Condition-dependent expression of virulence in a trypanosome infecting bumblebees

Parasite virulence affects both the temporal dynamics of host-parasite relationships and the degree to which parasites regulate host populations. If hosts can compensate for parasitism, then parasites may exhibit condition-dependent virulence, with high virulence being seen only when the host is under conditions of stress. Despite their usually low level of virulence, theory suggests that such parasites may still affect host population dynamics. We tested whether a trypanosome intestinal parasite of bumblebees, Crithidia bombi , expresses condition-dependent virulence. Hosts were infected with the parasite and then kept under either favourable or starvation (stressed) conditions. Under favourable conditions the infection caused no mortality, while when hosts were starved the infection increased the host mortality rate by 50%. In addition, we found a parasite-related change in host resource allocation patterns. Infected bees invested relatively more resources into their fat body and less into their reproductive system than did non-infected bees. Whether this reallocation is parasite-driven, to enhance transmission, or a host-response to parasitism, remains unknown.     

Non-native plants affect generalist pollinator diet overlap and foraging behavior indirectly,via impacts on native plant abundance

Flowering invasive plants can have dramatic effects on the resource landscape available to pollinators. Because many pollinators exhibit behavioral plasticity in response to competitor or resource density, this in turn can result in impacts on ecological processes such as pollination and plant reproduction. We examine how interactions between five common generalist eusocial bees change across an invasion gradient by examining how bee abundance and diet overlap changed with variation in both invasive plant abundance and competitor abundance in a temperate oak-savannah ecosystem. Specifically we focus on the bumblebees Bombus bifarius, B. mixtus, B. melanopygus and B. vosnesenskii, as well as the non-native honeybee Apis mellifera, and their interactions with the native flowering plants Camassia quamash, Camassia liechtlinii, and the invasive shrub Cytisus scoparius. We further examine whether changes in pollinator visits to the invasive and two common native plants can explain changes in diet overlap. Abundance of the invasive plant and other common floral resources had strong impacts on focal bee abundance, with certain species more likely to be present at highly invaded sites. This may be because highly invaded sites tended to be embedded in forested landscapes where those bees are common. Diet overlap was most affected by abundance of a common native plant, rather than the invasive plant, with diet overlap increasing non-linearly with abundance of the native plant. Furthermore, Apis mellifera, did not appear to have direct competitive effects on native bumblebees in this habitat. However, visit patterns suggest that bees most abundant at highly invaded sites may compete for access to native resources. Thus the impacts of this invasive plant on our focal bee species may be primarily indirect, via its’ competitive effects on native plants.     

Population genetics of wild and managed pollinators: implications for crop pollination and the genetic integrity of wild bees

Loss of habitat and chemical use associated with agriculture can cause population declines of wild pollinators. Less is known about the evolutionary consequences of interactions between species used in commercial agriculture and wild pollinators. Given population declines of many wild bee species, it is crucial to understand if commercial queens become established in natural areas, if wild bees visit agricultural fields and have the potential to interact with commercial bees, and if gene flow occurs between commercial and wild bees. We drew on a long-term data set that documents commercial bumble bee (Bombus impatiens) use in New England, and we conducted genetic analyses of foraging B. impatiens from areas with varying intensities of commercial bee use. In agricultural areas with a history of commercial bee use we also sampled bees directly from commercial hives. We found significant genetic differences among foraging B. impatiens and B. impatiens sampled directly from hives (average pairwise F′_ST = 0.14), but not among samples of foraging bees from natural areas (average F′_ST among foraging bees?=?0.002). Furthermore, Bayesian analysis of population structure revealed that foraging bees caught in areas with a history of commercial bee use grouped with samples from natural areas. These results document an agricultural setting where there was no widespread introgression of alleles from commercial bumble bees to wild bumble bees, commercial bumble bees did not become established in natural areas, and wild bees were providing pollination services to crops.     

