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Although the knowledge of the skeletal morphology of bees has progressed enormously, a corresponding advance has not happened for the muscular system. Most of the knowledge about bee musculature was generated over 50 years ago, well before the digital revolution for anatomical imaging, including the application of microcomputed tomography. This technique, in particular, has made it possible to dissect small insects digitally, document anatomy efficiently and in detail, and visualize these data three dimensionally. In this study, we document the skeletomuscular system of a cuckoo bee, Thyreus albomaculatus and, with that, we provide a 3D atlas of bee skeletomuscular anatomy. The results obtained for Thyreus are compared with representatives of two other bee families (Andrenidae and Halictidae), to evaluate the generality of our morphological conclusions. Besides documenting 199 specific muscles in terms of origin, insertion, and structure, we update the interpretation of complex homologies in the maxillolabial complex of bee mouthparts. We also clarify the complicated 3D structure of the cephalic endoskeleton, identifying the tentorial, hypostomal, and postgenal structures and their connecting regions. We describe the anatomy of the medial elevator muscles of the head, precisely identifying their origins and insertions as well as their homologs in other groups of Hymenoptera. We reject the hypothesis that the synapomorphic propodeal triangle of Apoidea is homologous with the metapostnotum, and instead recognize that this is a modification of the third phragma. We recognize two previously undocumented metasomal muscle groups in bees, clarifying the serial skeletomusculature of the metasoma and revealing shortcomings of Snodgrass' "internal-external" terminological system for the abdomen. Finally, we elucidate the muscular structure of the sting apparatus, resolving previously unclear interpretations. The work conducted herein not only provides new insights into bee morphology but also represents a source for future phenomic research on Hymenoptera., (© 2024 The Author(s). Journal of Morphology published by Wiley Periodicals LLC.)

The evolution of flowers that offer oils as rewards and are pollinated by specialized bees represents a distinctive theme in plant-pollinator co-diversification. Some plants that offer acetylated glycerols as floral oils emit diacetin, a volatile by-product of oil metabolism, which is utilized by oil-collecting bees as an index signal for the presence of floral oil. However, floral oils in the genus Krameria (Krameriaceae) contain β-acetoxy-substituted fatty acids instead of acetylated glycerols, making them unlikely to emit diacetin as an oil-bee attractant. We analyzed floral headspace composition from K. bicolor and K. erecta, native to the Sonoran Desert of southwestern North America, in search of alternative candidates for volatile index signals. Using solid-phase microextraction, combined with gas chromatography-mass spectrometry, we identified 26 and 45 floral volatiles, respectively, from whole flowers and dissected flower parts of these two Krameria species. As expected, diacetin was not detected. Instead, β-ionone emerged as a strong candidate for an index signal, as it was uniquely present in dissected oil-producing floral tissues (elaiophores) of K. bicolor, as well as the larval cells and provisions from its oil-bee pollinator, Centris cockerelli. This finding suggests that the floral oil of K. bicolor is perfused with β-ionone in its tissue of origin and retains the distinctive raspberry-like scent of this volatile after being harvested by C. cockerelli bees. In contrast, the elaiophores of K. erecta, which are not thought to be pollinated by C. cockerelli, produced a blend of anise-related oxygenated aromatics not found in the elaiophores of K. bicolor. Our findings suggest that β-ionone has the potential to impact oil-foraging by C. cockerelli bees through several potential mechanisms, including larval imprinting on scented provisions or innate or learned preferences by foraging adults., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Background: Xicotli data is the short name given to the dataset generated within the project framework "Integration of Biodiversity Data for the Management and Conservation of Wild Bee-Plant Interactions in Mexico (2021-2023)", as xicotli is the generic word for a bee in Nahuatl. The team comprised eco-informaticians, ecologists and taxonomists of both native bees and flora. The generated dataset contains so far 4,532 curated records of the plants, which are potential hosts of species of three focal families of bees native to Mexico: Apidae, Halictidae and Megachilidae and morphological and ecological data of the plant-bee interactions. This dataset was integrated and mobilised from citizen observations available at naturalista.mx (iNat), which were compiled through the iNaturalist project., New Information: The new information obtained with the Xicotli data project was: Taxonomic information about bee species curated by taxonomists based on the information contained in iNaturalist;Taxonomic identification of the host plants by a botanist from the photos compiled by the Xicotli Data project;Data on the ecomorphological traits of bees and plants based on expert knowledge and literature. All the data were integrated into the Xicotli Data Project via the creation of new "observation fields". The visibility of the information originally contained in iNaturalist was maximized and can be consulted directly on the iNaturalist platform., (Juan M Barrios, Brenda Y Bedolla-García, Paola A González-Vanegas, Andrés Lira-Noriega, Juan C López-Enriquez, Jorge A Mérida-Rivas, Daniel Madrigal-González, Pilar Rodríguez, Matthias Rös, Remy Vandame, Raúl Sierra-Alcocer, Carlos A. Cultid-Medina.)

Currently, only Apis mellifera is used in environmental regulation to evaluate the hazard of pesticides to pollinators. The low representativeness of pollinators and bee diversity in this approach may result in insufficient protection for the wild species. This scenario is intensified in tropical environments, where little is known about the effects of pesticides on solitary bees. We aimed to calculate the medium lethal dose (LD50) and medium lethal concentration (LC50) of the insecticide dimethoate in the Neotropical solitary bee Centris analis, a cavity-nesting, oil-collecting bee distributed from Brazil to Mexico. Males and females of C. analis were exposed orally to dimethoate for 48 h under laboratory conditions. Lethality was assessed every 24 h until 144 h after the beginning of the test. After the LD50 calculation, we compared the value with available LD50 values in the literature of other bee species using the species sensitivity distribution curve. In 48 h of exposure, males showed an LD50 value 1.33 times lower than females (32.78 and 43.84 ng active ingredient/bee, respectively). Centris analis was more sensitive to dimethoate than the model species A. mellifera and the solitary bee from temperate zones, Osmia lignaria. However, on a body weight basis, C. analis and A. mellifera had similar LD50 values. Ours is the first study that calculated an LD50 for a Neotropical solitary bee. Besides, the results are of crucial importance for a better understanding of the effects of pesticides on the tropical bee fauna and will help to improve the risk assessment of pesticides to bees under tropical conditions, giving attention to wild species, which are commonly neglected. Environ Toxicol Chem 2023;42:2758-2767. © 2023 SETAC., (© 2023 SETAC.)

Insecticides and climate change are among the multiple stressors that bees face, but little is known about their synergistic effects, especially for non- Apis bee species. In laboratory experiments, we tested whether the stingless bee Tetragonula hockingsi avoids insecticide in sucrose solutions and how T. hockingsi responds to insecticide and heat stress combined. We found that T. hockingsi neither preferred nor avoided sucrose solutions with either low (2.5 × 10 -4 ng µl -1 imidacloprid or 1.0 × 10 -4 ng µl -1 fipronil) or high (2.5 × 10 -3 ng µl -1 imidacloprid or 1.0 × 10 -3 ng µl -1 fipronil) insecticide concentrations when offered alongside sucrose without insecticide. In our combined stress experiment, the smallest dose of imidacloprid (7.5 × 10 -4 ng) did not significantly affect thermal tolerance (CT max ). However, CT max significantly reduced by 0.8°C (±0.16 SE) and by 0.5°C (±0.16 SE) when bees were fed as little as 7.5 × 10 -3 ng of imidacloprid or 3.0 × 10 -4 ng of fipronil, respectively, and as much as 1.5°C (±0.16 SE) and 1.2°C (±0.16 SE) when bees were fed 7.5 × 10 -2 ng of imidacloprid or 3.0 × 10 -2 ng of fipronil, respectively. Predictions of temperature increase, and increased insecticide use in the tropics suggest that T. hockingsi will be at increased risk of the effects of both stressors in the future., Competing Interests: We declare we have no competing interests., (© 2023 The Authors.)

