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Avocado (Persea americana Mill.) is a major horticultural crop that relies on insect mediated pollination. In avocado production, a knowledge gap exists as to the importance of insect pollination, especially in East African smallholder farms. In this study, conducted in a leading smallholder avocado production region in Kenya, we assessed the dependence of avocado fruit set on insect pollination and whether current smallholder production systems suffer from a deficit in pollination services. Furthermore, we assessed if supplementation with colonies of the Western honey bee (Apis mellifera L.) to farms mitigated potential pollination deficits. Our results revealed a very high reliance of avocado on insect pollinators, with a significantly lower fruit set observed for self- and wind-pollinated (17.4%) or self-pollinated flowers (6.4%) in comparison with insect-pollinated flowers (89.5%). We found a significant pollination deficit across farms, with hand-pollinated flowers on average producing 20.7% more fruits than non-treated open flowers prior to fruit abortion. This pollination deficit could be compensated by the supplementation of farms with A. mellifera colonies. Our findings suggest that pollination is limiting fruit set in avocado and that A. Mellifera supplementation on farms is a potential option to increase fruit yield.

Honey bees (genus Apis) are important pollinators in Asian tropical agricultural and natural ecosystems, yet the Asian species remain vastly understudied compared to the European honey bee, Apis mellifera. We studied the temporal and spatial foraging patterns of three co-occurring Asian honey bee species in Bangalore, India, to gain a better understanding of how they coexist. We found evidence for temporal resource partitioning, with Apis cerana having activity peaks in the early mornings, Apis florea initiating foraging later in the day, and Apis dorsata having the potential to do much of its foraging activity at night, even when the moon is not yet up. Apart from the established species differences in foraging ranges, we found limited evidence of spatial partitioning of the landscape. Although individual colonies foraging in parallel often focused their foraging effort on different patches, all three species preferred foraging in cultivated garden areas to seminatural or urban areas. These observations add to the growing evidence for a key role of gardens as foraging habitat for bees in cities.

Societal Impact Statement Pollen relates to many aspects of human and environmental health, which protection and improvement are endorsed by the United Nations Sustainable Development Goals. By highlighting these connections in the frame of current challenges in monitoring and research, we discuss the need of more integrative and multidisciplinary pollen research related to societal needs, improving health of humans and our ecosystems for a sustainable future. Summary Pollen is at once intimately part of the reproductive cycle of seed plants and simultaneously highly relevant for the environment (pollinators, vector for nutrients, or organisms), people (food safety and health), and climate (cloud condensation nuclei and climate reconstruction). We provide an interdisciplinary perspective on the many and connected roles of pollen to foster a better integration of the currently disparate fields of pollen research, which would benefit from the sharing of general knowledge, technical advancements, or data processing solutions. We propose a more interdisciplinary and holistic research approach that encompasses total environmental pollen diversity (ePD) (wind and animal and occasionally water distributed pollen) at multiple levels of diversity (genotypic, phenotypic, physiological, chemical, and functional) across space and time. This interdisciplinary approach holds the potential to contribute to pressing human issues, including addressing United Nations Sustainable Development Goals, fostering social and political awareness of these tiny yet important and fascinating particles.

Pollen beetles (Brassicogethes spp.) are the main pests of oilseed rape (OSR, Brassica napus) in Europe and responsible for massive yield losses. Upcoming pesticide resistances highlight the need for other means of crop protection, such as natural pest control. Sown flower fields aim to counteract the decrease of insect biodiversity in agricultural landscapes by providing resources to ecosystem service providers. However, the optimal age and size of flower fields to increase natural pest control is still unclear. We conducted experiments on 31 OSR fields located along a gradient of landscape-scale semi-natural habitat (SNH). OSR fields were located adjacent to flower fields which differed in age, continuity and size, or adjacent to crop fields or calcareous grasslands. Pesticide-free areas were established to examine interactive effects of pesticide use and flower field characteristics. The abundance of pollen beetle adults and larvae, parasitism and superparasitism rates in OSR were recorded at increasing distances to the adjacent sites. Flower fields and calcareous grasslands increased pollen beetle parasitism when compared to OSR fields neighbouring crop fields. The threshold for effective natural pest control of 35% could be reached in the pesticide-free areas of OSR fields adjacent to calcareous grasslands and flower fields maintained continuously for at least 6 years. In pesticide-sprayed areas, pollen beetle parasitism and superparasitism declined with increasing distance to the adjacent field. Furthermore, flower fields larger than 1.5 ha were able to improve pollen beetle parasitism more than smaller fields. Synthesis and applications. To promote natural pest control in oilseed rape (OSR), large flower fields should be maintained for several years, to create stable habitats for natural enemies. The continuous maintenance of flower fields should be preferred, as ploughing and resowing after 5–6 years decreased the positive effects of the flower fields on natural pest control in adjacent OSR fields. However, pesticide use can abrogate positive effects of flower fields on pollen beetle parasitism. This study highlights that sown flower fields have the potential to increase natural pest control in OSR, but this potential is depending on its age, continuity and size and can be hindered by pesticide use. © 2021 The Authors. Journal of Applied Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society

In their paper Hallmann et al. (1) show that total biomass decline of flying insects (2) is mirrored by diversity loss of hover flies (Diptera: Syrphidae). Furthermore, the authors highlight differences in persistence of common and rare hover fly species. While understanding the mechanisms of insect species decline is of utmost importance to halt biodiversity loss, we would like to make a cautionary note regarding the authors’ generalizing conclusions that “large insect biomass declines are predictive of insect diversity declines” (ref. 1, p. 1) and that “under large biomass drops, species richness … [↵][1] 1To whom correspondence may be addressed. Email: Sarah.redlich{at}uni-wuerzburg.de. [1]: #xref-corresp-1-1

Honey bee health is affected by multiple stressors, such as the exposure to plant protection products (PPPs), dietary limitation, monofloral diets and pressure of diseases and pathogens and their interactions. Here, we analysed the interacting effects of plant protection products and low nutritional pollen source on honey bee health under semi-field conditions. We established a healthy honey bee colony in each of 24 tents, planted either with monofloral maize, maize with a diverse flower strip or with monofloral Phacelia tanacetifolia. To evaluate the interaction between exposure to PPPs and nutritional status, a mixture of the insecticide thiacloprid and the fungicide prochloraz was applied. For each colony, we investigated brood capping rate as well as adult longevity, body and head weight, and enzyme activity of acetylcholinesterase and P450 reductase of newly hatched worker bees. We found a significant reduced capping rate in treated maize compared to flowering strips and Phacelia, but no interaction effect between pesticide treatment and nutritional status on capping rate. The response to treatment on the longevity of adults differed significantly between maize and Phacelia, with flower strips being intermediate, indicating interaction effects of PPP treatment and low pollen quality in maize compared to Phacelia and flowering strip treatments. Head weight of newly hatched worker bees showed significant interaction of nutritional status and treatment of PPPs. PPPs slightly increased body weight in all nutritional statuses, except for Phacelia. Enzyme activity of acetylcholinesterase and P450 reductase showed significant different responses between maize and Phacelia to PPP exposure, but not between maize and flowering strip. Our results support the hypothesis that higher pollen quality promotes development of larvae and pupae, longevity of adults and detoxification of PPPs.

Aim: While elevational gradients in species richness constitute some of the best depicted patterns in ecology, there is a large uncertainty concerning the role of food resource availability for the establishment of diversity gradients in insects. Here, we analysed the importance of climate, area, land use and food resources for determining diversity gradients of dung beetles along extensive elevation and land use gradients on Mt. Kilimanjaro, Tanzania. Location: Mt. Kilimanjaro, Tanzania. Taxon: Scarabaeidae (Coleoptera). Methods: Dung beetles were recorded with baited pitfall traps at 66 study plots along a 3.6 km elevational gradient. In order to quantify food resources for the dung beetle community in form of mammal defecation rates, we assessed mammalian diversity and biomass with camera traps. Using a multi‐model inference framework and path analysis, we tested the direct and indirect links between climate, area, land use and mammal defecation rates on the species richness and abundance of dung beetles. Results: We found that the species richness of dung beetles declined exponentially with increasing elevation. Human land use diminished the species richness of functional groups exhibiting complex behaviour but did not have a significant influence on total species richness. Path analysis suggested that climate, in particular temperature and to a lesser degree precipitation, were the most important predictors of dung beetle species richness while mammal defecation rate was not supported as a predictor variable. Main conclusions: Along broad climatic gradients, dung beetle diversity is mainly limited by climatic factors rather than by food resources. Our study points to a predominant role of temperature‐driven processes for the maintenance and origination of species diversity of ectothermic organisms, which will consequently be subject to ongoing climatic changes.

Recent studies have emphasized the role of the western honey bee, Apis mellifera, as a managed agricultural species worldwide, but also as a potential threat to endangered wild pollinators. This has resulted in the suggestion that honey bees should be regulated in natural areas to conserve wild pollinators. We argue that this perspective fails to appreciate the multifaceted nature of honey bees as native or introduced species with either managed or wild colonies. Wild populations of A. mellifera are currently imperiled, and natural areas are critical for the conservation of local subspecies and genotypes. We propose that a differentiation between managed and wild populations is required and encourage integrated conservation planning for all endangered wild bees, including A. mellifera.

