Bees display limited acclimation capacity for heat tolerance.

Bees are essential pollinators and understanding their ability to cope with extreme temperature changes is crucial for predicting their resilience to climate change, but studies are limited. We measured the response of the critical thermal maximum (CTMax) to short-term acclimation in foragers of six bee species from the Greek island of Lesvos, which differ in body size, nesting habit, and level of sociality. We calculated the acclimation response ratio as a metric to assess acclimation capacity and tested whether bees' acclimation capacity was influenced by body size and/or CTMax. We also assessed whether CTMax increases following acute heat exposure simulating a heat wave. Average estimate of CTMax varied among species and increased with body size but did not significantly shift in response to acclimation treatment except in the sweat bee Lasioglossum malachurum. Acclimation capacity averaged 9% among species and it was not significantly associated with body size or CTMax. Similarly, the average CTMax did not increase following acute heat exposure. These results indicate that bees might have limited capacity to enhance heat tolerance via acclimation or in response to prior heat exposure, rendering them physiologically sensitive to rapid temperature changes during extreme weather events. These findings reinforce the idea that insects, like other ectotherms, generally express weak plasticity in CTMax, underscoring the critical role of behavioral thermoregulation for avoidance of extreme temperatures. Conserving and restoring native vegetation can provide bees temporary thermal refuges during extreme weather events.

Intrafloral patterns of color and scent in Capparis spinosa L. and the ghosts of its selection past.

PREMISE: Capparis spinosa is a widespread charismatic plant, in which the nocturnal floral habit contrasts with the high visitation by diurnal bees and the pronounced scarcity of hawkmoths. To resolve this discrepancy and elucidate floral evolution of C. spinosa, we analyzed the intrafloral patterns of visual and olfactory cues in relation to the known sensory biases of the different visitor guilds (bees, butterflies, and hawkmoths). METHODS: We measured the intrafloral variation of scent, reflectance spectra, and colorimetric properties according to three guilds of known visitors of C. spinosa. Additionally, we sampled visitation rates using a motion-activated camera. RESULTS: Carpenter bees visited the flowers eight times more frequently than nocturnal hawkmoths, at dusk and in the following morning. Yet, the floral headspace of C. spinosa contained a typical sphingophilous scent with high emission rates of certain monoterpenes and amino-acid derived compounds. Visual cues included a special case of multisensory nectar guide and color patterns conspicuous to the visual systems of both hawkmoths and bees. CONCLUSIONS: The intrafloral patterns of sensory stimuli suggest that hawkmoths have exerted strong historical selection on C. spinosa. Our study revealed two interesting paradoxes: (a) the flowers phenotypically biased towards the more inconsistent pollinator; and (b) floral display demands an abundance of resources that seems maladaptive in the habitats of C. spinosa. The transition to a binary pollination system accommodating large bees has not required phenotypic changes, owing to specific eco-physiological adaptations, unrelated to pollination, which make this plant an unusual case in pollination ecology.

Risk to pollinators from anthropogenic electro-magnetic radiation (EMR): Evidence and knowledge gaps.

Worldwide urbanisation and use of mobile and wireless technologies (5G, Internet of Things) is leading to the proliferation of anthropogenic electromagnetic radiation (EMR) and campaigning voices continue to call for the risk to human health and wildlife to be recognised. Pollinators provide many benefits to nature and humankind, but face multiple anthropogenic threats. Here, we assess whether artificial light at night (ALAN) and anthropogenic radiofrequency electromagnetic radiation (AREMR), such as used in wireless technologies (4G, 5G) or emitted from power lines, represent an additional and growing threat to pollinators. A lack of high quality scientific studies means that knowledge of the risk to pollinators from anthropogenic EMR is either inconclusive, unresolved, or only partly established. A handful of studies provide evidence that ALAN can alter pollinator communities, pollination and fruit set. Laboratory experiments provide some, albeit variable, evidence that the honey bee Apis mellifera and other invertebrates can detect EMR, potentially using it for orientation or navigation, but they do not provide evidence that AREMR affects insect behaviour in ecosystems. Scientifically robust evidence of AREMR impacts on abundance or diversity of pollinators (or other invertebrates) are limited to a single study reporting positive and negative effects depending on the pollinator group and geographical location. Therefore, whether anthropogenic EMR (ALAN or AREMR) poses a significant threat to insect pollinators and the benefits they provide to ecosystems and humanity remains to be established.

Moderate fire severity is best for the diversity of most of the pollinator guilds in Mediterranean pine forests.

