An overview of proximate factors affecting the nesting behavior of solitary wasps and bees (Hymenoptera: Aculeata) in preexisting cavities in wood

Guilds of Aculeate solitary wasps and bees that nest in preexisting cavities in wood are important components of terrestrial ecosystems because they engage in several ecological interactions (e.g. predation and pollination) with other species of plants and animals. Spatial and temporal variations in richness and abundance of solitary wasps and bees can be related to changes in environmental structure and in the diversity of other groups of organisms. The nesting period of these Aculeata is their most critical life cycle stage. Females of solitary wasp and bee species invest relatively more time constructing and provisioning their nests than do females of social species. Differently from species that nest in the soil or construct exposed nests, the main factors affecting the reproductive success of solitary species nesting in preexisting wood holes are still unknown. Our objective is to provide an overview of the role of proximate causes of nesting failure or success among solitary wasps and bees (Aculeata), for designing effective conservation and management strategies for these Hymenoptera.     

Does bee or wasp mimicry by orchid flowers also deter herbivores?

General visual bee mimicry and specific chemical mimicry by flowers to solitary female bees or wasps are well known in several orchid genera, for example, the Mediterranean genus Ophrys, the Australian genera Cryptostylis and Chiloglottis, and the South-African Disa. This mimicry has been shown to attract solitary male bees or wasps, which are their species-specific pollinators. The visual and chemical signals are considered to be a type of deceptive pollination mechanism based on mimicry for the exploitation of perceptual biases of animals. We propose that in addition to this unique pollination mechanism, these plants exhibit another, rarely mentioned and practically forgotten, non-exclusive function of bee or wasp mimicry (Batesian mimicry). This mimicry may deter large mammalian herbivores, and possibly also insects from the plants and especially from their flowers by a type of visual and olfactory deceptive aposematism. While visiting the flowers, bees and wasps may add a Müllerian effect to this defense. We extend this hypothesis to many other rewarding flowers that are bee or wasp pollinated and propose that abundance of pollinating bees or wasps may deter herbivorous mammals and insects from the plants during their peak flowering season.     

Contrasting Patterns of Species Richness and Composition of Solitary Wasps and Bees (Insecta: Hymenoptera) According to Land‐use

Several species of arthropods inhabiting forest fragments interact with managed areas. The importance of such areas to biodiversity conservation, however, is not well established. Communities of solitary wasps and bees (Insecta: Hymenoptera) play a key role in agroecosystem functioning and they have been used in studies of biodiversity assessment in different land‐use types. We aimed to assess patterns of species richness and composition of solitary wasps and bees over a 1‐yr period in a gradient of decreasing land‐use intensity formed by pastures, alley croppings, young fallows, and old fallows using trap nests. Old fallows had the highest species richness of wasps and bees, harboring all bee species and 86 percent of wasp species occurring in the region, while the remaining land‐uses had similar species richness. Vegetation structure (tree richness) and relative humidity explained most of the variance for the species richness of wasps. For bees, however, there was no influence of environmental factors on the community among land‐use types, indicating better adaptability of this group to environmental variations related to land‐use. The composition of solitary wasp communities (but not those of bees) differed among land‐use types, and the occurrence of rare species in most cases was restricted to old fallow sites. In conclusion, the community of solitary wasps and bees is contingent on land‐use, with solitary wasps more sensitive to anthropized areas. For both groups, less anthropized areas harbor a greater richness and number of rare species while more intensively managed land‐use types harbor higher abundances.     

Trap-nest occupation by solitary wasps and bees (Hymenoptera: Aculeata) in a forest urban remanent

Temporal variation of solitary wasps and bees, nesting frequency, mortality, and parasitism were recorded from a remanent forest in Belo Horizonte, MG, Brazil. Wasps and bees were collected in trap-nests placed in areas with 25, 100, and 400 m2, from February to November 2004. The 137 trap-nests collected contained 11 species of wasps and bees. Wasps occupied most nests (75%). Occupation peaks occurred in March (25%) and September (26%); in June, the lowest occupation (2%) was observed. Except for Trypoxylon (Trypargilum) lactitarse Saussure, no significant correlation was found between number of occupied nests, and temperature and rainfall means. In the nests, 48% of the immature specimens died; 13% of the nests were parasitized. Total death and parasitism rates of wasps and bees differed significantly.     