The benefits of honey bee dance communication for colony performance in different resource environments are still not well understood. Here, we test the hypothesis that directional dance communication enables honey bee colonies to maintain a diverse pollen diet, especially in landscapes with low resource diversity. To test this hypothesis, we placed 24 Apis mellifera L. colonies with either intact or experimentally disrupted dance communication in eight agricultural landscapes that differed in the diversity of flowering plants and in the dominance of mass-flowering crops. Pollen from incoming foragers was collected and identified via DNA metabarcoding. Disrupting dance communication affected the way the diversity of honey bee pollen diets was impacted by the dominance of mass-flowering crops in available flower resources (p = .04). With increasing dominance of mass-flowering crops in resource environments, foragers of colonies with intact communication foraged on an increasing proportion of available plant genera (p = .01). This was not the case for colonies with disrupted dance communication (p = .5). We conclude that the honey bee dance communication benefits pollen foraging on diverse plant resources and thereby contributes to high quality nutrition in environments with low-resource diversity.

Increasingly frequent warm periods during winter, which are associated with climate change, may cause mismatches between the colony phenology of the western honey bee, Apis mellifera L., and their floral resources. Warmer winter periods can also affect colony brood rearing activity and consequently the reproduction of the invasive brood parasite Varroa destructor Anderson and Trueman. Until now little is known about the effects of climate change on biotic interactions in such a multitrophic system comprising flowering plants, a pollinator, and its parasite. We performed a reciprocal translocation experiment with honey bee colonies to simulate climate change-induced phenology shifts. Honey bee brood phenology was highly sensitive to environmental conditions in late winter. Colonies in which phenology was experimentally delayed had smaller worker populations in early spring and reduced amounts of stored honey during the following months. During summer, the varroa load in colonies with non-shifted phenology was three times higher than in colonies with delayed phenology. High varroa loads during summer were negatively correlated with worker population growth. Despite a remarkable resilience of colony development to phenology shifts, our results show that mismatches between the phenology of honey bee colonies and flowering plants can affect the build-up of resource stores. Further, an advanced onset of brood rearing activity after hibernation can reinforce the negative impact of the brood parasite V. destructor. We conclude that trade-offs between synchronisation with earlier flower phenology and prolonged brood phases with build-up of varroa populations might constrain the honey bees' capability to adapt to climate warming.

Land-use intensification is a major driver of biodiversity loss. However, understanding how different components of land use drive biodiversity loss requires the investigation of multiple trophic levels across spatial scales. Using data from 150 agricultural grasslands in central Europe, we assess the influence of multiple components of local- and landscape-level land use on more than 4,000 above- and belowground taxa, spanning 20 trophic groups. Plot-level land-use intensity is strongly and negatively associated with aboveground trophic groups, but positively or not associated with belowground trophic groups. Meanwhile, both above- and belowground trophic groups respond to landscape-level land use, but to different drivers: aboveground diversity of grasslands is promoted by diverse surrounding land-cover, while belowground diversity is positively related to a high permanent forest cover in the surrounding landscape. These results highlight a role of landscape-level land use in shaping belowground communities, and suggest that revised agroecosystem management strategies are needed to conserve whole-ecosystem biodiversity., Nature Communications, 12 (1), ISSN:2041-1723

Patterns of insect diversity along elevational gradients are well described in ecology. However, it remains little tested how variation in the quantity, quality, and diversity of food resources influence these patterns. Here we analyzed the direct and indirect effects of climate, food quantity (estimated by net primary productivity), quality (variation in the specific leaf area index, leaf nitrogen to phosphorus and leaf carbon to nitrogen ratio), and food diversity (diversity of leaf traits) on the species richness of phytophagous beetles along the broad elevation and land use gradients of Mt. Kilimanjaro, Tanzania. We sampled beetles at 65 study sites located in both natural and anthropogenic habitats, ranging from 866 to 4,550 m asl. We used path analysis to unravel the direct and indirect effects of predictor variables on species richness. In total, 3,154 phytophagous beetles representing 19 families and 304 morphospecies were collected. We found that the species richness of phytophagous beetles was bimodally distributed along the elevation gradient with peaks at the lowest (˜866 m asl) and upper mid-elevations (˜3,200 m asl) and sharply declined at higher elevations. Path analysis revealed temperature- and climate-driven changes in primary productivity and leaf trait diversity to be the best predictors of changes in the species richness of phytophagous beetles. Species richness increased with increases in mean annual temperature, primary productivity, and with increases in the diversity of leaf traits of local ecosystems. Our study demonstrates that, apart from temperature, the quantity and diversity of food resources play a major role in shaping diversity gradients of phytophagous insects. Drivers of global change, leading to a change of leaf traits and causing reductions in plant diversity and productivity, may consequently reduce the diversity of herbivore assemblages.

Fil: Senapathi, Deepa. University of Reading. School of Agriculture, Policy & Development. Centre for Agri-Environmental Research. Reino Unido. Fil: Fründ, Jochen. University of Freiburg. Faculty of Environment and Natural Resources. Biometry and Environmental System Analysis. Alemania. Fil: Albrecht, Matthias. Agroscope. Institute for Sustainability Sciences. Suiza. Fil: Garratt, Michael P. D. University of Reading. School of Agriculture, Policy & Development. Centre for Agri-Environmental Research. Reino Unido. Fil: Kleijn, David. Wageningen University. Plant Ecology and Nature Conservation Group. Países Bajos. Fil: Pickles, Brian J. University of Reading. School of Biological Sciences. Reino Unido. Fil: Potts, Simon G. University of Reading. School of Agriculture, Policy & Development. Centre for Agri-Environmental Research. Reino Unido. Fil: An, Jiandong. Chinese Academy of Agricultural Sciences. Institute of Apicultural Research. China. Fil: Andersson, Georg K. S. Universidad Nacional de Río Negro. Instituto de Investigaciones en Recursos Naturales, Agroecología y Desarrollo Rural. Río Negro, Argentina. Fil: Andersson, Georg K. S. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Investigaciones en Recursos Naturales, Agroecología y Desarrollo Rural. Río Negro, Argentina. Fil: Bänsch, Svenja. University of Göttingen. Department of Crop Sciences. Functional Agrobiodiversity. Alemania. Fil: Basu, Parthiba. University of Calcutta. Centre for Pollination Studies. India. Fil: Benjamin, Faye. The State University of New Jersey. Department of Ecology, Evolution and Natural Resources. Estados Unidos. Fil: Bezerra, Antonio Diego M. The State University of New Jersey. Department of Ecology, Evolution and Natural Resources. Estados Unidos. Fil: Bhattacharya, Ritam. University of Calcutta. Centre for Pollination Studies. India. Fil: Biesmeijer Jacobus C. Naturalis Biodiversity Centre. Países Bajos. Fil: Blaauw, Brett. University of Georgia. Department of Entomology. Estados Unidos. Fil: Blitzer, Eleanor J. Carroll College. Department of Biology. Estados Unidos. Fil: Brittain, Claire A. Jealott’s Hill International Research Centre. Syngenta. Reino Unido. Fil: Carvalheiro, Luísa G. Universidade Federal de Goiás. Departamento de Ecologia. Brasil. Fil: Cariveau, Daniel P. University of Minnesota. Department of Entomology. Estados Unidos. Fil: Chakraborty, Pushan. University of Calcutta. Centre for Pollination Studies. India. Fil: Chatterjee, Arnob. University of Calcutta. Centre for Pollination Studies. India. Fil: Chatterjee, Soumik. University of Calcutta. Centre for Pollination Studies. India. Fil: Cusser, Sarah. Michigan State University. W. K. Kellogg Biological Station. Estados Unidos. Fil: Danforth, Bryan N. Cornell University. Department of Entomology. Estados Unidos. Fil: Degani, Erika. University of Reading. School of Agriculture, Policy & Development. Centre for Agri-Environmental Research. Reino Unido. Fil: Freitas, Breno M. The State University of New Jersey. Department of Ecology, Evolution and Natural Resources. Estados Unidos. Fil: Garibaldi, Lucas A. Universidad Nacional de Río Negro. Instituto de Investigaciones en Recursos Naturales, Agroecología y Desarrollo Rural. Río Negro, Argentina. Fil: Garibaldi, Lucas A. Consejo Nacional de Investigaciones Científicas y Técnicas. Instituto de Investigaciones en Recursos Naturales, Agroecología y Desarrollo Rural. Río Negro, Argentina. Fil: Geslin, Benoit. Aix Marseille Univ, Avignon Université. IMBE. Francia. Fil: de Groot, G. Arjen. Wageningen University and Research. Wageningen Environmental Research. Países Bajos. Fil: Harrison, Tina. University of California Davis. Department of Entomology and Nematology. Estados Unidos. Fil: Howlett, Brad. The New Zealand Institute for Plant & Food Research Limited. Nueva Zelanda. Fil: Isaacs, Rufus. Michigan State University. Department of Entomology. Estados Unidos. Fil: Jha, Shalene. The University of Texas at Austin. Department of Integrative Biology. Estados Unidos. Fil: Klatt, Björn Kristian. University of Göttingen. Department of Crop Sciences. Agroecology. Alemania. Fil: Krewenka, Kristin. University of Heidelberg. Heidelberg Research Service. Alemania. Fil: Leigh, Samuel. University of Reading. School of Agriculture, Policy & Development. Centre for Agri-Environmental Research. Reino Unido. Fil: Lindström, Sandra A. M. Lund University. Department of Biology. Biodiversity. Suecia. Fil: Mandelik, Yael. The Hebrew University of Jerusalem. The Robert H. Smith Faculty of Agriculture, Food and Environment. Department of Entomology. Israel. Fil: McKerchar, Megan. University of Worcester. School of Science and Environment. Reino Unido. Fil: Park, Mia. Cornell University. Department of Entomology. Estados Unidos. Fil: Pisanty, Gideon. Canadian National Collection of Insects, Arachnids and Nematodes. Agriculture and Agri-Food Canada. Canada. Fil: Rader, Romina. University of New England. School of Environment and Rural Science. Australia. Fil: Reemer, Menno. Naturalis Biodiversity Centre. Países Bajos. Fil: Rundlöf, Maj. Lund University. Department of Biology. Biodiversity. Suecia. Fil: Smith, Barbara. University of Calcutta. Centre for Pollination Studies. India. Fil: Smith, Henrik G. Lund University. Centre of Environmental and Climate Research & Department of Biology. Suecia. Fil: Nunes Silva, Patrícia. Universidade do Vale do Rio dos Sinos. Programa de Pós-Graduação em Biologia. Brasil. Fil: Steffan-Dewenter, Ingolf. University of Würzburg. Department of Animal Ecology and Tropical Biology. Alemania. Fil: Tscharntke, Teja. University of Göttingen. Department of Crop Sciences. Agroecology. Alemania. Fil: Webber, Sean. University of Reading. School of Agriculture, Policy & Development. Centre for Agri-Environmental Research. Reino Unido. Fil: Westbury, Duncan B. University of Worcester. School of Science and Environment. Reino Unido. Fil: Westphal, Catrin. University of Göttingen. Department of Crop Sciences. Functional Agrobiodiversity. Alemania. Fil: Wickens, Jennifer B. University of Reading. School of Agriculture, Policy & Development. Centre for Agri-Environmental Research. Reino Unido. Fil: Wickens, Victoria J. University of Reading. School of Agriculture, Policy & Development. Centre for Agri-Environmental Research. Reino Unido. Fil: Winfree, Rachael. The State University of New Jersey. Department of Ecology, Evolution and Natural Resources. Estados Unidos. Fil: Zhang, Hong. Chinese Academy of Agricultural Sciences. Institute of Apicultural Research. China. Fil: Klein, Alexandra-Maria. University of Freiburg. Faculty of Environment and Natural Resources. Nature Conservation and Landscape Ecology. Alemania. While an increasing number of studies indicate that the range, diversity and abundance of many wild pollinators has declined, the global area of pollinator-dependent crops has significantly increased over the last few decades. Crop pollination studies to date have mainly focused on either identifying different guilds pollinating various crops, or on factors driving spatial changes and turnover observed in these communities. The mechanisms driving temporal stability for ecosystem functioning and services, however, remain poorly understood. Our study quantifies temporal variability observed in crop pollinators in 21 different crops across multiple years at a global scale. Using data from 43 studies from six continents, we show that (i) higher pollinator diversity confers greater inter-annual stability in pollinator communities, (ii) temporal variation observed in pollinator abundance is primarily driven by the three-most dominant species, and (iii) crops in tropical regions demonstrate higher inter-annual variability in pollinator species richness than crops in temperate regions. We highlight the importance of recognizing wild pollinator diversity in agricultural landscapes to stabilize pollinator persistence across years to protect both biodiversity and crop pollination services. Short-term agricultural management practices aimed at dominant species for stabilizing pollination services need to be considered alongside longer term conservation goals focussed on maintaining and facilitating biodiversity to confer ecological stability. .