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., (© The Author(s) 2023. Published by Oxford University Press on behalf of FEMS.)

Pollination is a vitally important function in nature and becomes an ecosystem service because it influences the food and nutritional security for people. However, the contribution of different functional traits of insects for pollen transport of plants is still poorly known. We explore the relationship between pollinator insect functional traits and the transport of pollen of sweet granadilla (Passiflora ligularis Juss) in eight crops. We sampled flower-visiting insects of this crop and recorded 10 functional traits (five by direct measurements and five from the literature) that were related to the amount of pollen carried by each insect. Bees (Apidae) were not only the most abundant insects but also the ones that loaded the highest amounts of pollen. Within these, the most abundant species was the exotic common honeybee (Apis mellifera (Linnaeus)) making up almost half of the specimens collected; however, this bee carried less pollen grains than other native bees. Bombus hortulanus (Smith) was one of the large-bodied native bees that carried more sweet granadilla pollen, despite not being an abundant species in the community. Body size was the most important trait determining the transport of sweet granadilla pollen, while the traits related to body hairs were not significant for the body's pollen load. None of the functional traits evaluated was influenced by taxonomy at species-level. Our results suggest that large body sizes in bees are the most important traits in granadilla pollen transport, regardless of other changes in composition and structure of pollinating insect assemblages in the crop., (© 2023. The Author(s).)

Background: Many insect species have shown dramatic declines over the last decades, as a result of man-related environmental changes. Many species which were formerly widespread are now rare. To document this trend with evidence, old records of collected specimens are vital., New Information: We provide here the data on 9752 bee (Hymenoptera: Apoidea) specimens hosted in several museums of south-east France: Musée des Confluences in Lyon, Muséum d'Histoire Naturelle de Marseille, Muséum d'Aix-en-Provence and the Muséum Départemental du Var in Toulon. Most of the specimens (9256) come from France and include data on 552 named species. For most of these specimens, the geographical location, including geographical coordinates, is based on the locality (town or village) where they were collected. The specimens were captured from the beginning of the nineteenth century to 2018. The identifications of 1377 specimens, mainly belonging to the genus Bombus , are considered reliable, as these were performed or been checked since 2009. All the other reported identifications are the original ones given by the original collectors., (Jean-Yves Meunier, Benoît Geslin, Mehdi Issertes, Gilles Mahé, Frédéric Vyghen, Harold Labrique, Yves Dutour, Vincent Poncet, Jérémy Migliore, Gabriel Nève.)

Brazil is one of the centers of diversity of Cactaceae, yet studies addressing both pollination biology and the breeding system in Brazilian cacti are scarce. We herein present a detailed analysis of two native species with economic relevance: Cereus hildmannianus and Pereskia aculeata . The first species produce edible, sweet, spineless fruits and the second species produces leaves with high protein content. Pollination studies were undertaken through fieldwork observations in three localities of Rio Grande do Sul, Brazil, over two flowering seasons, totaling over 130 observation hours. Breeding systems were elucidated utilizing controlled pollinations. Cereus hildmannianus is solely pollinated by nectar-gathering species of Sphingidae hawk moths. In contrast, the flowers of P. aculeata are pollinated by predominantly native Hymenoptera but also by Coleoptera and Diptera, which gather pollen and/or nectar. Both cacti species are pollinator-dependent; neither intact nor emasculated flowers turn into fruit, yet whereas C. hildmannianus is self-incompatible, P. aculeata is fully self-compatible. In sum, C. hildmannianus is more restrictive and specialized regarding its pollination and breeding system, whereas P. aculeata is more generalist. Understanding the pollination needs of these species is a necessary starting point towards their conservation but also for their proper management and eventual domestication.

The gut microbiota of bees affects nutrition, immunity and host fitness, yet the roles of diet, sociality and geographical variation in determining microbiome structure, including variant-level diversity and relatedness, remain poorly understood. Here, we use full-length 16S rRNA amplicon sequencing to compare the crop and gut microbiomes of two incipiently social carpenter bee species, Xylocopa sonorina and Xylocopa tabaniformis, from multiple geographical sites within each species' range. We found that Xylocopa species share a set of core taxa consisting of Bombilactobacillus, Bombiscardovia and Lactobacillus, found in >95% of all individual bees sampled, and Gilliamella and Apibacter were also detected in the gut of both species with high frequency. The crop bacterial community of X. sonorina comprised nearly entirely Apilactobacillus with occasionally abundant nectar bacteria. Despite sharing core taxa, Xylocopa species' microbiomes were distinguished by multiple bacterial lineages, including species-specific variants of core taxa. The use of long-read amplicons revealed otherwise cryptic species and population-level differentiation in core microbiome members, which was masked when a shorter fragment of the 16S rRNA (V4) was considered. Of the core taxa, Bombilactobacillus and Bombiscardovia exhibited differentiation in amplicon sequence variants among bee populations, but this was lacking in Lactobacillus, suggesting that some bacterial genera in the gut may be structured by different processes. We conclude that these Xylocopa species host a distinctive microbiome, similar to that of previously characterized social corbiculate apids, which suggests that further investigation to understand the evolution of the bee microbiome and its drivers is warranted., (© 2022 John Wiley & Sons Ltd.)

Pollen is an important source of nutrition for bumblebees to survive, reproduce, and raise their offspring. To explore the nutritional requirements for the egg laying and hatching of queenright Bombus breviceps colonies, camellia pollen, oilseed rape pollen, apricot pollen, and mixtures of two or three types of pollen in equal proportions were used to feed the queens in this study. The results showed that the camellia pollen with a higher essential amino acid content was superior to the pollen with a lower essential amino acid content in the initial egg-laying time ( p < 0.05), egg number ( p < 0.05), larval ejection ( p < 0.01), time of first worker emergence ( p < 0.05), and the average weight of workers in the first batch ( p < 0.01). It took less time for colonies under the camellia pollen and camellia-oilseed rape-apricot pollen mix treatments, both with a higher crude protein content, to reach ten workers in the colony ( p < 0.01). On the contrary, the queens fed apricot pollen never laid an egg, and larvae fed oilseed rape pollen were all ejected-both pollens with a lower essential amino acid content. The results emphasize that the diet should be rationally allocated to meet the nutritional needs of local bumblebees at various stages when guiding them to lay eggs, hatch, and develop a colony.

Bees and their microbes interact in complex networks in which bees form symbiotic relationships with their bacteria and fungi. Microbial composition and abundance affect bee health through nutrition, immunity, and fitness. In ever-expanding urban landscapes, land use development changes bee habitats and floral resource availability, thus altering the sources of microbes that wild bees need to establish their microbiome. Here, we implement metabarcoding of the bacterial 16S and fungal ITS regions to characterize the diversity and composition of the microbiome in 58 small carpenter bees, Ceratina calcarata, across urban land use gradients (study area 6,425 km 2 ). By categorizing land use development, green space, precipitation, and temperature variables as indicators of habitat across the city, we found that land use variables can predict microbial diversity. Microbial composition was also found to vary across urban land use gradients, with certain microbes such as Acinetobacter and Apilactobacillus overrepresented in less urban locations and Penicillium more abundant in developed areas. Environmental features may also lead to differences in microbe interactions, as co-occurrences between bacteria and fungi varied across percent land use development, exemplified by the correlation between Methylobacterium and Sphingomonas being more prevalent in areas of higher urban development. Surrounding landscapes change the microbial landscape in wild bees and alter the relationships they have with their microbiome. As such, urban centres should consider the impact of growing cities on their pollinators' health and protect wild bees from the effects of anthropogenic activities., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Nguyen and Rehan.)