Pervasiveness of parasites in pollinators

Many pollinator populations are declining, with large economic and ecological implications. Parasites are known to be an important factor in the some of the population declines of honey bees and bumblebees, but little is known about the parasites afflicting most other pollinators, or the extent of interspecific transmission or vectoring of parasites. Here we carry out a preliminary screening of pollinators (honey bees, five species of bumblebee, three species of wasp, four species of hoverfly and three genera of other bees) in the UK for parasites. We used molecular methods to screen for six honey bee viruses, Ascosphaera fungi, Microsporidia, and Wolbachia intracellular bacteria. We aimed simply to detect the presence of the parasites, encompassing vectoring as well as actual infections. Many pollinators of all types were positive for Ascosphaera fungi, while Microsporidia were rarer, being most frequently found in bumblebees. We also detected that most pollinators were positive for Wolbachia, most probably indicating infection with this intracellular symbiont, and raising the possibility that it may be an important factor in influencing host sex ratios or fitness in a diversity of pollinators. Importantly, we found that about a third of bumblebees (Bombus pascuorum and Bombus terrestris) and a third of wasps (Vespula vulgaris), as well as all honey bees, were positive for deformed wing virus, but that this virus was not present in other pollinators. Deformed wing virus therefore does not appear to be a general parasite of pollinators, but does interact significantly with at least three species of bumblebee and wasp. Further work is needed to establish the identity of some of the parasites, their spatiotemporal variation, and whether they are infecting the various pollinator species or being vectored. However, these results provide a first insight into the diversity, and potential exchange, of parasites in pollinator communities.     

The relationship between the abundances of bumblebees and honeybees in a native habitat

Abstract. 1. The western honeybee, Apis mellifera, has been introduced to many parts of the world and is sometimes purported to be detrimental to native bees because it reduces their food base. It is seldom viewed in this light in Europe; however, when beekeepers maintain very high bee densities, the species could also be displacing insects in its native European range by reducing the resource base. 2. In England, populations of bumblebees (Bombus Latr. Hym.) have been decreasing both in terms of diversity and abundance, mainly because of a loss of habitat resulting from agricultural intensification. The impact of competition from other flower feeders is largely unknown. 3. Nineteen dry lowland heaths in southern England were sampled once for honeybees and bumblebees. Honeybee abundance varied from 4 to 81 bees per 100 m² (mean = 30.89, median = 23), whereas bumblebees varied from 2 to 17 individuals per 100 m² (mean = 8.26, median = 7), belonging to between one and five species. There was a negative association between honeybee and bumblebee abundance but there was no apparent relationship between honeybee abundance and bumblebee diversity. 4. The Bray–Curtis coefficient was used to compare the similarity in honeybee and bumblebee floral host breadth at these 19 sites. The coefficient was negatively associated with honeybee abundance: thus where honeybees were most abundant, bumblebees were fewer and/or foraged on different flower species. 5. Foraging host breadth was also examined at four heathlands over a field season (April to September). No association between honeybee abundance and foraging host breadth was found for short‐tongued bees, although there was some evidence for a change in floral host breadth for long‐tongued bees. 6. It is concluded that the impact of honeybees on bumblebees is complex. Although competition between the two species cannot be ruled out, it is perhaps equally likely that bumblebees decline in response to other factors, and that honeybees move independently of this decline.     

传粉蜂在设施农业中的应用及挑战

昆虫为植物传粉是自然生态系统中的重要环节,在农业和自然生态系统的平衡与调控方面发挥着重要作用。以蜜蜂、熊蜂为代表的传粉蜂因其高效传粉及可人工饲养的特点,已成为设施农业中的优势传粉昆虫。本文总结了传粉蜂在设施农业中的应用现状,并从温湿度、农药、重金属等非生物因素和蜂种、病原生物、天敌、蜜源植物等生物因素两大方面讨论了传粉蜂在应用中面临的诸多挑战。此外,本文初步探讨了植物病虫害对传粉昆虫传粉效率的影响,并对传粉蜂未来的研究和应用方向进行了展望,旨在推动实现传粉蜂在农业中的高效授粉功能,为农产品增产增效服务。     

Effects of intrapopulation phenotypic traits of invasive crayfish on leaf litter processing

