Population dynamics of a stingless bee community in the seasonal dry lowlands of Costa Rica

This study examines the dynamics of a population of stingless bee colonies in the seasonal tropics of Guanacaste, Costa Rica. The community in a forest remnant was compared with that in surrounding deforested areas. During this 4-year study, a total of 192 wild stingless bee colonies were recorded, belonging to 14 species. Population dynamics were highly seasonal. Colony mortality peaked at the end of the wet season (October–November) while colony reproduction was most frequent during the dry season (December to April). Colony survival was not lower in founder colonies compared to established colonies. The most common species, T. angustula, had a much lower probability of annual survivorship in the forest (P = 0.74) than in deforested areas (P = 0.92). This results in an estimated colony life span for T. angustula of 3.8 years in the forest and 12.5 years in deforested areas. T. angustula should swarm once every two years to maintain its forest population, but only once every 12.5 years to maintain its population in the deforested areas. Survivorship of all other stingless bees was similar in the forest and deforested areas and did not significantly differ among the species. The average annual survivorship probability of these species was as high as 0.96, resulting in an estimated colony life span of 23.3 years. On average only one swarm per 20 years is needed to maintain their populations. Life history of the sympatric Africanised honey bee clearly differed from that of the stingless bees, with much lower annual survivorship probabilities for both founder (none survived) and established colonies (P = 0.33). These figures support the general idea that stingless bees invest more in colony survival rather than reproduction, but also show that life history is affected by both species and location. Received 27 October 2004; revised 8 March and 15 June 2005; accepted 5 July 2005.     

Anthropogenic hive movements are changing the genetic structure of a stingless bee (<Emphasis Type="Italic">Tetragonula carbonaria</Emphasis>) population along the east coast of Australia

Across the world, the keeping of stingless bees is increasingly popular, providing commercial pollination, high-value honey and a rewarding pass time. The popularity of stingless beekeeping has resulted in large-scale anthropogenic movements of nests, sometimes from outside their native range. Colony movement has the potential to impact local populations via transfer of parasites and pathogens and gene flow across unnaturally large geographic scales. Tetragonula carbonaria is the most widespread and commonly kept stingless bee species in Australia. Concerns have been raised that large-scale artificial propagation of T. carbonaria colonies by Sydney beekeepers, from a small number of colonies that originated in south-east Queensland, may have two consequences. First, the managed population may be becoming increasingly inbred. Second, the wild population may be experiencing significant introgression of south-east Queensland genotypes, potentially diluting local adaptations to the Sydney environment and resulting in the loss of local alleles. Here we show, based on microsatellite and mitochondrial markers, that both the managed and wild Sydney populations are significantly different from the south-east Queensland population. Nonetheless there is evidence that introgression of south-east Queensland alleles is impacting the genetic structure of both wild and managed Sydney populations. The two Sydney populations are indistinguishable, suggesting two-way gene flow in Sydney consistent with expectations of gene flow via male dispersal. All populations have low inbreeding coefficients, suggesting that they are genetically healthy.     

Beekeeping practices and geographic distance,not land use,drive gene flow across tropical bees

Across the globe, wild bees are threatened by ongoing natural habitat loss, risking the maintenance of plant biodiversity and agricultural production. Despite the ecological and economic importance of wild bees and the fact that several species are now managed for pollination services worldwide, little is known about how land use and beekeeping practices jointly influence gene flow. Using stingless bees as a model system, containing wild and managed species that are presumed to be particularly susceptible to habitat degradation, here we examine the main drivers of tropical bee gene flow. We employ a novel landscape genetic approach to analyse data from 135 populations of 17 stingless bee species distributed across diverse tropical biomes within the Americas. Our work has important methodological implications, as we illustrate how a maximum‐likelihood approach can be applied in a meta‐analysis framework to account for multiple factors, and weight estimates by sample size. In contrast to previously held beliefs, gene flow was not related to body size or deforestation, and isolation by geographic distance (IBD) was significantly affected by management, with managed species exhibiting a weaker IBD than wild ones. Our study thus reveals the critical importance of beekeeping practices in shaping the patterns of genetic differentiation across bee species. Additionally, our results show that many stingless bee species maintain high gene flow across heterogeneous landscapes. We suggest that future efforts to preserve wild tropical bees should focus on regulating beekeeping practices to maintain natural gene flow and enhancing pollinator‐friendly habitats, prioritizing species showing a limited dispersal ability.     