It is increasingly recognised that intraspecific variation in traits, such as morphology, behaviour, or diet is both ubiquitous and ecologically important. While many species of predators and herbivores are known to display high levels of between-individual diet variation, there is a lack of studies on pollinators. It is important to fill in this gap because individual-level specialisation of flower-visiting insects is expected to affect their efficiency as pollinators with consequences for plant reproduction. Accordingly, the aim of our study was to quantify the level of individual-level specialisation and foraging preferences, as well as interspecific resource partitioning, in three co-occurring species of bees of the genus Ceratina (Hymenoptera: Apidae: Xylocopinae), C. chalybea , C. nigrolabiata , and C. cucurbitina . We conducted a field experiment where we provided artificial nesting opportunities for the bees and combined a short-term mark-recapture study with the dissection of the bees' nests to obtain repeated samples from individual foraging females and complete pollen provisions from their nests. We used DNA metabarcoding based on the ITS2 locus to identify the composition of the pollen samples. We found that the composition of pollen carried on the bodies of female bees and stored in the brood provisions in their nests significantly differed among the three co-occurring species. At the intraspecific level, individual females consistently differed in their level of specialisation and in the composition of pollen carried on their bodies and stored in their nests. We also demonstrate that higher generalisation at the species level stemmed from larger among-individual variation in diets, as observed in other types of consumers, such as predators. Our study thus reveals how specialisation and foraging preferences of bees change from the scale of individual foraging bouts to complete pollen provisions accumulated in their nests over many days. Such a multi-scale view of foraging behaviour is necessary to improve our understanding of the functioning of plant-flower visitor communities., Competing Interests: The authors declare there are no competing interests., (©2022 Klečka et al.)

Unravelling the evolutionary origins of eusocial life is a longstanding endeavour in the field of evolutionary-developmental biology. Descended from solitary ancestors, eusocial insects such as honeybees have evolved ontogenetic division of labour in which short-lived workers perform age-associated tasks, while a long-lived queen produces brood. It is hypothesized that (i) eusocial caste systems evolved through the co-option of deeply conserved genes and (ii) longevity may be tied to oxidative damage mitigation capacity. To date, however, these hypotheses have been examined primarily among only obligately eusocial corbiculate bees. We present brain transcriptomic data from a Japanese small carpenter bee, Ceratina japonica (Apidae: Xylocopinae), which demonstrates both solitary and eusocial nesting in sympatry and lives 2 or more years in the wild. Our dataset captures gene expression patterns underlying first- and second-year solitary females, queens and workers, providing an unprecedented opportunity to explore the molecular mechanisms underlying caste-antecedent phenotypes in a long-lived and facultatively eusocial bee. We find that C. japonica 's queens and workers are underpinned by divergent gene regulatory pathways, involving many differentially expressed genes well-conserved among other primitively eusocial bee lineages. We also find support for oxidative damage reduction as a proximate mechanism of longevity in C. japonica .

Stingless bees are important pollinators in the tropics. The tremendous variation in body size makes them an excellent group to study how miniaturization affects vision and visual behaviours. Using direct measurements and micro-CT, we reconstructed the eye structure, estimated anatomical spatial resolution and optical sensitivity of the stingless bee Tetragonula iridipennis. T. iridipennis is similar in size to the Australian stingless bee Tetragonula carbonaria and is smaller than honeybees. It has correspondingly small eyes (area = 0.56 mm 2 ), few ommatidia (2451 ± 127), large inter-facet (3.0 ± 0.6°) and acceptance angles (2.8°). Theoretical estimates suggest that T. iridipennis has poorer spatial resolution (0.17 cycles degree -1 ) than honeybees, bumblebees, and T. carbonaria. Its optical sensitivity (0.08 µm 2 sr), though higher than expected, is within the range of diurnal bees. This may provide them with greater contrast sensitivity, which is likely more relevant than the absolute sensitivity in this diurnal bee. Behaviourally determined detection thresholds for single targets using y-maze experiments were 11.5° for targets that provide chromatic contrast alone and 9.1° for targets providing chromatic and achromatic contrast. Further studies into microhabitat preferences and behaviour are required to understand how miniaturization influences its visual ecology., (© 2021. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Sown wildflower areas are increasingly recommended as an agri-environmental intervention measure, but evidence for their success is limited to particular insect groups or hampered by the challenges of establishing seed mixes and maintaining flower abundance over time. We conducted a replicated experiment to establish wildflower areas to support insect pollinators in apple orchards. Over three years, and across 23 commercial UK orchards with and without sown wildflowers, we conducted 828 transect surveys across various non-crop habitats. We found that the abundance of flower-visiting solitary bees, bumblebees, honeybees, and beetles was increased in sown wildflower areas, compared with existing non-crop habitats in control orchards, from the second year following floral establishment. Abundance of hoverflies and other non-syrphid flies was increased in wildflower areas from the first year. Beyond the effect of wildflower areas, solitary bee abundance was also positively related to levels of floral cover in other local habitats within orchards, but neither local nor wider landscape-scale context affected abundance of other studied insect taxa within study orchards. There was a change in plant community composition on the sown wildflower areas between years, and in patterns of flowering within and between years, showing a succession from unsown weedy species towards a dominance of sown species over time. We discuss how the successful establishment of sown wildflower areas and delivery of benefits for different insect taxa relies on appropriate and reactive management practices as a key component of any such agri-environment scheme., Competing Interests: The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (© 2021 The Authors.)

Understanding whether pollinator behaviors and species richness drive crop yields is a key area of investigation in pollination ecology. Using sunflower as a study species we describe variation in mean floral visitation times among bee taxa and test how interactions between bee richness and the proportion of bumblebees in localized communities impact seed yield. Seven bee genera commonly visited sunflower including Agapostemon, Bombus, Halictus, Lasioglossum, Megachile, Melissodes, and Svastra. Mean visitation times to sunflower varied across genera and Bombus and Halictus spp. spent the most time foraging on inflorescences, but the number of visits by Bombus spp. was the only parameter associated with increased yields. Experimental pollination deficit reduced seed development and yields, and these effects were stronger in stands of wild-type sunflower in the field compared to a confection variety grown in the greenhouse. Relationships between bee richness and pollination services differed for potted and wild sunflower: when bees had short-term access to potted sunflower, bee richness and relative Bombus abundances were not associated with pollination quotients. When bees had long-term access to wild sunflower, relative Bombus abundances predicted pollination services but were modified by site-level bee richness: as richness increased, the effects of Bombus abundance decreased. Our studies demonstrate that bee species richness is not always a clear predictor of pollination services; instead, our results underscore the importance of specific taxa when species richness is low (here, bumblebees), and show that the effects of bee functional groups important for pollination may be modified by changes in site-level species richness., (© The Author(s) 2021. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Extensively managed and flower-rich mountain hay meadows, hotspots of Europe's biodiversity, are subject to environmental and climatic gradients linked to altitude. While the shift of pollinators from bee- to fly-dominated communities with increasing elevation across vegetation zones is well established, the effect of highland altitudinal gradients on the community structure of pollinators within a specific habitat is poorly understood. We assessed wild bee and hoverfly communities, and their pollination service to three plant species common in mountain hay meadows, in eighteen extensively managed yellow oat grasslands ( Trisetum flavescens ) with an altitudinal gradient spanning approx. 300 m. Species richness and abundance of pollinators increased with elevation, but no shift between hoverflies and wild bees (mainly bumblebees) occurred. Seedset of the woodland cranesbill ( Geranium sylvaticum ) increased with hoverfly abundance, and seedset of the marsh thistle ( Cirsium palustre ) increased with wild bee abundance. Black rampion ( Phyteuma nigrum ) showed no significant response. The assignment of specific pollinator communities, and their response to altitude in highlands, to different plant species underlines the importance of wild bees and hoverflies as pollinators in extensive grassland systems., Competing Interests: All authors declared no conflict of interest., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