Flower-feeding specialization in wild bee and wasp communities in seasonal neotropical habitats

Summary Flower visitation records were obtained for 192 species of bees and 138 species of wasps in three neotropical habitats. Four indices of feeding specialization were calculated for common species, and similar patterns were observed using each index. Variation in specialization could be attributed to several factors: the habitat and season in which species were found, the degree of sociality of species, and whether species were bees or wasps. Important differences among habitats and seasons were the number of coexisting consumer species and the species richness and phenology of resource plants. Differences in average consumer specialization among habitats and seasons may have been the result of differences in competitive pressures and to a lesser extent, differences in the flowering strategies of local plants. Eusocial bees and wasps were much more generalized in feeding than solitary species, and bees were more generalized than wasps. Bees in these neotropical communities were much less specialized than their north temperate counterparts. This pattern is reflected by the near absence of species that feed at only one or two plant species in Guanacaste, as compared to a very high frequency of these extreme specialist species in north temperate areas. Temperate zone species of bees have less even visitation rates to their resources than tropical bees. Consequently, temperate zone bees are more specialized in both the variety of plant species visited and the dominance of visitation to each set of resource plants. Several hypotheses are examined as possible explanations for this apparent qualitative difference in community structure between neotropical and temperate communities.     

Ecosystem services provided by aculeate wasps

The aculeate wasps are one of the most diverse and speciose insect taxa; they are omnipresent across ecosystems and exhibit diverse co-evolutionary and exploitative associations with other organisms. There is widespread conjecture that aculeate wasps are likely to perform essential ecological and economic services of importance to the health, well-being and nutritional needs of our planet. However, the scope and nature of the ecosystem services they provide are not well understood relative to other insect groups (e.g. bees, butterflies, beetles); an appreciation of their value is further tarnished by their public reputation as pointless pests. Here, we conduct the first comprehensive review of how aculeate wasps contribute to the four main areas of ecosystem services: regulatory, provisioning, supporting and cultural services. Uniting data from a large but previously disconnected literature on solitary and social aculeate wasps, we provide a synthesis on how these insects perform important ecosystem services as parasites, predators, biological indicators, pollinators, decomposers and seed dispersers; and their additional services as a sustainable alternative to meat for human consumption, and medicinal potential as sources of research leads for anti-microbials and cancer treatments. We highlight how aculeate wasps offer substantial, but largely overlooked, economic benefits through their roles in natural pest management and biological control programs. Accordingly, we provide data-driven arguments for reasons to consider the ecosystem service value of aculeate wasps on a par with other ‘useful’ insects (e.g. bees). Finally, we provide a research roadmap identifying the key areas of research required to capitalise better on the services provided by these important insects.     

Death of an order: a comprehensive molecular phylogenetic study confirms that termites are eusocial cockroaches

Termites are instantly recognizable mound-builders and house-eaters: their complex social lifestyles have made them incredibly successful throughout the tropics. Although known as 'white ants', they are not ants and their relationships with other insects remain unclear. Our molecular phylogenetic analyses, the most comprehensive yet attempted, show that termites are social cockroaches, no longer meriting being classified as a separate order (Isoptera) from the cockroaches (Blattodea). Instead, we propose that they should be treated as a family (Termitidae) of cockroaches. It is surprising to find that a group of wood-feeding cockroaches has evolved full sociality, as other ecologically dominant fully social insects (e.g. ants, social bees and social wasps) have evolved from solitary predatory wasps.     

Comparative analysis of thermoregulation in the nests of certain species of social insects

Studies have been made on thermal regulation in the nests of families of the honey bee Apis mellifera, wasp Dolihovespula silvestris and bumblebees Bombus terrestris, B. agrorum and B. lapidaris during their maximum development. It was shown that thermoregulation significantly stimulated the brood in the nest. Among the species investigated, the highest thermoregulatory capacities are exhibited by honey bees, this fact being associated with the large number of individuals in their colonies. During cooling, bees group around the brood and their bodies make a cover of thermal insulation. Sheltering of the nests plays the main role in heat preservation of wasps and bumblebees. The latter, using contact method of heating of cells with the brood, provide rather constant temperature for its development. Honey bees, wasps and bumblebees react to overheating in the nests essentially in a similar way, i. e. by active aeration of their nests by vigorous wing beatings. The frequency of beatings increases with the increase in temperature.     