Workers' sons rescue genetic diversity at the sex locus in an invasive honey bee population

The hallmark of eusociality is the division of labour between reproductive (queen) and nonreproductive (worker) females. Yet in many eusocial insects, workers retain the ability to produce haploid male offspring from unfertilized eggs. The reproductive potential of workers has well‐documented consequences for the structure and function of insect colonies, but its implications at the population level are less often considered. We show that worker reproduction in honey bees can have an important role in maintaining genetic diversity at the sex locus in invasive populations. The honey bee sex locus is homozygous‐lethal, and, all else being equal, a higher allele number in the population lead to higher mean brood survival. In an invasive population of the honey bee Apis cerana in Australia, workers contribute significantly to male production: 38% of male‐producing colonies are queenless, and these contribute one‐third of all males at mating congregations. Using a model, we show that such male production by queenless workers will increase the number of sex alleles retained in nascent invasive populations following founder events, relative to a scenario in which only queens reproduce. We conclude that by rescuing sex locus diversity that would otherwise be lost, workers' sons help honey bee populations to minimize the negative effects of inbreeding after founder events and so contribute to their success as invaders.     

Outbreeding and lack of temporal genetic structure in a drone congregation of the neotropical stingless bee Scaptotrigona mexicana

Drone aggregations are a widespread phenomenon in many stingless bee species (Meliponini), but the ultimate and proximate causes for their formation are still not well understood. One adaptive explanation for this phenomenon is the avoidance of inbreeding, which is especially detrimental for stingless bees due to the combined effects of the complementary sex-determining system and the small effective population size caused by eusociality and monandry. We analyzed the temporal genetic dynamics of a drone aggregation of the stingless bee Scaptotrigona mexicana with microsatellite markers over a time window of four weeks. We estimated the drones of the aggregation to originate from a total of 55 colonies using sibship re-construction. There was no detectable temporal genetic differentiation or sub-structuring in the aggregation. Most important, we could exclude all colonies in close proximity of the aggregation as origin of the drones in the aggregation, implicating that they originate from more distant colonies. We conclude that the diverse genetic composition and the distant origin of the drones of the S. mexicana drone congregation provides an effective mechanism to avoid mating among close relatives.     

Isolation and characterization of ten microsatellite loci in stingless bee Trigona spinipes (Apidae: Meliponini)

Stingless bees are the most abundant pollinators of Brazilian tropical flora. Trigona spinipes has some of the largest colonies of any stingless bee species found in several types of environment. This work describes the isolation and characterization of microsatellite loci for this species. A microsatellite‐enriched genomic library was constructed and ten primer pairs were designed for T. spinipes. The primers were tested in 20 unrelated individuals. The mean number of alleles was 8.10 and mean observed and expected heterozygosity were 0.655 and 0.680, respectively. Primers were also tested in cross‐species amplification and five loci were successfully amplified in Trigona chanchamayoensis, Trigona hyalinata, Tetragonisca angustula, Partamona mulata and Frieseomelitta varia. The microsatellite primers described herein will be useful for evaluating genetic variability and gaining a better understanding of the population structure of T. spinipes as well as other species of stingless bees.     

