Variation in floral morphology, histochemistry, and floral visitors of three sympatric morning glory species.

Three morning glory species in the genus Argyreia Lour., A. lycioides (Choisy) Traiperm & Rattanakrajang, A. mekongensis Gagnep & Courchet, and A. versicolor (Kerr) Staples & Traiperm, were found co-occurring and co-flowering. Argyreia mekongensis and A. versicolor are rare, while A. lycioides is near threatened and distributed throughout Myanmar and Thailand. We investigated key floral characters (floral morphology and phenology, as well as the micromorphology of the floral nectary disc and staminal trichomes) and screened for important chemical compounds hypothesized to contribute to pollinator attraction. Our findings demonstrate that some aspects of floral morphology (e.g., corolla size, limb presence, and floral color) of the three studied congeners exhibit significant differences. Moreover, pollinator composition appears to be influenced by floral shape and size; morning glory species with wider corolla tubes were pollinated by larger bees. The morphology of the floral nectary disc was similar in all species, while variation in staminal trichomes was observed across species. Glandular trichomes were found in all three species, while non-glandular trichomes were found only in A. versicolor. Histochemical results revealed different compounds in the floral nectary and staminal trichomes of each species, which may contribute to both floral attraction and defense. These findings demonstrate some segregation of floral visitors among sympatric co-flowering morning glory species, which appears to be influenced by the macro- and micromorphology of flowers and their chemical compounds. Moreover, understanding the floral morphology and chemical attractants of these sympatric co-flowering Argyreia species may help to maintain their common pollinators in order to conserve these rare and endangered species, especially A. versicolor.

Bee morphology: A skeletomuscular anatomy of Thyreus (Hymenoptera: Apidae).

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.

Body-size-dependent effects of landscape-level resource energetics on pollinator abundance in woodland remnants.

Land use change alters floral resource availability, thereby contributing to declines in important pollinators. However, the severity of land use impact varies by species, influenced by factors such as dispersal ability and resource specialization, both of which can correlate with body size. Here. we test whether floral resource availability in the surrounding landscape (the 'matrix') influences bee species' abundance in isolated remnant woodlands, and whether this effect varies with body size. We sampled quantitative flower-visitation networks within woodland remnants and quantified floral energy resources (nectar and pollen calories) available to each bee species both within the woodland and the matrix. Bee abundance in woodland increased with floral energy resources in the surrounding matrix, with strongest effects on larger-bodied species. Our findings suggest important but size-dependent effects of declining matrix floral resources on the persistence of bees in remnant woodlands, highlighting the need to incorporate landscape-level floral resources in conservation planning for pollinators in threatened natural habitats.

Deep learning for identifying bee species from images of wings and pinned specimens.

One of the most challenging aspects of bee ecology and conservation is species-level identification, which is costly, time consuming, and requires taxonomic expertise. Recent advances in the application of deep learning and computer vision have shown promise for identifying large bumble bee (Bombus) species. However, most bees, such as sweat bees in the genus Lasioglossum, are much smaller and can be difficult, even for trained taxonomists, to identify. For this reason, the great majority of bees are poorly represented in the crowdsourced image datasets often used to train computer vision models. But even larger bees, such as bumble bees from the B. vagans complex, can be difficult to separate morphologically. Using images of specimens from our research collections, we assessed how deep learning classification models perform on these more challenging taxa, qualitatively comparing models trained on images of whole pinned specimens or on images of bee forewings. The pinned specimen and wing image datasets represent 20 and 18 species from 6 and 4 genera, respectively, and were used to train the EfficientNetV2L convolutional neural network. Mean test precision was 94.9% and 98.1% for pinned and wing images respectively. Results show that computer vision holds great promise for classifying smaller, more difficult to identify bees that are poorly represented in crowdsourced datasets. Images from research and museum collections will be valuable for expanding classification models to include additional species, which will be essential for large scale conservation monitoring efforts.

Influence of honey bee (Apis mellifera) breeding on wing venation in Serbia and neighbouring countries.

In order to improve the productivity of honey bees (Apis mellifera), some of their traits are selected by breeding. On one hand, breeding is mainly based on the natural geographical variation of this species; on the other hand, mass production and distribution of artificially selected queens can significantly affect the natural geographic variation of honey bees. In this study, we have compared honey bee wings originating from breeding and non-breeding populations in Serbia. In the comparison, we have also used data from a large area of south-eastern Europe. The wings were measured using the 19 landmarks indicated on the wing images. The coordinates were analysed using the methodology of geometric morphometrics. We found that honey bees obtained from honey bee queen breeder differed in wing venation from surrounding populations, which are under natural selection. Therefore, we argue against including populations under artificial selection in the analysis of the natural geographical variation of honey bees. In our analysis of non-breeding samples, we found that in south-eastern Europe there is continuous variation in wing venation and no clear boundaries between A. m. carnica, A. m. cecropia, and A. m. macedonica.