Agri-environmental schemes (AES) aim to restore biodiversity and biodiversity-mediated ecosystem services in landscapes impoverished by modern agriculture. However, a systematic, empirical evaluation of different AES types across multiple taxa and functional groups is missing. Within one orthogonal design, we studied sown flowering AES types with different temporal continuity, size, and landscape context and used calcareous grasslands as seminatural reference habitat. We measured species richness of 12 taxonomic groups (vascular plants, cicadas, orthopterans, bees, butterflies, moths, hoverflies, flower visiting beetles, parasitoid wasps, carabid beetles, staphylinid beetles, and birds) representing 5 trophic levels. A total of 54,955 specimens were identified using traditional taxonomic methods, and bulk arthropod samples were identified through DNA metabarcoding, resulting in a total of 1,077 and 2,110 taxa, respectively. Species richness of most taxonomic groups, as well as multidiversity and richness of pollinators, increased with temporal continuity of AES types. Some groups responded to size and landscape context, but multidiversity and richness of pollinators and natural enemies were not affected. AES flowering fields supported different species assemblages than calcareous grasslands, but assemblages became more similar to those in seminatural grasslands with increasing temporal continuity. Our results indicate that AES flowering fields and seminatural grasslands function synergistically. Flowering fields support biodiversity even when they are relatively small and in landscapes with few remaining seminatural habitats. We therefore recommend a network of smaller, temporally continuous AES flowering fields of different ages, combined with permanent seminatural grasslands, to maximize benefits for biodiversity conservation and ecosystem service delivery in agricultural landscapes.

“Bee pollen” is pollen collected from flowers by honey bees. It is used by the bees to nourish themselves, mainly by providing royal jelly and brood food, but it is also used for human nutrition. For the latter purpose, it is collected at the hive entrance as pellets that the bees bring to the hive. Bee pollen has diverse bioactivities, and thus has been used as a health food, and even as medication in some countries. In this paper, we provide standard methods for carrying out research on bee pollen. First, we introduce a method for the production and storage of bee pollen which assures quality of the product. Routine methods are then provided for the identification of the pollen’s floral sources, and determination of the more important quality criteria such as water content and content of proteins, carbohydrates, fatty acids, vitamins, alkaloids, phenolic and polyphenolic compounds. Finally, methods are described for the determination of some important bioactivities of bee pollen such as its antioxidant, anti-inflammatory, antimicrobial and antimutagenic properties. Métodos estándar Para la investigación del polen El "polen de abeja" es el polen recogido de las flores por las abejas melíferas. El polen de abeja es utilizado para nutrir a las propias abejas, principalmente para proporcionar jalea real y alimento para las crías, pero también se utiliza para la nutrición humana. Para este último fin, se recoge en la entrada de la colmena en forma de gránulos que las abejas llevan a la colmena. El polen de abeja tiene diversas bioactividades, por lo que se hautilizado como alimento para la salud, e incluso como medicamento en algunos países. En este artículo, proporcionamos métodos estándar para llevar a cabo investigaciones sobre el polen de abeja. En primer lugar, presentamos un método de producción y almacenamiento de polen de abeja que garantiza la calidad del producto. A continuación, se ofrecen métodos de rutina para la identificación de las fuentes florales del polen y la determinación de los criterios de calidad más importantes, como el contenido de agua y de proteínas, carbohidratos, ácidos grasos, vitaminas, alcaloides y compuestos fenólicos y polifenólicos. Por último, se describen métodos para la determinación de algunas bioactividades importantes del polen de abeja, como sus propiedades antioxidantes, antiinflamatorias, antimicrobianas y antimutagénicas. The COLOSS (Prevention of honey bee COlony LOSSes) Association aims to explain and prevent massive honey bee colony losses. It was funded through the COST Action FA0803. COST (European Cooperation in Science and Technology) is a unique means for European researchers to jointly develop their own ideas and new initiatives across all scientific disciplines through trans-European networking of nationally funded research activities. Based on a pan-European intergovernmental framework for cooperation in science and technology, COST has contributed since its creation more than 40 years ago to closing the gap between science, policy makers and society throughout Europe and beyond. COST is supported by the EU Seventh Framework Program for research, technological development and demonstration activities (Official Journal L 412, 30 December 2006). The European Science Foundation as implementing agent of COST provides the COST Office through an EC Grant Agreement. The Council of the European Union provides the COST Secretariat. The COLOSS network is now supported by the Ricola Foundation – Nature & Culture. Figures 26–28 are reproduced from Sawyer (1981) with the permission of the publishers University College Cardiff Press and Northern Bee Books. Lidia Barreto and J Nordi wish to thank the Apiculture Research Center of Taubate University (UNITAU-SP/Brazil) and Agriculture Secretary of Bahia State (SEAGRI-BA/ BRAZIL). Maria Campos wishes to thank (UI0204): UIDB/ 00313/2020, Center of Chemistry from Faculty of Sciences and Technology of University of Coimbra, Portugal. Of elia Anjos wishes to thank to Forest Research Centre, a research unit funded by Fundac¸~ao para a Ci^encia e a Tecnologia I.P. (FCT), Portugal (UIDB/00239/2020), and to Centro de Biotecnologia de Plantas da Beira Interior for the OPUS software availability. Norma Almaraz Abarca thanks to the Instituto Politecnico Nacional for financial and logistic support. Manuel Chica and Pascual Campoy wish to thank the APIFRESH project. APIFRESH has been co-funded by the European Commission under the R4SMEs 7th Framework Program. Olena Lokutova thanks the Austrian Institute of beekeeping, Dr H Pehhacker and the same members of the Institute H Hagel and E H€uttinger for conducting photomicroscopic studies and pollen analysis Ukrainian samples of pollen loads, which were the basis of the atlas of pollen “honey plants” of Ukraine. Olena is grateful also to Polish colleagues Z Warakomska (Department of Botany University of Lublin) and D Teper (Polish Institute of beekeeping) for their professionalism and consultations to determine the botanical origin of some Ukrainian honey. Also thanks to their scientific advisers’ academician G Bogdanov (National Academy of Agrarian Sciences of Ukraine), Prof. V Polishuk (Department of beekeeping National University of Life and Environmental Sciences of Ukraine) and O Martynyuk (M.G. Kholodny Institute of Botany, Kiev, Ukraine) for his helpful co-operation in the field of beekeeping and palinology. Janka Nozkova wishes to thank the Operational Program Research and Development of the European Regional Development Fund in the frame of the project “Support of technologies innovation for special bio-food products for human healthy nutrition” (ITMS 26220220115) and also by the Excellence Center for Agrobiodiversity Conservation and Benefit – project implemented under the Operational Program Research and Development financed by European Fund for Regional Development ITMS 26220120015 (Slovak Republic) and all colleagues from Institute of Biodiversity and Biosafety, Slovak University of Agriculture in Nitra for their help with image analysis. Ananias Pascoal, Georgina Tolentino and Let ıcia Estevinho would like to thank Fundac¸~ao para a Ci^encia e Tecnologia (FCT), Programa Operacional Pontencial Humano (POPH) and European Union (EU) for his Postdoctoral grant SFRH/BPD/91380/2012. Wiebke Sickel, Markus Ankenbrand, Gudrun Grimmer, Frank F€orster, Ingolf Steffan-Dewenter and Alexander Keller thank the financial support by the DFG Collaborative Research Center 1047, Insect Timing. MJA was further supported by a grant of the German Excellence Initiative to the Graduate School of Life Sciences of the University of W€urzburg. They are grateful to the members of the Departments of Animal Ecology and Tropical Biology; Bioinformatics; and Human Genetics, University of W€urzburg, for constructive input on the design of the workflow. Additionally thank to the Department of Human Genetics, especially S. Rost, for granting access to the Illumina MiSeq device. Zivoslav Te si c, Mirjana Mosi c, Aleksandar Kosti c, Mirjana Pe si c, Du sanka Milojkovi c-Opsenica thank the Ministry of Education, Science and Technological Development of Serbia, Grants 172017 and TR 31069. Gina Tolentino would like to thank the Mountain Research Center (CIMO), Agricultural College of Bragança, Polytechnic Institute of Braganc¸a for his research grant in the project titled " Development of new bee products in biological production way.” info:eu-repo/semantics/publishedVersion

