Socially induced brain development in a facultatively eusocial sweat bee Megalopta genalis (Halictidae)

Changes in the relative size of brain regions are often dependent on experience and environmental stimulation, which includes an animal''s social environment. Some studies suggest that social interactions are cognitively demanding, and have examined predictions that the evolution of sociality led to the evolution of larger brains. Previous studies have compared species with different social organizations or different groups within obligately social species. Here, we report the first intraspecific study to examine how social experience shapes brain volume using a species with facultatively eusocial or solitary behaviour, the sweat bee Megalopta genalis. Serial histological sections were used to reconstruct and measure the volume of brain areas of bees behaving as social reproductives, social workers, solitary reproductives or 1-day-old bees that are undifferentiated with respect to the social phenotype. Social reproductives showed increased development of the mushroom body (an area of the insect brain associated with sensory integration and learning) relative to social workers and solitary reproductives. The gross neuroanatomy of young bees is developmentally similar to the advanced eusocial species previously studied, despite vast differences in colony size and social organization. Our results suggest that the transition from solitary to social behaviour is associated with modified brain development, and that maintaining dominance, rather than sociality per se, leads to increased mushroom body development, even in the smallest social groups possible (i.e. groups with two bees). Such results suggest that capabilities to navigate the complexities of social life may be a factor shaping brain evolution in some social insects, as for some vertebrates.     

Social and ecological contexts of trophallaxis in facultatively social sweat bees, Megalopta genalis and M. ecuadoria (Hymenoptera, Halictidae)

Exchange of liquid food among adults (trophallaxis) is documented for the first time in New World sweat bees (Halictinae). Megalopta genalis and M. ecuadoria are facultatively social, and in social groups foragers regularly give food to the oldest resident female bee, which dominates social interactions. In turn, the oldest resident sometimes re-distributes this food, and shares it with younger foragers. Food is sometimes offered freely, but often the dominant bee exhibits escalating aggressive behavior until she is fed, whereupon she immediately ceases to be aggressive. The occurrence of trophallaxis in a species with mass-provisioned larvae provides an opportunity to examine the ritualization of social behavior. Trophallaxis also increases survivorship of males and females by almost 50% under experimental conditions, suggesting the behavior is also important in ecological contexts. Received 25 July 2005; revised 22 November 2005; accepted 23 December 2005.     

Characterization of 12 polymorphic microsatellite markers for a facultatively eusocial sweat bee (Megalopta genalis)

We developed a library of twelve polymorphic di- and tri-nucleotide microsatellite markers for Megalopta genalis, a facultatively eusocial sweat bee. We tested each locus in a panel of 23 unrelated females and found 7-20 alleles per locus. Observed and expected heterozygosities ranged from 0.65 to 0.96 and from 0.69 to 0.95 respectively. None of the loci deviated from Hardy-Weinberg equilibrium proportions or was found to be in gametic disequilibrium.     

The population genetics of a solitary oligolectic sweat bee, Lasioglossum (Sphecodogastra) oenotherae (Hymenoptera: Halictidae)

Strong evidence exists for global declines in pollinator populations. Data on the population genetics of solitary bees, especially diet specialists, are generally lacking. We studied the population genetics of the oligolectic bee Lasioglossum oenotherae, a specialist on the pollen of evening primrose (Onagraceae), by genotyping 455 females from 15 populations across the bee's North American range at six hyper-variable microsatellite loci. We found significant levels of genetic differentiation between populations, even at small geographic scales, as well as significant patterns of isolation by distance. However, using multilocus genotype assignment tests, we detected 11 first-generation migrants indicating that L. oenotherae's sub-populations are experiencing ongoing gene flow. Southern populations of L. oenotherae were significantly more likely to deviate from Hardy-Weinberg equilibrium and from genotypic equilibrium, suggesting regional differences in gene flow and/or drift and inbreeding. Short-term N(e) estimated using temporal changes in allele frequencies in several populations ranged from approximately 223 to 960. We discuss our findings in terms of the conservation genetics of specialist pollinators, a group of considerable ecological importance.     

Genetic differentiation across the social transition in a socially polymorphic sweat bee,Halictus rubicundus

Eusociality is widely considered a major evolutionary transition. The socially polymorphic sweat bee Halictus rubicundus, solitary in cooler regions of its Holarctic range and eusocial in warmer parts, is an excellent model organism to address this transition, and specifically the question of whether sociality is associated with a strong barrier to gene flow between phenotypically divergent populations. Mitochondrial DNA (COI) from specimens collected across the British Isles, where both solitary and social phenotypes are represented, displayed limited variation, but placed all specimens in the same European lineage; haplotype network analysis failed to differentiate solitary and social lineages. Microsatellite genetic variability was high and enabled us to quantify genetic differentiation among populations and social phenotypes across Great Britain and Ireland. Results from conceptually different analyses consistently showed greater genetic differentiation between geographically distant populations, independently of their social phenotype, suggesting that the two social forms are not reproductively isolated. A landscape genetic approach revealed significant isolation by distance (Mantel test r = 0.622, P < 0.001). The Irish Sea acts as physical barrier to gene flow (partial Mantel test r = 0.453, P < 0.01), indicating that geography, rather than expression of solitary or social behaviour (partial Mantel test r = −0.238, P = 0.053), had a significant effect on the genetic structure of H. rubicundus across the British Isles. Although we cannot reject the hypothesis of a genetic underpinning to differences in solitary and eusocial phenotypes, our data clearly demonstrate a lack of reproductive isolation between the two social forms.     

