Projected decline in European bumblebee populations in the twenty-first century.

Habitat degradation and climate change are globally acting as pivotal drivers of wildlife collapse, with mounting evidence that this erosion of biodiversity will accelerate in the following decades1-3. Here, we quantify the past, present and future ecological suitability of Europe for bumblebees, a threatened group of pollinators ranked among the highest contributors to crop production value in the northern hemisphere4-8. We demonstrate coherent declines of bumblebee populations since 1900 over most of Europe and identify future large-scale range contractions and species extirpations under all future climate and land use change scenarios. Around 38-76% of studied European bumblebee species currently classified as 'Least Concern' are projected to undergo losses of at least 30% of ecologically suitable territory by 2061-2080 compared to 2000-2014. All scenarios highlight that parts of Scandinavia will become potential refugia for European bumblebees; it is however uncertain whether these areas will remain clear of additional anthropogenic stressors not accounted for in present models. Our results underline the critical role of global change mitigation policies as effective levers to protect bumblebees from manmade transformation of the biosphere.

Towards unbiased interpretations of interactive effects in ecotoxicological studies.

Ecotoxicological research has increasingly focused on the interactive effects of chemical mixtures on biological models, emphasising additive, synergistic, or antagonistic interactions. However, these combination studies often test chemicals at unique concentrations (e.g. x:y), limiting our understanding of the effects across the full spectrum of possible combinations. Evidence from human toxicology suggests that interactive effects among chemicals can vary significantly with total concentration (e.g. x:y vs. 2x:2y), their ratio (e.g. x:2y vs. 2x:y), and the magnitude of the tested effect (e.g. LC10vs. LC50). Our non-exhaustive review of studies on binary mixtures in bee ecotoxicology reveals that such parameters are frequently neglected. Of the 60 studies we examined, only two utilised multiple total concentrations and ratios, thus exploring a broad range of possible combinations. In contrast, 26 studies tested only a single concentration of each chemical, resulting in incomplete interpretations of the potential interactive effects. Other studies utilised various concentrations and/or ratios but failed to capture a broad spectrum of possible combinations. We also discuss potential discrepancies in interactive effects based on different metrics and exposure designs. We advocate for future ecotoxicological studies to investigate a wider spectrum of chemical combinations, including various concentrations and ratios, and to address different levels of effects.

Essential role of papillae flexibility in nectar capture by bees.

Many bees possess a tongue resembling a brush composed of a central rod (glossa) covered by elongated papillae, which is dipped periodically into nectar to collect this primary source of energy. In vivo measurements show that the amount of nectar collected per lap remains essentially constant for sugar concentrations lower than 50% but drops significantly for a concentration around 70%. To understand this variation of the ingestion rate with the sugar content of nectar, we investigate the dynamics of fluid capture by Bombus terrestris as a model system. During the dipping process, the papillae, which initially adhere to the glossa, unfold when immersed in the nectar. Combining in vivo investigations, macroscopic experiments with flexible rods, and an elastoviscous theoretical model, we show that the capture mechanism is governed by the relaxation dynamics of the bent papillae, driven by their elastic recoil slowed down through viscous dissipation. At low sugar concentrations, the papillae completely open before the tongue retracts out of nectar and thus, fully contribute to the fluid capture. In contrast, at larger concentrations corresponding to the drop of the ingestion rate, the viscous dissipation strongly hinders the papillae opening, reducing considerably the amount of nectar captured. This study shows the crucial role of flexible papillae, whose aspect ratio determines the optimal nectar concentration, to understand quantitatively the capture of nectar by bees and how physics can shed some light on the degree of adaptation of a specific morphological trait.

A new exposure protocol adapted for wild bees reveals species-specific impacts of the sulfoximine insecticide sulfoxaflor.