Tropospheric ozone is a highly oxidative pollutant with the potential to alter plant metabolism. The direct effects of ozone on plant phenotype may alter interactions with other organisms, such as pollinators, and, consequently, affect plant reproductive success. In a set of greenhouse experiments, we tested whether exposure of plants to a high level of ozone affected their phenological development, their attractiveness to four different pollinators (mason bees, honeybees, hoverflies and bumblebees) and, ultimately, their reproductive success. Exposure of plants to ozone accelerated flowering, particularly on plants that were growing in autumn, when light and temperature cues, that commonly promote flowering, were weaker. Simultaneously, there was a tendency for ozone-exposed plants to disinvest in vegetative growth. Plant exposure to ozone did not substantially affect pollinator preference, but bumblebees had a tendency to visit more flowers on ozone-exposed plants, an effect that was driven by the fact that these plants tended to have more open flowers, meaning a stronger attraction signal. Honeybees spent more time per flower on ozone-exposed plants than on control plants. Acceleration of flower production and the behavioural responses of pollinators to ozone-exposed plants resulted in retained reproductive fitness of plants pollinated by bumblebees, honeybees and mason bees, despite the negative effects of ozone on plant growth. Plants that were pollinated by hoverflies had a reduction in reproductive fitness in response to ozone. In a natural setting, acceleration of flowering by ozone might foster desynchronization between plant and pollinator activities. This can have a strong impact on plants with short flowering periods and on plants that, unlike wild mustard, lack compensatory mechanisms to cope with the absence of pollinator activity in the beginning of flowering.

Wild bee populations are threatened by current agricultural practices in many parts of the world, which may put pollination services and crop yields at risk. Loss of pollination services can potentially be predicted by models that link bee abundances with landscape-scale land-use, but there is little knowledge on the degree to which these statistical models are transferable across time and space. This study assesses the transferability of models for wild bee abundance in a mass-flowering crop across space (from one region to another) and across time (from one year to another). The models used existing data on bumblebee and solitary bee abundance in winter oilseed rape fields, together with high-resolution land-use crop-cover and semi-natural habitats data, from studies conducted in five different regions located in four countries (Sweden, Germany, Netherlands and the UK), in three different years (2011, 2012, 2013). We developed a hierarchical model combining all studies and evaluated the transferability using cross-validation. We found that both the landscape-scale cover of mass-flowering crops and permanent semi-natural habitats, including grasslands and forests, are important drivers of wild bee abundance in all regions. However, while the negative effect of increasing mass-flowering crops on the density of the pollinators is consistent between studies, the direction of the effect of semi-natural habitat is variable between studies. The transferability of these statistical models is limited, especially across regions, but also across time. Our study demonstrates the limits of using statistical models in conjunction with widely available land-use crop-cover classes for extrapolating pollinator density across years and regions, likely in part because input variables such as cover of semi-natural habitats poorly capture variability in pollinator resources between regions and years.

BackgroundHoneybees rely on nectar as their main source of carbohydrates [1]. Sucrose, glucose and fructose are the main components of plant nectars [2] [3]. Intriguingly, honeybees express only three putative sugar receptors (AmGr1, AmGr2 and AmGr3) [4], which is in stark contrast to many other insects and vertebrates. The sugar receptors are only partially characterized [5] [6]. AmGr1 detects different sugars including sucrose and glucose. AmGr2 is assumed to act as a co-receptor only, while AmGr3 is assumedly a fructose receptor.ResultsWe show that honeybee gustatory receptor AmGr3 is highly specialized for fructose perception when expressed inXenopusoocytes. When we introduced nonsense mutations to the respectiveAmGr3gene using CRISPR/Cas9 in eggs of female workers, the resulting mutants displayed almost a complete loss of responsiveness to fructose. In contrast, responses to sucrose were normal. Nonsense mutations introduced by CRISPR/Cas9 in honeybees can thus induce a measurable behavioural change and serve to characterize the function of taste receptorsin vivo.ConclusionCRISPR/Cas9 is an excellent novel tool for characterizing honeybee taste receptorsin vivo. Biophysical receptor characterisation inXenopusoocytes and nonsense mutation ofAmGr3in honeybees unequivocally demonstrate that this receptor is highly specific for fructose.Graphical AbstractFigure 0

Solitary bees are subject to a variety of pressures that cause severe population declines. Currently, habitat loss, temperature shifts, agrochemical exposure, and new parasites are identified as major threats. However, knowledge about detrimental bacteria is scarce, although they may disturb natural microbiomes, disturb nest environments, or harm the larvae directly. To address this gap, we investigated 12 Osmia bicornis nests with deceased larvae and 31 nests with healthy larvae from the same localities in a 16S ribosomal RNA (rRNA) gene metabarcoding study. We sampled larvae, pollen provisions, and nest material and then contrasted bacterial community composition and diversity in healthy and deceased nests. Microbiomes of pollen provisions and larvae showed similarities for healthy larvae, whilst this was not the case for deceased individuals. We identified three bacterial taxa assigned to Paenibacillus sp. (closely related to P. pabuli/amylolyticus/xylanexedens), Sporosarcina sp., and Bacillus sp. as indicative for bacterial communities of deceased larvae, as well as Lactobacillus for corresponding pollen provisions. Furthermore, we performed a provisioning experiment, where we fed larvae with untreated and sterilized pollens, as well as sterilized pollens inoculated with a Bacillus sp. isolate from a deceased larva. Untreated larval microbiomes were consistent with that of the pollen provided. Sterilized pollen alone did not lead to acute mortality, while no microbiome was recoverable from the larvae. In the inoculation treatment, we observed that larval microbiomes were dominated by the seeded bacterium, which resulted in enhanced mortality. These results support that larval microbiomes are strongly determined by the pollen provisions. Further, they underline the need for further investigation of the impact of detrimental bacterial acquired via pollens and potential buffering by a diverse pollen provision microbiome in solitary bees.