Vibration and sound communication in solitary bees and wasps

ABSTRACT. Females of solitary bees ( Colletes cunicularius L.) and of digger wasps ( Bembix rostrata L.) produce buzzing sounds and are known to secrete volatile odours when digging their way from the subterranean nests to the soil surface. The odours allow patrolling males to determine the approximate position of the digging virgin female. The buzzes are measured as substrate-borne sound (soil buzz vibrations) and as air-borne sound (soil buzz sounds). Play-back experiments suggest that the soil buzzes are used by the males as additional cues for localization. Faint buzz sounds are emitted regularly by the male during genital contact in copulation. They may serve to change the receptivity of the female. Intense and broadband buzz sounds are produced by bees of either sex, if restrained from moving, perhaps serving to deter predators.     

Multitrophic effects of experimental changes in plant diversity on cavity-nesting bees, wasps, and their parasitoids

Plant diversity changes can impact the abundance, diversity, and functioning of species at higher trophic levels. We used an experimental gradient in grassland plant diversity ranging from 1 to 16 plant species to study multitrophic interactions among plants, cavity-nesting bees and wasps, and their natural enemies, and analysed brood cell density, insect diversity (species richness), and bee and wasp community similarity over two consecutive years. The bee and wasp communities were more similar among the high (16 species) diversity plots than among plots of the lower diversity levels (up to 8 species), and a more similar community of bees and wasps resulted in a more similar community of their parasitoids. Plant diversity, which was closely related to flower diversity, positively and indirectly affected bee diversity and the diversity of their parasitoids via increasing brood cell density of bees. Increasing plant diversity directly led to higher wasp diversity. Parasitism rates of bees and wasps (hosts) were not affected by plant diversity, but increased with the diversity of their respective parasitoids. Decreases in parasitism rates of bees arose from increasing brood cell density of bees (hosts), whereas decreasing parasitism rates of wasps arose from increasing wasp diversity (hosts). In conclusion, decreases in plant diversity propagated through different trophic levels: from plants to insect hosts to their parasitoids, decreasing density and diversity. The positive relationship between plant diversity and the community similarity of higher trophic levels indicates a community-stabilising effect of high plant diversity.     

Restore it,and they will come: trap-nesting bee and wasp communities (Hymenoptera: Aculeata) are recovered by restoration of riparian forests

Riparian forests have been greatly affected by anthropogenic actions with formerly continuous riparian forests being slowly converted into small and isolated patches. Riparian forests are extremely important habitats for many groups of insects, including bees and wasps, because they are sources of shelter and food for them and their offspring. There is a growing body of evidence of success in the restoration of riparian forest plant communities; however, little research has been done on the associated invertebrate communities. We test whether restoring plant communities is sufficient for restoring the taxonomic composition of trap-nesting bees and wasps and which functional traits are favored in different sites. We predict that species richness, abundance, and community composition of trap-nesting bees and wasps of riparian sites undergoing restoration will converge on the “target” of a reference site with increasing time, since restoration increases habitat complexity. We also predict that the width of restored patches will also influence the species richness, abundance and community composition of trap-nesting bees and wasps. Bee richness and abundance, and wasp richness, were strongly related to fragment width, but not to age since restoration. Our results indicate that although restored sites are relatively small and scattered in a fragmented landscape, they provide suitable habitat for re-colonization by community assemblages of trap-nesting bees and wasps and the traits selected captured the responses to the habitat restoration. Hence, restored riparian areas can be considered important habitats for invertebrates, thus contributing to an increase in local biodiversity and, possibly, the restoration of some of the ecosystem services they originally provided.     

Insights from measuring pollen deposition: quantifying the pre-eminence of bees as flower visitors and effective pollinators