Wild bees are crucial pollinators of flowering plants and concerns are rising about their decline associated with pesticide use. Interspecific variation in wild bee response to pesticide exposure is expected to be related to variation in their morphology, physiology, and ecology, though there are still important knowledge gaps in its understanding. Pesticide risk assessments have largely focussed on the Western honey bee sensitivity considering it protective enough for wild bees. Recently, guidelines for Bombus terrestris and Osmia bicornis testing have been developed but are not yet implemented at a global scale in pesticide risk assessments. Here, we developed and tested a new simplified method of pesticide exposure on wild bee species collected from the field in Belgium. Enough specimens of nine species survived in a laboratory setting and were exposed to oral and topical acute doses of a sulfoximine insecticide. Our results confirm significant variability among wild bee species. We show that Osmia cornuta is more sensitive to sulfoxaflor than B. terrestris, whereas Bombus hypnorum is less sensitive. We propose hypotheses on the mechanisms explaining interspecific variations in sensitivity to pesticides. Future pesticide risk assessments of wild bees will require further refinement of protocols for their controlled housing and exposure.

Microbiota and pathogens in an invasive bee: Megachile sculpturalis from native and invaded regions.

The present study aimed to characterise the bacterial, fungal and parasite gut community of the invasive bee Megachile sculpturalis sampled from native (Japan) and invaded (USA and France) regions via 16S rRNA and ITS2 amplicon sequencing and PCR detection of bee microparasites. The bacterial and fungal gut microbiota communities in bees from invaded regions were highly similar and differed strongly from those obtained in Japan. Core amplicon sequence variants (ASVs) within each population represented environmental micro-organisms commonly present in bee-associated niches that likely provide beneficial functions to their host. Although the overall bacterial and fungal communities of the invasive M. sculpturalis in France and the co-foraging native bees Anthidium florentinum and Halictus scabiosae, were significantly different, five out of eight core ASVs were shared suggesting common environmental sources and potential transmission. None of the 46 M. sculpturalis bees analysed harboured known bee pathogens, while microparasite infections were common in A. florentinum, and rare in H. scabiosae. A common shift in the gut microbiota of M. sculpturalis in invaded regions as a response to changed environmental conditions, or a founder effect coupled to population re-establishment in the invaded regions may explain the observed microbial community profiles and the absence of parasites. While the role of pathogen pressure in shaping biological invasions is still debated, the absence of natural enemies may contribute to the invasion success of M. sculpturalis.

Convivina is a specialised core gut symbiont of the invasive hornet Vespa velutina.

We provide a culturomics analysis of the cultivable bacterial communities of the crop, midgut and hindgut compartments, as well as the ovaries, of the invasive insect Vespa velutina, along with a cultivation-independent analysis of samples of the same nest through 16S rRNA amplicon sequencing. The Vespa velutina bacterial symbiont community was dominated by the genera Convivina, Fructobacillus, Lactiplantibacillus, Lactococcus, Sphingomonas and Spiroplasma. Lactococcus lactis and Lactiplantibacillus plantarum represented generalist core lactic acid bacteria (LAB) symbionts, while Convivina species and Fructobacillus fructosus represented highly specialised core LAB symbionts with strongly reduced genome sizes. Sphingomonas and Spiroplasma were the only non-LAB core symbionts but were not isolated. Convivina bacteria were particularly enriched in the hornet crop and included Convivina intestini, a species adapted towards amino acid metabolism, and Convivina praedatoris sp. nov. which was adapted towards carbohydrate metabolism.

The wild solitary bees Andrena vaga, Anthophora plumipes, Colletes cunicularius, and Osmia cornuta microbiota are host specific and dominated by endosymbionts and environmental microorganisms.

We characterized the microbial communities of the crop, midgut, hindgut, and ovaries of the wild solitary bees Andrena vaga, Anthophora plumipes, Colletes cunicularius, and Osmia cornuta through 16S rRNA gene and ITS2 amplicon sequencing and a large-scale isolation campaign. The bacterial communities of these bees were dominated by endosymbionts of the genera Wolbachia and Spiroplasma. Bacterial and yeast genera representing the remaining predominant taxa were linked to an environmental origin. While only a single sampling site was examined for Andrena vaga, Anthophora plumipes, and Colletes cunicularius, and two sampling sites for Osmia cornuta, the microbiota appeared to be host specific: bacterial, but not fungal, communities generally differed between the analyzed bee species, gut compartments and ovaries. This may suggest a selective process determined by floral and host traits. Many of the gut symbionts identified in the present study are characterized by metabolic versatility. Whether they exert similar functionalities within the bee gut and thus functional redundancy remains to be elucidated.

