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Wild pollinators and their microbiota are sensitive to land use changes from anthropogenic activities that disrupt landscape and environmental features. As urbanization and agriculture affect bee habitats, human-led disturbances are driving changes in bee microbiomes, potentially leading to dysbiosis detrimental to bee fitness. This study examines the bacterial, fungal, and plant compositions of the small carpenter bee, Ceratina calcarata , and its pollen provisions across an urban–rural divide. We performed metabarcoding of C. calcarata and provisions in Toronto by targeting the 16S rRNA, ITS, and rbcL regions. Despite similar plant composition and diversity across bees and their provisions, there was a greater microbial diversity in pollen provisions than in bees. By characterizing the differences in land use, climate, and pesticide residues that differentiate urban and rural landscapes, we find that urban areas support elevated levels of microbial diversity and more complex networks between microbes and plants than rural areas. However, urban areas may lead to lower relative abundances of known beneficial symbionts and increased levels of pathogens, such as Ascosphaera and Alternaria fungi. Further, rural pollen provisions indicate elevated pesticide residues that may dysregulate symbiosis. As anthropogenic activities continue to alter land use, ever changing environments threaten microbiota crucial in maintaining bee health. [ABSTRACT FROM AUTHOR]

Facultatively social insects are an optimal model group for the study of the emergence of cooperation between individuals. Factors influencing the fitness benefits of social nesting are still debated. Nonreproductive subordinates can benefit from indirect fitness benefits due to increasing reproductive success of related individuals or direct fitness benefits due to direct future reproduction. Here, we studied the costs and benefits of social nesting in the small carpenter bee Ceratina albosticta. From demographic data and within-nest relatedness, we obtained key parameters for assessing the fitness of solitary females, social primaries, and social secondaries. C. albosticta were found to usually mate with one male and multiple mating is not common although exists at low frequencies. Social nests usually contain two females, which were found to be related (full sisters), but also some females were unrelated to each other. Patterns of parentage from microsatellite loci revealed that only one female reproduces in social nests. Our results show that relatedness, per capita brood productivity, and offspring survival strongly increase the fitness benefits of social nesting strategies. Social secondaries, when related to the social primaries, have higher inclusive fitness than solitary females, but unrelated social nesting females had no indirect fitness and much reduced inclusive fitness compared to solitary females. Interestingly, average fitness benefits of the social secondary were higher than solitary females. This study provides important empirical data on the costs and benefits of sociality in a facultatively social bee and sets the stage for future comparative studies. [ABSTRACT FROM AUTHOR]

Rudimentary social systems have the potential to both advance our understanding of how complex sociality may have evolved and our understanding of how changes in social environment may influence gene expression and cooperation. Recently, studies of primitively social Hymenoptera have greatly expanded empirical evidence for the role of social environment in shaping behavior and gene expression. Here, we compare brain gene expression profiles of foragers across social contexts in the small carpenter bee, Ceratina calcarata. We conducted experimental manipulations of field colonies to examine gene expression profiles among social contexts including foraging mothers, regular daughters, and worker-like dwarf eldest daughters in the presence and absence of mother. Our analysis found significant differences in gene expression associated with female age, reproductive status, and social environment, including circadian clock gene dyw , hexamerin, and genes involved in the regulation of juvenile hormone and chemical communication. We also found that candidate genes differentially expressed in our study were also associated with division of labor, including foraging, in other primitively and advanced eusocial insects. Our results offer evidence for the role of the regulation of key developmental hormones and circadian rhythms in producing cooperative behavior in rudimentary insect societies. [ABSTRACT FROM AUTHOR]