Heat and desiccation tolerances predict bee abundance under climate change.

Climate change could pose an urgent threat to pollinators, with critical ecological and economic consequences. However, for most insect pollinator species, we lack the long-term data and mechanistic evidence that are necessary to identify climate-driven declines and predict future trends. Here we document 16 years of abundance patterns for a hyper-diverse bee assemblage1 in a warming and drying region2, link bee declines with experimentally determined heat and desiccation tolerances, and use climate sensitivity models to project bee communities into the future. Aridity strongly predicted bee abundance for 71% of 665 bee populations (species × ecosystem combinations). Bee taxa that best tolerated heat and desiccation increased the most over time. Models forecasted declines for 46% of species and predicted more homogeneous communities dominated by drought-tolerant taxa, even while total bee abundance may remain unchanged. Such community reordering could reduce pollination services, because diverse bee assemblages typically maximize pollination for plant communities3. Larger-bodied bees also dominated under intermediate to high aridity, identifying body size as a valuable trait for understanding how climate-driven shifts in bee communities influence pollination4. We provide evidence that climate change directly threatens bee diversity, indicating that bee conservation efforts should account for the stress of aridity on bee physiology.

Unexpected worker mating and colony-founding in a superorganism.

The emergence of caste-differentiated colonies, which have been defined as 'superorganisms', in ants, bees, and wasps represents a major transition in evolution. Lifetime mating commitment by queens, pre-imaginal caste determination and lifetime unmatedness of workers are key features of these animal societies. Workers in superorganismal species like honey bees and many ants have consequently lost, or retain only vestigial spermathecal structures. However, bumble bee workers retain complete spermathecae despite 25-40 million years since their origin of superorganismality, which remains an evolutionary mystery. Here, we show (i) that bumble bee workers retain queen-like reproductive traits, being able to mate and produce colonies, underlain by queen-like gene expression, (ii) the social conditions required for worker mating, and (iii) that these abilities may be selected for by early queen-loss in these annual species. These results challenge the idea of lifetime worker unmatedness in superorganisms, and provide an exciting new tool for the conservation of endangered bumble bee species.

Anatomical, phenological and genetic aspects of the host-parasite relationship between Andrena vaga (Hymenoptera) and Stylops ater (Strepsiptera).

Stylops ater is an endoparasite of the mining bee Andrena vaga with extreme sexual dimorphism and hypermetamorphosis. Its population structure, parasitization mode, genetic diversity and impact on host morphology were examined in nesting sites in Germany to better understand this highly specialized host­parasite interaction. The shift in host emergence due to stylopization was proven to be especially strong in A. vaga. Around 10% of bees hosted more than 1 Stylops, with at maximum 4. A trend in Stylops' preference for hosts of their own sex and a sex-specific position of extrusion from the host abdomen was found. Invasion of Andrena eggs by Stylops primary larvae was depicted for the first time. Cephalothoraces of female Stylops were smaller in male and pluristylopized hosts, likely due to lower nutrient supply. The genes H3, 18S and cytochrome c oxidase subunit 1 were highly conserved, revealing near-absence of local variation within Stylops. Ovaries of hosts with male Stylops contained poorly developed eggs while those of hosts with female Stylops were devoid of visible eggs, which might be due to a higher protein demand of female Stylops. Male Stylops, which might have a more energy-consuming development, led to a reduction in head width of their hosts. Host masculinization was present in the leaner shape of the metabasitarsus of stylopized females and is interpreted as a by-product of manipulation of the host's endocrine system to shift its emergence. Stylopization intensified tergal hairiness, most strongly in hosts with female Stylops, near the point of parasite extrusion, hinting towards substance-induced host manipulation.

The potential and realized foraging movements of bees are differentially determined by body size and sociality.