A case study of the diet-microbiota-parasite interplay in bumble bees.

AIMS: Diets and parasites influence the gut bacterial symbionts of bumble bees, but potential interactive effects remain overlooked. The main objective of this study was to assess the isolated and interactive effects of sunflower pollen, its phenolamides, and the widespread trypanosomatid Crithidia sp. on the gut bacterial symbionts of Bombus terrestris males. METHODS AND RESULTS: Bumble bee males emerged in microcolonies fed on either (i) willow pollen (control), (ii) sunflower pollen, or (iii) willow pollen spiked with phenolamide extracts from sunflower pollen. These microcolonies were infected by Crithidia sp. or were pathogen-free. Using 16S rRNA amplicon sequencing (V3-V4 region), we observed a significant alteration of the beta diversity but not of the alpha diversity in the gut microbial communities of males fed on sunflower pollen compared to males fed on control pollen. Similarly, infection by the gut parasite Crithidia sp. altered the beta diversity but not the alpha diversity in the gut microbial communities of males, irrespective of the diet. By contrast, we did not observe any significant alteration of the beta or alpha diversity in the gut microbial communities of males fed on phenolamide-enriched pollen compared to males fed on control pollen. Changes in the beta diversity indicate significant dissimilarities of the bacterial taxa between the treatment groups, while the lack of difference in alpha diversity demonstrates no significant changes within each treatment group. CONCLUSIONS: Bumble bees harbour consistent gut microbiota worldwide, but our results suggest that the gut bacterial communities of bumble bees are somewhat shaped by their diets and gut parasites as well as by the interaction of these two factors. This study confirms that bumble bees are suitable biological surrogates to assess the effect of diet and parasite infections on gut microbial communities.

Evaluation of Tandem Mass Spectrometry Experiments in the Negative Ionization Mode for Phenolamide Regioisomer Characterization.

Phenolamides are abundant specialized metabolites found in nature and consist of hydroxycinnamic acids mono- or polyconjugated with polyamines. Their participation in flower development is well-documented, and their presence in pollen raises the question of their role in pollen/pollinator interactions. The structural characterization of phenolamides is complicated by the presence of positional isomers and stereoisomers. Liquid chromatography coupled to tandem mass spectrometry in the positive ionization mode is becoming very popular in phenolamide structural characterization. However, collision-induced transamidation processes that cause the swapping of side chains have been detected, making it difficult to distinguish regioisomers with this technique. In the present report, we explore the dissociation processes undergone by the [M - H]- ions of spermidine-based phenolamides as model compounds. We describe two original competitive dissociation routes, namely, the phenolate and imidate pathways, to account for the observed fragmentation reactions undergone by collisional activated standard phenolamide anions. Whereas the phenolate pathway is regioselective at the central position for spermidine, the imidate pathway, requiring a deprotonated amide, only occurs at the extremities. Tandem mass spectrometry experiments on negatively charged phenolamide ions may then outperform their positive ionization mode counterparts for the distinction between phenolamide regioisomers and globally for the identification of phenolamides in natural extracts.

Nutritional stress exacerbates impact of a novel insecticide on solitary bees' behaviour, reproduction and survival.