Using our accumulated datasets from Kenyan savanna, Mediterranean garigue, UK gardens and heathland, involving 76 plants from 30 families, we present detailed data to quantify the superiority of bees as pollinators of most flowering plants when compared with other flower visitors. Bees provided the majority of visits to study species at all sites, and 33 of the 76 plants received more than 90% of their visits from bees. Furthermore, pollen deposition onto stigmas from single-visit events (SVD, a measure of pollination effectiveness) was significantly higher for bees than non-bees at all the four sites where a major proportion of the flora was sampled. Solitary bees, and also bumblebees in temperate habitats, were the best potential pollinators for most plants in this respect, and significantly out-performed honeybees. Only a few plants were well served by bombyliid flies, and fewer again by larger hoverflies, butterflies, or solitary wasps. Bees also achieved better matches of their visit timing to peak pollen availability (measured indirectly as peak SVD), and made much shorter visits to flowers than did non-bees, permitting a substantially greater visit frequency. Additionally, they deposited significantly lower levels of potentially deleterious heterospecific pollen on stigmas in heathland and Mediterranean garigue, though not in the UK garden with densely clustered high-diversity flowering, or in the Kenyan savanna site with particularly dispersed flowering patches and some specialist non-bee flowers. Our data provide a novel and quantified characterisation of the specific advantages of bees as flower visitors, and underline the need to conserve diverse bee communities.     

Reproductive plasticity in Polistes paper wasp workers and the evolutionary origins of sociality

Regulatory pathways in solitary species provide the raw materials for the evolution of sociality. Therefore, comparing the mechanisms that mediate reproductive plasticity in social species and their solitary ancestors can provide insight into the evolutionary origin of sociality. In many solitary insects, the effect of juvenile hormone (JH) on fertility is mediated through the fat body; individuals in good physical condition show a stronger fertility response to JH than individuals in poor physical condition. Here, we test whether a similar, condition-dependent JH response mediates fertility in workers of the primitively eusocial Polistes dominulus wasps. We test how body weight, JH, and adult nutrition influence worker ovarian development. Both JH-treatment and adult nutrition dramatically increased ovarian development. Body weight also influenced ovarian development, as large workers developed more eggs than smaller workers. Body weight and fat are strongly linked in P. dominulus workers, so these results suggest that the fat-dependent JH responsiveness common in solitary insects is conserved in social wasps. The simple, ancestral relationship between reproductive investment and physical condition may facilitate cooperation by allowing workers to adaptively allocate energy into reproduction based on their probability of successfully becoming a queen.     

Experience-dependent plasticity in the mushroom bodies of the solitary bee Osmia lignaria (Megachilidae)

All members of the solitary bee species Osmia lignaria (the orchard bee) forage upon emergence from their natal nest cell. Conversely, in the honey bee, days-to-weeks of socially regulated behavioral development precede the onset of foraging. The social honey bee's behavioral transition to foraging is accompanied by neuroanatomical changes in the mushroom bodies, a region of the insect brain implicated in learning. If these changes were general adaptations to foraging, they should also occur in the solitary orchard bee. Using unbiased stereological methods, we estimated the volume of the major compartments of the mushroom bodies, the neuropil and Kenyon cell body region, in adult orchard bees. We compared the mushroom bodies of recently emerged bees with mature bees that had extensive foraging experience. To separate effects of general maturation from field foraging, some orchard bees were confined to a cage indoors. The mushroom body neuropil of experienced field foragers was significantly greater than that of both recently emerged and mature caged orchard bees, suggesting that, like the honey bee, this increase is driven by outdoor foraging experience. Unlike the honey bee, where increases in the ratio of neuropil to Kenyon cell region occur in the worker after emerging from the hive cell, the orchard bee emerged from the natal nest cell with a ratio that did not change with maturation and was comparable to honey-bee foragers. These results suggest that a common developmental endpoint may be reached via different development paths in social and solitary species of foraging bees.     

Social complexity in bees is not sufficient to explain lack of reversions to solitary living over long time scales

Background  

The major lineages of eusocial insects, the ants, termites, stingless bees, honeybees and vespid wasps, all have ancient origins (≥ 65 mya) with no reversions to solitary behaviour. This has prompted the notion of a 'point of no return' whereby the evolutionary elaboration and integration of behavioural, genetic and morphological traits over a very long period of time leads to a situation where reversion to solitary living is no longer an evolutionary option.     