Fire, a frequent disturbance in the Mediterranean, affects pollinator communities. We explored the response of major pollinator guilds to fire severity, across a fire-severity gradient at different spatial scales. We show that the abundance of all pollinator groups responded to fire severity, and that bees and beetles showed in addition a significant species-diversity response. Bees, sawflies, and wasps responded to fire severity at relatively small spatial scales (250-300 m), whereas flies and beetles responded at larger spatial scales. The response of bees, sawflies, and wasps was unimodal, as predicted by the intermediate disturbance hypothesis, whereas flies and beetles showed a negative response. A possible explanation is that the observed patterns (spatial scale and type of response) are driven by taxa-specific ecological and life-history traits, such as nesting preference and body size, as well as the availability of resources in the postfire landscape. Our observational study provides an insight into the effect of fire severity on pollinators. However, future research exploring the explicit link between the pre- and postfire landscape structure and pollinator traits and responses is required for further establishment and understanding of cause-effect relationships.

Fluorescent Pan Traps Affect the Capture Rate of Insect Orders in Different Ways.

To monitor and quantify the changes in pollinator communities over time, it is important to have robust survey techniques of insect populations. Pan traps allow for the assessment of the relative insect abundance in an environment and have been promoted by the Food and Agricultural Organization (FAO) as an efficient data collection methodology. It has been proposed that fluorescent pan traps are particularly useful, as it has been suggested that they capture high numbers of insects in an unbiased fashion. We use a simultaneous presentation of fluorescent and non-fluorescent pan trap colours to assess how flower-visiting insects of different orders respond to visual stimuli and reveal a significant interaction between trap fluorescence and captured insect type. In particular, Coleoptera (beetles) and Lepidoptera (butterflies and moths) were captured significantly more frequently by fluorescent traps, whilst Dipterans (flies) were captured significantly less frequently by this type of pan trap. Hymenopterans (bees and wasps) showed no significant difference in their preference for fluorescent or non-fluorescent traps. Our results reveal that the use of fluorescent pan traps may differently bias insect capture rates when compared to the typical experience of colour flower-visiting insects in natural environments. Correction factors may, therefore, be required for interpreting insect pan trap data collected with different methodologies.

Differential Effects of Climate Warming on the Nectar Secretion of Early- and Late-Flowering Mediterranean Plants.

Floral nectar is a vital resource for pollinators, thus having a very important role in ecosystem functioning. Ongoing climate warming could have a negative effect on nectar secretion, particularly in the Mediterranean, where a strong temperature rise is expected. In turn, decreased nectar secretion, together with shifts in flowering phenology can disrupt plant-pollinator interactions and consequently affect the entire ecosystem. Under fully controlled conditions, we tested how temperature influenced nectar secretion (through nectar volume, sugar concentration, sugar content, and number of flowers produced) in six Mediterranean plant species flowering from winter to summer (viz. Asphodelus ramosus, Ballota acetabulosa, Echium plantagineum, Lavandula stoechas, Rosmarinus officinalis, and Teucrium divaricatum). We compared the changes in nectar secretion under temperatures expected by the end of the century and estimated the effect of climate warming on nectar secretion of plants flowering in different seasons. We found a significant effect of temperature on nectar secretion, with a negative effect of very high temperatures in all species. Optimal temperatures for nectar secretion were similar to the mean temperatures in the recent past (1958-2001) during the respective flowering time of each species. Increasing temperatures, however, will affect differently the early-flowering (blooming in winter and early spring) and late-flowering species (blooming in late spring and early summer). Temperature rise expected by the end of the century will shift the average temperature beyond the optimal range for flower production and the sugar produced per plant in late-flowering species. Therefore, we expect a future decrease in nectar secretion of late-flowering species, which could reduce the amount of nectar resources available for their pollinators. Early-flowering plants will be less affected (optimal temperatures were not significantly different from the future projected temperatures), and may in some cases even benefit from rising temperatures. However, as many earlier studies have found that early-flowering species are more prone to shifts in phenology, the plant-pollinator interactions could instead become affected in a different manner. Consequently, climate warming will likely have a distinctive effect on both plant and pollinator populations and their interactions across different seasons.

Disentangling the role of floral sensory stimuli in pollination networks.