Pesticide exposure and food stress are major threats to bees, but their potential synergistic impacts under field-realistic conditions remain poorly understood and are not considered in current pesticide risk assessments. We conducted a semi-field experiment to examine the single and interactive effects of the novel insecticide flupyradifurone (FPF) and nutritional stress on fitness proxies in the solitary bee Osmia bicornis. Individually marked bees were released into flight cages with monocultures of buckwheat, wild mustard or purple tansy, which were assigned to an insecticide treatment (FPF or control) in a crossed design. Nutritional stress, which was high in bees foraging on buckwheat, intermediate on wild mustard and low on purple tansy, modulated the impact of insecticide exposure. Within the first day after application of FPF, mortality of bees feeding on buckwheat was 29 times higher compared with control treatments, while mortality of FPF exposed and control bees was similar in the other two plant species. Moreover, we found negative synergistic impacts of FPF and nutritional stress on offspring production, flight activity, flight duration and flower visitation frequency. These results reveal that environmental policies and risk assessment schemes that ignore interactions among anthropogenic stressors will fail to adequately protect bees and the pollination services they provide.

Rapid assessment of insect pollination services to inform decision-making.

Pollinator declines have prompted efforts to assess how land-use change affects insect pollinators and pollination services in agricultural landscapes. Yet many tools to measure insect pollination services require substantial landscape-scale data and technical expertise. In expert workshops, 3 straightforward methods (desk-based method, field survey, and empirical manipulation with exclusion experiments) for rapid insect pollination assessment at site scale were developed to provide an adaptable framework that is accessible to nonspecialist with limited resources. These methods were designed for TESSA (Toolkit for Ecosystem Service Site-Based Assessment) and allow comparative assessment of pollination services at a site of conservation interest and in its most plausible alternative state (e.g., converted to agricultural land). We applied the methods at a nature reserve in the United Kingdom to estimate the value of insect pollination services provided by the reserve. The economic value of pollination services provided by the reserve ranged from US$6163 to US$11,546/year. The conversion of the reserve to arable land would provide no insect pollination services and a net annual benefit from insect-pollinated crop production of approximately $1542/year (US$24∙ha-1 ∙year-1 ). The methods had wide applicability and were readily adapted to different insect-pollinated crops: rape (Brassica napus) and beans (Vicia faba) crops. All methods were rapidly employed under a low budget. The relatively less robust methods that required fewer resources yielded higher estimates of annual insect pollination benefit.

Diversidad y Conservación de Gasterópodos Subterráneos de Agua Dulce en los Estados Unidos y en México Resumen Las declinaciones de los polinizadores han impulsado los esfuerzos por evaluar cómo el cambio del uso de suelo afecta a los insectos polinizadores y los servicios de polinización en los paisajes agrícolas. Aun así, muchas de las herramientas para medir los servicios de los insectos polinizadores requieren datos sustanciales a escala de paisaje y el conocimiento de expertos. Desarrollamos tres métodos sencillos (método de gabinete, censo de campo y manipulación empírica con experimentos de exclusión) durante algunos talleres de expertos para la evaluación rápida de la polinización por insectos a escala de sitio con el objetivo de proporcionar un marco de trabajo adaptable y accesible para quienes no son especialistas y cuentan con recursos limitados. Estos métodos fueron diseñados para TESSA (Toolkit for Ecosystem Service Site-Based Assessment, en inglés) y permiten la evaluación comparativa de los servicios de polinización en los sitios de interés para la conservación y su estado alternativo más plausible (p. ej.: convertido a suelo agrícola). Aplicamos los métodos en una reserva natural del Reino Unido para estimar el valor de los servicios de polinización por insectos que proporciona la reserva. El valor económico de los servicios de polinización que proporciona la reserva varió desde US$6,163 a US$11,546 al año-1 . La conversión de la reserva a suelo arable no proporcionaría servicios de polinización por insectos, pero sí un beneficio anual neto a partir de la producción de cultivos polinizados por insectos de aproximadamente $1,542 al año-1 (US$24 ha-1 año-1 ). Los métodos tuvieron una aplicabilidad generalizada y estaban ya adaptados a los diferentes cultivos polinizados por insectos: cultivos de colza (Brassica napus) y habas (Vicia faba). Todos los métodos pudieron usarse con bajo presupuesto. Los métodos relativamente menos robustos que requirieron menos recursos produjeron estimados más elevados del beneficio anual de la polinización por insectos.

Bumblebee resilience to climate change, through plastic and adaptive responses.

