Reduced pollinator service and elevated pollen limitation at the geographic range limit of an annual plant

Mutualisms are well known to influence individual fitness and the population dynamics of partner species, but little is known about whether they influence species distributions and the location of geographic range limits. Here, we examine the contribution of plant-pollinator interactions to the geographic range limit of the California endemic plant Clarkia xantiana ssp. xantiana. We show that pollinator availability declined from the center to the margin of the geographic range consistently across four years of study. This decline in pollinator availability was caused to a greater extent by variation in the abundance of generalist rather than specialist bee pollinators. Climate data suggest that patterns of precipitation in the current and previous year drove variation in bee abundance because of its effects on cues for bee emergence in the current year and the abundance of floral resources in the previous year. Experimental floral manipulations showed that marginal populations had greater outcross pollen limitation of reproduction, in parallel with the decline in pollinator abundance. Although plants are self-compatible, we found no evidence that autonomous selfing contributes to reproduction, and thus no evidence that it alleviates outcross pollen limitation in marginal populations. Furthermore, we found no association between the distance to the range edge and selfing rate, as estimated from sequence and microsatellite variation, indicating that the mating system has not evolved in response to the pollination environment at the range periphery. Overall, our results suggest that dependence on pollinators for reproduction may be an important constraint limiting range expansion in this system.     

Consequences of vegetative herbivory for maintenance of intermediate outcrossing in an annual plant

Given the occurrence of mixed mating systems among plants, a general mechanism explaining the evolution and maintenance of this condition is needed. Although numerous theoretical models predict mixed mating to be evolutionarily stable, conditions favoring intermediate selfing are often stringent and have limited applicability. Here we investigated the role of vegetative herbivory, a ubiquitous biotic factor limiting plant reproduction, in the mating system expression of Impatiens capensis (Balsaminaceae), a species with an obligate mixed-mating system (individuals produce both selfing, cleistogamous, and facultatively outcrossing, chasmogamous flowers). Herbivory reduced proportional chasmogamous reproduction partially, but not entirely, through a reduction in plant size and the strength of this effect varied among replicates. Herbivory decreased geitonogamous selfing in chasmogamous flowers via several mechanisms including reduced chasmogamous flower display size and pollinator visitation rate and altered pollinator composition. Overall, herbivory caused a decrease in whole-plant outcrossing, indicating that the effects of herbivory on proportional chasmogamous reproduction, which favor selfing, outweigh the effects on chasmogamous outcrossing rate, which favor outcrossing. Not only do our findings unravel the mechanisms underlying herbivore-mediated changes in the mating system, but they also point to the role of natural enemies in contributing to the maintenance of a mixed mating system.     

Evaluating the Quality of Citizen‐Scientist Data on Pollinator Communities

Abstract: Concerns about pollinator declines have grown in recent years, yet the ability to detect changes in abundance, taxonomic richness, and composition of pollinator communities is hampered severely by the lack of data over space and time. Citizen scientists may be able to extend the spatial and temporal extent of pollinator monitoring programs. We developed a citizen‐science monitoring protocol in which we trained 13 citizen scientists to observe and classify floral visitors at the resolution of orders or super families (e.g., bee, wasp, fly) and at finer resolution within bees (superfamily Apoidea) only. We evaluated the protocol by comparing data collected simultaneously at 17 sites by citizen scientists (observational data set) and by professionals (specimen‐based data set). The sites differed with respect to the presence and age of hedgerows planted to improve habitat quality for pollinators. We found significant, positive correlations among the two data sets for higher level taxonomic composition, honey bee (Apis mellifera) abundance, non‐Apis bee abundance, bee richness, and bee community similarity. Results for both data sets also showed similar trends (or lack thereof) in these metrics among sites differing in the presence and age of hedgerows. Nevertheless, citizen scientists did not observe approximately half of the bee groups collected by professional scientists at the same sites. Thus, the utility of citizen‐science observational data may be restricted to detection of community‐level changes in abundance, richness, or similarity over space and time, and citizen‐science observations may not reliably reflect the abundance or frequency of occurrence of specific pollinator species or groups.     