Partitioning of pollen resources by two stingless bee species in the north Bahia,Brazil

Analysing the pollen stored by stingless bees allows identification of the diversity of flowering plants visited by these bees while collecting resources during a given period. However, few studies have focused on investigating the sharing of resources between species of native bees that coexist in transitional plant formations, which support conservation efforts and increase colony productivity. This study aims to describe the partitioning of pollen resources among colonies of two species of stingless bees in the state of the Bahia (Brazil) through the analysis of stored pollen. The study was conducted in neighbouring vegetation zones of the relevant species Melipona quadrifasciata anthidioides and Melipona scutellaris by collecting monthly pollen samples stored by three colonies of each species over one year. The collected samples were treated using the acetolysis technique, and the partitioning of pollen resources between the two species was investigated through quali-quantitative analysis of the treated pollen. The results show that among the 16 pollen types belonging to a particular species or group identified in the Melipona quadrifasciata anthidioides samples and the 18 identified in the Melipona scutellaris samples, 13 were shared by both species. The two most common pollen types stored by both species were from Mimosa arenosa (Fabaceae, Mimosoidae) and Eucalyptus sp. (Myrtaceae). These results demonstrate the niche similarity of pollen pasture between these two species and suggest that conservation efforts for these bee species should focus on the plant families most visited by bees.     

Imidacloprid-induced impairment of mushroom bodies and behavior of the native stingless bee Melipona quadrifasciata anthidioides

Declines in pollinator colonies represent a worldwide concern. The widespread use of agricultural pesticides is recognized as a potential cause of these declines. Previous studies have examined the effects of neonicotinoid insecticides such as imidacloprid on pollinator colonies, but these investigations have mainly focused on adult honey bees. Native stingless bees (Hymenoptera: Apidae: Meliponinae) are key pollinators in neotropical areas and are threatened with extinction due to deforestation and pesticide use. Few studies have directly investigated the effects of pesticides on these pollinators. Furthermore, the existing impact studies did not address the issue of larval ingestion of contaminated pollen and nectar, which could potentially have dire consequences for the colony. Here, we assessed the effects of imidacloprid ingestion by stingless bee larvae on their survival, development, neuromorphology and adult walking behavior. Increasing doses of imidacloprid were added to the diet provided to individual worker larvae of the stingless bee Melipona quadrifasciata anthidioides throughout their development. Survival rates above 50% were only observed at insecticide doses lower than 0.0056 μg active ingredient (a.i.)/bee. No sublethal effect on body mass or developmental time was observed in the surviving insects, but the pesticide treatment negatively affected the development of mushroom bodies in the brain and impaired the walking behavior of newly emerged adult workers. Therefore, stingless bee larvae are particularly susceptible to imidacloprid, as it caused both high mortality and sublethal effects that impaired brain development and compromised mobility at the young adult stage. These findings demonstrate the lethal effects of imidacloprid on native stingless bees and provide evidence of novel serious sublethal effects that may compromise colony survival. The ecological and economic importance of neotropical stingless bees as pollinators, their susceptibility to insecticides and the vulnerability of their larvae to insecticide exposure emphasize the importance of studying these species.     

Key factors influencing forager distribution across macadamia orchards differ among species of managed bees

To achieve maximised and sustainable crop productivity, it is critical that we develop crop-specific strategies for managing pollination. Honey bees (Apis mellifera) and stingless bees (Tetragonula carbonaria) are considered effective pollinators of macadamia (Macadamia integrifolia). The introduction of managed honey bee or stingless bee hives into orchards is likely to boost the numbers of these insects visiting flowers; however, there is a lack of published information and consensus regarding their management for pollination. Here, we identify factors that affect the distribution of both honey bees and stingless bees across cultivated macadamia, and establish whether increased flower visitation leads to higher nut set. A gradient of bee visitation rates was created by placing colonies on the ends of a four-hectare block, and mixed-effect models were applied to assess forager abundance and nut set with respect to distance from hive, time of day, cultivar, and floral display size. Distance from colony had a strong effect on stingless bee numbers, with >96% of individuals recorded within 100 metres of colonies, whereas the distribution of honey bees was more closely related to daily floral display: trees with greater numbers of flowers attracted more honey bees. Simplified surveys conducted in a further 17 macadamia blocks confirm that these are broadly occurring distribution patterns. Bee abundance alone did not significantly predict nut production; however, an indirect effect of bee visits to flowers is inferred, as nut production increased with size of floral display. To encourage a more even distribution of bees and uniform pollination, we recommend placement of stingless bee hives to maximise their distribution through a block (e.g. at 100-m intervals) and management practices that promote even distributions of flowers across trees.     