Bumblebees are ubiquitous, cold-adapted eusocial bees found worldwide from subarctic to tropical regions of the world. They are key pollinators in most temperate and boreal ecosystems, and both wild and managed populations are significant contributors to agricultural pollination services. Despite their broad ecological niche at the genus level, bumblebee species are threatened by climate change, particularly by rising average temperatures, intensifying seasonality and the increasing frequency of extreme weather events. While some temperature extremes may be offset at the individual or colony level through temperature regulation, most bumblebees are expected to exhibit specific plastic responses, selection in various key traits, and/or range contractions under even the mildest climate change. In this review, we provide an in-depth and up-to-date review on the various ways by which bumblebees overcome the threats associated with current and future global change. We use examples relevant to the fields of bumblebee physiology, morphology, behaviour, phenology, and dispersal to illustrate and discuss the contours of this new theoretical framework. Furthermore, we speculate on the extent to which adaptive responses to climate change may be influenced by bumblebees' capacity to disperse and track suitable climate conditions. Closing the knowledge gap and improving our understanding of bumblebees' adaptability or avoidance behaviour to different climatic circumstances will be necessary to improve current species climate response models. These models are essential to make correct predictions of species vulnerability in the face of future climate change and human-induced environmental changes to unfold appropriate future conservation strategies.

Global patterns in bumble bee pollen collection show phylogenetic conservation of diet.

Bumble bees (Bombus) are a group of eusocial bees with a strongly generalised feeding pattern, collecting pollen from many different botanical families. Though predominantly generalists, some bumble bee species seem to have restricted dietary choices. It is unclear whether restricted diets in bumble bees are inherent or a function of local conditions due to a lack of data for many species across different regions. The objective of this study was to determine whether bumble bee species displayed specific patterns of pollen collection, and whether patterns were influenced by phylogenetic relatedness or tongue length, a trait known to be associated with structuring floral visitation. Bumble bee pollen collection patterns were quantified from 4,132 pollen loads taken from 58 bumble bee species, representing 24% of the pollen-collecting diversity of this genus. Phylogenetic trait mapping showed a conserved pattern of dietary dissimilarity across species, but not for dietary breadth. Dietary dissimilarity was driven by collection of Fabaceae, with the most similar species collecting around 50%-60% of their diet from this botanical family. The proportion of the diet collected from Fabaceae also showed a conserved phylogenetic signal. Greater collection of Fabaceae was associated with longer tongue lengths, with shorter tongued species focusing on alternative botanical families. However, this result was largely driven by phylogenetic relatedness, not tongue length per se. These results demonstrate that, though generalists, bumble bees are still subject to dietary restrictions that constrain their foraging choices. These dietary constraints have implications for their persistence should their core resources decline in abundance.

Global effects of extreme temperatures on wild bumblebees.

Climate plays a key role in shaping population trends and determining the geographic distribution of species because of limits in species' thermal tolerance. An evaluation of species tolerance to temperature change can therefore help predict their potential spatial shifts and population trends triggered by ongoing global warming. We assessed inter- and intraspecific variations in heat resistance in relation to body mass, local mean temperatures, and evolutionary relationships in 39 bumblebee species, a major group of pollinators in temperate and cold ecosystems, across 3 continents, 6 biomes, and 20 regions (2386 male specimens). Based on experimental bioassays, we measured the time before heat stupor of bumblebee males at a heatwave temperature of 40 °C. Interspecific variability was significant, in contrast to interpopulational variability, which was consistent with heat resistance being a species-specific trait. Moreover, cold-adapted species are much more sensitive to heat stress than temperate and Mediterranean species. Relative to their sensitivity to extreme temperatures, our results help explain recent population declines and range shifts in bumblebees following climate change.