In Sardinia, the second largest Mediterranean island, 316 species of bees are known. Here, for the first time, the following 20 taxa are reported: Colletes cunicularius (Linnaeus, 1761), and C. eous Morice, 1904 (Colletidae); Andrena humilis Imhoff, 1832, A. granulosa Pérez, 1902, A. cineraria (Linnaeus, 1758), A . pallitarsis Pérez, 1903, A. rugulosa Stöckhert, 1935, A. savignyi Spinola, 1838, and A. tenuistriata Pérez, 1895 (Andrenidae); Sphecodes reticulatus Thomson, 1870 (Halictidae); Lithurgus tibialis Morawitz, 1875, Chelostoma emarginatum (Nylander, 1856), Dioxys cinctus (Jurine, 1807), Coelioxys caudatus Spinola, 1838, C. obtusus Pérez, 1884, and Megachile ericetorum (Lepeletier, 1841) (Megachilidae); and Nomada melathoracica Imhoff, 1834, N. pulchra Arnold, 1888, Eucera proxima Morawitz, 1875 and Tetralonia malvae (Rossi, 1790) (Apidae). N. pulchra is reported for the first time in Italy.

Bumble bees are among the most effective pollinators in orchards during the blooming period, yet they are often threatened by the high levels of pesticide use in apple production. This study aimed to evaluate the influence of landscape enhancements (e.g., hedgerows, flower strips) on bumble bee queens in apple orchards. Bumble bee queens from 12 orchards in southern Québec (Canada) were marked, released, and recaptured in the springs and falls of 2017 to 2019. Half of the 12 orchards had landscape enhancements. Apples were harvested in 2018 and 2019 to compare their quality (weight, diameter, sugar level, and seed number) in sites with and without landscape enhancements. Species richness, as well as the occurrence of three species out of eight, was higher in orchards with landscape enhancements than in orchards without such structures. The occurrence of Bombus ternarius was lower in orchards with high levels of pesticide use. Apples had fewer seeds when collected in orchards with landscape enhancements and were heavier in orchards that used more pesticides. Our work provides additional evidence that landscape enhancements improve bumble bee presence in apple orchards and should therefore be considered as a means to enhance pollination within farms.

In this study, we report the complete mitochondrial genomes (mtgenome) of Thyreus decorus , Ceratina okinawana and Amegilla calceifera , which are the first time of mtgenome report also for the genera Thyreus , Ceratina and Amegilla in the family Apidae. They contain 15,237, 15,207, and 17,728 bp, with AT content of 84.97%, 79.30%, and 84.63%, respectively. Each mtgenome includes 13 protein-coding genes (PCGs), 22 transfer RNA (tRNAs), two ribosomal RNA (16S and 12S rRNA) and an AT-rich control region (CR). The phylogenetic relationships of 45 species in the family were constructed using Bayesian Inference based on concatenated nucleotide sequences of 13 PCGs. Our study suggests that the subfamily Apinae is a paraphyletic group, with the genus Eucera claded into the subfamily Xylocopinae and the genera Amegilla and Thyreus into the subfamily Nomadinae. In Apinae, the genera Melipona and Bombus are significantly sister group, and the genus Apis is the sister group with Melipona + Bombus ., Competing Interests: The authors declare no conflict of interest. The authors alone are responsible for the content and writing of the paper., (© 2021 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group.)

Declining pollinator populations worldwide are attributed to multiple stressors, including the loss of quality forage. Habitat management in agricultural areas often targets honey bees ( Apis mellifera L.) specifically, with the assumption that native bees will benefit from an 'umbrella species' strategy. We tested this theory using a conservation physiology approach to compare the effects of landscape composition and floral dietary composition on the physiological status of honey bees and Melissodes desponsa in eastern South Dakota, USA. The total glycogen, lipid and protein concentrations were quantified from field collected bees. Next-generation sequencing of the trn L chloroplast gene from bee guts was used to evaluate dietary composition. The effects of landscape and dietary composition on macronutrient concentrations were compared between bee species. As the mean land-use patch area increased, honey bee glycogen levels increased, though M. desponsa experienced a decrease in glycogen. Protein levels decreased in honey bees as the largest patch index, a measure of single patch dominance, increased versus M. desponsa . Lipids in both species were unaffected by the measured landscape variables. Dietary analysis revealed that honey bees foraged preferentially on weedy non-native plant species, while M. desponsa sought out native and rarer species, in addition to utilizing non-native plants. Both species foraged on Asteraceae, Oleaceae and Fabaceae, specifically Melilotus sp. and Medicago sp. Dietary composition was not predictive of the macronutrients measured for either species. Together, these data highlight the management importance of including patch area in conservation recommendations, as bee species may have divergent physiological responses to landscape characteristics. While solitary bees may forage on weedy introduced plants in agricultural areas, robust strategies should also reincorporate native plant species, though they may not be preferred by honey bees, to maximize overall health and diversity of pollinator communities., (© The Author(s) 2020. Published by Oxford University Press and the Society for Experimental Biology.)

Meliponiculture, the keeping of domesticated stingless bees such as Geniotrigona thoracica (Smith, 1857) (Hymenoptera: Apidae), is an increasingly popular agricultural industry in Malaysia. This study reports the soldier fly (Diptera: Stratiomyidae) species of the genus Hermetia colonizing stingless bee colonies in Malaysia. The larvae were reared in the laboratory to the adult stage and identified through molecular and morphological approaches. Hermetia illucens (Linnaeus, 1758) and Hermetia fenestrata de Meijere, 1904 (Diptera: Stratiomyidae) were identified from the sample provided. Earlier records of stratiomyids in stingless bee nests were misidentified as H. illucens . This paper represents the first identified record of H. fenestrata colonizing a "spoiled" stingless bee colony. In addition, adult and larval morphological differences between both species and the roles of both species in bee nest decomposition are discussed., Competing Interests: The authors declare no conflict of interest.

Worldwide pollinator declines have dramatically increased our need to survey and monitor pollinator distributions and abundances. The giant honey bee, Apis laboriosa , is one of the important pollinators at higher altitudes of the Himalayas. This species has a restricted distribution along the Himalayas and neighbouring mountain ranges of Asia. Previous assessments of its distribution, published more than 20 years ago, were based on museum specimens. Since then, 244 additional localities have been revealed through field trips by the authors, publications, and websites. We present a revised distribution for A. laboriosa that better defines its range and extends it eastward to the mountains of northern Vietnam, southward along the Arakan Mountains to west-central Myanmar, into the Shillong Hills of Meghalaya, India, and northwestward in Uttarakhand, India. This species is generally found at elevations between 1000-3000 m a.s.l.. In northeastern India A. laboriosa colonies occur during summer at sites as low as 850 m a.s.l. and some lower elevation colonies maintain their nests throughout the winter. Finally, we report three regions in Arunachal Pradesh, India, and nine locations in northern Vietnam, where we observed workers of A. laboriosa and A. dorsata foraging sympatrically; their co-occurrence supports the species status of Apis laboriosa .