Despite progress in understanding pollination network structure, the functional roles of floral sensory stimuli (visual, olfactory) have never been addressed comprehensively in a community context, even though such traits are known to mediate plant-pollinator interactions. Here, we use a comprehensive dataset of floral traits and a novel dynamic data-pooling methodology to explore the impacts of floral sensory diversity on the structure of a pollination network in a Mediterranean scrubland. Our approach tracks transitions in the network behaviour of each plant species throughout its flowering period and, despite dynamism in visitor composition, reveals significant links to floral scent, and/or colour as perceived by pollinators. Having accounted for floral phenology, abundance and phylogeny, the persistent association between floral sensory traits and visitor guilds supports a deeper role for sensory bias and diffuse coevolution in structuring plant-pollinator networks. This knowledge of floral sensory diversity, by identifying the most influential phenotypes, could help prioritize efforts for plant-pollinator community restoration.

Landscape spatial configuration is a key driver of wild bee demographics.

The majority of studies investigating the effects of landscape composition and configuration on bee populations have been conducted in regions of intensive agricultural production, ignoring regions which are dominated by seminatural habitats, such as the islands of the Aegean Archipelago. In addition, research so far has focused on the landscape impacts on bees sampled in cropped fields while the landscape effects on bees inhabiting seminatural habitats are understudied. Here, we investigate the impact of the landscape on wild bee assemblages in 66 phryganic (low scrubland) communities on 8 Aegean islands. We computed landscape metrics (total area and total perimeter-area ratio) in 4 concentric circles (250, 500, 750, and 1000 m) around the center of each bee sampling site including 3 habitat groups (namely phrygana, cultivated land, and natural forests). We further measured the local flower cover in 25 quadrats distributed randomly at the center of each sampling site. We found that the landscape scale is more important than the local scale in shaping abundance and species richness of bees. Furthermore, habitat configuration was more important than the total area of habitats, probably because it affects bees' movement across the landscape. Phrygana and natural forests had a positive effect on bee demographics, while cultivated land had a negative effect. This demonstrates that phryganic specialists drive bee assemblages in these seminatural landscapes. This finding, together with the shown importance of landscape scale, should be considered for the management of wild bees with special emphasis placed on the spatial configuration of seminatural habitats.

Moderation is best: effects of grazing intensity on plant--flower visitor networks in Mediterranean communities.

The structure of pollination networks is an important indicator of ecosystem stability and functioning. Livestock grazing is a frequent land use practice that directly affects the abundance and diversity of flowers and pollinators and, therefore, may indirectly affect the structure of pollination networks. We studied how grazing intensity affected the structure of plant-flower visitor networks along a wide range of grazing intensities by sheep and goats, using data from 11 Mediterranean plant-flower visitor communities from Lesvos Island, Greece. We hypothesized that intermediate grazing might result in higher diversity as predicted by the Intermediate Disturbance Hypothesis, which could in turn confer more stability to the networks. Indeed, we found that networks at intermediate grazing intensities were larger, more generalized, more modular, and contained more diverse and even interactions. Despite general responses at the network level, the number of interactions and selectiveness of particular flower visitor and plant taxa in the networks responded differently to grazing intensity, presumably as a consequence of variation in the abundance of different taxa with grazing. Our results highlight the benefit of maintaining moderate levels of livestock grazing by sheep and goats to preserve the complexity and biodiversity of the rich Mediterranean communities, which have a long history of grazing by these domestic animals.

Lessons from Red Data Books: Plant Vulnerability Increases with Floral Complexity.

The architectural complexity of flower structures (hereafter referred to as floral complexity) may be linked to pollination by specialized pollinators that can increase the probability of successful seed set. As plant-pollinator systems become fragile, a loss of such specialized pollinators could presumably result in an increased likelihood of pollination failure. This is an issue likely to be particularly evident in plants that are currently rare. Using a novel index describing floral complexity we explored whether this aspect of the structure of flowers could be used to predict vulnerability of plant species to extinction. To do this we defined plant vulnerability using the Red Data Book of Rare and Threatened Plants of Greece, a Mediterranean biodiversity hotspot. We also tested whether other intrinsic (e.g. life form, asexual reproduction) or extrinsic (e.g. habitat, altitude, range-restrictedness) factors could affect plant vulnerability. We found that plants with high floral complexity scores were significantly more likely to be vulnerable to extinction. Among all the floral complexity components only floral symmetry was found to have a significant effect, with radial-flower plants appearing to be less vulnerable. Life form was also a predictor of vulnerability, with woody perennial plants having significantly lower risk of extinction. Among the extrinsic factors, both habitat and maximum range were significantly associated with plant vulnerability (coastal plants and narrow-ranged plants are more likely to face higher risk). Although extrinsic and in particular anthropogenic factors determine plant extinction risk, intrinsic traits can indicate a plant's proneness to vulnerability. This raises the potential threat of declining global pollinator diversity interacting with floral complexity to increase the vulnerability of individual plant species. There is potential scope for using plant-pollinator specializations to identify plant species particularly at risk and so target conservation efforts towards them.