Efectos Globales de las Temperaturas Extremas sobre Abejorros Silvestres Resumen El clima juega un papel importante en la configuración de las tendencias poblacionales y en la determinación de la distribución geográfica de las especies debido a los límites de la tolerancia térmica y al agua que tiene cada especie. Por lo tanto, una evaluación de la tolerancia de las especies al cambio térmico puede ayudar a predecir los potenciales cambios espaciales y las tendencias poblacionales detonadas por el calentamiento global en curso. Evaluamos las variaciones inter- e intraespecíficas de la resistencia al calor en relación con la masa corporal, temperaturas locales promedio y las relaciones evolutivas para 39 especies de abejorros, un grupo primordial de polinizadores en ecosistemas templados y fríos, en tres continentes, seis biomas y 20 regiones (2,386 especímenes machos). Con base en bioanálisis experimentales, medimos el tiempo previo al letargo por calor de los abejorros machos a temperaturas de 40 °C, típicas de una ola de calor. La variabilidad interespecífica fue significativa, en contraste con la variabilidad interpoblacional, lo cual fue consistente con el hecho de que la resistencia al calor sea una característica específica de la especie. Además, las especies adaptadas al frío son mucho más sensibles al estrés por calor que las especies de clima templado o Mediterráneo. En relación con la sensibilidad a las temperaturas extremas, nuestros resultados ayudan a explicar las recientes declinaciones poblacionales y cambios en la distribución de los abejorros posteriores al cambio climático.

Impact of intraspecific variation on measurements of thermal tolerance in bumble bees.

Climate change is an important driver of bee decline despite the fact that many species might respond to climate change differently. One method to predict how a species will respond to climate change is to identify its thermal tolerance limits. However, differences in thermal tolerance might also occur among distant populations of the same species based on their local environment or even among castes of social insects. Here, we investigated intraspecific differences in thermal tolerance among subspecies of the large earth bumble bee, Bombus terrestris (Apidae). We determined the critical thermal minima and maxima (CTmin and CTmax, respectively) of workers and queens from three lab-reared B. terrestris subspecies (B. t. terrestris, B. t. audax, and B. t. canariensis) which originated from different thermal environments. Our results showed that caste has an influence on critical thermal minima, with queens being most cold-tolerant, but the values of critical thermal maxima were not correlated to caste or size. The thermal tolerance of workers did not differ among the subspecies. Although heat tolerance was similar in queens, B. t. canariensis queens (originating from the warmest environments) were the least cold tolerant. Overall, we showed that B. terrestris may be generally robust against climate warming, but that particular subspecies and/or populations may be more vulnerable to extreme temperature variability. Future research should focus on responses of B. terrestris populations to short, extreme thermal events.

Long-term effects of global change on occupancy and flight period of wild bees in Belgium.

Global change affects species by modifying their abundance, spatial distribution, and activity period. The challenge is now to identify the respective drivers of those responses and to understand how those responses combine to affect species assemblages and ecosystem functioning. Here we correlate changes in occupancy and mean flight date of 205 wild bee species in Belgium with temporal changes in temperature trend and interannual variation, agricultural intensification, and urbanization. Over the last 70 years, bee occupancy decreased on average by 33%, most likely because of agricultural intensification, and flight period of bees advanced on average by 4 days, most likely because of interannual temperature changes. Those responses resulted in a synergistic effect because species which increased in occupancy tend to be those that have shifted their phenologies earlier in the season. This leads to an overall advancement and shortening of the pollination season by 9 and 15 days respectively, with lower species richness and abundance compared to historical pollinator assemblages, except at the early start of the season. Our results thus suggest a strong decline in pollination function and services.

Shift in size of bumblebee queens over the last century.

Species can respond differently when facing environmental changes, such as by shifting their geographical ranges or through plastic or adaptive modifications to new environmental conditions. Phenotypic modifications related to environmental factors have been mainly explored along latitudinal gradients, but they are relatively understudied through time despite their importance for key ecological interactions. Here we hypothesize that the average bumblebee queen body size has changed in Belgium during the last century. Based on historical and contemporary databases, we first tested if queen body sizes changed during the last century at the intraspecific level among four common bumblebee species and if it could be linked to global warming and/or habitat fragmentation as well as by the replacement by individuals from new populations. Then, we assessed body size changes at the community level, among 22 species, taking into account species population trends (i.e. increasing, stable or decreasing relative abundance). Our results show that the average queen body size of all four bumblebee species increased over the last century. This size increase was significantly correlated to global warming and habitat fragmentation, but not explained by changes in the population genetic structure (i.e. colonization). At the community level, species with stable or increasing relative abundance tend to be larger than declining species. Contrary to theoretical expectations from Bergmann's rule (i.e. increasing body size in colder climates), temperature does not seem to be the main driver of bumblebee body size during the last century as we observed the opposite body size trend. However, agricultural intensification and habitat fragmentation could be alternative mechanisms that shape body size clines. This study stresses the importance of considering alternative global change factors when assessing body size change.