Reversing biodiversity declines requires a better understanding of organismal mobility, as movement processes dictate the scale at which species interact with the environment. Previous studies have demonstrated that species foraging ranges, and therefore, habitat use increases with body size. Yet, foraging ranges are also affected by other life-history traits, such as sociality, which influence the need of and ability to detect resources. We evaluated the effect of body size and sociality on potential and realized foraging ranges using a compiled dataset of 383 measurements for 81 bee species. Potential ranges were larger than realized ranges and increased more steeply with body size. Highly eusocial species had larger realized foraging ranges than primitively eusocial or solitary taxa. We contend that potential ranges describe species movement capabilities, whereas realized ranges depict how foraging movements result from interactions between species traits and environmental conditions. Furthermore, the complex communication strategies and large colony sizes in highly eusocial species may facilitate foraging over wider areas in response to resource depletion. Our findings should contribute to a greater understanding of landscape ecology and conservation, as traits that influence movement mediate species vulnerability to habitat loss and fragmentation.

Do the different life history strategies of ants and honeybees determine fat body morphology?

The separation of two sister groups such as ants and bees in the Cretaceous involved the development of distinctive characteristics to occupy separate ecological niches. From the point of view of biology and ecology, it is important to see how different life history strategies affect the physiology of these insects. The fat body is the most metabolically important tissue in the organism of each insect. Therefore, we conducted a comparative analysis of the morphological image of the subcuticular fat body in different localisation/segments in Formica (Seviformica) cinerea and Apis mellifera mellifera foragers, because of the similarity of their functions in colonies. We observed that the fat bodies of ants and bees were composed of the same cell types: trophocytes and oenocytes. However, in each of the segments, the fat body cells in ants were bigger and there were fewer of them in comparison with bees. The dorsal part of the fat body of ants had a bilayer structure, where the outer layer was formed by binucleated oenocytes. Binucleated oenocytes were also found in the inner layer near the heart and tracheole. In bees, the fat body was unilayered and the trophocytes and oenocytes were present side by side. The similarities and, in particular, the differences in the structure of the fat body are the adaptation of these sister groups to life in a diverse environment.

A new bee species of the genus Dasypoda Latreille (Hymenoptera, Apoidea) from Northwest Africa with comparative remarks on the subgenus Microdasypoda Michez.

North Africa, with its vast array of ecosystems and reliefs, constitutes a remarkable place to explore and describe the diversity of wild bees. In this paper, a new bee species of the genus Dasypoda Latreille (Hymenoptera, Apoidea, Melittidae), D. schwarzi Radchenko et Michez sp. nov., is described from the Atlas Mountains area (Morocco and Tunisia). This species belongs to the subgenus Microdasypoda Michez and is phenotypically related to D. brevicornis Pérez, but differs from all other species of this subgenus by the structure of the male genitals, the metasomal sterna, and by its overall hair colour. A detailed comparative diagnosis of D. schwarzi with the other four species of this subgenus is provided, as well as a key to the males of Microdasypoda, and a correction to the diagnosis of the subgenus. This new species is the fortieth described Dasypoda species and should be looked for in other mountain regions of Northwest Africa, such as in the Algerian Atlas where it could be present.

Taxonomic studies on Dasyproctus Lepeletier & Brullé (Hymenoptera: Crabronidae) from North Vietnam with description of two new species.

Seven species of Dasyproctus Lepeletier & Brullé, 1835 (Hymenoptera: Crabronidae) are recorded from North Vietnam. Of these, D. vietnamensis and D. longi are new species, and D. idrieus (Cameron) and D. pentheri Leclercq are recorded for the first time from the country. Keys to both sexes of Dasyproctus from North Vietnam based on morphological characters are given, and new distributional records are also presented.

Effect of comb age on cell measurements and worker body size.

The honey bees (Apis mellifera L.) mainly use beeswax (comb) for brood rearing and food storage. Changes in the color and cell dimensions occur due to repeated food storage and brood rearing in the comb. The study aimed to determine the changes in comb cell measurements and worker body sizes in relation to comb age. For this purpose, the cell measurements of combs at age zero (wax foundation), 1, 2, 3, 4, 5, and 6 years and the body size of workers reared in them were estimated. The weight of the comb, the height of the cell base, and the weight of accumulated substances in the cell significantly increased with time. Comb age had negative effects on the cell diameter, cell depth, cell volume, cell honey or pollen capacity, and newly emerged worker body weight. Significant negative correlations were observed between the accumulated substances in a cell and the cell diameter, cell depth, and cell size, while significant positive correlations were observed among the cell volume, cell diameter, cell depth, cell honey capacity, cell pollen capacity, and worker body weight. It can be concluded that the dimensions of the comb cells and worker body size changed with the age of the comb. The obtained results recommend beekeepers to replace combs aged more than 3 years with a new comb to allow large workers to gather more nectar and pollen, rear a larger brood, and store more honey.