Detecting Insect Pollinator Declines on Regional and Global Scales

Recently there has been considerable concern about declines in bee communities in agricultural and natural habitats. The value of pollination to agriculture, provided primarily by bees, is >$200 billion/year worldwide, and in natural ecosystems it is thought to be even greater. However, no monitoring program exists to accurately detect declines in abundance of insect pollinators; thus, it is difficult to quantify the status of bee communities or estimate the extent of declines. We used data from 11 multiyear studies of bee communities to devise a program to monitor pollinators at regional, national, or international scales. In these studies, 7 different methods for sampling bees were used and bees were sampled on 3 different continents. We estimated that a monitoring program with 200–250 sampling locations each sampled twice over 5 years would provide sufficient power to detect small (2–5%) annual declines in the number of species and in total abundance and would cost U.S.$2,000,000. To detect declines as small as 1% annually over the same period would require >300 sampling locations. Given the role of pollinators in food security and ecosystem function, we recommend establishment of integrated regional and international monitoring programs to detect changes in pollinator communities. Detección de Declinaciones de Insectos Polinizadores a Escalas Regional y Global     

Population differentiation in female sex pheromone and male preferences in a solitary bee

Population differentiation in female mating signals and associated male preferences can drive reproductive isolation among segregated populations. We tested this assumption by investigating intraspecific variation in female sex pheromone and associated male odour preferences among distant populations in the solitary bee Colletes cunicularius (L.) by using quantitative gas chromatography and by performing field bioassays with synthetic blends of key sex pheromone compounds. We found significant differences in sex pheromone blends among the bee populations, and the divergence in odour blends correlated positively with geographic distance, suggesting that genetic divergence among distant populations can affect sex pheromone chemistry. Our behavioural experiments, however, demonstrate that synthetic copies of allopatric female sex pheromones were cross-attractive to patrolling males from distant populations, making reproductive isolation by non-recognition of mating signals among populations unlikely. Our data also show that patrolling male bees from different populations preferred odour types from allopatric populations at the two sites of bioassays. These male preferences are not expected to select for changes in the female sex pheromone, but may influence the evolution of floral odour in sexually deceptive orchids of the genus Ophrys that are pollinated by C. cunicularius males.     

Bee community shifts with landscape context in a tropical countryside.

The ongoing scientific controversy over a putative "global pollination crisis" underscores the lack of understanding of the response of bees (the most important taxon of pollinators) to ongoing global land-use changes. We studied the effects of distance to forest, tree management, and floral resources on bee communities in pastures (the dominant land-use type) in southern Costa Rica. Over two years, we sampled bees and floral resources in 21 pastures at three distance classes from a large (approximately 230-ha) forest patch and of three common types: open pasture; pasture with remnant trees; and pasture with live fences. We found no consistent differences in bee diversity or abundance with respect to pasture management or floral resources. Bee community composition, however, was strikingly different at forest edges as compared to deforested countryside only a few hundred meters from forest. At forest edges, native social stingless bees (Apidae: Meliponini) comprised approximately 50% of the individuals sampled, while the alien honeybee Apis mellifera made up only approximately 5%. Away from forests, meliponines dropped to approximately 20% of sampled bees, whereas Apis increased to approximately 45%. Meliponine bees were also more speciose at forest edge sites than at a distance from forest, their abundance decreased with continuous distance to the nearest forest patch, and their species richness was correlated with the proportion of forest cover surrounding sample sites at scales from 200 to 1200 m. Meliponines and Apis together comprise the eusocial bee fauna of the study area and are unique in quickly recruiting foragers to high-quality resources. The diverse assemblage of native meliponine bees covers a wide range of body sizes and flower foraging behavior not found in Apis, and populations of many bee species (including Apis), are known to fluctuate considerably from year to year. Thus, the forest-related changes in eusocial bee communities we found may have important implications for: (1) sustaining a diverse bee fauna in tropical countryside; (2) ensuring the effective pollination of a diverse native plant community; and (3) the efficiency and stability of agricultural pollination, particularly for short-time-scale, mass-flowering crops such as coffee.     