Elucidating bee response to urbanization is essential to promoting pollinator diversity in cities especially considering such landscapes are projected to expand to support future global populations. To determine how bee community composition and plant–pollinator interactions respond to urbanization, 29 sites representing three urban categories (high, medium and low urbanization) were monitored biweekly from May through early October in Toronto, Canada. Bees were collected passively using pan and blue vane traps as well as actively using aerial nets and vacuums to compare community structure and plant–pollinator networks among urban categories. Functional traits such as dietary breadth, behaviour, nesting substrate and native or non-native status were also examined to determine how landscape influences bee community assemblages. In total, 5477 bees, comprising 26 genera and 164 species, were represented in this study. The urban landscape was largely supportive of species within the family Apidae and Halictidae as well as ground nesting, native and generalist species. Overall, community composition was affected by urban landscape characteristics such as percent tree cover and impervious (i.e. paved and built) surface surrounding sites; however, bee richness and abundance were significantly influenced by plant richness and not by landscape variables. A total of 3267 interactions were observed throughout the study region with characteristics of plant–pollinator networks remaining consistent along the urban gradient with a few floral host plants such as Solidago dominating interactions in certain urban contexts. These results provide important land use and floral host information for targeting pollinator conservation and habitat restoration. [ABSTRACT FROM AUTHOR]

The evolution of eusociality requires that individuals forgo some or all their own reproduction to assist the reproduction of others in their group, such as a primary egg-laying queen. A major open question is how genes and genetic pathways sculpt the evolution of eusociality, especially in rudimentary forms of sociality—those with smaller cooperative nests when compared with species such as honeybees that possess large societies. We lack comprehensive comparative studies examining shared patterns and processes across multiple social lineages. Here we examine the mechanisms of molecular convergence across two lineages of bees and wasps exhibiting such rudimentary societies. These societies consist of few individuals and their life histories range from facultative to obligately social. Using six species across four independent origins of sociality, we conduct a comparative meta-analysis of publicly available transcriptomes. Standard methods detected little similarity in patterns of differential gene expression in brain transcriptomes among reproductive and non-reproductive individuals across species. By contrast, both supervised machine learning and consensus co-expression network approaches uncovered sets of genes with conserved expression patterns among reproductive and non-reproductive phenotypes across species. These sets overlap substantially, and may comprise a shared genetic "toolkit" for sociality across the distantly related taxa of bees and wasps and independently evolved lineages of sociality. We also found many lineage-specific genes and co-expression modules associated with social phenotypes and possible signatures of shared life-history traits. These results reveal how taxon-specific molecular mechanisms complement a core toolkit of molecular processes in sculpting traits related to the evolution of eusociality. [ABSTRACT FROM AUTHOR]

Male hymenopterans do not typically provide help with nest construction or maintenance. This is thought to be due to the decreased relatedness of males to their siblings compared to sisters, and selection for outbreeding resulting in male dispersal from natal nesting sites. However, some instances of male 'helping' behaviors have been observed and can usually be explained by increased access to mating with resident females. Here we report on the first observations of cohabiting males within the nests of reproductive females of the facultatively social small carpenter bee, Ceratina australensis. Social nesting in C. australensis occurs at a consistently low rate across populations. We used microsatellites markers to determine relatedness, combined with 3 yr of nest demographic data collected across three populations, to assess the relative fitness of reproductive, nonreproductive, and male individuals cohabiting in reproductive nests. We found that males were brothers of reproductive females, both remaining in their natal nest. However, there was no evidence that they were mating with their sisters across all nests observed. Males in reproductive nests did not gain any direct or indirect fitness benefits as they did not sire any brood and their presence did not increase brood productivity or survivorship. It is possible that males were waiting to mate with nieces who had not yet emerged. Why males were tolerated remains unknown. Mating biology is an important consideration in social theory which requires additional empirical studies. Future long-term studies are needed to capture unusual social behaviors including male nesting behaviors. [ABSTRACT FROM AUTHOR]

How do social insects expand and adapt to new ranges and how does sociality per se contribute to their success (or failure)? These questions can become tractable with the use of population genomics. We explored the population genomics of the socially polymorphic small carpenter bee, Ceratina australensis , across its range in eastern and southern Australia to search for evidence of selection and identify loci associated with social nesting. We sampled and sequenced fully the genomes of 54 socially and solitarily nesting C. australensis within Queensland, Victoria and South Australia, yielding 2 061 234 single nucleotide polymorphisms across the genome. We found strong evidence of population-specific selection and evidence of genetic variants associated with social nesting behaviour. Both the sets of associated loci and differentially expressed 'social' genes had evidence of positive selection, suggesting that alleles at genes associated with social nesting might provide different fitness benefits. [ABSTRACT FROM AUTHOR]