Local adaptation across a climatic gradient despite small effective population size in the rare sapphire rockcress

When assigning conservation priorities in endangered species, two common management strategies seek to protect remnant populations that (i) are the most genetically divergent or (ii) possess the highest diversity at neutral genetic markers. These two approaches assume that variation in molecular markers reflects variation in ecologically important traits and ignore the possibility of local adaptation among populations that show little divergence or variation at marker loci. Using common garden experiments, we demonstrate that populations of the rare endemic plant Arabis fecunda are physiologically adapted to the local microclimate. Local adaptation occurs despite (i) the absence of divergence at almost all marker loci and (ii) very small effective population sizes, as evidenced by extremely low levels of allozyme and DNA sequence polymorphism. Our results provide empirical evidence that setting conservation priorities based exclusively on molecular marker diversity may lead to the loss of locally adapted populations.     

The effects of temperature and body size on the mating pattern of a gregariously nesting bee, Colletes cunicularius (Hymenoptera: Colletidae)

Abstract. 1. In a 3-year study of the solitary bee Colletes cunicularius L. in Sweden, average body size and population density fluctuated greatly between years.
2. In this protandrous population, females mated just once and the sex ratio was slightly male biased. Males were smaller than females.
3. Size assortative mating (homogamy), associated with an increase in population density during the central days of female emergence and mating, was observed in two out of three years. Homogamy was also observed in pairs with remating males.
4. Most of the mating males had emerged the day they mated, but 42% were older. We found no support for a general large-male mating advantage.
5. Weight of emerging females and mating males were negatively correlated with ground temperature, indicating thermoregulatory influence on the process of sexual selection in this species.     

The effect of pollen protein concentration on body size in the sweat bee Lasioglossum zephyrum (Hymenoptera: Apiformes)

Adult bees and wasps provide all the food their offspring require to grow from egg to adult. For a given diet, offspring body size generally increases with an increase in the amount of food consumed as a larva, but the extent to which body size is influenced by the type of food consumed is poorly known. Pollen ranges from 2–60% protein among plant species, and bees are extremely efficient at assimilating nitrogen; therefore, it seems likely that either parent bees adjust the size of larval provisions to compensate for differences in pollen protein concentration or bee offspring attain different body size depending on the pollen type(s) consumed as a larva. We presented the generalist sweat bee Lasioglossum zephyrum with pollen diets that differed in protein content and monitored offspring body size during two experiments. In a protein supplementation experiment, diets ranged from 20–66% protein and consisted of Typha pollen amended with soy protein. On a pollen/soy diet, offspring body size increased 25% with a shift from 20–37% protein, but did not increase further at greater protein concentrations. In a multiple pollen experiment, pollen diets ranged from 20–39% protein and consisted of eight pollens that differed naturally in protein concentration. The largest offspring arose from the most protein-rich pollens, whereas much smaller bees developed on protein-poor pollens. Provision size only predicted offspring size when pollen type, and therefore protein quantity, was considered. Adult foragers did not adjust provision size to compensate for pollen protein. Therefore, offspring body size appears to result from a combination of controlled (provision size) and uncontrolled (pollen quality) factors that arise out of bee foraging decisions.     

Persistent size variation in the anthophorine bee Centris pallida (Apidae) despite a large male mating advantage

Abstract.

• 1 Despite apparent directional sexual selection in favour of large body size, males of the anthophorine bee Centris pallida remain highly variable in body size.
• 2 One possible cause of persistent size variation among males is geographic variation in the extent of the large male mating advantage. However, a study of a population in an area not previously investigated revealed that the large male mating advantage was as strong here as it has been elsewhere in other years.
• 3 Although the reproductive benefits of being large were consistent in populations separated spatially and temporally, the intensity of bird predation on mate-searching males varied greatly between locations.
• 4 The bee-killing birds focused exclusively on bees which were digging down to meet emerging females or fighting on the ground, never on flying males. Males which were collected on the ground by hand (to simulate avian predation) were significantly larger on average than flying males collected by sweep netting.
• 5 Therefore, in some location in some years, sexual selection in favour of large body size may be opposed by natural selection exerted by predators, perhaps contributing to the maintenance of size variation in this bee.

     

Regulation of brood size in a burying beetle,Nicrophorus tomentosus (Silphidae)

Regulation of brood size in a biparental burying beetle, Nicrophorus tomentosusWeber, was studied by providing pairs with one of two sizes of mouse carcasses in the laboratory. For a given carcass size, there was an inverse relationship between number and mass offspring in a brood. The requirement for regulation was that brood size was adjusted such that mean mass of individual larvae was constant for carcasses of different size. Brood size was regulated if parents were present but regulation did not occur if parents were removed prior to hatching of larvae. Pairs bred in quick succession on two carcasses raised fewer than the regulated number of young in the second reproductive attempt. Reasons for regulation of brood size in this genus are discussed.     