Climate change reduces nectar secretion in two common Mediterranean plants.

Global warming can lead to considerable impacts on natural plant communities, potentially inducing changes in plant physiology and the quantity and quality of floral rewards, especially nectar. Changes in nectar production can in turn strongly affect plant-pollinator interaction networks-pollinators may potentially benefit under moderate warming conditions, but suffer as resources reduce in availability as elevated temperatures become more extreme. Here, we studied the effect of elevated temperatures on nectar secretion of two Mediterranean Lamiaceae species-Ballota acetabulosa and Teucrium divaricatum. We measured nectar production (viz. volume per flower, sugar concentration per flower and sugar content per flower and per plant), number of open and empty flowers per plant, as well as biomass per flower under a range of temperatures selected ad hoc in a fully controlled climate chamber and under natural conditions outdoors. The average temperature in the climate chamber was increased every 3 days in 3 °C increments from 17.5 to 38.5 °C. Both study species showed a unimodal response of nectar production (volume per flower, sugar content per flower and per plant) to temperature. Optimal temperature for sugar content per flower was 25-26 °C for B. acetabulosa and 29-33 °C for T. divaricatum. According to our results, moderate climate warming predicted for the next few decades could benefit nectar secretion in T. divaricatum as long as the plants are not water stressed, but have a moderate negative effect on B. acetabulosa. Nevertheless, strong warming as predicted by climate change models for the end of the 21st century is expected to reduce nectar secretion in both species and can thus significantly reduce available resources for both wild bees and honeybees in Mediterranean systems.

The potential for indirect effects between co-flowering plants via shared pollinators depends on resource abundance, accessibility and relatedness.

Co-flowering plant species commonly share flower visitors, and thus have the potential to influence each other's pollination. In this study we analysed 750 quantitative plant-pollinator networks from 28 studies representing diverse biomes worldwide. We show that the potential for one plant species to influence another indirectly via shared pollinators was greater for plants whose resources were more abundant (higher floral unit number and nectar sugar content) and more accessible. The potential indirect influence was also stronger between phylogenetically closer plant species and was independent of plant geographic origin (native vs. non-native). The positive effect of nectar sugar content and phylogenetic proximity was much more accentuated for bees than for other groups. Consequently, the impact of these factors depends on the pollination mode of plants, e.g. bee or fly pollinated. Our findings may help predict which plant species have the greatest importance in the functioning of plant-pollination networks.

Contribution of insect pollinators to crop yield and quality varies with agricultural intensification.

Background. Up to 75% of crop species benefit at least to some degree from animal pollination for fruit or seed set and yield. However, basic information on the level of pollinator dependence and pollinator contribution to yield is lacking for many crops. Even less is known about how insect pollination affects crop quality. Given that habitat loss and agricultural intensification are known to decrease pollinator richness and abundance, there is a need to assess the consequences for different components of crop production. Methods. We used pollination exclusion on flowers or inflorescences on a whole plant basis to assess the contribution of insect pollination to crop yield and quality in four flowering crops (spring oilseed rape, field bean, strawberry, and buckwheat) located in four regions of Europe. For each crop, we recorded abundance and species richness of flower visiting insects in ten fields located along a gradient from simple to heterogeneous landscapes. Results. Insect pollination enhanced average crop yield between 18 and 71% depending on the crop. Yield quality was also enhanced in most crops. For instance, oilseed rape had higher oil and lower chlorophyll contents when adequately pollinated, the proportion of empty seeds decreased in buckwheat, and strawberries' commercial grade improved; however, we did not find higher nitrogen content in open pollinated field beans. Complex landscapes had a higher overall species richness of wild pollinators across crops, but visitation rates were only higher in complex landscapes for some crops. On the contrary, the overall yield was consistently enhanced by higher visitation rates, but not by higher pollinator richness. Discussion. For the four crops in this study, there is clear benefit delivered by pollinators on yield quantity and/or quality, but it is not maximized under current agricultural intensification. Honeybees, the most abundant pollinator, might partially compensate the loss of wild pollinators in some areas, but our results suggest the need of landscape-scale actions to enhance wild pollinator populations.