Vibrations play an important role in insect behaviour. In bees, vibrations are used in a variety of contexts including communication, as a warning signal to deter predators and during pollen foraging. However, little is known about how the biomechanical properties of bee vibrations vary across multiple behaviours within a species. In this study, we compared the properties of vibrations produced by Bombus terrestris audax (Hymenoptera: Apidae) workers in three contexts: during flight, during defensive buzzing, and in floral vibrations produced during pollen foraging on two buzz-pollinated plants ( Solanum , Solanaceae). Using laser vibrometry, we were able to obtain contactless measures of both the frequency and amplitude of the thoracic vibrations of bees across the three behaviours. Despite all three types of vibrations being produced by the same power flight muscles, we found clear differences in the mechanical properties of the vibrations produced in different contexts. Both floral and defensive buzzes had higher frequency and amplitude velocity, acceleration and displacement than the vibrations produced during flight. Floral vibrations had the highest frequency, amplitude velocity and acceleration of all the behaviours studied. Vibration amplitude, and in particular acceleration, of floral vibrations has been suggested as the key property for removing pollen from buzz-pollinated anthers. By increasing frequency and amplitude velocity and acceleration of their vibrations during vibratory pollen collection, foraging bees may be able to maximise pollen removal from flowers, although their foraging decisions are likely to be influenced by the presumably high cost of producing floral vibrations., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2020. Published by The Company of Biologists Ltd.)

The High Arctic bumble bee fauna is rather poorly known, while a growing body of recent molecular research indicates that several Arctic species may represent endemic lineages with restricted ranges. Such local endemics are in need of special conservation efforts because of the increasing anthropogenic pressure and climate changes. Here, we re-examine the taxonomic and biogeographic affinities of bumble bees from Novaya Zemlya using historical samples and recently collected materials (1895-1925 vs. 2015-2017). Three bumble bee species inhabit the Yuzhny (Southern) Island and the southern edge of Severny (Northern) Island of this archipelago: Bombusglacialis Friese, 1902, B.hyperboreus Schönherr, 1809, and B.pyrrhopygus Friese, 1902. Bombusglacialis shares three unique COI haplotypes that may indicate its long-term (pre-glacial) persistence on Novaya Zemlya. In contrast, Bombushyperboreus and B.pyrrhopygus share a rather low molecular divergence from mainland populations, with the same or closely related haplotypes as those from Arctic Siberia and Norway. A brief re-description of Bombuspyrrhopygus based on the newly collected topotypes is presented. Habitats, foraging plants and life cycles of bumble bees on Novaya Zemlya are characterized, and possible causes of extremely low bumble bee abundance on the archipelago are discussed. The species-poor bumble bee fauna of Novaya Zemlya is compared with those in other areas throughout the Arctic. The mean bumble bee species richness on the Arctic Ocean islands is three times lower than that in the mainland Arctic areas (3.1 vs. 8.6 species per local fauna, respectively). General linear models (GLMs) indicate that this difference can be explained by specific environmental conditions of insular areas. Our findings highlight that the insularity is a significant factor sharply decreasing species richness in bumble bee assemblages on the Arctic Ocean archipelagoes through colder climate (lower summer temperatures), prevalence of harsh Arctic tundra landscapes with poor foraging resources, and in isolation from the mainland.

Meliponiculture - the management of stingless bee colonies - is an expanding activity in Brazil with economic, social and environmental potential. However, unlike in apiculture, the pathogens that impact on meliponiculture remain largely unknown. In southern Brazil, every year at the end of the summer, managed colonies of the stingless bee Melipona quadrifasciata manifest a syndrome that eventually leads to collapse. Here we characterize the M. quadrifasciata virome using high-throughput sequencing, with the aim of identifying potentially pathogenic viruses, and test whether they are related to the syndrome outbreaks. Two paired viromes are explored, one from healthy bees and another from unhealthy ones. Each virome is built from metagenomes assembled from sequencing reads derived either from RNA or DNA. A total of 40 621 reads map to viral contigs of the unhealthy bees' metagenomes, whereas only 11 reads map to contigs identified as viruses of healthy bees. The viruses showing the largest copy numbers in the virome of unhealthy bees belong to the family Dicistroviridae - common pathogenic honeybee viruses - as well as Parvoviridae and Circoviridae, which have never been reported as being pathogenic in insects. Our analyses indicate that they represent seven novel viruses associated with stingless bees. PCR-based detection of these viruses in individual bees (healthy or unhealthy) from three different localities revealed a statistically significant association between viral infection and symptom manifestation in one meliponary. We conclude that although viral infections may contribute to colony collapses in the annual syndrome in some meliponaries, viruses spread opportunistically during the outbreak, perhaps due to colony weakness.

The East Himalaya is one of the world's most biodiverse ecosystems. However, very little is known about the abundance and distribution of many plant and animal taxa in this region. Bumble bees are a group of cold-adapted and high elevation insects that fulfil an important ecological and economical function as pollinators of wild and agricultural flowering plants and crops. The Himalayan mountain range provides ample suitable habitats for bumble bees. Systematic study of Himalayan bumble bees began a few decades ago and the main focus has centred on the western region, while the eastern part of the mountain range has received little attention and only a few species have been verified. During a three-year survey, more than 700 bumble bee specimens of 21 species were collected in Arunachal Pradesh, the largest of the north-eastern states of India. The material included a range of species that were previously known from a limited number of collected specimens, which highlights the unique character of the East Himalayan ecosystem. Our results are an important first step towards a future assessment of species distribution, threat, and conservation. Clear elevation patterns of species diversity were observed, which raise important questions about the functional adaptations that allow bumble bees to thrive in this particularly moist region in the East Himalaya.

Large carpenter bees are charismatic and ubiquitous flower visitors in the tropics and sub-tropics. Unlike honeybees and bumblebees that have been popular subjects of extensive studies on their neuroethology, behaviour and ecology, carpenter bees have received little attention. This review integrates what is known about their foraging behaviour as well as sensory, physiological and cognitive adaptations and is motivated by their versatility as flower visitors and pollinators. This is evident from their extremely generalist foraging and adeptness at handling diverse flower types as legitimate pollinators and as illegitimate nectar robbers. They purportedly use traplining to forage between isolated patches and are long-distance flyers over several kilometres suggesting well-developed spatial learning, route memory and navigational capabilities. They have a broad range of temperature tolerance and thermoregulatory capabilities which are likely employed in their forays into crepuscular and nocturnal time periods. Such temporal extensions into dim-light periods invoke a suite of visual adaptations in their apposition optics. Thus, we propose that carpenter bees are an excellent though understudied group for exploring the complex nature of plant-pollinator mutualisms from ecological and mechanistic perspectives.