Spatial mismatch between wild bee diversity hotspots and protected areas

Wild bees are critical for multiple ecosystem functions but are currently threatened. Understanding the determinants of the spatial distribution of wild bee diversity is a major research gap for their conservation. We modeled wild bee α and β taxonomic and functional diversity in Switzerland to uncover countrywide diversity patterns and determine the extent to which they provide complementary information, assess the importance of the different drivers structuring wild bee diversity, identify hotspots of wild bee diversity, and determine the overlap between diversity hotspots and the network of protected areas. We used site-level occurrence and trait data from 547 wild bee species across 3343 plots and calculated community attributes, including taxonomic diversity metrics, community mean trait values, and functional diversity metrics. We modeled their distribution with predictors describing gradients of climate, resource availability (vegetation), and anthropogenic influence (i.e., land-use types and beekeeping intensity). Wild bee diversity changed along gradients of climate and resource availability; high-elevation areas had lower functional and taxonomic α diversity, and xeric areas harbored more diverse bee communities. Functional and taxonomic β diversities diverged from this pattern, with high elevations hosting unique species and trait combinations. The proportion of diversity hotspots included in protected areas depended on the biodiversity facet, but most diversity hotspots occurred in unprotected land. Climate and resource availability gradients drove spatial patterns of wild bee diversity, resulting in lower overall diversity at higher elevations, but simultaneously greater taxonomic and functional uniqueness. This spatial mismatch among distinct biodiversity facets and the degree of overlap with protected areas is a challenge to wild bee conservation, especially in the face of global change, and calls for better integrating unprotected land. The application of spatial predictive models represents a valuable tool to aid the future development of protected areas and achieve wild bee conservation goals.     

Modeling scale up of anthropogenic impacts from individual pollinator behavior to pollination systems

Understanding how anthropogenic disturbances affect plant–pollinator systems has important implications for the conservation of biodiversity and ecosystem functioning. Previous laboratory studies show that pesticides and pathogens, which have been implicated in the rapid global decline of pollinators over recent years, can impair behavioral processes needed for pollinators to adaptively exploit floral resources and effectively transfer pollen among plants. However, the potential for these sublethal stressor effects on pollinator–plant interactions at the individual level to scale up into changes to the dynamics of wild plant and pollinator populations at the system level remains unclear. We developed an empirically parameterized agent-based model of a bumblebee pollination system called SimBee to test for effects of stressor-induced decreases in the memory capacity and information processing speed of individual foragers on bee abundance (scenario 1), plant diversity (scenario 2), and bee–plant system stability (scenario 3) over 20 virtual seasons. Modeling of a simple pollination network of a bumblebee and four co-flowering bee-pollinated plant species indicated that bee decline and plant species extinction events could occur when only 25% of the forager population showed cognitive impairment. Higher percentages of impairment caused 50% bee loss in just five virtual seasons and system-wide extinction events in less than 20 virtual seasons under some conditions. Plant species extinctions occurred regardless of bee population size, indicating that stressor-induced changes to pollinator behavior alone could drive species loss from plant communities. These findings indicate that sublethal stressor effects on pollinator behavioral mechanisms, although seemingly insignificant at the level of individuals, have the cumulative potential in principle to degrade plant–pollinator species interactions at the system level. Our work highlights the importance of an agent-based modeling approach for the identification and mitigation of anthropogenic impacts on plant–pollinator systems.     

A mark-recapture study of male<Emphasis Type="Italic"> Colletes cunicularius</Emphasis> bees: implications for pollination by sexual deception