Attack of the dark clones the genetics of reproductive and color traits of South African honey bees (Apis mellifera spp.).

The traits of two subspecies of western honey bees, Apis mellifera scutellata and A.m. capensis, endemic to the Republic of South Africa (RSA), are of biological and commercial relevance. Nevertheless, the genetic basis of important phenotypes found in these subspecies remains poorly understood. We performed a genome wide association study on three traits of biological relevance in 234 A.m. capensis, 73 A.m. scutellata and 158 hybrid individuals. Thirteen markers were significantly associated to at least one trait (P ≤ 4.28 × 10-6): one for ovariole number, four for scutellar plate and eight for tergite color. We discovered two possible causative variants associated to the respective phenotypes: a deletion in GB46429 or Ebony (NC_007070.3:g.14101325G>del) (R69Efs*85) and a nonsense on GB54634 (NC_007076.3:g.4492792A>G;p.Tyr128*) causing a premature stop, substantially shortening the predicted protein. The mutant genotypes are significantly associated to phenotypes in A.m. capensis. Loss-of-function of Ebony can cause accumulation of circulating dopamine, and increased dopamine levels correlate to ovary development in queenless workers and pheromone production. Allelic association (P = 1.824 x 10-5) of NC_007076.3:g.4492792A>G;p.Tyr128* to ovariole number warrants further investigation into function and expression of the GB54634 gene. Our results highlight genetic components of relevant production/conservation behavioral phenotypes in honey bees.

Body size allometry impacts flight-related morphology and metabolic rates in the solitary bee Megachile rotundata.

Body size is related to many aspects of life history, including foraging distance and pollination efficiency. In solitary bees, manipulating the amount of larval diet produces intraspecific differences in adult body size. The goal of this study was to determine how body size impacts metabolic rates, allometry, and flight-related morphometrics in the alfalfa leafcutting bee, Megachile rotundata. By restricting or providing excess food, we produced a range of body sizes, which allowed us to test the effect of body size on allometry, the power required for flight, and amount of energy produced, as measured indirectly through CO2 emission. The power required during flight was predicted using the flight biomechanical formulas for wing loading and excess power index. We found larger bees had higher absolute metabolic rates at rest and during flight, but smaller bees had higher mass-specific metabolic rates at rest. During flight, bees did not have size-related differences in mass-specific metabolic rate. As bees increase in size, their thorax and abdomens become disproportionately larger, while their wings (area, and length) become disproportionately smaller. Smaller bees had more power available during flight as demonstrated by flight biomechanical formulas. Smaller body size was advantageous because of a reduced power requirement for flight with no metabolic cost.

Can plants fool artificial intelligence? Using machine learning to compare between bee orchids and bees.

Bee orchids have long been an excellent example of how dishonest signal works in plant-animal interaction. Many studies compared the flower structures that resemble female bees, leading toward pseudo-copulation of the male bees on the flower. Using Machine Learning, we tested whether nature is capable of besting artificial intelligence. A total of 2000 images of related bees, wasps, and Ophrys sp. were collected from the Google Image Repository. Unsuitable images were later filtered out manually, leaving a total of 995 images in the final selection. 80% of these images were used to build a supervised model using Logistic Regression, while the model accuracy was tested using 20% of the remaining images. Based on our results using Wolfram Mathematica, the Ophrys is not capable of fooling artificial intelligence. The accuracy, accuracy baseline, mean cross-entropy, Area Under ROC (receiver operating characteristic curve) curve (AUC) and the confusion matrix gave excellent image classification. However, we can now show the key points and highlights of the images and how the structures closely resemble actual bees using the SURF method. Rather than just a descriptive method, ML learning has enabled a more quantitative approach. Since this is a simple test, we encourage other scientists to adopt our approach using a larger dataset and better database samples.

Body and Wing Allometries Reveal Flight-Fecundity Tradeoff in Response to Larval Provisioning in Osmia lignaria (Hymenoptera: Megachilidae).