The pharyngeal plate is a morphological complex with extensive anatomical variation among bees and, therefore, potential as a source of phylogenetic information. The pharyngeal plate of bees is divided into four morphologically distinct regions: sitophore, hypopharyngeal lobe, pharyngeal rods, and median oral plate. In this work we illustrate and document in detail for the first time the pharyngeal plate of 43 bee species, providing descriptions of the morphological variation and contrasting these findings with representatives of apoid wasps (Crabronidae and Sphecidae). We evaluate and discuss the potential of this structure as a rich source of morphological information in the context of bee phylogeny and any research potentially impacted by comparative morphological data. The shape of the hypopharyngeal lobe is highly variable among suprageneric taxa of bees and can be readily employed to characterise taxa at various levels. We argue that the global patterns in the variation of the pharyngeal plate can provide information for phylogenetic inference within bees and constructed and coded 10 characters that encompass the most noticeable morphological differences discussed herein., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Honey bees are globally distributed and have received increased attention due to their high economic and ecological value for pollination, their exceptional eusocial lifestyle and complex behavioral repertoire. Interestingly, most research on learning and memory in honey bees has been performed in the Western honey bee, Apis mellifera L., and other honey bee species were largely neglected. In the current study, we thus compared visual learning performance of A. mellifera and the Eastern honey bee, A. cerana Fabr., using the proboscis extension response (PER) paradigm. Workers of A. mellifera and A. cerana were differentially conditioned to two monochromatic light stimuli, with peak maxima at 435 and 528 nm. Both honey bee species were able to form an association between the color stimulus and a sugar reward and significantly distinguished between the two color stimuli in a differential discrimination test. However, besides similar performance levels during visual learning, A. cerana showed a reduced mid-term memory (tested after 2 h) compared to A. mellifera. Finally, performance of the visual PER conditioning in our study reached similar levels as found in olfactory PER conditioning, and we thus recommend the visual PER conditioning approach in addition to olfactory conditioning as a useful tool for studying species-specific learning and memory capabilities in honey bees under controlled laboratory conditions., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

The risk of ecosystem function degradation with biodiversity loss has emerged as a major scientific concern in recent years. Possible relationships between taxonomic diversity and magnitude and stability of ecosystem processes build upon species' functional characteristics, which determine both susceptibility to environmental change and contribution to ecosystem properties. The functional diversity within communities thus provides a potential buffer against environmental disturbance, especially for properties emerging from interactions among species. In complex plant-pollinator networks, distantly related taxa spanning a great trait diversity shape network architecture. Here, we address the question of whether network properties are maintained after habitat loss by complementary contributions of phylogenetically distant pollinator taxa. We quantified contributions of wild bees and hoverflies to network structure (connectance, network specialization, specialization asymmetry) in 32 calcareous grassland fragments varying in size. Although hoverflies are often regarded less susceptible to environmental change than wild bees, species richness of both taxa was similarly affected by habitat loss. The associated loss of 80% of interactions resulted in small and tightly connected networks, which was more strongly attributed to wild bee loss than hoverfly loss. Networks in small fragments were less specialized due to equivalent losses of species and interactions in both pollinators and plants. Because wild bee and hoverfly loss contributed similarly to declining network specialization, we conclude that trait diversity among distantly related pollinators does not necessarily provide insurance against functional homogenization during community disassembly following habitat loss., (© 2018 by the Ecological Society of America.)

Two increasingly popular approaches to reconstruct the Tree of Life involve whole transcriptome sequencing and the target capture of ultraconserved elements (UCEs). Both methods can be used to generate large, multigene datasets for analysis of phylogenetic relationships in non-model organisms. While targeted exon sequencing across divergent lineages is now a standard method, it is still not clear if UCE data can be readily combined with published transcriptomes. In this study, we evaluate the combination of UCEs and transcriptomes in a single analysis using genome-, transcriptome-, and UCE data for 79 bees in the largest and most biologically diverse bee family, Apidae. Using existing tools, we first developed a workflow to assemble phylogenomic data from different sources and produced two large nucleotide matrices of combined data. We then reconstructed the phylogeny of the Apidae using concatenation- and coalescent-based methods, and critically evaluated the resulting phylogenies in the context of previously published genetic, genomic, and morphological data sets. Our estimated phylogenetic trees are robustly supported and largely congruent with previous molecular hypotheses, from deep nodes to shallow species-level phylogenies. Moreover, the combined approach allows us to resolve controversial nodes of the apid Tree of Life, by clarifying the relationships among the genera of orchid bees (Euglossini) and the monophyly of the Centridini. Additionally, we present novel phylogenetic evidence supporting the monophyly of the diverse clade of cleptoparasitic Apidae and the placement of two enigmatic, oil-collecting genera (Ctenoplectra and Tetrapedia). Lastly, we propose a revised classification of the family Apidae that reflects our improved understanding of apid higher-level relationships., (Copyright © 2018 The Authors. Published by Elsevier Inc. All rights reserved.)

We reassessed species diversity and genetic structure in Korean bumble bees using DNA barcode analyses of 484 cytochrome c oxidase subunit I (COI) sequences from 24 morphospecies. Based on COI, all of the Korean species formed distinct clades in the phylogenetic trees, except for Bombus (Megabombus) koreanus in the maximum likelihood tree. Five species exhibited low interspecific genetic distances (range: 1.2-2.7%), indicating that they are recently diverged species. COI data could not be used to identify bumble bees at the subspecies level. For the dominant species, most local populations in Korea were panmictic and were more closely related to continental populations than to allopatric populations. Furthermore, sympatric haplotypes within Korea could be distinguished. We detected B. (Megabombus) diversus in South Korea for the first time. Our results demonstrate that DNA barcoding is a useful technique for species recognition and for allopatric and sympatric haplotype detection in bumble bees.

This paper introduces both a hardware and a software system designed to allow low-cost electronic monitoring of social insects using RFID tags. Data formats for individual insect identification and their associated experiment are proposed to facilitate data sharing from experiments conducted with this system. The antennas' configuration and their duty cycle ensure a high degree of detection rates. Other advantages and limitations of this system are discussed in detail in the paper., Competing Interests: The authors declare no conflict of interest.

Background: The native bee fauna of Mississippi, USA has been historically poorly sampled, but is of particular relevance to determine range limits for species that occur in the southern United States. Currently published literature includes 184 species of bees that occur within the state of Mississippi. Additions to the list of native bees known for Mississippi are reported with notes on range, ecology and resources for identification., New Information: The geographic ranges of seven additional species are extended into the state of Mississippi: Andrena (Melandrena) obscuripennis Smith, 1853, Anthemurgus passiflorae Robertson, 1902, Dieunomia bolliana (Cockerell 1910), Diadasia (Diadasia) enavata (Cresson 1872), Peponapis crassidentata (Cockerell 1949), Triepeolus subnitens Cockerell and Timberlake, 1929 and Brachynomada nimia (Snelling and Rozen 1987). These records raise the total number of published species known from the state to 191. Anthemurgus and Brachynomada are also genera new to Mississippi.

Floral volatile organic compounds (VOCs) play important roles in plant-pollinator interactions. We investigated the reproductive ecology and floral VOCs of Zygopetalinae orchids to understand the relationship between floral scents and pollinators. We performed focal observations, phenological censuses and breeding system experiments in eight species in southeast Brazil. Floral scents were collected and analysed using SPME/GC-MS. We performed multivariate analyses to group species according to affinities of their VOCs and define compounds associated to each plant. Dichaea cogniauxiana was pollinated by weevils which use their developing ovules, while D. pendula was pollinated by the same weevils and perfume-collecting male euglossine bees. The other species were deceit-pollinated by bees. Zygopetalum crinitum was pollinated by carpenter bees, while W. warreana, Z. mackayi and Z. maxillare were bumblebee-pollinated. The latter was also pollinated by Centris confusa. Breeding system varied widely with no association to any pollinator group. Most VOCs are common to other floral scents. Zygopetalum crinitum presented an exclusive blend of VOCs, mainly composed of benzenoids. The scents of Pabstia jugosa, Promenaea xanthina and the Zygopetalum spp. were similar. The bumblebee-pollinated species have flowering periods partially overlapped, thus neither phenology nor pollinators constitute hybridization barriers among these species. Euglossines are not the only pollinators of Zygopetalinae. Different VOCs, size and lifespan of flowers are associated with distinct pollinators. A distinctive VOC bouquet may determine specialisation in carpenter bees or male euglossines within bee-pollinated flowers. Finally, visitation of deceit-pollinated flowers by perfume-collecting euglossines allows us to hypothesise how pollination by this group of bees had evolved.