An unusual pollination strategy is pollination by sexual deception in which orchids sexually attract male insects as pollinators. One gap in knowledge concerns the pattern and extent of pollinator movement among these sexually deceptive flowers and how this translates to pollen and gene flow. Our aim was to use mark and recapture techniques to investigate the behavior and movement of male Colletes cunicularius, an important bee pollinator of Ophrys. Our study site was located in northern Switzerland where a large population of the bees was nesting. Within two plots, (10×40 m), we marked bees with different colors and numbered tags. Seventeen percent of the 577 marked bees were recaptured over a period of 1 to a maximum of 11 days. However, the number of recaptures dropped dramatically after 3–5 days, suggesting an average lifetime of less than 10 days. Mark-recapture distances varied from 0 to 50 m, with a mean of 5 m. Our findings show that individual male bees patrol a specific and restricted region of the nesting area in search of mates. This mark-recapture study provides the first clues about the potential movement of pollen within populations of Ophrys orchids. We predict that orchid-pollen movements mediated by bees will be similar to the mark-recapture distances in this study. Parallel studies within orchid populations, including direct studies of pollen movement, are now required to better understand how pollinator mate-searching behavior translates to pollination success and pollen movement within sexually deceptive orchid populations.Communicated by R.F.A. Moritz     

Density-dependent effects of ants on selection for bumble bee pollination in Polemonium viscosum

Mutualisms are commonly exploited by cheater species that usurp rewards without providing reciprocal benefits. Yet most studies of selection between mutualist partners ignore interactions with third species and consequently overlook the impact of cheaters on evolution in the mutualism. Here, we explicitly investigate how the abundance of nectar-thieving ants (cheaters) influences selection in a pollination mutualism between bumble bees and the alpine skypilot, Polemonium viscosum. As suggested in past work with this species, bumble bees accounted for most of the seed production (78% +/- 6% [mean +/- SE]) in our high tundra study population and, in the absence of ants, exerted strong selection for large flowers. We tested for indirect effects of ant abundance on seed set through bumble bee pollination services (pollen delivery and pollen export) and a direct effect through flower damage. Ants reduced seed set per flower by 20% via flower damage. As ant density increased within experimental patches, the rate of flower damage rose, but pollen delivery and export did not vary significantly, showing that indirect effects of increased cheater abundance on pollinator service are negligible in this system. To address how ants affect selection for plant participation in the pollination mutualism we tested the impact of ant abundance on selection for bumble bee-mediated pollination. Results show that the impact of ants on fitness (seed set) accruing under bumble bee pollination is density dependent in P. viscosum. Selection for bumble bee pollination declined with increasing ant abundance in experimental patches, as predicted if cheaters constrain fitness returns of mutualist partner services. We also examined how ant abundance influences selection on flower size, a key component of plant investment in bumble bee pollination. We predicted that direct effects of ants would constrain bumble bee selection for large flowers. However, selection on flower size was significantly positive over a wide range of ant abundance (20-80% of plants visited by ants daily). Although high cheater abundance reduces the fitness returns of bumble bee pollination, it does not completely eliminate selection for bumble bee attraction in P. viscosum.     

A Hierarchical View of Genetic Structure in the Rare Annual Plant Clarkia springvillensis

Genetic structure at several spatial scales was examined in the rare California annual, Clarkia springvillensis . Using seven isozyme-encoding loci as genetic markers, we assessed the amount and distribution of genetic variation among three populations and eight subpopulations. Total genetic variation was lower than in species with similar life history traits but equivalent to that of other endemic plants. Spatial autocorrelation showed some evidence for very limited differentiation within subpopulations at a scale of 1–2 m. The subpopulations, separated by tens of meters, were found to be more differentiated from each other ( F _sp = 0.084) on average than were populations ( F,pt = 0.017). This local genetic differentiation was not correlated with physical distance between subpopulations. The low F_pt estimates suggest that substantial gene flow is occurring among populations. However, the lack of correlation between genetic and geographic distances and the significant differentiation of subpopulations suggest that genetic drift is occurring within populations. Therefore, we believe the apparent homogeneity of populations is due to each population's gene frequencies' being an average of several divergent subpopulations. If drift is causing differentiation within populations, it may eventually cause differentiation between populations. The importance of using a hierarchical approach to evaluating genetic structure is clear. Patterns occurring at one spatial scale may not be evident at others. One should not necessarily conclude that gene flow is substantial and that the risk of genetic erosion via drift is negligible just because differentiation between populations is small; the system may not be at equilibrium. This lesson is particularly important when recent changes in climate or land use are apparent.     