Landscape-level crop diversification is a promising tool for ecological intensification, whereby biodiversity and ecosystem services are enhanced, and pesticide applications reduced. Yet evidence for the effects of crop diversity at multiple scales and in different landscape contexts is lacking. Here, we investigate the potential benefits and context dependencies of multiscale crop diversity on natural enemies and overall biological control in winter wheat. Simultaneously, we examine the mediating effects of bird predation on aphid regulation in this system. Eighteen conventional winter wheat fields were selected along two independent gradients of crop diversity and semi-natural habitat cover (SNH). We assessed biological control using a natural enemy exclusion experiment (“Open Treatment,” “Bird Exclosure,” “Full Exclosure”). Biological control, predator and parasitoid densities within cages were analysed as functions of landscape (crop diversity × SNH), bird predation (yes/no) and temporal change (three surveys) on six spatial scales (100–3,000 m). Crop diversity rather than SNH enhanced aphid regulation in our study system. Biological control in fields with high landscape-level crop diversity was 8%–33% higher than in low-diversity landscapes, with main effects observed on scales

Mass-flowering crops lead to spatial redistributions of pollinators and to transient shortages within nearby semi-natural grasslands, but the impacts on plant–pollinator interactions remain largely unexplored. Here, we characterised which pollinator species are attracted by oilseed rape and how this affected the structure of plant–pollinator networks in nearby grasslands. We surveyed 177 networks from three countries (Germany, Sweden and United Kingdom) in 24 landscapes with high crop cover, and compared them to 24 landscapes with low or no oilseed rape during and after crop blooming. On average 55% of grassland pollinator species were found on the crop, which attracted 8–35% of individuals away from grasslands. However, networks in the grasslands were resistant to these reductions, since mainly abundant and highly mobile species were attracted. Nonetheless, simulations indicated that network structural changes could be triggered if > 50% of individuals were attracted to the crop (a value well-above that found in our study system), which could affect community stability and resilience to further disturbance., This project was funded by the EU FP7 STEP project ‘Status and Trends of European Pollinators’ (244 090, < www.STEP‐project.net>) and Biodiversa‐FACCE project ‘Enhancing biodiversity‐based ecosystem services to crops through optimized densities of green infrastructure in agricultural landscapes’ (PCIN‐2014‐048, < www.cec.lu.se/research/ecodeal >). AM, MV, JG‐V and IB acknowledge funding from the Spanish Severo Ochoa Program (SEV‐2012‐0262). AM acknowledges funding from the Juan de la Cierva program. JBW, VJW and SGP acknowledge the support of Insect Pollinators Initiative research funded jointly by a grant from BBSRC, Defra, NERC, the Scottish Government and the Wellcome Trust, under the Living with Environmental Change Partnership. HGS acknowledges the support of the Formas project SAPES for landscape analyses. AH received funding from DFG‐Collaborative Research Center 1047, Insect Timing.

Managed and wild pollinators of different functional groups can provide pollination services in agricultural landscapes. These pollinators differ in their resource requirements and response to the amount and arrangement of different habitat types, that is, landscape composition and configuration. Most current approaches to test landscape effects on pollinators and pollination services are either applied to central individual crop fields or other landscape elements but rarely consider that pollinators depend on and make use of multiple habitat elements in an entire landscape. To capture these complex spatial and temporal interactions between different pollinators and habitat elements at the landscape scale, we propose to apply a combination of experimental and observational approaches across multiple habitat types and seasons.

Resource availability in agricultural landscapes has been disturbed for many organisms, including pollinator species. Abundance and diversity in flower availability benefit bee populations; however, little is known about which of protein or carbohydrate resources may limit their growth and reproductive performance. Here, we test the hypothesis of complementary resource limitation using a supplemental feeding approach. We applied this assumption with bumble bees (Bombus terrestris), assuming that colony growth and reproductive performance should depend on the continuous supply of carbohydrates and proteins, through the foraging for nectar and pollen, respectively. We placed wild-caught bumble bee colonies along a landscape gradient of seminatural habitats, and monitored the colonies' weight, foraging activity, and reproductive performance during the whole colony cycle. We performed supplemental feeding as an indicator of landscape resource limitation, using a factorial design consisting of the addition of sugar water (carbohydrate, supplemented or not) crossed by pollen (protein, supplemented or not). Bumble bee colony dynamics showed a clear seasonal pattern with a period of growth followed by a period of stagnation. Higher abundance of seminatural habitats resulted in reducing the proportion of pollen foragers relative to all foragers in both periods, and in improving the reproductive performance of bumble bees. Interestingly, the supplemental feeding of sugar water positively affected the colony weight during the stagnation period, and the supplemental feeding of pollen mitigated the landscape effect on pollen collection investment. Single and combined supplementation of sugar water and pollen increased the positive effect of seminatural habitats on reproductive performance. This study reveals a potential colimitation in pollen and nectar resources affecting foraging behavior and reproductive performance in bumble bees, and indicates that even in mixed agricultural landscapes with higher proportions of seminatural habitats, bumble bee populations face resource limitations. We conclude that the seasonal management of floral resources must be considered in conservation to support bumble bee populations and pollination services in farmlands.

Aim: Temperature, food resources and top‐down regulation by antagonists are considered as major drivers of insect diversity, but their relative importance is poorly understood. Here, we used cavity‐nesting communities of bees, wasps and their antagonists to reveal the role of temperature, food resources, parasitism rate and land use as drivers of species richness at different trophic levels along a broad elevational gradient. Location: Mt. Kilimanjaro, Tanzania. Taxon: Cavity‐nesting Hymenoptera (Hymenoptera: Apidae, Colletidae, Megachilidae, Crabronidae, Sphecidae, Pompilidae, Vespidae). Methods: We established trap nests on 25 study sites that were distributed over similar large distances in terms of elevation along an elevational gradient from 866 to 1788 m a.s.l., including both natural and disturbed habitats. We quantified species richness and abundance of bees, wasps and antagonists, parasitism rates and flower or arthropod food resources. Data were analysed with generalized linear models within a multi‐model inference framework. Results: Elevational species richness patterns changed with trophic level from monotonically declining richness of bees to increasingly humped‐shaped patterns for caterpillar‐hunting wasps, spider‐hunting wasps and antagonists. Parasitism rates generally declined with elevation but were higher for wasps than for bees. Temperature was the most important predictor of both bee and wasp host richness patterns. Antagonist richness patterns were also well predicted by temperature, but in contrast to host richness patterns, additionally by resource abundance and diversity. The conversion of natural habitats through anthropogenic land use, which included biomass removal, agricultural inputs, vegetation structure and percentage of surrounding agricultural habitats, had no significant effects on bee and wasp communities. Main conclusions: Our study underpins the importance of temperature as a main driver of diversity gradients in ectothermic organisms and reveals the increasingly important role of food resources at higher trophic levels. Higher parasitism rates at higher trophic levels and at higher temperatures indicated that the relative importance of bottom‐up and top‐down drivers of species richness change across trophic levels and may respond differently to future climate change.

Accumulating reports of global bee declines have drawn much attention to the bee microbiota and its importance. Most research has focused on social bees, while solitary species have received scant attention despite their enormous biodiversity, ecological importance, and agroeconomic value. We review insights from several recent studies on diversity, function, and drivers of the solitary-bee microbiota, and compare these factors with those relevant to the social-bee microbiota. Despite basic similarities, the social-bee model, with host-specific core microbiota and social transmission, is not representative of the vast majority of bee species. The solitary-bee microbiota exhibits greater variability and biodiversity, with a strong impact of environmental acquisition routes. Our synthesis identifies outstanding questions that will build understanding of these interactions, responses to environmental threats, and consequences for health.

Despite decades of scientific effort, there is still no consensus on the determinants of broad-scale gradients of animal diversity. We argue that general drivers of diversity are unlikely to be found among the narrowly defined taxa which are typically analyzed in studies of broad-scale diversity gradients because ecological niches evolve largely conservatively. This causes constraints in the use of available niche space leading to systematic differences in diversity gradients among taxa. We instead advocate studies of phylogenetically diverse animal communities along broad environmental gradients. Such multi-taxa communities are less constrained in resource use and diversification and may be better targets for testing major classical hypotheses on diversity gradients. Besides increasing the spatial scale in analyses, expanding the phylogenetic coverage may be a second way to achieve higher levels of generality in studies of broad-scale diversity gradients.

1.Physiological and energetic mechanisms have been proposed to constrain body sizes of organisms along climatic gradients; however, these provide contrasting predictions. While Bergmann′s rule predicts increases of body sizes in cooler climates resulting from physiological constraints, energy-based community assembly rules suggest declines in the mean body size of species caused by increased extinction probabilities for large-bodied species in low-energy habitats. 2.We tested these contrasting hypotheses by quantifying trait distributions in bee communities along a 3.6 km elevational gradient at Mt. Kilimanjaro. Traditionally, intra- and interspecific trait shifts along environmental gradients have been investigated in isolation. However, a surge of theoretical approaches and studies on plants demonstrated that the explicit integration of trait variation among and within species can be essential for identifying the mechanisms that shape traits and related ecosystem functions along environmental gradients. We therefore studied variation of body size and related morphological traits at both the intra- and interspecific level. 3.We found support for both physiological constraints and energy-based community assembly rules as drivers of trait distribution in bee communities along elevational gradients, which, however, affected different levels of biotic organization, i.e. the population and community level. While the number of bee species with large body sizes declined with elevation, individuals within species became on average larger, resulting in contrasting trends in morphometric parameters at the community versus population level. 4.Furthermore, body size within bee communities became less variable at higher elevations, largely as a result of a non-random, directive loss of species, but paralleled by a decline in intraspecific variance, suggesting intensified filtering effects with increasing elevation. Similar patterns were found for other functional traits related to the foraging ecology of bees (tongue length, relative forewing length). 5.We conclude that along climatic gradients both physiological and energetic constraints shape trait distributions of pollinators, but at different levels of biological organization. This article is protected by copyright. All rights reserved.

Semi-natural grasslands in Europe are insect biodiversity hotspots and important source habitats delivering ecosystem services to adjacent agricultural land by species spillover. However, this spillover might also occur in the opposite direction, affecting the diversity of semi-natural grasslands. This opposite spillover has got little attention in scientific literature even though generalist species penetrating into the grasslands can affect local biotic interactions, community composition and the conservation value of grassland habitats. In this study, we examined spillover effects from two different adjacent habitat types on carabid beetle assemblages in 20 semi-natural calcareous grasslands. The grasslands were either adjacent to a cereal crop field or to a coniferous forest. We found distinct differences in carabid beetle assemblages in calcareous grasslands depending on adjacent habitat type. Species richness and activity density were higher, but the evenness was lower in calcareous grasslands adjacent to crop fields compared with calcareous grasslands adjacent to coniferous forests. Further, we found a strong spillover of carabid beetles from adjacent crop fields after crop harvest, which may result in transiently increased predation pressure and resource competition in calcareous grasslands. Our results highlight that species composition, diversity and presumably ecosystem functions within semi-natural habitats are affected by the type and management of surrounding habitats. This needs to be considered by nature conservation measures, which aim to protect the unique insect communities of semi-natural European grasslands.