Variation in body size has important implications for physical performance and fitness. For insects, adult size and morphology are determined by larval growth and metamorphosis. Female blue orchard bees, Osmia lignaria, (Say) provision a finite quantity of food to their offspring. In this study, we asked how provision-dependent variation in size changes adult morphology. We performed a diet manipulation in which some larvae were starved in the final instar and some were given unlimited food. We examined the consequences on adult morphology in two ways. First, allometric relationships between major body regions (head, thorax, abdomen) and total body mass were measured to determine relative growth of these structures. Second, morphometrics that are critical for flight (wing area, wing loading, and extra flight power index) were quantified. Head and thorax mass had hyperallometric relationships with body size, indicating these parts become disproportionately large in adults when larvae are given copious provisions. However, abdominal mass and wing area increased hypoallometrically with body size. Thus, large adults had disproportionately lighter abdomens and smaller wing areas than smaller adults. Though both males and females followed these general patterns, allometric patterns were affected by sex. For flight metrics, small adults had reduced wing loading and an increased extra flight power index. These results suggest that diet quantity alters development in ways that affect the morphometric trait relationships in adult O. lignaria and may lead to functional differences in performance.

Neuroanatomical differentiation associated with alternative reproductive tactics in male arid land bees, Centris pallida and Amegilla dawsoni.

Alternative reproductive tactics (ARTs) occur when there is categorical variation in the reproductive strategies of a sex within a population. These different behavioral phenotypes can expose animals to distinct cognitive challenges, which may be addressed through neuroanatomical differentiation. The dramatic phenotypic plasticity underlying ARTs provides a powerful opportunity to study how intraspecific nervous system variation can support distinct cognitive abilities. We hypothesized that conspecific animals pursuing ARTs would exhibit dissimilar brain architecture. Dimorphic males of the bee species Centris pallida and Amegilla dawsoni use alternative mate location strategies that rely primarily on either olfaction (large-morph) or vision (small-morph) to find females. This variation in behavior led us to predict increased volumes of the brain regions supporting their primarily chemosensory or visual mate location strategies. Large-morph males relying mainly on olfaction had relatively larger antennal lobes and relatively smaller optic lobes than small-morph males relying primarily on visual cues. In both species, as relative volumes of the optic lobe increased, the relative volume of the antennal lobe decreased. In addition, A. dawsoni large males had relatively larger mushroom body lips, which process olfactory inputs. Our results suggest that the divergent behavioral strategies in ART systems can be associated with neuroanatomical differentiation.

A revision of the bee genus Ancyloscelis (Hymenoptera: Apidae) in Argentina.

A revision of the emphorine bee genus Ancyloscelis in Argentina is presented. The genus extends from the United States to the center of Argentina, but the maximum diversity of morphological structures is found among South American species. Its species form two distinctive groups regarding their mouthparts: one of them with hooked setae on the proboscis, and the other one with simple or plumose setae on the proboscis. To the first group belong in Argentina, A. bonariensis Brèthes, A. halictoides (Holmberg), A. mesopotamica (Holmberg), and the new species A. holmergi. To the second group belong in Argentina A. apiformis (Fabricius), A. romeroi (Holmberg), A. saltensis Rodríguez Roig-Alsina, and A. nigricornis Rodríguez Roig-Alsina. Lectotype designations are made for Leptergatis bonariensis Brèthes and Leptergatis mesopotamica Holmberg. Ancyloscelis turmalis Vachal is a junior new synonym of Ancyloscelis mesopotamica. The synonymy of Ancyloscelis gigas Friese as junior synonym of Ancyloscelis halictoides is confirmed. Descriptions, illustrations, and a key to the species are provided.

Hairy-Layer Friction Reduction Mechanism in the Honeybee Abdomen.

Abdominal sections of honeybees undergo numerous reciprocating motions during their lifetime. However, the overlapped contact areas adjacent to the abdominal sections have a shallow wear extent, a physical mechanism that remains obscure to date. Therefore, this study explored a biofrictional reduction model based on a solid surface texture and the hairy surface of the honeybee abdomen. We collected honeybee samples and observed their abdomens using a camera (Zeiss Stemi 508). Subsequently, we sliced these samples using a microtome and detected their microscopic friction. The exterior surface of the honeybee abdomen was not smooth but was distributed with a dense microvilli structure, which played a vital role in adjusting the friction reduction characteristics between the abdominal sections. When the adjacent abdominal sections moved relatively to each other, their upper and lower surfaces were not in direct rigid contact. Briefly, this study shows that the microscale hair arrays on the surface of the posterior abdominal segment can significantly reduce real contact area and friction, which considerably decreases wear or abrasion. The friction reduction mechanism alleviates the abrasion during the relative bending movement and saves a large amount of energy, which is essential for the honeybees' daily activities. This microtexture compliance friction reduction characteristic could be used to fabricate hierarchical surfaces for long-lasting friction reduction mechanisms, which increase the life of soft devices, including soft actuators and hinges.