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.     

Bees for Development: Brazilian Survey Reveals How to Optimize Stingless Beekeeping

Stingless bees are an important asset to assure plant biodiversity in many natural ecosystems, and fulfill the growing agricultural demand for pollination. However, across developing countries stingless beekeeping remains an essentially informal activity, technical knowledge is scarce, and management practices lack standardization. Here we profited from the large diversity of stingless beekeepers found in Brazil to assess the impact of particular management practices on productivity and economic revenues from the commercialization of stingless bee products. Our study represents the first large-scale effort aiming at optimizing stingless beekeeping for honey/colony production based on quantitative data. Survey data from 251 beekeepers scattered across 20 Brazilian States revealed the influence of specific management practices and other confounding factors over productivity and income indicators. Specifically, our results highlight the importance of teaching beekeepers how to inspect and feed their colonies, how to multiply them and keep track of genetic lineages, how to harvest and preserve the honey, how to use vinegar traps to control infestation by parasitic flies, and how to add value by labeling honey containers. Furthermore, beekeeping experience and the network of known beekeepers were found to be key factors influencing productivity and income. Our work provides clear guidelines to optimize stingless beekeeping and help transform the activity into a powerful tool for sustainable development.     

Integrative population genetics and metagenomics reveals urbanization increases pathogen loads and decreases connectivity in a wild bee

As urbanization continues to increase, it is expected that two-thirds of the human population will reside in cities by 2050. Urbanization fragments and degrades natural landscapes, threatening wildlife including economically important species such as bees. In this study, we employ whole genome sequencing to characterize the population genetics, metagenome and microbiome, and environmental stressors of a common wild bee, Ceratina calcarata. Population genomic analyses revealed the presence of low genetic diversity and elevated levels of inbreeding. Through analyses of isolation by distance, resistance, and environment across urban landscapes, we found that green spaces including shrubs and scrub were the most optimal pathways for bee dispersal, and conservation efforts should focus on preserving these land traits to maintain high connectivity across sites for wild bees. Metagenomic analyses revealed landscape sites exhibiting urban heat island effects, such as high temperatures and development but low precipitation and green space, had the highest taxa alpha diversity across all domains even when isolating for potential pathogens. Notably, the integration of population and metagenomic data showed that reduced connectivity in urban areas is not only correlated with lower relatedness among individuals but is also associated with increased pathogen diversity, exposing vulnerable urban bees to more pathogens. Overall, our combined population and metagenomic approach found significant environmental variation in bee microbiomes and nutritional resources even in the absence of genetic differentiation, as well as enabled the potential early detection of stressors to bee health.     

Socialized medicine: Individual and communal disease barriers in honey bees

Honey bees are attacked by numerous parasites and pathogens toward which they present a variety of individual and group-level defenses. In this review, we briefly introduce the many pathogens and parasites afflicting honey bees, highlighting the biology of specific taxonomic groups mainly as they relate to virulence and possible defenses. Second, we describe physiological, immunological, and behavioral responses of individual bees toward pathogens and parasites. Third, bees also show behavioral mechanisms for reducing the disease risk of their nestmates. Accordingly, we discuss the dynamics of hygienic behavior and other group-level behaviors that can limit disease. Finally, we conclude with several avenues of research that seem especially promising for understanding host–parasite relationships in bees and for developing breeding or management strategies for enhancing honey bee health. We discuss how human efforts to maintain healthy colonies intersect with similar efforts by the bees, and how bee management and breeding protocols can affect disease traits in the short and long term.     