Multiple stressors on biotic interactions: how climate change and alien species interact to affect pollination.

Global change may substantially affect biodiversity and ecosystem functioning but little is known about its effects on essential biotic interactions. Since different environmental drivers rarely act in isolation it is important to consider interactive effects. Here, we focus on how two key drivers of anthropogenic environmental change, climate change and the introduction of alien species, affect plant-pollinator interactions. Based on a literature survey we identify climatically sensitive aspects of species interactions, assess potential effects of climate change on these mechanisms, and derive hypotheses that may form the basis of future research. We find that both climate change and alien species will ultimately lead to the creation of novel communities. In these communities certain interactions may no longer occur while there will also be potential for the emergence of new relationships. Alien species can both partly compensate for the often negative effects of climate change but also amplify them in some cases. Since potential positive effects are often restricted to generalist interactions among species, climate change and alien species in combination can result in significant threats to more specialist interactions involving native species.

Invasive plant integration into native plant-pollinator networks across Europe.

The structure of plant-pollinator networks has been claimed to be resilient to changes in species composition due to the weak degree of dependence among mutualistic partners. However, detailed empirical investigations of the consequences of introducing an alien plant species into mutualistic networks are lacking. We present the first cross-European analysis by using a standardized protocol to assess the degree to which a particular alien plant species (i.e. Carpobrotus affine acinaciformis, Impatiens glandulifera, Opuntia stricta, Rhododendron ponticum and Solanum elaeagnifolium) becomes integrated into existing native plant-pollinator networks, and how this translates to changes in network structure. Alien species were visited by almost half of the pollinator species present, accounting on average for 42 per cent of the visits and 24 per cent of the network interactions. Furthermore, in general, pollinators depended upon alien plants more than on native plants. However, despite the fact that invaded communities received more visits than uninvaded communities, the dominant role of alien species over natives did not translate into overall changes in network connectance, plant linkage level and nestedness. Our results imply that although supergeneralist alien plants can play a central role in the networks, the structure of the networks appears to be very permeable and robust to the introduction of invasive alien species into the network.

Potential contribution of natural enemies to patterns of local adaptation in plants.

Genetic differentiation among plant populations and adaptation to local environmental conditions are well documented. However, few studies have examined the potential contribution of plant antagonists, such as insect herbivores and pathogens, to the pattern of local adaptation. Here, a reciprocal transplant experiment was set up at three sites across Europe using two common plant species, Holcus lanatus and Plantago lanceolata. The amount of damage by the main above-ground plant antagonists was measured: a rust fungus infecting Holcus and a specialist beetle feeding on Plantago, both in low-density monoculture plots and in competition with interspecific neighbours. Strong genetic differentiation among provenances in the amount of damage by antagonists in both species was found. Local provenances of Holcus had significantly higher amounts of rust infection than foreign provenances, whereas local provenances of Plantago were significantly less damaged by the specialist beetle than the foreign provenances. The presence of surrounding vegetation affected the amount of damage but had little influence on the ranking of plant provenances. The opposite pattern of population differentiation in resistance to local antagonists in the two species suggests that it will be difficult to predict the consequences of plant translocations for interactions with organisms of higher trophic levels.

Ultrastructure and motility pattern of the spermatozoa of Aleochara curtula (Coleoptera, Staphylinidae).

Ultrastructure and motility pattern of spermatozoa of the rove beetle Aleochara curtula were examined using electron and light microscopic methods. The spermatozoon is about 100 microm long and filiform. The head piece comprises a 5 microm long triple layered acrosome and 10 microm long nucleus. The flagellum consists of a 9+9+2 axoneme, two accessory bodies and two mitochondrial derivatives about equal in size but of different shape in their cross sections. In both derivatives there are paracrystalline inclusions. The flagellum is attached to the head by a 2 microm long centriole adjunct which is characterized by its electron dense material that forms a three layered folded lamellar structure. When liberated in buffer solution the sperm flagella assume a coiled hook-like form with the excentric stiff head protruding in front. The spermatozoa are driven through the medium by a small helicoidal wave of high frequency superimposed to the bent flagella. The maximum speed measured was 15.2 microm/s. The sperm architecture of A. curtula is similar to that of other Aleochara species but differs in total length and dimensions of the mitochondrial derivatives. For that reason Aleochara sperm can certainly prove useful to study the effect of the mitochondrial derivatives on sperm motility.