Habitat structure components are effective predictors of trap-nesting Hymenoptera diversity

Habitat heterogeneity can be the major factor affecting species diversity in a community and measuring bee and wasp community habitat preferences in natural systems may provide insights for biodiversity management and conservation. In the present study, we investigate the effects of habitat structure components on solitary bee and wasp species richness and abundance. The research was conducted in an urban forest remnant in southeast Brazil. Our main questions were: (1) is similarity in habitat structure mirrored by similarity in Aculeate assemblage composition? and (2) what are the vegetation features that could be used as predictors of solitary bee and wasp richness and abundance? Aculeate bees and wasps were sampled using trap nests from February to November 2004. Trap nests were placed in sampling units located in 6 ha of secondary mesophytic forest. One hundred and thirty-seven trap nests were occupied by four species of wasps and seven species of bees. Altogether, our sampling units had a mean capture rate (relative to expected richness) of 72% during all the study period. The more similar sampling units were in terms of vegetation structure, the more similar they were in solitary bee and wasp species composition. The variance of tree abundance, shrub height and the abundance of wood logs were good predictors of solitary bee and wasp species richness and abundance in the study area. We demonstrate that even in a small scale it is possible to detect significant influences of habitat features on alpha diversity and that some of them are effective as predictors of trap-nesting Hymenoptera richness and abundance.     

SEX-RATIO MANIPULATION IN RESPONSE TO HOST SIZE BY THE PARASITOID WASP SPALANGIA CAMERONI: A LABORATORY STUDY

The prediction of Charnov et al.'s (1981) host-size model that there should be a negative relationship between host size and wasp sex ratio (proportion sons) was supported for Spalangia cameroni, a solitary parasitoid wasp. The relationship was shown to be a result of offspring sex manipulation by females in response to host size rather than a result of differential mortality of the sexes. A major assumption of the host-size model is that host size has a greater effect on the ultimate reproductive success of emerging female wasps than of males. This assumption was not supported. Host size had a positive effect on the size of both male and female S. cameroni. However, neither host size nor wasp size affected longevity, production of offspring by females, or ability of males to compete for mates. Host size may differentially affect the reproductive success of female and male wasps through effects on other aspects of reproductive success. Tests of the assumptions of offspring sex-ratio manipulation hypotheses are scarce but critical, not only for parasitoid wasps, but also for other organisms.     

Solitary Floral Specialists Do Not Respond to Cryptic Flower-Occupying Predators

The impacts of predators on bee foraging behavior are varied, but have been suggested to depend on both the type of predator (namely their hunting strategy) and also risk assessment by the prey (i.e., ability to perceive predators and learn to avoid them). However, nearly all studies have explored these impacts using social bees, despite the fact that solitary bees are extremely diverse, often specialized in their floral interactions, and may exhibit different behaviors in response to flower-occupying predators. In this study, we examined foraging behaviors of wild solitary long-horned bees (Melissodes spp.) in response to a cryptic predator, the ambush bug (Phymata americana) on the bees’ primary floral host, the prairie sunflower (Helianthus petiolaris). We found sex-specific differences in foraging behaviors of bees, but little evidence that ambush bugs affected either pre-landing or post-landing foraging behaviors. Male bees visited flowers three times more often than females but female bees were five times more likely to land than males. Ambush bugs did not reduce visitation in either sex. Spectral analysis through a bee vision model indicated that ambush bug dorsal coloration was indistinguishable from the disc flowers of sunflowers, suggesting that ambush bugs are indeed cryptic and likely rarely detected by solitary bees. We discuss the implications of these findings for the perceived risk of predation in solitary bees and compare them to other studies of social bees.     

The behaviour of wasps (Vespula germanica L. andV. Vulgaris L.) when foraging

Summary While foraging on dead adult and pupal honeybees, individual wasps showed little conformity in the order in which they dismembered their prey; they attempted to take as large loads as possible and preferred abdomens and thoraces to heads.Although wasps always removed some of the appendages from thoraces, especially the hind and middle legs, the presence of these appendages did not encourage foraging or facilitate recognition of the prey.Pupae were preferred to newly emerged bees, and newly emerged bees to old bees, probably because of the difference in hardness of the cuticle. Whereas wasps learnt to divide adult bodies at the neck and waist, they showed considerable adaptability when dismembering pupae and when confronted with unusual situations. Although individuals tended to become conditioned to collecting one type of prey, some changed from collecting adults to collecting pupae.Wasps could easily be enticed away from meat by offering sugar syrup, but the change from syrup to meat was much more difficult, although it happened occasionally.Some wasps attempted to defend a supply of food against other would-be collectors. Despite their wariness of each other, wasps were attracted to the sight of others at a food source. The frequency with which a wasp continues to visit a site that has ceased to yield food depends on its previous foraging experience there.