Environmental gradients and competition influence aquatic macrophyte distribution in estuaries. The competition-to-stress hypothesis states that some species are excluded from lower estuaries (high salinity) due to abiotic stress and others from upper estuaries (low salinity) by competition. The growth of Crinum americanum L. and Spartina alterniflora Loisel. in monoculture (10:0/0:10) and mixed culture (5:5) under different salinity levels (4/12/26) was analysed by a laboratory experiment (3 cultures × 3 sediment types × 3 replicate) to understand the role of competition and salinity on the distribution of these species in a tropical estuary as well as to verify whether the competition-to-stress hypothesis explains their zonation. We tested the hypothesis that S. alterniflora is not established in the upper estuary due to the effect of competition with C. americanum, whereas the latter presents restrictions to high salinity and has greater competitive ability in the upper estuary. Our data confirm the competition-to-stress hypothesis but not as proposed originally. We conclude that abiotic stress (low nutrient availability) is responsible for the absence of S. alterniflora in the upper estuary and that the competition between the two species is responsible for the absence of C. americanum in the lower estuary.     

Honey bees are the dominant diurnal pollinator of native milkweed in a large urban park

In eastern North America, the field milkweed, Asclepias syriaca L. (Asclepiadaceae), is used in planting schemes to promote biodiversity conservation for numerous insects including the endangered monarch butterfly, Danaus plexippus (Linnaeus) (Nymphalidae). Less is known about its pollinators, and especially in urban habitats where it is planted often despite being under increasing pressure from invasive plant species, such as the related milkweed, the dog‐strangling vine (DSV), Vincetoxicum rossicum (Kleopow) Barbar. (Asclepiadaceae). During the A. syriaca flowering period in July 2016, we surveyed bees in open habitats along a DSV invasion gradient and inspected 433 individuals of 25 bee species in 12 genera for pollinia: these were affixed to bees that visited A. syriaca for nectar and contain pollen packets that are vectored (e.g., transferred) between flowers. Of all bees sampled, pollinia were found only on the nonindigenous honeybee, Apis mellifera (43% of all bees identified), as well as one individual bumblebee, Bombus impatiens Cresson. Pollinia were recorded from 45.2% of all honeybees collected. We found no relationship between biomass of DSV and biomass of A. syriaca per site. There was a significant positive correlation between A. syriaca biomass and the number of pollinia, and the proportion vectored. No relationship with DSV biomass was detected for the number of pollinia collected by bees but the proportion of vectored pollinia declined with increasing DSV biomass. Although we find no evidence of DSV flowers attracting potential pollinators away from A. syriaca and other flowering plants, the impacts on native plant–pollinator mutualisms relate to its ability to outcompete native plants. As wild bees do not appear to visit DSV flowers, it could be altering the landscape to one which honeybees are more tolerant than native wild bees.     

Thinking beyond Western commercial honeybee hives: towards improved conservation of honey bee diversity

A decline of wild pollinators, along with a decline of bee diversity, has been a cause of concern among academics and governmental organizations. According to IPBES, a lack of wild pollinator data contributes to difficulties in comprehensively analyzing the regional status of wild pollinators in Africa, Latin America, Asia and Oceania. It may have also contributed to the prevailing lack of awareness of the diversity of honey bees, of which the managed Apis mellifera is often considered as “the (only) honey bee,” despite the fact that there are eight other honey bee species extant in Asia. A survey of 100 journal articles published in 2016 shows that 57% of the studies still identified A. mellifera as “the honey bee.” In total, 80% of studies were conducted solely on A. mellifera. This focus on A. mellifera has also caused the honey standard of Codex Alimentarius and the European Union to be based solely on A. mellifera, causing improper evaluation of honeys from other species. We recommend adapting current standards to reflect the diversity of honey bees and in the process correct failures in the honey market and pave the way towards improved protection of honey bee species and their habitats.     