Modification of natural ecosystems has threatened biodiversity worldwide, with forests suffering especially. Strategies aimed at mitigating such loss in forests often include enrichment of deadwood, a critical resource for many decomposer species. However, it remains unclear how deadwood can best be enriched to most effectively promote the diversity of saproxylic species. In this study, we investigated saproxylic beetle diversity in experimentally exposed deadwood logs of 13 different tree species across 30 forests in three regions of Germany. We tested whether gamma-diversity differs between tree species and whether the alpha-diversity within an individual log depended on whether logs were placed in unmanaged beech forests, managed beech forests, or managed conifer forests. We found significant differences in gamma- and alpha-diversity of saproxylic beetles among tree species, but the ranking of tree species differed between regions, suggesting differences in regional beetle species pools. Randomization tests aiming to identify how many and which deadwood logs would need to be exposed to best conserve saproxylic beetle diversity, showed that the overall diversity of beetles increased with the number of tree species exposed, due to turnover of beetle species between tree species. However, some species (e.g. Carpinus) and species combinations (e.g. Carpinus-Picea) reached exceptionally high beetle diversity. Alpha-diversity was higher in conifer than in beech forests, but did not differ between managed and unmanaged beech forests. Canopy cover above logs and average stand temperature strongly influenced alpha-diversity, suggesting that environmental conditions that may be affected by management act as habitat filters for species assemblages. We conclude that deadwood enrichment strategies would be most effective when combining particular tree species that support highest diversity.

Mass-flowering crops (MFCs) are increasingly cultivated and might influence pollinator communities in MFC fields and nearby semi-natural habitats (SNHs). Across six European regions and 2 years, we assessed how landscape-scale cover of MFCs affected pollinator densities in 408 MFC fields and adjacent SNHs. In MFC fields, densities of bumblebees, solitary bees, managed honeybees and hoverflies were negatively related to the cover of MFCs in the landscape. In SNHs, densities of bumblebees declined with increasing cover of MFCs but densities of honeybees increased. The densities of all pollinators were generally unrelated to the cover of SNHs in the landscape. Although MFC fields apparently attracted pollinators from SNHs, in landscapes with large areas of MFCs they became diluted. The resulting lower densities might negatively affect yields of pollinator-dependent crops and the reproductive success of wild plants. An expansion of MFCs needs to be accompanied by pollinator-supporting practices in agricultural landscapes., The study was funded by the EU FP7 project “Status and Trends of European Pollinators” (244 090, www.STEP-project.net) and Biodiversa‐FACCE project “Enhancing biodiversity‐based ecosystem services to crops through optimised densities of green infrastructure in agricultural landscapes” (PCIN‐2014‐048, http://www.cec.lu.se/research/ecodeal). JPGV and MV acknowledge support from the Severo Ochoa Program for Centres of Excellence in R+D+I (SEV‐2012‐0262), and FLORMAS (CGL 2012‐33801). JBW, VJW, SGP and SPMR acknowledge the support of Insect Pollinators Initiative research funded jointly by a grant from BBSRC, Defra, NERC, the Scottish Government and the Wellcome Trust, under the Living with Environmental Change Partnership. HGS and RB were supported by the Swedish Research Council FORMAS. JS and DK were supported by the Dutch Ministry of Economic Affairs (BO‐11‐011.01‐011 and KB‐14‐003‐006), AH and ISD were supported by the DFG Collaborative Research Centre 1047 Insect timing, and ISD and MD through the 2013–2014 BiodivERsA/FACCE – JPI project Ecodeal with national funding from BMBF.

Many experiments have shown that loss of biodiversity reduces the capacity of ecosystems to provide the multiple services on which humans depend. However, experiments necessarily simplify the complexity of natural ecosystems and will normally control for other important drivers of ecosystem functioning, such as the environment or land use. In addition, existing studies typically focus on the diversity of single trophic groups, neglecting the fact that biodiversity loss occurs across many taxa and that the functional effects of any trophic group may depend on the abundance and diversity of others. Here we report analysis of the relationships between the species richness and abundance of nine trophic groups, including 4,600 above- and below-ground taxa, and 14 ecosystem services and functions and with their simultaneous provision (or multifunctionality) in 150 grasslands. We show that high species richness in multiple trophic groups (multitrophic richness) had stronger positive effects on ecosystem services than richness in any individual trophic group; this includes plant species richness, the most widely used measure of biodiversity. On average, three trophic groups influenced each ecosystem service, with each trophic group influencing at least one service. Multitrophic richness was particularly beneficial for 'regulating' and 'cultural' services, and for multifunctionality, whereas a change in the total abundance of species or biomass in multiple trophic groups (the multitrophic abundance) positively affected supporting services. Multitrophic richness and abundance drove ecosystem functioning as strongly as abiotic conditions and land-use intensity, extending previous experimental results to real-world ecosystems. Primary producers, herbivorous insects and microbial decomposers seem to be particularly important drivers of ecosystem functioning, as shown by the strong and frequent positive associations of their richness or abundance with multiple ecosystem services. Our results show that multitrophic richness and abundance support ecosystem functioning, and demonstrate that a focus on single groups has led to researchers to greatly underestimate the functional importance of biodiversity.

Chronobiological studies of individual activity rhythms in social insects can be constrained by the artificial isolation of individuals from their social context. We present a new experimental set-up that simultaneously measures the temperature rhythm in a queen-less but brood raising mini colony and the walking activity rhythms of singly kept honey bees that have indirect social contact with it. Our approach enables monitoring of individual bees in the social context of a mini colony under controlled laboratory conditions. In a pilot experiment, we show that social contact with the mini colony improves the survival of monitored young individuals and affects locomotor activity patterns of young and old bees. When exposed to conflicting Zeitgebers consisting of a light–dark (LD) cycle that is phase-delayed with respect to the mini colony rhythm, rhythms of young and old bees are socially synchronized with the mini colony rhythm, whereas isolated bees synchronize to the LD cycle. We conclude that the social environment is a stronger Zeitgeber than the LD cycle and that our new experimental set-up is well suited for studying the mechanisms of social entrainment in honey bees.

Aim Identifying traits that enhance the success of animals in unhospitable climates helps to understand biogeographical patterns and to predict the consequences of climatic changes. On temperate mountains, traits that increase cold performance potentially allow species to colonize higher elevations. We first tested the importance of morphological traits for the cold performance of bees. Second, we assessed the relevance of these traits for species distributional patterns by analysing intraspecific and interspecific trait shifts along elevational gradients. Location National Park Berchtesgaden, Germany; Wurzburg, Germany. Methods We determined thermal activity thresholds by measuring signs of immobility along temperature gradients under laboratory conditions and related them to morphometric traits. We sampled bee communities at 34 sites along an elevation gradient in the northern Alps, measured for all species morphometric traits and thermal activity limits in the field, and analysed how trait measures changed with elevation within and across species. Results Laboratory and field experiments revealed that bees with larger body size, longer hair and relatively short wings tolerated lower temperatures before reaching immobility and started being active in the field at lower temperatures than small, short-haired and long-winged species. In correspondence to these results, we found that morphometric traits increased (body size, hair length) or decreased (relative wing length) with increasing elevation in communities and, in case of body size, also within species. Main conclusions Results of laboratory and field experiments and their reflectance in community- and species-level trends of morphological traits along elevational gradients underscore the adaptive nature of large body size, long hair and short wings for bees’ thermal performance. Morphological traits expanding lower thermal limits probably contribute to the fitness of bees in cold environments and may simultaneously be under selection in a warming world.

Legumes are important crops for food security, consumed by millions of people especially in Africa where they are an essential protein source and provide key vitamins and minerals. Most legumes depend on insect pollination and natural pest regulation for sufficient yields, however, there is emerging evidence that yield gaps caused by lack of pollination and/or pest pressure may be common. Here we review the literature reporting on pollinators and natural enemies contributing to legume crop yields, and the impact of land-use change on the services provided by these beneficial organisms. We identify strategies for enhancing the benefits of pollination and natural pest control in legumes and propose policy and practice interventions for better utilization of pollinators and natural enemies in legume cropping systems in Africa.

Efficient communication is highly important for the evolutionary success of social animals. Honeybees (genusApis) are unique in that they communicate the spatial information of resources using a symbolic ‘language’, the waggle dance. Different honeybee species differ in foraging ecology but it remains unknown whether this shaped variation in the dance. We studied distance dialects—interspecific differences in how waggle duration relates to flight distance—and tested the hypothesis that these evolved to maximize communication precision over the bees' foraging ranges. We performed feeder experiments withApis cerana,A. floreaandA. dorsatain India and found thatA. ceranahad the steepest dialect, i.e. a rapid increase in waggle duration with increasing feeder distance,A. floreahad an intermediate, andA. dorsatahad the lowest dialect. By decoding dances for natural food sites, we inferred that the foraging range was smallest inA. cerana, intermediate inA. floreaand largest inA. dorsata. The inverse correlation between foraging range and dialect was corroborated when comparing six (sub)species across the geographical range of the genus including previously published data. We conclude that dance dialects constitute adaptations resulting from a trade-off between the spatial range and the spatial accuracy of communication.