Estimates of outcrossing rates and of inbreeding depression in a population of the marine angiosperm Zostera marina

Despite the great diversity of pollination and fertilization mechanisms observed in marine plants, little is known about the causes or maintenance of this variation. In this study, I estimated outcrossing rates and levels of inbreeding depression in Zostera marina L. (eelgrass), providing the first empirical test of hypotheses about the evolution of breeding systems in plants with submerged flowers. This study also addressed temporal separation of female and male flowering (dichogamy) in eelgrass as a mechanism promoting an outcrossing mating system, and whether the mating system in eelgrass is related to the degree of dichogamy in the field. Outcrossing rates (0<t<1) estimated from two polymorphic allozyme loci indicate that the Z. marina population in intertidal and subtidal habitats in False Bay, Washington, USA, was highly outcrossing in both 1991 (intertidal t=0.905, subtidal t=1.0) and 1992 (intertidal t=0.775, subtidal t=0.611). The outcrossing rates were positively associated with the degree of dichogamy in 1992; intertidal plants exhibited a greater temporal separation of female and male flowering and a higher outcrossing rate than did subtidal plants. Inbreeding depression at seed set was estimated from hand pollinations (self- and outcross) on 20 reproductive individuals from the False Bay population. Averaged across all maternal parents, a greater proportion of outcrossed flowers set seed than selfed flowers; i.e., inbreeding depression was detected. Plants exhibited genetic variation for inbreeding depression, detected as a significant pollination treatment × maternal family interaction in a log-likelihood analysis. By the end of the seed-maturation period (7 mo after intial seed set) some families showed outbreeding depression, i.e., greater fitness in progeny derived from selfing than in progeny from outcrossing. The inbreeding depression in the False Bay population may be an important selective factor contributing to the maintenance of dichogamy and an outcrossing mating system, as proposed for aquatic plants.     

Tracing impacts of partner abundance in facultative pollination mutualisms: from individuals to populations

Partner abundance affects costs and benefits in obligate mutualisms, but its role in facultative partnerships is less clear. We address this gap in a pollination web consisting of two clovers (Trifolium) that differ in specialization on a bumble bee pollinator Bombus balteatus. We examine how pollination niche breadth affects plant responses to pollinator abundance, comparing early-flowering (specialized) and late-flowering (generalized) cohorts of T. parryi and early T. parryi to T. dasyphyllum, a pollination generalist. Co-pollinators disrupt the link between B. halteatus visitation and pollination rate for both clovers. Only for early-flowering T. parryi do visitation, pollination, and seed set increase with density of B. balteatus. Bumble bee density also alters timing of seed germination in T. parryi, with seeds from plants receiving augmented B. balteatus germinating sooner than seeds of open-pollinated counterparts. Benefits saturate at intermediate bumble bee densities. Despite strong effects of B. balteatus density on individual plant fitness components, population models suggest little impact of B. balteatus density on lamda in T. parryi or T. dasyphyllum. Findings show that functional redundancy in a pollinator guild mediates host-plant responses to partner density. Unexpected effects of pollinator density on life history schedule have implications for recruitment under pollinator decline.     

Selfing in <Emphasis Type="Italic">Parvulastra exigua</Emphasis>: an asterinid sea star with benthic development

Parvulastra exigua, a species that lacks a dispersive larva, is paradoxically one of the most widely distributed and abundant sea stars in Australia. The potential that self-fertilization and successful development of self-generated progeny might contribute to population maintenance in this species was investigated through induction of spawning in isolated individuals collected from five populations near Sydney, New South Wales (34°S, 151°E), during the spawning period (August–October 2010, 2011). Evidence of selfing differed significantly among the five populations but the level of selfing within individual egg masses did not. Overall, there was a mean of 8.9% fertilization in egg masses with embryos. Some clutches developed to the juvenile stage while others arrested development before hatching. The presence of miniscule testis tissue in gonads that appeared to be ovaries indicated that hermaphroditism may be more common in some populations than indicated by anatomy. Mixed mating (selfing + outcrossing), pseudocopulation during egg laying and developmental success of self-fertilized embryos may contribute to the ecological success of P. exigua. These might influence population genetic structure and would facilitate establishment of founder populations by a small number of migrants.     

