population, approximately one third of M. genalis nests were eusocial and the remaining nests never produced workers. Though they do not differ in survival, nest-founding females who do not attempt to produce workers (which we refer to as the solitary phenotype) are significantly smaller and become reproductive later than females who attempt to recruit workers (the social phenotype). Females with the social phenotype are more likely to produce additional broods but at a cost of having some of their first offspring become nonreproductive workers. The likelihood of eusocial organization varies with body size across females of the social phenotype. Thus, fitness consequences associated with size-based plasticity in foundress behavior has colony level effects on eusociality. The potential for size-based polyphenisms among reproductive females may be an important factor to consider in the evolutionary origins of eusociality.     

康滇地轴东缘铅锌矿床铅硫同位素地球化学研究

康滇地轴东缘不同时代碳酸盐地层中铅锌矿床的铅同位素研究表明 ,其2 0 7Pb/2 0 4 Pb与2 0 6 Pb/ 2 0 4 Pb呈良好线性关系 ,2 0 7Pb/ 2 0 6 Pb和2 0 8Pb/ 2 0 6 Pb为一常数。结合对本区矿床的稀土元素及成矿流体地球化学的研究 ,判定不同层位铅锌矿床是在同一个成矿体系同时形成的 ,一次成矿 ,其成矿年龄为 2 4 5Ma(峨眉运动 ) ;同时说明不同矿床成矿金属有相同的来源 ,主要来自上地幔。成矿硫以来自地层中的硫酸盐还原硫为主 ,幔源硫次之。本区铅锌矿床的成矿作用与峨眉山玄武岩的喷流关系密切 ,这是与著名的MVT型铅锌矿床的显著区别。     

Predator recognition and evasive behavior by sweat bees, <Emphasis Type="Italic"> Lasioglossum umbripenne </Emphasis>(Hymenoptera: Halictidae), in response to predation by ants, <Emphasis Type="Italic"> Ectatomma ruidum </Emphasis>(Hymenoptera: Formicidae)

Ectatomma ruidum is an abundant soil-nesting Neotropical ant, which displays extensive behavioral flexibility during foraging activities. We studied here one unusual element of their behavioral repertoire: ambush predation. A worker of E. ruidum waits near a nest of a social sweat bee, Lasioglossum umbripenne, lunging at incoming bees, or less frequently, at departing bees. However, bees detected ambushing ants and modified their behavior. Dead ants placed at bees' nest entrances significantly decreased bee activity, indicating that bees recognized dead ants as potential predators. Neither simple black models (square and rectangle) nor olfactory cues had any effect on overall bee activity. A returning bee usually approached her entrance and immediately entered, but if an ant was waiting at the nest, a bee was significantly more likely to abort the first approach flight and then to re-approach the nest on the side opposite the ant's position. As models became increasingly ant-like, returning bees more frequently aborted their first approach flight, expressing other behaviors before entering nests. These behaviors included withdrawal followed by an approach from a different direction; zigzagging flights, either from a distance or close to the entrance or even a close inspection; landing a short distance from the nest, then approaching on foot or waiting for several seconds before entering. Ants responded with effective counter-behaviors. Behavioral flexibility in nest entering/exiting by L. umbripenne and in hunting strategy by E. ruidum shows the complexity of this predator-prey relationship, and illustrates the importance of information processing by both species involved in determining the outcome of the interspecific interaction.     

Attraction and learning in mate-finding by solitary bees,Lasioglossum (Dialictus) figueresi Wcislo and Nomia triangulifera Vachal (Hymenoptera : Halictidae)

Summary Males of the solitary sweat bee, Nomia triangulifera, patrol over large areas where thousands of females emerge, searching for receptive females. The daily operational sex ratio is strongly male-biased. Males contact dead, frozen (untreated) females more frequently than they contact females which were washed in hexane, showing that olfactory cues are utilized in mate-finding. A major source of female sex pheromone is in the head. Male pouncing on females is temporally non-random, indicative of group stimulation. Bioassays show that newly emerged females are more attractive to males than are older pollen-collecting females. Female odors are individually distinctive, based on male responses, and there is much variation among females in their attractive properties. Male responses to female odors suggest that learning is important for mating in natural populations. In contrast, the following hypotheses are unlikely to account for the observed behavior: (1) dissipation of female odors; (2) site learning and avoidance behavior by males; (3) decay of male motivation; or (4) male-produced repellents effective against other males. Laboratory and field studies show that female Lasioglossum figueresi produce individually-distinctive odors, which are attractive to males. There is considerable inter-individual variation among females in their attractiveness to males among sexually immature females. Male responses to female odors decay over the course of the presentation, suggesting the importance of learning in natural populations, although several alternatives could not be tested.     

Assured fitness returns favor sociality in a mass-provisioning sweat bee,Megalopta genalis (Hymenoptera: Halictidae)

Assured fitness returns models for the evolution of sociality emphasize the selective value of ensuring that offspring receive adequate parental care to reach maturity. If a member of a social group dies, it can accrue returns on investment in offspring through the efforts of surviving social partners. We provide evidence that in the mass-provisioning, facultatively social sweat bee Megalopta genalis, adult presence in the nest throughout brood development provides protection from ant predation. Nests with adults present were well protected, and brood in nests with adults removed suffered higher predation. Females in observation nests showed effective defensive behavior against experimentally introduced ants, and bees in natural nests repulsed naturally occurring ant raids. Megalopta nest architecture and behavior are such that the brood of several cooperating females can be defended with little additional cost relative to solitary nesting. The benefits of cooperative defense may favor group living in mass provisioning bees. Our observations and experiments suggest that parental care throughout brood development can be adaptive in mass provisioning species, supporting the predictions of assured fitness returns models.