The effects of habitat loss and fragmentation on wild bee populations are still not fully understood. Availability of resources, parasitism and habitat connectivity might influence population sizes of bees. Whereas floral resources for wild bees have been considered in research and conservation, nesting resources have been largely neglected as these are challenging to investigate. Snail shells are the exclusive nesting cavities for several Osmia species and provide a good - but so far unused - tool to study factors driving bee population dynamics. We investigated the effects of habitat area, connectivity and management of semi-natural grasslands on populations of snail-shell nesting bees. On 23 calcareous grasslands, we monitored snail shell colonization by providing empty snail shells as nesting resources, and recorded flower-visitor interactions. Five snail-shell nesting Osmia species were found on the grasslands, which made almost a quarter of recorded bee flower visits. Three species colonized the offered snail shells with high variation between study sites, Osmia species, and snail shell size. Habitat area had a positive effect on the population size of the habitat specialist Osmia aurulenta, whereas the more generalist species Osmia bicolor was positively influenced by habitat connectivity. Destruction rates of snail shells increased with sheep grazing intensity, leading to an estimated loss of more than a third of all bee nests. We conclude that large and connected habitats benefit bee populations in fragmented landscapes, while conservation management regimes should take into account potential negative effects of grazing on specific nesting resources of specialized bee species.

Despite their diversity and their large functional and cultural importance, the patterns and predictors of large mammal diversity along elevational gradients on tropical mountains remain poorly understood. Today, large mammals are threatened by human disturbances such as habitat destruction and hunting and may increasingly depend on the conservation of protected areas. Here, we use field data on the diversity of large mammals along a 3.6 km elevational gradient on Mt. Kilimanjaro to evaluate the importance of climate, net primary productivity and human impact for the distribution, species richness and community biomass of wild mammals. Mammal species richness was explored with camera traps on 66 study plots along an elevational gradient from 870 to 4,550 m a.s.l.. We applied path analysis and variance partitioning analysis to unravel the direct and indirect effects of temperature, precipitation, primary productivity, land use, land area, the protection of habitats and the occurrence of domestic mammals on the diversity of wild mammals. Both species richness and community biomass of wild mammals showed a unimodal distribution with elevation, peaking in the montane zone of Mt. Kilimanjaro. However, the peak shifted significantly to lower elevations when only protected habitats were considered. Wild mammal diversity increased with net primary productivity, protection of habitats and temperature. Our study underscores the importance of energy resources for the establishment of diversity gradients in large mammals. While temperature has been revealed as a direct predictor of diversity in most ectothermic taxa, in endothermic organisms temperature has stronger indirect effects, via a modulation of net primary productivity. Moreover, our study reveals how patterns of diversity on tropical mountains are influenced by human impact, pointing to the pivotal role of protected areas for the long-term conservation of mountain biodiversity.

Temperature, primary productivity, plant functional traits, and herbivore abundances are considered key predictors of leaf herbivory but their direct and indirect contributions to community-level herbivory are not well understood along broad climatic gradients. Here, we determined elevational herbivory patterns and used a path analytical approach to disentangle the direct and indirect effects of climate, land use, net primary productivity (NPP), herbivore abundance, and plant functional traits on community-level invertebrate herbivory along the extensive elevational and land use gradients at Mt. Kilimanjaro, Tanzania. We recorded standing leaf herbivory caused by leaf chewers, leaf miners and leaf gallers on 55 study sites distributed in natural and anthropogenic habitats along a 3,060 m elevation gradient. We related the total community-level herbivory to climate (temperature and precipitation), NPP, plant functional traits (specific leaf area, leaf carbon-to-nitrogen [CN] ratio and leaf nitrogen-to-phosphorus [NP] ratio) and herbivore abundances. Leaf herbivory ranged from 5% to 11% along the elevation gradient. Total leaf herbivory showed unimodal pattern in natural habitats but a strongly contrasting bimodal pattern in anthropogenic habitats. We also detected some variation in the patterns of leaf herbivory along environmental gradients across feeding guilds with leaf chewers being responsible for a disproportionally large part of herbivory. Path analyses indicated that the variation in leaf herbivory was mainly driven by changes in leaf CN and NP ratios which were closely linked to changes in NPP in natural habitats. Similarly, patterns of leaf herbivory in anthropogenic habitats were best explained by variation in leaf CN ratios and a negative effect of land use. Our study elucidates the strong role of leaf nutrient stoichiometry and its linkages to climate and NPP for explaining the variation in leaf herbivory along broad climatic gradients. Furthermore, the study suggests that climatic changes and nutrient inputs in the course of land use change may alter leaf herbivory and consequently energy and nutrient fluxes in terrestrial habitats.

In order to save resources, honey bee (Apis mellifera) colonies in the temperate zones stop brood rearing during winter. Brood rearing is resumed in late winter to build up a sufficient worker force that allows to exploit floral resources in upcoming spring. The timing of brood onset in hibernating colonies is crucial and a premature brood onset could lead to an early depletion of energy reservoirs. However, the mechanisms underlying the timing of brood onset and potential risks of mistiming in the course of ongoing climate change are not well understood. To assess the relative importance of ambient temperature and photoperiod as potential regulating factors for brood rearing activity in hibernating colonies, we overwintered 24 honey bee colonies within environmental chambers. The colonies were assigned to two different temperature treatments and three different photoperiod treatments to disentangle the individual and interacting effects of temperature and photoperiod. Tracking in-hive temperature as indicator for brood rearing activity revealed that increasing ambient temperature triggered brood onset. Under cold conditions, photoperiod alone did not affect brood onset, but the light regime altered the impact of higher ambient temperature on brood rearing activity. Further the number of brood rearing colonies increased with elapsed time which suggests the involvement of an internal clock. We conclude that timing of brood onset in late winter is mainly driven by temperature but modulated by photoperiod. Climate warming might change the interplay of these factors and result in mismatches of brood phenology and environmental conditions.

International audience; AbstractIn agricultural landscapes all over the world, honey bees are exposed to genetically modified (GM) pollen. Stacked Bt-maize varieties combine multiple Bt proteins against different insect taxa with herbicide resistance. To test for potential effects on non-target organisms, we conducted an in vitro larvae-rearing experiment where we fed increasing amounts of stacked Bt pollen to honey bee larvae. Bt pollen was equally well digested as two control maize varieties. No dose-dependent effects of Bt maize were detected in terms of survival and developmental delay. However, for prepupal weight we did find a dose-dependent response, suggesting a pleiotropic effect. Comparing this finding with the literature we conclude that the found effect is not likely to occur in a field situation. We could however show that dose-dependent effects are detectable. Our results underline the importance of testing dose-depending effects of GM plant material in an environmental risk assessment (ERA).

Summary Modern agricultural landscapes are dynamic systems with interannually changing proportions of different crop types. However, the effects of spatiotemporal changes in crop area on crop yields and crop–herbivore–antagonist interactions have been rarely considered, in contrast to documented beneficial effects of semi-natural habitats on biological pest control at local and landscape scales. In this study, we examined how the proportion of oilseed rape (OSR) fields in a landscape and the annual increase or decrease in OSR cover due to crop rotation affect OSR herbivores, their natural enemies and crop yield. During two study years, we examined the abundance of adult and larval pollen beetles, parasitism of pollen beetle larvae by a parasitic ichneumonid and crop yields (seed weight per plant) in the edge and centre of 36 OSR fields. The fields differed in the proportion of OSR in the surrounding landscape at 1-km radius and in the interannual change in the proportion of OSR from the previous year to the respective study year. Adult pollen beetle abundance decreased in one study year and larval pollen beetle abundance and parasitism rates decreased in both study years with spatially increasing OSR proportion in the landscape, indicating dilution effects on both trophic levels. Crop yield was positively affected by spatially increasing proportions of OSR. An interannual increase in the proportion of OSR led to the dilution of pest species, but had no significant effect on yield or parasitism rate. Synthesis and applications. The negative effects of pollen beetles on yields despite regular insecticide applications underpin the need for improved techniques for controlling this pest. Our results emphasize the potential to enhance crop yields by the management of spatiotemporal crop cover dynamics within landscapes. We recommend that future management schemes should coordinate the spatial aggregation and annual dynamics of oilseed rape (OSR) cover in a landscape as a potential way to reduce pest impacts in intensively managed OSR fields.

Summary 1. Among saproxylic beetles, many early colonizers prefer particular host species. Ranking of preferred hosts of local saproxylic beetle communities is critical for effective dead-wood management in forests, but is rarely done because experiments with numerous tree species are labour and cost intensive. 2. We analysed the host preference of local saproxylic beetle communities on logs of 13 tree species in relation to management (unmanaged and managed beech stands, conifer plantations on natural beech sites) in three regions of Germany during the most critical period for host specificity, that is the first two years after harvesting. Hosts were ranked quantitatively based on the ordinal ranking of hosts by single beetle species, which in turn was based on the variation in beetle abundance. First, we employed a Bradley–Terry model in which ranking was derived from paired comparisons of host trees. Then, a model-based recursive partitioning of the Bradley–Terry model tested whether host preference of beetle communities is affected by stand management, region and decay progress of dead wood. 3. Our results indicated that beetle communities overall avoided logs of Fraxinus, Pseudotsuga, Larix and Tilia, and Carpinus ranked highest in preference. Carpinus also ranked highest for communities of broadleaf specialists; Picea abies ranked highest for communities of conifer specialists. Model-based recursive partitioning revealed that ranking of local hosts in conifer stands differed from that of broadleaf stands, and that ranking of hosts in broadleaf stands differed between regions, but only in the first year for both. 4. Synthesis and applications. Early-colonizing saproxylic beetle communities vary locally in their choice of host trees. Therefore, forest managers should focus on the enrichment of dead wood of regional tree species and tree species of the local stand to successfully promote earlycolonizing beetle.