Impacts of a pesticide on pollinator species richness at different spatial scales

Pesticides are an important potential cause of biodiversity and pollinator decline. Little is known about the impacts of pesticides on wild pollinators in the field. Insect pollinators were sampled in an agricultural system in Italy with the aim of detecting the impacts of pesticide use. The insecticide fenitrothion was over 150 times greater in toxicity than other pesticides used in the area, so sampling was set up around its application. Species richness of wild bees, bumblebees and butterflies were sampled at three spatial scales to assess responses to pesticide application: (i) the ‘field’ scale along pesticide drift gradients; (ii) the ‘landscape’ scale sampling in different crops within the area and (iii) the ‘regional’ scale comparing two river basins with contrasting agricultural intensity. At the field scale, the interaction between the application regime of the insecticide and the point in the season was important for species richness. Wild bee species richness appeared to be unaffected by one insecticide application, but declined after two and three applications. At the landscape scale, the species richness of wild bees declined in vine fields where the insecticide was applied, but did not decline in maize or uncultivated fields. At the regional scale, lower bumblebee and butterfly species richness was found in the more intensively farmed basin with higher pesticide loads. Our results suggest that wild bees are an insect pollinator group at particular risk from pesticide use. Further investigation is needed on how the type, quantity and timing of pesticide application impacts pollinators.     

How <Emphasis Type="Italic">Apis mellifera</Emphasis> Behaves with its Invasive Hornet Predator <Emphasis Type="Italic">Vespa velutina</Emphasis>?

Invasive species are now recognized as a major cause of native biodiversity loss worldwide. In the current deleterious context for pollinators, the invasive yellow-legged hornet, Vespa velutina, represents an additional threat to the domestic honeybee, Apis mellifera, in Europe. Therefore, understanding the impact of this predator on honeybee colonies is of major importance. In the present study, we tried to assess the impact of V. velutina on the honeybee foraging and defence behaviour based on the video monitoring of two hives. Balling behaviour is reported here for the first time under natural conditions in A. mellifera against V. velutina in Europe. Although these results are preliminary and should be carefully considered, we found that the number of hornets impacted honeybee foraging and defence behaviours. More defensive behaviours were notified in the hive, which survives slightly longer. This may suggest that selecting for more defensive colonies may provide an interesting perspective.     

Effects of alien invasion by <Emphasis Type="Italic">Bombus terrestris</Emphasis> L. (Apidae) on the visitation patterns of native bumblebees in coastal plants in northern Japan

When alien pollinator species enter a native community of pollinators in which resource partitioning has been established, the pollination network between plants and pollinators may be modified through the interactions between the pollinators over the use of floral resources. We observed the floral-use patterns of native (Bombus hypocrita and B. deuteronymus) and alien (B. terrestris) bumblebee species in a coastal grassland in northern Japan. We analyzed the factors determining resource partitioning patterns. B. hypocrita tended to visit flowers with shallow or wide open corollas, such as Rosa rugosa, whereas B. deuteronymus visited flowers with complex or deeper corollas, such as Lathyrus japonicus. Given the wider floral preference of B. terrestris, floral use by the alien bumblebees consistently overlapped with that of native bumblebees. The visitation of B. terrestris to R. rugosa flowers was positively correlated with that of B. hypocrita. These bumblebee species frequently used similar floral resources, in part because of the large overlap in the seasonality of their foraging activity. The visitation frequency of B. deuteronymus to L. japonicus flowers was independent of the visitation frequency of other bumblebee species. The major visitation periods of the bumblebees to L. japonicus flowers reciprocally differed between B. deuteronymus and B. terrestris, suggesting phenological resource partitioning between these species. Our study suggests that phenological niche partitioning is more common in specialized flowers (L. japonicus) than in generalized flowers (R. rugosa).     

Intersexual mimicry and flowering phenology facilitate pollination in a dioecious habitat specialist species, <Emphasis Type="Italic">Myristica fatua</Emphasis> (Myristicaceae)

Myristica fatua is a dioecious specialist species restricted to the endangered, freshwater Myristica swamp forests in the Western Ghats, India. Earlier studies have alluded to pollination by deception in members of the Myristica genus, and thus we examined the pollination ecology comprising floral biology, flower production, flower visitors, and reproductive success in M. fatua and inferred the potential strategies that could permit such deception in this habitat specialist tree. Male flowers provide pollen rewards for an extended period of time while female flowers are rewardless and both sexes are visited by generalist insects, mainly by honeybees and stingless bees. Bee visits were significantly more frequent and longer on male than on female flowers as bees collected pollen from male flowers. We found that flower production patterns create a preponderance of males compared to females in the swamp populations. Using a model of honeybee color vision, we found the distance between the color loci of male and female flowers and based on minimum visual angle subtended by these flowers, we suggest that the two floral sexes cannot be discriminated by bees. Bees are likely deceived by the perceptual similarity of rewardless female flowers to pollen-offering male flowers and pollination is the consequence of foraging errors made by pollinators that encounter largely male–rarely female flower mosaics as they forage among clump-distributed M. fatua trees in the swamp habitat.     