In urban landscapes, gardens provide refuges for bee diversity, but conservation potential may depend on local and landscape features. Foraging and population persistence of bee species, as well as overall pollinator community structure, may be supported by the abundance, richness, and spatial distribution of floral resources. Floral resources strongly differ in urban gardens. Using hand netting and pan traps to survey bees, we examined whether abundance, richness, and spatial distribution of floral resources, as well as ground cover and garden landscape surroundings influence bee abundance, species richness, and diversity on the central coast of California. Differences in floral abundance and spatial distribution, as well as urban cover in the landscape, predicted different bee community variables. Abundance of all bees and of honeybees (Apis mellifera) was lower in sites with more urban land cover surrounding the gardens. Honeybee abundance was higher in sites with patchy floral resources, whereas bee species richness and bee diversity was higher in sites with more clustered floral resources. Surprisingly, bee species richness and bee diversity was lower in sites with very high floral abundance, possibly due to interactions with honeybees. Other studies have documented the importance of floral abundance and landscape surroundings for bees in urban gardens, but this study is the first to document that the spatial arrangement of flowers strongly predicts bee abundance and richness. Based on these findings, it is likely that garden managers may promote bee conservation by managing for floral connectivity and abundance within these ubiquitous urban habitats.

Nosema ceranae and Nosema apis are microsporidia which play an important role in the epidemiology of honeybee microsporidiosis worldwide. Nosemiasis reduces honeybee population size and causes significant losses in honey production. To the best of our knowledge, limited information is available about the prevalence of nosemiasis in Italy. In this research, we determined the occurrence of Nosema infection in Central Italy. Thirty-eight seemingly healthy apiaries (2 to 4 hives each) were randomly selected and screened from April to September 2014 ( n  = 11) or from May to September 2015 ( n  = 27). The apiaries were located in six areas of Central Italy, including Lucca ( n  = 11), Massa Carrara ( n  = 9), Pisa ( n  = 9), Leghorn ( n  = 7), Florence ( n  = 1), and Prato ( n  = 1) provinces. Light microscopy was carried out according to current OIE recommendations to screen the presence of microsporidiosis in adult worker honeybees. Since the morphological characteristics of N. ceranae and N. apis spores are similar and can hardly be distinguished by optical microscopy, all samples were also screened by multiplex polymerase chain reaction (M-PCR) assay based on 16S rRNA - gene - targeted species-specific primers to differentiate N. ceranae from N. apis. Furthermore, PCR-positive samples were also sequenced to confirm the species of amplified Nosema DNA. Notably, Nosema spores were detected in samples from 24 out of 38 (63.2%, 95% CI: 47.8-78.5%) apiaries. Positivity rates in single provinces were 10/11, 8/9, 3/9, 1/7, or 1/1 ( n  = 2). A full agreement (Cohen's Kappa = 1) was assessed between microscopy and M-PCR. Based on M-PCR and DNA sequencing results, only N. ceranae was found. Overall, our results highlighted that N. ceranae infection occurs frequently in the cohort of honeybee populations that was examined despite the lack of clinical signs. These findings suggest that colony disease outbreaks might result from environmental factors that lead to higher susceptibility of honeybees to this microsporidian.

The taxonomy of the genus Xylocopa Latreille, 1802, subgenus Xenoxylocopa Hurd & Moure, 1963, is reviewed. There is a single valid species in this subgenus, Xylocopa (Xenoxylocopa) inconstans Smith, 1874, which is widely distributed throughout sub-Saharan Africa, from Senegal to Ethiopia and south to northern Republic of South Africa. Synonyms of Xylocopa inconstans include Xylocopa abyssinica Radoszkowski, 1899, proposed for a male specimen from Ethiopia, as well as three names proposed for females with yellow (rather than white) dorsal pubescence: Mesotrichia chiyakensis Cockerell, 1908 (new synonym), Xylocopa inconstans var. flavescens Vachal, 1899, and Xylocopa inconstans var. flavocincta Friese, 1909. Quantitative analyses of body measurements and examination of male reproductive structures support the new synonymy of Mesotrichia chiyakensis with Xylocopa inconstans . Males and females of Xylocopa (Xenoxylocopa) inconstans are illustrated, along with male reproductive structures, and diagnostic characters and keys are provided to separate the males and females of Xylocopa (Xenoxylocopa) inconstans from those of species in other closely-allied African subgenera of the genus Xylocopa .

We report 18 species of wild bees plus the domesticated honeybee from the Azores, which adds nine species to earlier lists. One species, Hylaeus azorae , seems to be a single island endemic, and three species are possibly native ( Colletes eous , Halictus villosulus , and Hylaeus pictipes ). All the remaining bee species are most likely accidental introductions that arrived after human colonization of the archipelago in the 15 th century. Bee diversity in the Azores is similar to bee diversity of Madeira and Cape Verde but nearly ten times lower than it is in the Canary Islands.

Numerous species of solitary bees and wasps build linear nests with only one entrance. Developing insects must orient themselves inside their nest to choose the correct direction in which to emerge. Misorientation and chewing towards the dead end of the nest can result in significant mortality. Most insects position themselves towards the nest entrance during cocoon construction; however, some individuals are misoriented. We tested whether imagines can examine and possibly correct their orientation after emerging from their cocoons. Males were usually able to correct their misoriented position based on the shape of the cell wall and emerged through the correct entrance, whereas most females pursued the direction that they faced in their cocoons. We suggest that there can be more than one time point during development when bees can control their position in relation to the nest entrance and that the importance of these time points varies between sexes., (© The Author 2017. Published by Oxford University Press on behalf of the Entomological Society of America.)

We identified 41 species of native bees from a total of 1,083 specimens collected at cultivated highbush blueberry plantings throughout Rhode Island in 2014 and 2015. Andrena spp., Bombus spp., and Xylocopa virginica (L.) were collected most often. Bombus griseocollis (DeGeer), B. impatiens Cresson, B. bimaculatus Cresson, B. perplexus Cresson, and Andrena vicina Smith collected the largest mean numbers of blueberry pollen tetrads. The largest mean percent blueberry pollen loads were carried by the miner bees Andrena bradleyi Viereck (91%), A. carolina Viereck (90%), and Colletes validus Cresson (87%). The largest mean total pollen grain loads were carried by B. griseocollis (549,844), B. impatiens (389,558), X. virginica (233,500), and B. bimaculatus (193,132). Xylocopa virginica was the fourth and fifth most commonly collected bee species in 2014 and 2015, respectively. They exhibit nectar robbing and females carried relatively low blueberry pollen loads (mean 33%). Overall, we found 10 species of bees to be the primary pollinators of blueberries in Rhode Island., (© The Authors 2016. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For Permissions, please email: journals.permissions@oup.com.)