Species’ functional traits set the blueprint for pair-wise interactions in ecological networks. Yet, it is unknown to what extent the functional diversity of plant and animal communities controls network assembly along environmental gradients in real-world ecosystems. Here we address this question with a unique dataset of mutualistic bird–fruit, bird–flower and insect–flower interaction networks and associated functional traits of 200 plant and 282 animal species sampled along broad climate and land-use gradients on Mt. Kilimanjaro. We show that plant functional diversity is mainly limited by precipitation, while animal functional diversity is primarily limited by temperature. Furthermore, shifts in plant and animal functional diversity along the elevational gradient control the niche breadth and partitioning of the respective other trophic level. These findings reveal that climatic constraints on the functional diversity of either plants or animals determine the relative importance of bottom-up and top-down control in plant–animal interaction networks., Differential responses of plant and animal functional diversity to climatic variation could affect trait matching in mutualistic interactions. Here, Albrecht et al. show that network structure varies across an elevational gradient owing to bottom-up and top-down effects of functional diversity.

Summary 1.The decline of managed honeybees and the rapid expansion of mass-flowering crops increase the risk of pollination limitation in crops and raise questions about novel management approaches for wild pollinators in agroecosystems. Adding artificial nesting sites, such as trap nests, can promote cavity-nesting bees in agroecosystems, but effectiveness could be limited by the availability of floral resources in the surrounding landscape and by natural antagonists. 2.In two European regions, we exposed artificial trap nests in paired field boundaries adjacent to oilseed rape (OSR) fields or non-flowering crops for two years within 32 landscapes covering two independent gradients of OSR cover and semi-natural habitat (SNH) cover in the landscape. We analysed the effects of local and landscape-wide floral resource availability, land-use intensity, landscape complexity and natural antagonists on community composition and population dynamics of trap-nesting bees. 3.Number of brood cells showed a strong, three-fold increase in response to the additional nesting sites. Species richness and abundance of cavity-nesting bees that were active during OSR flowering increased significantly with increasing amount of early-season landscape-wide floral resource availability, such as the cultivation of OSR. Later foraging species benefited instead from the availability of late-season alternative flower resources or SNH cover once the mass-flowering had ceased. Density-dependent parasitism increased following mass-flowering, while no density-dependent effect was found during mass-flowering. 4.Structural equation modelling revealed that the influence of floral resource availability on community growth rate was mediated by community size. Community size showed a strong negative effect on community growth rate. Despite positive density-dependent parasitism, antagonists had only weak regulating effects on community growth rate. 5.Synthesis and applications. Trap-nesting bee populations grow markedly with the increasing availability of food resources in the landscape and effectiveness of trap nests is only marginally limited by natural antagonists. Thus, trap nests could be a simple pollinator-supporting strategy to accompany the current expansion of mass-flowering crops, and to ensure pollination services for insect-pollinated crops. Trap nests benefit not only early season active generalist bees during oilseed rape flowering but also species with later phenology if accompanied by other pollinator-supporting practices. This article is protected by copyright. All rights reserved.

Aim Understanding the mechanisms controlling variation in species richness along environmental gradients is one of the most important objectives in ecology. Resource availability is often considered as the major driver of animal diversity. However, in ectotherms, temperature might play a predominant role as it modulates metabolic rates and the access of animals to resources. Here, we investigate the relative importance of resource availability and temperature in determining the diversity pattern of bees along a 3.6-km elevational gradient. Location Mount Kilimanjaro, Tanzania. Methods We assessed bee species richness and abundance with pan traps and floral resources with transect records on 60 study sites which were equally distributed over six near-natural and six disturbed habitat types along an elevational gradient from 870 to 4550 m a.s.l. We used path analysis to disentangle the effects of temperature, precipitation, floral resource abundance, bee abundance and land use on bee species richness. In addition, we monitored flower visitation rates during transect walks at different elevations to evaluate the temperature dependence of bee–flower interactions. Results Bee species richness continuously declined with elevation in natural and disturbed habitats. While the abundance of floral resources had a significant but only weak effect on species richness, the effect of temperature was strong. Temperature had a strong positive effect on species richness that was not mediated by bee abundance and an indirect effect via bee abundances. We observed higher levels of bee–flower interactions at higher temperatures, supporting the hypothesis that temperature limits diversity by constraining resource exploitation in ectotherms. Main conclusions Temperature and the availability of resources shape species richness patterns along environmental gradients. In ectothermic organisms like bees temperature seems to have the more important role, as it both limits the access to resources (abundance-mediated effect) and accelerates other (abundance-independent) ecological and evolutionary processes that drive the maintenance and origination of diversity.

BackgroundArtificial rearing of honey bee larvae is an established method which enables to fully standardize the rearing environment and to manipulate the supplied diet to the brood. However, there are no studies which compare learning performance or neuroanatomic differences of artificially-reared (in-lab) bees in comparison with their in-hive reared counterparts.MethodsHere we tested how different quantities of food during larval development affect body size, brain morphology and learning ability of adult honey bees. We used in-lab rearing to be able to manipulate the total quantity of food consumed during larval development. After hatching, a subset of the bees was taken for which we made 3D reconstructions of the brains using confocal laser-scanning microscopy. Learning ability and memory formation of the remaining bees was tested in a differential olfactory conditioning experiment. Finally, we evaluated how bees reared with different quantities of artificial diet compared to in-hive reared bees.ResultsThorax and head size of in-lab reared honey bees, when fed the standard diet of 160 µl or less, were slightly smaller than hive bees. The brain structure analyses showed that artificially reared bees had smaller mushroom body (MB) lateral calyces than their in-hive counterparts, independently of the quantity of food they received. However, they showed the same total brain size and the same associative learning ability as in-hive reared bees. In terms of mid-term memory, but not early long-term memory, they performed even better than the in-hive control.DiscussionWe have demonstrated that bees that are reared artificially (according to the Aupinel protocol) and kept in lab-conditions perform the same or even better than their in-hive sisters in an olfactory conditioning experiment even though their lateral calyces were consistently smaller at emergence. The applied combination of experimental manipulation during the larval phase plus subsequent behavioral and neuro-anatomic analyses is a powerful tool for basic and applied honey bee research.

Habitat fragmentation is a primary threat to biodiversity, but how it affects the structure and stability of ecological networks is poorly understood. Here, we studied plant–pollinator and host–parasitoid networks on 32 calcareous grassland fragments covering a size gradient of several orders of magnitude and with amounts of additional habitat availability in the surrounding landscape that varied independent of fragment size. We find that additive and interactive effects of habitat fragmentation at local (fragment size) and landscape scales (1,750 m radius) directly shape species communities by altering the number of interacting species and, indirectly, their body size composition. These, in turn, affect plant–pollinator, but not host–parasitoid, network structure: the nestedness and modularity of plant–pollinator networks increase with pollinator body size. Moreover, pollinator richness increases modularity. In contrast, the modularity of host–parasitoid networks decreases with host richness, whereas neither parasitoid richness nor body size affects network structure. Simulating species coextinctions also reveals that the structure–stability relationship depends on species’ sensitivity to coextinctions and their capacity for adaptive partner switches, which differ between mutualistic and antagonistic interaction partners. While plant–pollinator communities may cope with future habitat fragmentation by responding to species loss with opportunistic partner switches, past effects of fragmentation on the current structure of host–parasitoid networks may strongly affect their robustness to coextinctions under future habitat fragmentation. Analysing the structure of both plant–pollinator and host–parasitoid networks in calcareous grasslands, the authors reveal scale-dependent responses to habitat fragmentation in the structure and stability of different network types.

The increasing demand for insect pollinated crops and high recent losses of honey bee colonies raise concerns about food security. Systemic insecticides are recognized as one of the drivers of worldwide honey and wild bee declines. Particularly honey bees in agricultural environments are exposed to pesticides when they collect crop pollen and nectar. However, landscape scale studies which analyze pollen use and foraging distances of honey bees on mass-flowering crops like maize to evaluate potential exposure risks are currently lacking. In an experimental approach on a landscape scale we took advantage of intra-colonial dance communication to gather information about the location of utilized pollen resources. During maize flowering, four observation hives were placed in and rotated between 11 different landscapes which covered a gradient from low to high maize acreage. A higher frequency of dances for foraging locations on maize fields compared to other land use types shows that maize is an intensively used pollen resource for honey bee colonies. Mean foraging distances were significantly shorter for maize pollen than for other pollen origins. The percentage of maize pollen foragers did not increase with maize acreage in the landscape. The proportion of grassland area providing alternative pollen sources did not reduce the percentage of maize pollen foragers. Our findings allow estimating the distance-related exposure risk of honey bee colonies to pollen from surrounding maize fields treated with systemic insecticides. Similarly, the results can be used to estimate the exposure to transgenic maize pollen, which is relevant for honey production in European countries. Provision of alternative pollen resources within agri-environmental schemes could potentially reduce exposure risk to pesticide contaminated crop pollen.