Effects of Invasive Parasites on Bumble Bee Declines

Abstract: Bumble bees are a group of pollinators that are both ecologically and economically important and declining worldwide. Numerous mechanisms could be behind this decline, and the spread of parasites from commercial colonies into wild populations has been implicated recently in North America. Commercial breeding may lead to declines because commercial colonies may have high parasite loads, which can lead to colonization of native bumble bee populations; commercial rearing may allow higher parasite virulence to evolve; and global movement of commercial colonies may disrupt spatial patterns in local adaptation between hosts and parasites. We assessed parasite virulence, transmission mode, and infectivity. Microparasites and so‐called honey bee viruses may pose the greatest threat to native bumble bee populations because certain risk factors are present; for example, the probability of horizontal transmission of the trypanosome parasite Crithidia bombi is high. The microsporidian parasite Nosema bombi may play a role in declines of bumble bees in the United States. Preliminary indications that C. bombi and the neogregarine Apicystis bombi may not be native in parts of South America. We suggest that the development of molecular screening protocols, thorough sanitation efforts, and cooperation among nongovernmental organizations, governments, and commercial breeders might immediately mitigate these threats.     

Resource distributions among habitats determine solitary bee offspring production in a mosaic landscape.

Within mosaic landscapes, many organisms depend on attributes of the environment that operate over scales ranging from a single habitat patch to the entire landscape. One such attribute is resource distribution. Organisms' reliance on resources from within a local patch vs. those found among habitats throughout the landscape will depend on local habitat quality, patch quality, and landscape composition. The ability of individuals to move among complementary habitat types to obtain various resources may be a critical mechanism underlying the dynamics of animal populations and ultimately the level of biodiversity at different spatial scales. We examined the effects that local habitat type and landscape composition had on offspring production and survival of the solitary bee Osmia lignaria in an agri-natural landscape in California (U.S.A.). Female bees were placed on farms that did not use pesticides (organic farms), on farms that did use pesticides (conventional farms), or in seminatural riparian habitats. We identified pollens collected by bees nesting in different habitat types and matched these to pollens of flowering plants from throughout the landscape. These data enabled us to determine the importance of different plant species and habitat types in providing food for offspring, and how this importance changed with landscape and local nesting-site characteristics. We found that increasing isolation from natural habitat significantly decreased offspring production and survival for bees nesting at conventional farms, had weaker effects on bees in patches of seminatural habitat, and had little impact on those at organic farm sites. Pollen sampled from nests showed that females nesting in both farm and seminatural habitats relied on pollen from principally native plant species growing in seminatural habitat. Thus connectivity among habitats was critical for offspring production. Females nesting on organic farms were buffered to isolation effects by switching to floral resources growing at the farm site when seminatural areas were too distant. Overall local habitat conditions (farm management practices) can help bolster pollinators, but maintaining functional connectivity among habitats will likely be critical for persistence of pollinator populations as natural habitats are increasingly fragmented by human activities.     

The early bee catches the flower - circadian rhythmicity influences learning performance in honey bees, <Emphasis Type="Italic">Apis mellifera</Emphasis>

Circadian rhythmicity plays an important role for many aspects of honey bees’ lives. However, the question whether it also affects learning and memory remained unanswered. To address this question, we studied the effect of circadian timing on olfactory learning and memory in honey bees Apis mellifera using the olfactory conditioning of the proboscis extension reflex paradigm. Bees were differentially conditioned to odours and tested for their odour learning at four different “Zeitgeber” time points. We show that learning behaviour is influenced by circadian timing. Honey bees perform best in the morning compared to the other times of day. Additionally, we found influences of the light condition bees were trained at on the olfactory learning. This circadian-mediated learning is independent from feeding times bees were entrained to, indicating an inherited and not acquired mechanism. We hypothesise that a co-evolutionary mechanism between the honey bee as a pollinator and plants might be the driving force for the evolution of the time-dependent learning abilities of bees.     