Intraspecific queen parasitism in a highly eusocial bee

Insect societies are well-known for their advanced cooperation, but their colonies are also vulnerable to reproductive parasitism. Here, we present a novel example of an intraspecific social parasitism in a highly eusocial bee, the stingless bee Melipona scutellaris. In particular, we provide genetic evidence which shows that, upon loss of the mother queen, many colonies are invaded by unrelated queens that fly in from unrelated hives nearby. The reasons for the occurrence of this surprising form of social parasitism may be linked to the fact that unlike honeybees, Melipona bees produce new queens in great excess of colony needs, and that this exerts much greater selection on queens to seek alternative reproductive options, such as by taking over other nests. Overall, our results are the first to demonstrate that queens in highly eusocial bees can found colonies not only via supersedure or swarming, but also by infiltrating and taking over other unrelated nests.     

Conservation study of an endangered stingless bee (Melipona capixaba—Hymenoptera: Apidae) with restricted distribution in Brazil

Melipona capixaba, popularly known as “uruçu preta”, is a stingless bee restricted to the mountainous Atlantic Rainforest areas of Espírito Santo State, Brazil. Due to the endemism and small population size, this species discovered in 1994 is now considered “vulnerable to extinction”. Using ISSR, PCR–RFLP and microsatellites markers, we studied the genetic variability and structure of M. capixaba from 88 colonies collected throughout the distribution area of the species within Espírito Santo State. The microsatellite, ISSR and mitochondrial haplotype analyses showed that M. capixaba has low genetic variability compared to other insect species. The molecular analyses also indicated a high genetic similarity among the M. capixaba samples, with no clear pattern of structuring. The analyses of molecular variance results indicated that most of the total genetic variation in M. capixaba was explained by the genetic diversity within local populations. Results suggest that the analyzed samples could be treated as a single population for preservation purposes. Thus, given its endemism, local adaptation and low number of natural colonies, efforts for the conservation of M. capixaba should focus on preservation and increasing the number of colonies in the wild, so that M. capixaba can support constant captures and the effects of habitat deforestation in Espírito Santo State.     

Genetic Differentiation of the Stingless Bee Tetragonula pagdeni in Thailand Using SSCP Analysis of a Large Subunit of Mitochondrial Ribosomal DNA

Genetic diversity and population differentiation of the stingless bee Tetragonula pagdeni (Schwarz) was assessed using single-strand conformational polymorphism (SSCP) analysis of a large subunit of the ribosomal RNA gene (16S rRNA). High levels of genetic variation among individuals within each population (North, Northeast, Central, Prachuap Khiri Khan, Chumphon, and Peninsular Thailand) of T. pagdeni were observed. Analysis of molecular variance indicated significant genetic differentiation among the six geographic populations (Φ (PT)?=?0.28, P?相似文献   

栖息条件对中华蜜蜂遗传多样性的影响

为了评估不同栖息条件对中华蜜蜂(简称中蜂)种群遗传多样性的影响,本研究运用微卫星标记的方法,对皖南山区4个不同采样点120个中华蜜蜂群体的2400只工蜂样本进行了分析.结果表明:与天然林野生中蜂群相比,单一茶树种植结构下,皖南中蜂群的等位基因数、平均期望杂合度和多态信息含量的差异均不显著;人工驯养中蜂群的等位基因数、多态信息含量差异显著,平均期望杂合度差异不显著;2006年采集的野生中蜂群与天然林野生中蜂群的等位基因数、平均期望杂合度和多态信息含量均差异显著.各种群间存在极显著的遗传分化,平均分化系数为0.32.说明皖南地区茶树单一化种植不会影响中蜂的遗传结构与资源保护.     