Despite a strong history of theoretical work on the mechanisms of social evolution, relatively little is known of the molecular genetic changes that accompany transitions from solitary to eusocial forms. Here, we provide the first genome of an incipiently social bee that shows both solitary and social colony organization in sympatry, the Australian carpenter bee Ceratina australensis. Through comparative analysis, we provide support for the role of conserved genes and cis-regulation of gene expression in the phenotypic plasticity observed in nest-sharing, a rudimentary form of sociality. Additionally, we find that these conserved genes are associated with caste differences in advanced eusocial species, suggesting these types of mechanisms could pave the molecular pathway from solitary to eusocial living. Genes associated with social nesting in this species show signatures of being deeply conserved, in contrast to previous studies in other bees showing novel and faster-evolving genes are associated with derived sociality. Our data provide support for the idea that the earliest social transitions are driven by changes in gene regulation of deeply conserved genes. [ABSTRACT FROM AUTHOR]

Social organisms are some of the most pervasive on earth, with the origin of sociality considered a major evolutionary transition. Previous studies suggest a role for both genetic and environmental factors in the transition from solitary to social living, with the relative contributions of these factors varying among taxa. Eusociality has arisen up to 11 times in the Hymenoptera. Four of these origins occurred within the bees, plus many probable reversions, making them ideal to understand the influence of genes and environment on social behaviour. We used a well-supported phylogeny with broad taxonomic coverage of the globally distributed bee genus Ceratina to test whether climate and sociality are correlated. Ceratina was most probably social ancestrally and originated in tropical Africa, with subsequent dispersals into temperate regions corresponding to shifts to solitary living. These findings highlight the utility of facultatively social lineages, such as Ceratina, for assessing the relative importance of phylogeny and ecology in the evolution of social complexity. [ABSTRACT FROM AUTHOR]

The Australian small carpenter bee, Ceratina australensis , is the sole member of the small carpenter bees, genus Ceratina , in Australia. Ceratina australensis is found throughout eastern Australia in dead, broken stems of weedy plant species, where it makes its nests. Here we developed eight microsatellite loci and used these markers to characterize the genetic structure of three populations across three distinct ecoregions that are connected by the Murray–Darling River basin. We genotyped 57 female bees and found significant and geographically consistent variation in allelic diversity and heterozygosity between populations. Through comparisons with the results of mitochondrial DNA screening from a previous study, we infer the possibilities of male-biased dispersal and limited interpopulation migration. Based on available distribution data, the expansion of C. australensis into arid regions appears to follow the Murray–Darling River basin and associated waterways. This basin has undergone severe anthropogenic disturbance since European settlement of Australia, with large-scale changes to native vegetation communities. Although it is unknown how these changes have affected local insect communities, C. australensis does not rely on a single plant species for nesting habitat. We provide evidence for the hypothesis that expansion of C. australensis into Australia might have been aided by the introduction of non-native pithy stemmed plants. [ABSTRACT FROM AUTHOR]

Kin selection theory is foundational in helping to explain the evolution of sociality; however, the degree to which indirect fitness benefits may underlie helping behavior in species of early stage sociality has received relatively little empirical attention. Facultatively social bees, which demonstrate multiple forms of social organization, provide prime systems in which to empirically test hypotheses regarding the evolutionary origins of sociality. The subsocial small carpenter bee, Ceratina calcarata, may establish a social nest by manipulating brood provisions to rear a worker daughter, which then assists in critical late-season alloparental care. Here, we combine nest demographic and behavioral data with genetic relatedness estimates to calculate the relative inclusive fitness of both subsocial and social reproductive strategies in C. calcarata. Social mothers benefit from improved likelihood of brood survivorship and have higher fitness than subsocial mothers. Worker daughters have low indirect fitness on average, and will not produce their own offspring. Among-sibling relatedness is significantly higher in social nests than subsocial nests, though mothers of either reproductive strategy may mate multiply. Though this study corroborates the ultimate role of indirect fitness and assured fitness returns in the evolution of social traits, it also offers additional support for maternal manipulation as the proximate mechanism underlying evolutionary transitions in early stage insect societies. [ABSTRACT FROM AUTHOR]