Pesticide exposure affects flight dynamics and reduces flight endurance in bumblebees

The emergence of agricultural land use change creates a number of challenges that insect pollinators, such as eusocial bees, must overcome. Resultant fragmentation and loss of suitable foraging habitats, combined with pesticide exposure, may increase demands on foraging, specifically the ability to collect or reach sufficient resources under such stress. Understanding effects that pesticides have on flight performance is therefore vital if we are to assess colony success in these changing landscapes. Neonicotinoids are one of the most widely used classes of pesticide across the globe, and exposure to bees has been associated with reduced foraging efficiency and homing ability. One explanation for these effects could be that elements of flight are being affected, but apart from a couple of studies on the honeybee (Apis mellifera), this has scarcely been tested. Here, we used flight mills to investigate how exposure to a field realistic (10 ppb) acute dose of imidacloprid affected flight performance of a wild insect pollinator—the bumblebee, Bombus terrestris audax. Intriguingly, observations showed exposed workers flew at a significantly higher velocity over the first ¾ km of flight. This apparent hyperactivity, however, may have a cost because exposed workers showed reduced flight distance and duration to around a third of what control workers were capable of achieving. Given that bumblebees are central place foragers, impairment to flight endurance could translate to a decline in potential forage area, decreasing the abundance, diversity, and nutritional quality of available food, while potentially diminishing pollination service capabilities.     

A field experiment on the effect of Nosema bombi in colonies of the bumblebee Bombus terrestris

Abstract 1. Nosema bombi is a microsporidian parasite of bumblebees that is of potentially great ecological and economic importance. Nosema bombi has been shown to have detrimental fitness effects on the bumblebee Bombus terrestris under laboratory conditions. The present study investigates whether N. bombi also has negative fitness effects under natural conditions. 2. A field experiment was conducted with B. terrestris spring queens caught from the wild. In the laboratory these queens were assigned to either infected or control treatment groups. On the emergence of workers, colonies were placed in the field and a number of fitness and life‐history traits were measured. 3. The size of colonies produced by infected queens was significantly smaller than those of control queens. None of the 14 infected colonies produced sexual offspring, whereas four of the 14 control colonies reached the male production stage. Results of the field experiment are in agreement with prior laboratory‐based results, and even imply that such effects are amplified under natural conditions. 4. The evidence suggests that, on entering the colony early, N. bombi has a highly detrimental effect on the fitness of its host. It would seem that this parasite reduces its opportunities to transmit into the next host generation.     

Hunting for healthy microbiomes: determining the core microbiomes of <Emphasis Type="Italic">Ceratina,Megalopta</Emphasis>, and <Emphasis Type="Italic">Apis</Emphasis> bees and how they associate with microbes in bee collected pollen

Social corbiculate bees such as honey bees and bumble bees maintain a specific beneficial core microbiome which is absent in wild bees. It has been suggested that maintaining this microbiome can prevent disease and keep bees healthy. The main aim of our study was to identify if there are any core bacterial groups in the non-corbiculate bees Ceratina and Megalopta that have been previously overlooked. We additionally test for associations between the core bee microbes and pollen provisions to look for potential transmission between the two. We identify three enterotypes in Ceratina samples, with thirteen core bacterial phylotypes in Ceratina females: Rosenbergiella, Pseudomonas, Gilliamella, Lactobacillus, Caulobacter, Snodgrassella, Acinetobacter, Corynebacterium, Sphingomonas, Commensalibacter, Methylobacterium, Massilia, and Stenotrophomonas, plus 19 in pollen (6 of which are shared by bees). Unlike Apis bees, whose gut microbial community differs compared to their pollen, Ceratina adults and pollen largely share a similar microbial composition and enterotype difference was largely explained by pollen age. Megalopta displays a highly diverse composition of microbes throughout all adults, yet Lactobacillus and Saccharibacter were prevalent in 90% of adults as core bacteria. Only Lactobacillus was both a core bee and pollen provision microbe in all three species. The consequences of such diversity in core microbiota between bee genera and their associations with pollen are discussed in relation to identifying potentially beneficial microbial taxa in wild bees to aid the conservation of wild, understudied, non-model bee species.     