Fragmentation in the clouds? The population genetics of the native bee <Emphasis Type="Italic">Partamona bilineata</Emphasis> (Hymenoptera: Apidae: Meliponini) in the cloud forests of Guatemala

Habitat fragmentation and loss are important drivers of genetic differentiation, often leading to a decrease in genetic diversity. Yet, natural populations of tropical bees often show a lack of differentiation, even in fragmented landscapes, suggesting resilience to deal with unfavourable land use. It is not clear what leads to this lack of differentiation, but large population sizes, high rates of dispersal and stable demography likely play important roles. Here, we investigate the population genetic structure and infer the present and historic demography of the eusocial stingless bee Partamona bilineata from tropical montane cloud forests in Guatemala. We used microsatellites and mitochondrial DNA to test for genetic differentiation, to infer migration rates, and to evaluate the effects of landscape. We also used demographic modelling to trace population sizes over time. We found that six populations of P. bilineata exhibited only subtle differentiation, with the exception of one site at the edge of the cloud forest, which was clearly distinct from all others. Effective population sizes (number of colonies) appeared to be rather small (18?±?6 colonies) compared to the original sample size (N?=?51?±?9), but stable over time, and inferred rates of gene flow were low; yet, no genetic bottleneck was detected. A statistical model including elevation was the best in explaining the observed pattern of differentiation. We find that P. bilineata does not exhibit strong genetic structure, making it a resilient species for provision of pollination services. But, at the same time, our data point to the potential vulnerability of this and similar species, as effective population sizes appear to be low and hence populations may be easily affected by future environmental change. As such, P. bilineata may be representative of many other tropical stingless bees, for which lack of differentiation has been invoked.     

Genetic structure of nest aggregations and drone congregations of the southeast Asian stingless bee Trigona collina

In stingless bees, sex is determined by a single complementary sex-determining locus. This method of sex determination imposes a severe cost of inbreeding because an egg fertilized by sperm carrying the same sex allele as the egg results in a sterile diploid male. To explore how reproductive strategies may be used to avoid inbreeding in stingless bees, we studied the genetic structure of a population of 27 colonies and three drone congregations of Trigona collina in Chanthaburi, Thailand. The colonies were distributed across six nest aggregations, each aggregation located in the base of a different fig tree. Genetic analysis at eight microsatellite loci showed that colonies within aggregations were not related. Samples taken from three drone congregations showed that the males were drawn from a large number of colonies (estimated to be 132 different colonies in our largest swarm). No drone had a genotype indicating that it could have originated from the colony that it was directly outside. Combined, these results suggest that movements of drones and possibly movements of reproductive swarms among colony aggregations provide two mechanisms of inbreeding avoidance.     

Morphological and molecular characterization of the Landes honey bee (<Emphasis Type="Italic">Apis mellifera</Emphasis> L.) ecotype for genetic conservation

A population of honey bees (Apis mellifera mellifera L.) with an annual colony brood cycle adapted to a locally abundant floral source in the Landes region of Southwest France is the subject of genetic conservation efforts. This population is maintained by local beekeepers in an area that experiences both an annual seasonal influx of non-local colonies and the permanent culture of imported stock. However, some colonies native to the Landes do not have the adapted brood cycle and their status as ecotypic are in question. The present study used morphology, mitochondrial DNA and microsatellites to characterize the endemic population and suggests further genetic conservation strategies. These methods yielded different degrees of discrimination of native and imported colonies and provided a powerful suite of tools for local resource managers. Colonies from the Landes could be differentiated from non-local French A. m. mellifera populations using morphometric analysis, and from non-native and reference populations using mtDNA and microsatellites. Seven morphological characters were identified by discriminant analysis as informative for delineating the Landes ecotype from other A. m. mellifera populations. Mitochondrial haplotypes for the population were characterized and five microsatellite loci were found to be informative in characterizing the Landes population. Asymmetric gene flow detected with microsatellite alleles was observed to be 5.5–5.9% from imported to native stocks of honey bees while introgression of native microsatellite alleles into imported colonies was 21.6%.