Population Size and Reproduction in Phlox pilosa

Abstract: We analyzed the relationships between population size and reproductive characteristics in the perennial prairie forb Phlox pilosa , an obligate outcrossing butterfly-pollinated species. We examined 27 populations ranging in size from 9 to over 75,000 flowering ramets in two regions of the state of Iowa (eastcentral and northwest) in 1993 and 1994. We collected flowers from each population and scored them for pollen arrival to stigmas and number of pollen tubes. We collected fruiting ramets from each population at the end of the Phlox growing season and scored them for height, biomass, and reproductive variables, including the number of flowers initiated and opened and the number of capsules initiated and matured. In both years, population size was significantly correlated with the number of capsules matured per ramet. Differences between populations in capsule production were set primarily at the pollination stage. In 1993, pollen arrival to stigmas was significantly lower than in 1994 and was correlated with population size in eastcentral Iowa populations, indicating that lower reproduction in small populations that year was at least partially due to inadequate amounts of pollen being moved. In 1993, weather conditions likely depressed pollinator activity, but absolute capsule formation was high because of high flower production per ramet and high population densities. In 1994, when pollen arrival to stigmas was relatively high and unrelated to population size, outcross pollen movement was greater in larger populations. Increased efficacy of outcross pollen movement in 1994 may have resulted from lower flower production and less dense populations forcing greater pollinator movement between ramets or from variation between years in fine-scale spatial genetic substructuring of populations. Our results indicate that the viability of Phlox pilosa can be best ensured by protecting and creating populations of at least 1000–2000 flowering ramets.     

The evolution of polyandry by queens in social Hymenoptera: the significance of the timing of removal of diploid males

Summary Multiple mating by queens in social Hymenoptera with single locus sex determination may be an adaptation to reduce the effect of genetic load caused by the production of diploid males, if there is a concave relationship between queen fitness and the proportion of diploid male offspring in the colony. In this situation queens should be selected to reduce the variance in the production of diploid male offspring by multiple mating. It has been suggested that this concave relationship occurs in species such as the honey bee, Apis mellifera, in which reproduction occurs near the peak of colony population. This paper suggests that the timing of diploid male removal may influence mating frequency, with early removal of diploid males favoring multiple mating and late removal of diploid males favoring single mating. This idea is explored in two ways. A mathematical model shows that cell use in the brood area of species that rear young in cells will be more efficient with multiple mating. This would favor multiple mating in species, such as the honey bee, in which brood rearing is constrained by the usable area of the brood chamber. Secondly, comparison of polyandrous honey bees (early removal of diploid males as young larvae) with monandrous fire ants, Solenopsis invicta, and Melipona bees (non-removal of immature diploid males) suggests that in the species without diploid male removal, variance reduction may reduce queen fitness. Suggestions are made for testing this hypothesis.     

Landscape-scale resources promote colony growth but not reproductive performance of bumble bees

Variation in the availability of food resources over space and time is a likely driver of how landscape structure and composition affect animal populations. Few studies, however, have directly assessed the spatiotemporal variation in resource availability that arises from landscape pattern, or its effect on populations and population dynamic parameters. We tested the effect of floral resource availability at the landscape scale on the numbers of worker, male, and queen offspring produced by bumble bee, Bombus vosnesen?kii, colonies experimentally placed within complex agricultural-natural landscapes. We quantified flower densities in all land use types at different times of the season and then used these data to calculate spatially explicit estimates of floral resources surrounding each colony. Floral availability strongly correlated with landscape structure, and different regions of the landscape showed distinct seasonal patterns of floral availability. The floral resources available in the landscape surrounding a colony positively affected the number of workers and males it produced. Production was more sensitive to early- than to later-season resources. Floral resources did not significantly affect queen production despite a strong correlation between worker number and queen number among colonies. No landscape produced high floral resources during both the early and late season, and seasonal consistency is likely required for greater queen production. Floral resources are important determinants of colony growth and likely affect the pollination services provided by bumble bees at a landscape scale. Spatiotemporal variation in floral resources across the landscape precludes a simple relationship between resources and reproductive success as measured by queens, but nonetheless likely influences the total abundance of bumble bees in our study region.