Social polymorphism in the sweat bee <Emphasis Type="Italic">Lasioglossum (Evylaeus) baleicum</Emphasis> (Cockerell) (Hymenoptera,Halictidae) in Hokkaido,northern Japan

Summary The life cycle and social behaviour of the sweat bee Lasioglossum (Evylaeus) baleicum (Cockerell) was investigated in two geographically separate populations in Hokkaido, northern Japan. Colonies were excavated throughout the brood rearing season from an aggregation in Nishioka forest park, Sapporo, and near Kawakita in eastern Hokkaido during 2000 and 2001. The Nishioka population produced two discrete broods during the year and was weakly eusocial; 57% of workers were mated and 28% exhibited some ovarian development, 12–16% of the first brood was male, and workers were on average 4.5% smaller than their respective queen. In contrast, the population at Kawakita was solitary, and produced a single brood per year with an unbiased sex ratio. In addition however, there were some solitary colonies in the Nishioka population and evidence of a partial second brood in some colonies at Kawakita, suggesting differences between the populations are not fixed and that this species is truly socially polymorphic. L. (E.) baleicum is a member of the fulvicorne species group, which includes other examples of social polymorphic species as well as solitary and eusocial species, though this is the only species of this group so far known to exhibit a solitary/non-delayed eusocial polymorphism. Recent studies suggest that social polymorphism has both genetic and environmental influences, raising questions as to the relative import of each.     

Genetic differentiation across a behavioural boundary in a primitively eusocial bee, <Emphasis Type=Italic>Halictus poeyi</Emphasis> Lepeletier (Hymenoptera,Halictidae)

Summary. In Florida, the primitively eusocial bee, Halictus poeyi Lepeletier exhibits two different colony cycles: populations in northern and central Florida are univoltine with an annually brooded colony cycle, while populations in southern Florida, including the Florida Keys, are multivoltine with a continuously brooded colony cycle. In this study, we test the hypothesis of reduced gene flow between the colony types by comparing levels of genetic differentiation among populations having similar colony phenologies with that among populations having different colony phenologies. We use allozyme markers to estimate F_ST, a robust measure of genetic differentiation. We found that genetic differentiation is not significantly higher between populations having different colony phenologies than between populations having similar colony phenologies. Environmental conditions thus may play an important role in influencing the expression of the alternative colony types of H. poeyi populations in Florida.     

Effects of reproductive timing and colony size on the survival, offspring colony size and drone production in the honey bee (Apis mellifera)

ABSTRACT. 1. The effects of colony size and time of reproduction on the survival and size of offspring colonies and on drone production were examined for honey bees, Apis mellifera L. Drone and worker production and survival of parental and offspring colonies were monitored following swarming. Also, the temporal patterns of drone emergence and availability of unmated queens were examined.
2. Colony size at swarming was positively correlated with the number of workers invested in offspring colonies and the number of queens produced. However, colony size at swarming was not correlated with the number of offspring colonies produced.
3. Swarm size was positively correlated with drone and worker production after swarms were hived. Worker production of hived swarms was positively correlated with colony survival. Offspring queens which inherited a parental nest survived longer than queens in either primary swarms or afterswarms, presumably due to the advantage of inheriting a nest.
4. Drone emergence peaked just prior to swarming, the time when unmated queens were available. High drone production by colonies initiated by swarms probably reflected an attempt to reproduce prior to winter. The probabilities of a second swarming cycle within the same year and of surviving the winter were low for colonies initiated from swarms.     

Relationship between body size and flying-related structures in Neotropical social wasps (Polistinae,Vespidae, Hymenoptera)

Body size influences wing shape and associated muscles in flying animals which is a conspicuous phenomenon in insects, given their wide range in body size. Despite the significance of this, to date, no detailed study has been conducted across a group of species with similar biology allowing a look at specific relationship between body size and flying structures. Neotropical social vespids are a model group to study this problem as they are strong predators that rely heavily on flight while exhibiting a wide range in body size. In this paper we describe the variation in both wing shape, as wing planform, and mesosoma muscle size along the body size gradient of the Neotropical social wasps and discuss the potential factors affecting these changes. Analyses of 56 species were conducted using geometric morphometrics for the wings and lineal morphometrics for the body; independent contrast method regressions were used to correct for the phylogenetic effect. Smaller vespid species exhibit rounded wings, veins that are more concentrated in the proximal region, larger stigmata and the mesosoma is proportionally larger than in larger species. Meanwhile, larger species have more elongated wings, more distally extended venation, smaller stigmata and a proportionally smaller mesosoma. The differences in wing shape and other traits could be related to differences in flight demands caused by smaller and larger body sizes. Species around the extremes of body size distribution may invest more in flight muscle mass than species of intermediate sizes.