Collection of nectar by bumblebees: how the physics of fluid demonstrates the prominent role of the tongue's morphology.

Bumblebees and some other tiny animals feed on nectar by visiting flowers in their neighborhood. Some bee species appear to be highly specialized, their tongue being adapted to specific flowers. Bombus terrestris in contrast is able to feed on a wide variety of flowers and can thus be considered as a kind of universal nectar catcher. Since plant nectars show highly variable sugar content, Bombus terrestris have developed a capture mechanism that works for almost any fluid viscosity. Their tongues are decorated with very elongated papillae forming a hairy coating surrounding a rod-like main stalk. When settled on a flower, Bombus rapidly dip their tongue into the inflorescence to catch the highly sought-after nectar. To determine the physical mechanism at the origin of this outstanding ability, the capture dynamics was followed from videos recorded during viscous fluid ingestion. Surprisingly, the volume per lap and the lapping frequency are independent of the fluid viscosity over three orders of magnitude. To explain this observation, we designed a physical model of viscous dipping with structured rods. Predictions of the model compared to observations for bees showed that the nectar is not captured with the help of viscous drag, as proposed in the Landau-Levich-Derjaguin model, but thanks to the hairy structure that traps the viscous fluid, capillary forces drastically limiting the drainage. Our approach can be transposed to others nectar foragers such as bats and hummingbirds.

Long-legged bees make adaptive leaps: linking adaptation to coevolution in a plant-pollinator network.

Adaptation is evolution in response to natural selection. Hence, an adaptation is expected to originate simultaneously with the acquisition of a particular selective environment. Here we test whether long legs evolve in oil-collecting Rediviva bees when they come under selection by long-spurred, oil-secreting flowers. To quantify the selective environment, we drew a large network of the interactions between Rediviva species and oil-secreting plant species. The selective environment of each bee species was summarized as the average spur length of the interacting plant species weighted by interaction frequency. Using phylogenetically independent contrasts, we calculated divergence in selective environment and evolutionary divergence in leg length between sister species (and sister clades) of Rediviva We found that change in the selective environment explained 80% of evolutionary change in leg length, with change in body size contributing an additional 6% of uniquely explained variance. The result is one of four proposed steps in testing for plant-pollinator coevolution.

Playing with extremes: Origins and evolution of exaggerated female forelegs in South African Rediviva bees.

Despite close ecological interactions between plants and their pollinators, only some highly specialised pollinators adapt to a specific host plant trait by evolving a bizarre morphology. Here we investigated the evolution of extremely elongated forelegs in females of the South African bee genus Rediviva (Hymenoptera: Melittidae), in which long forelegs are hypothesised to be an adaptation for collecting oils from the extended spurs of their Diascia host flowers. We first reconstructed the phylogeny of the genus Rediviva using seven genes and inferred an origin of Rediviva at around 29MYA (95% HPD=19.2-40.5), concurrent with the origin and radiation of the Succulent Karoo flora. The common ancestor of Rediviva was inferred to be a short-legged species that did not visit Diascia. Interestingly, all our analyses strongly supported at least two independent origins of long legs within Rediviva. Leg length was not correlated with any variable we tested (ecological specialisation, Diascia visitation, geographic distribution, pilosity type) but seems to have evolved very rapidly. Overall, our results indicate that foreleg length is an evolutionary highly labile, rapidly evolving trait that might enable Rediviva bees to respond quickly to changing floral resource availability.