The genetic mechanisms behind aggressive behaviors are important for understanding the formation of dominance hierarchies, and thus social systems in general. Studies into the effects of social experience and agonistic contest outcomes have shown significant changes in brain gene expression resulting from repeated winning and losing, as well as changing dominance rank, primarily in obligately social species. However, our knowledge of the genetic underpinnings of behavior in subsocial organisms is relatively poor, yet understanding the behavioral genetics of this simplest form of sociality provides the basis for understanding all other forms of social living. Here, we measured the effects of aggression on brain gene expression in the subsocial bee, Ceratina calcarata, in individuals that had experienced repeated winning, repeated losing, or a change in rank during repeated encounters. Consistent winning accounted for the majority of variation in brain gene expression, followed by changing rank over maintaining rank. Candidate genes for aggression are identified through comparative transcriptomics across 21 invertebrate and 6 vertebrate taxa. Lastly, we identified significantly over-represented cis-regulatory elements, namely transcription factor binding motifs deeply conserved across a wide range of taxa and broadly implicated in differential regulation of genes related to aggressive/dominant behavior. We present evidence for both genetic and cis-regulatory mechanisms for aggression that may have broad importance to social evolution. [ABSTRACT FROM AUTHOR]

Changes in gene regulation that underlie phenotypic evolution can be encoded directly in the DNA sequence or mediated by chromatin modifications such as DNA methylation. It has been hypothesized that the evolution of eusocial division of labor is associated with enhanced gene regulatory potential, which may include expansions in DNA methylation in the genomes of Hymenoptera (bees, ants, wasps, and sawflies). Recently, this hypothesis garnered support from analyses of a commonly used metric to estimate DNA methylation in silico, CpG content. Here, we test this hypothesis using direct, nucleotide-level measures of DNAmethylation across nine species of Hymenoptera. In doing so, we generated new DNA methylomes for three species of interest, including one solitary and one facultatively eusocial halictid bee and a sawfly. We demonstrate that the strength of correlation between CpG content and DNA methylation varies widely among hymenopteran taxa, highlighting shortcomings in the utility of CpG content as a proxy for DNA methylation in comparative studies of taxa with sparse DNA methylomes. We observed strikingly high levels of DNAmethylation in the sawfly relative to other investigated hymenopterans. Analyses of molecular evolution suggest the relatively distinct sawfly DNAmethylome may be associatedwith positive selection on functional DNMT3 domains. Sawflies are an outgroup to all ants, bees, and wasps, and no sawfly species are eusocial. We find no evidence that either global expansions or variation within individual ortholog groups in DNA methylation are consistently associated with the evolution of social behavior. [ABSTRACT FROM AUTHOR]

Many wild bee species are in global decline, yet much is still unknown about their diversity and contemporary distributions. National parks and forests offer unique areas of refuge important for the conservation of rare and declining species populations. Here we present the results of the first biodiversity survey of the bee fauna in the White Mountain National Forest (WMNF). More than a thousand specimens were collected from pan and sweep samples representing 137 species. Three species were recorded for the first time in New England and an additional seven species were documented for the first time in the state of New Hampshire. Four introduced species were also observed in the specimens collected. A checklist of the species found in the WMNF, as well as those found previously in Strafford County, NH, is included with new state records and introduced species noted as well as a map of collecting locations. Of particular interest was the relatively high abundance of Bombus terricola Kirby 1837 found in many of the higher elevation collection sites and the single specimen documented of Bombus fervidus (Fabricius 1798). Both of these bumble bee species are known to have declining populations in the northeast and are categorized as vulnerable on the International Union for Conservation of Nature's Red List. [ABSTRACT FROM AUTHOR]