In face of a dramatic decline of wild bee species in many rural landscapes, potential conservation functions of urban areas gain importance. Yet effects of urbanization on pollinators, and in particular on wild bees, remain ambiguous and not comprehensively understood. This is especially true for amenity grassland and extensively managed wastelands within large-scale residential housing areas. Using Berlin as a study region, we aimed to investigate (a) if these greenspaces are accepted by wild bee assemblages as foraging habitats; (b) how assemblage structure of bees and individual bee species are affected by different habitat (e.g., management, flower density) and urban matrix variables (e.g., isolation, urbanization); and (c) to what extent grassland restoration can promote bees in urban environments. In summer 2012, we collected 62 bee species belonging to more than 20% of the taxa known for Berlin. Urbanization significantly affected species composition of bees; 18 species were affiliated to different levels of urbanization. Most bee species were not affected by any of the environmental variables tested, and urbanization had a negative effect only for one bee species. Further, we determined that restoration of diverse grasslands positively affected bee species richnesss in urban environments. We conclude that differently structured and managed greenspaces in large-scale housing areas can provide additional foraging habitats and refuges for pollinators. This supports approaches towards a biodiversity friendly management within urban regions and may be of particular importance given that anthropogenic pressure is increasing in many rural landscapes., Competing Interests: The authors declare there are no competing interests.

As part of a research project on the food deception strategy in Orchis militaris (L.), the objective of this study was to identify insect visitors and potential pollinators of this orchid species in Belgium. In 2013, insects were collected over 2 d per site in five localities distributed in Southern Belgium (Wallonia). A total of 104 insects belonging to 49 species were caught. Dipterans were the most abundant visitors (50% of total specimens), followed by Hymenopterans (32%). Rhingia campestris Meigen, Bombylius venosus Mikan, Apis mellifera (L.), and Bombus lapidarius (L.) were the most abundant species. Only five specimens bore one to more than 10 pollinia: four honeybees (A. mellifera) and one bumblebee worker (B. lapidarius). These two species should be considered as potential pollinators in the study area, but probably not confirmed ones., (© The Authors 2016. Published by Oxford University Press on behalf of Entomological Society of America.)

Parental care is a behavior that increases the growth and survival of offspring, often at a cost to the parents' own survival and/or future reproduction. In this study, we focused on nest guarding, which is one of the most important types of extended parental care; we studied this behavior in two solitary bee species of the genus Ceratina with social ancestors. We performed the experiment of removing the laying female, who usually guards the nest after completing its provisioning, to test the effects of nest guarding on the offspring survival and nest fate. By dissecting natural nests, we found that Ceratina cucurbitina females always guarded their offspring until the offspring reached adulthood. In addition, the females of this species were able to crawl across the nest partitions and inspect the offspring in the brood cells. In contrast, several Ceratina chalybea females guarded their nests until the offspring reached adulthood, but others closed the nest entrance with a plug and deserted the nest. Nests with a low number of provisioned cells were more likely to be plugged and abandoned than nests with a higher number of cells. The female removal experiment had a significantly negative effect on offspring survival in both species. These nests frequently failed due to the attacks of natural enemies (e.g., ants, chalcidoid wasps, and other competing Ceratina bees). Increased offspring survival is the most important benefit of the guarding strategy. The abandonment of a potentially unsuccessful brood might constitute a benefit of the nest plugging behavior. The facultative nest desertion strategy is a derived behavior in the studied bees and constitutes an example of an evolutionary reduction in the extent of parental care.

Resource concentration effects occur when high resource density patches attract and support more foragers than low density patches. In contrast, resource dilution effects can occur if high density patches support fewer consumers. In this study, we examined the foraging rates of pollinators and seed predators on two perennial plant species (Rudbeckia triloba and Verbena stricta) as functions of resource density. Specifically, we examined whether resource-dense patches (densities of flower and seeds on individual plants) resulted in greater visitation and seed removal rates, respectively. We also examined whether foraging rates were context-dependent by conducting the study in two sites that varied in resource densities. For pollinators, we found negative relationships between the density of flowers per plant and visitation rates, suggesting dilution effects. For seed predators, we found positive relationships consistent with concentration effects. Saturation effects and differences in foraging behaviors might explain the opposite relationships; most of the seed predators were ants (recruitment-based foragers), and pollinators were mostly solitary foragers. We also found that foraging rates were site-dependent, possibly due to site-level differences in resource abundance and consumer densities. These results suggest that these two plant species may benefit from producing as many flowers as possible, given high levels of pollination and low seed predation.

The cost of an uncontrolled incursion of invasive alien species (IAS) arising from undetected entry through ports can be substantial, and knowledge of port-specific risks is needed to help allocate limited surveillance resources. Quantifying the establishment likelihood of such an incursion requires quantifying the ability of a species to enter, establish, and spread. Estimation of the approach rate of IAS into ports provides a measure of likelihood of entry. Data on the approach rate of IAS are typically sparse, and the combinations of risk factors relating to country of origin and port of arrival diverse. This presents challenges to making formal statistical inference on establishment likelihood. Here we demonstrate how these challenges can be overcome with judicious use of mixed-effects models when estimating the incursion likelihood into Australia of the European (Apis mellifera) and Asian (A. cerana) honeybees, along with the invasive parasites of biosecurity concern they host (e.g., Varroa destructor). Our results demonstrate how skewed the establishment likelihood is, with one-tenth of the ports accounting for 80% or more of the likelihood for both species. These results have been utilized by biosecurity agencies in the allocation of resources to the surveillance of maritime ports., (© 2015 Society for Risk Analysis.)

Sexual selection predicts that individuals competing for access to sexual partners should maximize their chances of mating by looking for sites where the chances of finding partners are more likely to occur. However, males of stingless bees have been observed sharing nonspecific reproductive aggregations. This uncommon behavior appears to confer no obvious increase of individual fitness. It has been suggested that this reproductive strategy is due to the similarity between male odors common to different stingless bee species. Cuticular hydrocarbons (CHCs) are candidate odors of interest because their nonvolatile pheromone nature allows them to play an important role in sexual behavior and species recognition. Here, we review the literature to evaluate whether any phylogenetic patterns exist among male stingless bees that aggregate with closely or distantly related species. We also compared the CHC profiles of males of Neotropical stingless bee species (Plebeia sp. Schwarz, Trigona spinipes (F.), Tetragona clavipes (F.), Nannotrigona testaceicornis (Lepeletier), Scaptotrigona aff. depilis (Moure), Tetragonisca angustula (Latreille), and Melipona subnitida (Ducke) to reveal any chemical similarities among their male odors. We found males of 21 stingless bee species involved in interspecific interactions mainly from Neotropical and Indo-Malayan/Australasian regions. Alien males did not necessarily visit host aggregations of closely related species. Furthermore, the CHC profiles of different studied species were very distinct from each other and do not overlapped at all. It is unclear yet why this apparently nonadaptive behavior carried out by some stingless bee males., (© The Author 2015. Published by Oxford University Press on behalf of the Entomological Society of America.)

As part of an ongoing research project on the pollination networks in European heathlands, the objective of this study was to assess the insect visitor guild on Calluna vulgaris (L.) Hull (Ericaceae). We focused the study on a region renowned for its largely well-preserved heathlands, the Cévennes National Park, Southern France. In 2013, flower visitors were observed over 3 d per site, in four heathland sites at mont Lozère. Honeybees (Apis mellifera L.) were the main visitors (62-88% of total visitors). Besides honeybees, a high diversity of visitors was detected with 57 different species identified (42 Diptera and 15 Hymenoptera). Hoverflies (Syrphidae, Diptera) visitors were abundant and diverse, especially individuals belonging to the genera Eristalis and Episyrphus. The reported diversity of visitors was probably due to the preservation of large heathland areas at mont Lozère and to the generalist pollination system of C. vulgaris., (© The Author 2015. Published by Oxford University Press on behalf of the Entomological Society of America.)