熊蜂生物学及种群影响因素研究进展

熊蜂是膜翅目蜜蜂科熊蜂属内物种的统称,全球约有250种,中国已知125种,是全球熊蜂资源最丰富的国家。该类昆虫是众多野生植物和农作物的重要传粉者,对维持自然生态系统和农业粮食生产极为重要。一些群势强、易于人工饲养的熊蜂物种被开发利用,为多种目标作物授粉。本文介绍了熊蜂的生物学特性和授粉应用现状,综述了栖息地丧失、气候变化、病原体传播、外来物种入侵及化学农药等多重因素对熊蜂种群的影响,并从熊蜂的应用基础研究、资源保护及授粉经济价值评估等多方面作了展望,旨在为中国本土熊蜂的保护、应用和生态功能研究提供参考。     

Foraging habitats and foraging distances of bumblebees, Bombus spp. (Hym., Apidae), in an agricultural landscape

Abstract: In selected foraging habitats of an agricultural landscape flower visits of bumblebees and community structure of foraging bumblebees were studied, with special regard to the role of crops as super-abundant resources. Most crops represent temporal foraging habitats with high abundance of bumblebees but mainly with low diversity in the bumblebee forage community, in contrast to permanent foraging habitats such as, for example, a hedgerow. The high numbers of bumblebees in the monoculture of crop plantations consisted mainly of short-tongued bumblebee species. The role of foraging distances for the visitation rate of foraging habitats was studied by performing capture–recapture experiments with natural nests of Bombus terrestris , Bombus lapidarius and Bombus muscorum . Differences were found on the species as well as the individual level. The foraging distances of B. muscorum were more restricted to the neighbourhood of the nesting habitat than the foraging activity of B. terrestris and B. lapidarius . High percentages of B. terrestris workers were recaptured while foraging on super-abundant resources in distances up to 1750 m from the nest. Isolated patches of highly rewarding forage crops, in agricultural landscapes, are probably only accessed by bumblebee species with large mean foraging distances, such as the short-tongued B. terrestris . Species like the rare, long-tongued B. muscorum depend on a close connection between nesting and foraging habitat. A restricted foraging radius might be one important factor of bumblebee species loss and potential pollinator limitation in modern agricultural landscapes. Furthermore, long-distance flights of bumblebee pollinators have to be considered in the present discussion on gene flow from transgenic plant species on a landscape scale.     

Increase in ant density promotes dual effects on bee behaviour and plant reproductive performance

Floral rewards do not only attract pollinators, but also herbivores and their predators. Ants are attracted by extrafloral nectaries (EFNs), situated near flowers, and may interfere with the efficiency and behaviour of pollinators. We tested the hypothesis that the impacts of ant–pollinator interactions in plant–pollinator systems are dependent on (1) the seasonal activity of EFNs, which increase ant abundance closer to flowers; (2) consequently, an ant effect, where ants decrease the temporal niche overlap of bees due to predator avoidance; and (3) ant density, where higher densities may negatively affect plant–pollinator interactions and plant performance. We studied two ant–plant–pollinator systems based on Banisteriopsis campestris and Banisteriopsis malifolia plant species. The periods of high ant abundance coincided with plant species blooming. The presence of ants around flowers reduced the visitation rates of the smaller bees and the temporal niche overlap between bee species was not higher than randomly expected when ants had free access. Additionally, we observed variable ant effects on fruit set and duration of bee visits to both Malpighiaceae species when ant density was experimentally kept constant on branches, especially on B. campestris. Our goal was to show the dual role of ant density effects, especially because the different outcomes are not commonly observed in the same plant species. We believe that reduced temporal niche overlap between floral visitors due to ant presence provides an opportunity for smaller bees to improve compatible pollination behaviour. Additionally, we concluded that ant density had variable effects on floral visitor behaviours and plant reproductive performance.