Understanding the evolution of animal societies, considered to be a major transition in evolution, is a key topic in evolutionary biology. Recently, new gateways for understanding social evolution have opened up due to advances in genomics, allowing for unprecedented opportunities in studying social behavior on a molecular level. In particular, highly eusocial insect species (caste-containing societies with nonreproductives that care for siblings)have taken center stage instudies of the molecular evolution of sociality. Despite advances in genomic studies of both solitary and eusocial insects, we still lack genomic resources for early insect societies. To study the genetic basis of social traits requires comparison of genomes from a diversity of organisms ranging from solitary to complex social forms. Here we present the genome of a subsocial bee, Ceratina calcarata. This study begins to address the types of genomic changes associated with the earliest origins of simple sociality using the small carpenter bee. Genes associated with lipid transport and DNA recombination have undergone positive selection in C. calcarata relative to other bee lineages. Furthermore, we provide the first methylome of a noneusocial bee. Ceratina calcarata contains the complete enzymatic toolkit for DNA methylation. As in the honey bee and many other holometabolous insects, DNA methylation is targeted to exons. The addition of this genome allows for new lines of research into the genetic and epigenetic precursors to complex social behaviors. [ABSTRACT FROM AUTHOR]

Bees fulfil a critical ecological role as pollinators, significantly contributing to the reproductive success of myriad angiosperm species. Although increasingly appreciated for their agricultural contributions, relatively little is yet known about the natural history of the vast majority of the more than 20 000 wild bee species worldwide. The small carpenter bee genus Ceratina occurs globally, and is represented in North America by its most recently diverged subgenera, Ceratinula and Zadontomerus. Recent genetic analysis of eastern Ceratina ( Zadontomerus) supports the existence of five closely related, yet genetically distinct species living in sympatry. This phylogeographical study employs molecular barcoding of the most comprehensive specimen collection yet assembled to confirm the identities of these recently diverged eastern North American Ceratina ( Zadontomerus) species. Delineation of extant population structure, evolutionary history and known range of this emerging model native pollinator are greatly improved by this study. We consider ecological and behavioural factors potentially contributing to the maintenance of genetic identity among these sympatric species. [ABSTRACT FROM AUTHOR]

The small carpenter bee, Ceratina calcarata (Robertson), is a widespread native pollinator across eastern North America. The behavioral ecology and nesting biology of C. calcarata has been relatively well-studied and the species is emerging as a model organism for both native pollinator and social evolution research. C. calcarata is subsocial: reproductively mature females provide extended maternal care to their brood. As such, studies of C. calcarata may also reveal patterns of relatedness and demography unique to primitively social Hymenoptera. Here, we present 21 microsatellite loci, isolated from the recently completed C. calcarata genome. Screening in 39 individuals across their distribution revealed that no loci were in linkage disequilibrium, nor did any deviate significantly from Hardy-Weinberg following sequential Bonferroni correction. Allele count ranged from 2 to 14, and observed and expected heterozygosities ranged from 0.08 to 0.82 (mean 0.47) and 0.26 to 0.88 (mean 0.56), respectively. These markers will enable studies of population-wide genetic structuring across C. calcarata's distribution. Such tools will also allow for exploration of between and within-colony relatedness in this subsocial native pollinator. [ABSTRACT FROM AUTHOR]

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. In the present study, we assess the fitness consequences of these three ecological factors on reproductive success of solitary nests and social colonies in the socially polymorphic small carpenter bee, Ceratina australensis, based on 982 nests collected over four reproductive periods. Nest site limitation was predicted to decrease opportunities for independent nest initiation and increase the frequency of social nesting. Nest sites were not limiting in this species and the frequency of social nesting was consistent across the four brood-rearing periods studied. Unfavourable weather was predicted to lower the frequency of female dispersal from their natal nests and to limit the brood-rearing season; this would increase the frequency and fitness of social colonies. Daily temperature and precipitation accumulation varied between seasons but were not correlated with reproductive success in this bee. Increased parasite pressure is predicted to increase the frequency and fitness of social colonies because solitary bees must leave the nest unattended during foraging bouts and are less able to defend the nest against parasites. Severe parasitism by a chalcid wasp ( Eurytoma sp.) resulted in low reproductive success and total nest failure in solitary nests. Social colonies had higher reproductive success and were never extirpated by parasites. The high frequency of solitary nests suggests that this is the optimal strategy. However, social colonies have a selective advantage over solitary nesting females during periods of extreme parasite pressure, and we suggest that social nesting represents a form of bet-hedging against unpredictable fluctuations in parasite number. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, , 57-67. [ABSTRACT FROM AUTHOR]