Faster-growing parasites threaten host populations via patch-level population dynamics and higher virulence; a case study in Varroa mites (Mesostigmata: Varroidae) and honey bees (Hymenoptera: Apidae).

Honey bee parasites remain a critical challenge to management and conservation. Because managed honey bees are maintained in colonies kept in apiaries across landscapes, the study of honey bee parasites allows the investigation of spatial principles in parasite ecology and evolution. We used a controlled field experiment to study the relationship between population growth rate and virulence (colony survival) of the parasite Varroa destructor (Anderson and Trueman). We used a nested design of 10 patches (apiaries) of 14 colonies to examine the spatial scale at which Varroa population growth matters for colony survival. We tracked Varroa population size and colony survival across a full year and found that Varroa populations that grow faster in their host colonies during the spring and summer led to larger Varroa populations across the whole apiary (patch) and higher rates of neighboring colony loss. Crucially, this increased colony loss risk manifested at the patch scale, with mortality risk being related to spatial adjacency to colonies with fast-growing Varroa strains rather than with Varroa growth rate in the colony itself. Thus, within-colony population growth predicts whole-apiary virulence, demonstrating the need to consider multiple scales when investigating parasite growth-virulence relationships.

Plants, pollinators and their interactions under global ecological change: The role of pollen DNA metabarcoding.

Anthropogenic activities are triggering global changes in the environment, causing entire communities of plants, pollinators and their interactions to restructure, and ultimately leading to species declines. To understand the mechanisms behind community shifts and declines, as well as monitoring and managing impacts, a global effort must be made to characterize plant-pollinator communities in detail, across different habitat types, latitudes, elevations, and levels and types of disturbances. Generating data of this scale will only be feasible with rapid, high-throughput methods. Pollen DNA metabarcoding provides advantages in throughput, efficiency and taxonomic resolution over traditional methods, such as microscopic pollen identification and visual observation of plant-pollinator interactions. This makes it ideal for understanding complex ecological networks and their responses to change. Pollen DNA metabarcoding is currently being applied to assess plant-pollinator interactions, survey ecosystem change and model the spatiotemporal distribution of allergenic pollen. Where samples are available from past collections, pollen DNA metabarcoding has been used to compare contemporary and past ecosystems. New avenues of research are possible with the expansion of pollen DNA metabarcoding to intraspecific identification, analysis of DNA in ancient pollen samples, and increased use of museum and herbarium specimens. Ongoing developments in sequencing technologies can accelerate progress towards these goals. Global ecological change is happening rapidly, and we anticipate that high-throughput methods such as pollen DNA metabarcoding are critical for understanding the evolutionary and ecological processes that support biodiversity, and predicting and responding to the impacts of change.

Differences in individual flowering time change pollen limitation and seed set in three montane wildflowers.

PREMISE: Changes to flowering time caused by climate change could affects plant fecundity, but studies that compare the individual-level responses of phenologically distinct, co-occurring species are lacking. We assessed how variation in floral phenology affects the fecundity of individuals from three montane species with different seasonal flowering times, including in snowmelt acceleration treatments to increase variability in phenology. METHODS: We collected floral phenology and seed set data for individuals of three montane plant species (Mertensia fusiformis, Delphinium nuttallianum, Potentilla pulcherrima). To examine the drivers of seed set, we measured conspecific floral density and conducted pollen limitation experiments to isolate pollination function. We advanced the phenology of plant communities in a controlled large-scale snowmelt acceleration experiment. RESULTS: Differences in individual phenology relative to the rest of the population affected fecundity in our focal species, but effects were species-specific. For our early-season species, individuals that bloomed later than the population peak bloom had increased fecundity, while for our midseason species, simply blooming before or after the population peak increased individual fecundity. For our late-season species, blooming earlier than the population peak increased fecundity. The early and midseason species were pollen-limited, and conspecific density affected seed set only for our early-season species. CONCLUSIONS: Our study shows that variation in individual phenology affects fecundity in three phenologically distinct montane species, and that pollen limitation may be more influential than conspecific density. Our results suggest that individual-level changes in phenology are important to consider for understanding plant reproductive success.

Loss of endangered frugivores from seed dispersal networks generates severe mutualism disruption.

Many tropical seed-dispersing frugivores are facing extinction, but the consequences of the loss of endangered frugivores for seed dispersal is not well understood. We investigated the role of frugivore endangerment status via robustness-to-coextinction simulations (in this context, more accurately described as robustness-to-partner-loss simulations) using data from the Brazilian Atlantic Forest biodiversity hotspot. By simulating the extinction of endangered frugivores, we found a rapid and disproportionate loss of tree species with dispersal partners in the network, and this surprisingly surpassed any other frugivore extinction scenario, including the loss of the most generalist frugivores first. A key driver of this pattern is that many specialist plants rely on at-risk frugivores as seed-dispersal partners. Moreover, interaction compensation in the absence of endangered frugivores may be unlikely because frugivores with growing populations forage on fewer plant species than frugivores with declining populations. Therefore, protecting endangered frugivores could be critical for maintaining tropical forest seed dispersal, and their loss may have higher-than-expected functional consequences for tropical forests, their regeneration processes, and the maintenance of tropical plant diversity.

Upper-limit agricultural dietary exposure to streptomycin in the laboratory reduces learning and foraging in bumblebees.

In the past decade, the broadcast-spray application of antibiotics in US crops has increased exponentially in response to bacterial crop pathogens, but little is known about the sublethal impacts on beneficial organisms in agroecosystems. This is concerning given the key roles that microbes play in modulating insect fitness. A growing body of evidence suggests that insect gut microbiomes may play a role in learning and behaviour, which are key for the survival of pollinators and for their pollination efficacy, and which in turn could be disrupted by dietary antibiotic exposure. In the laboratory, we tested the effects of an upper-limit dietary exposure to streptomycin (200 ppm)-an antibiotic widely used to treat bacterial pathogens in crops-on bumblebee (Bombus impatiens) associative learning, foraging and stimulus avoidance behaviour. We used two operant conditioning assays: a free movement proboscis extension reflex protocol focused on short-term memory formation, and an automated radio-frequency identification tracking system focused on foraging. We show that upper-limit dietary streptomycin exposure slowed training, decreased foraging choice accuracy, increased avoidance behaviour and was associated with reduced foraging on sucrose-rewarding artificial flowers flowers. This work underscores the need to further study the impacts of antibiotic use on beneficial insects in agricultural systems.

Conserving alpha and beta diversity in wood-production landscapes.

International demand for wood and other forest products continues to grow rapidly, and uncertainties remain about how animal communities will respond to intensifying resource extraction associated with woody bioenergy production. We examined changes in alpha and beta diversity of bats, bees, birds, and reptiles across wood production landscapes in the southeastern United States, a biodiversity hotspot that is one of the principal sources of woody biomass globally. We sampled across a spatial gradient of paired forest land-uses (representing pre and postharvest) that allowed us to evaluate biological community changes resulting from several types of biomass harvest. Short-rotation practices and residue removal following clearcuts were associated with reduced alpha diversity (-14.1 and -13.9 species, respectively) and lower beta diversity (i.e., Jaccard dissimilarity) between land-use pairs (0.46 and 0.50, respectively), whereas midrotation thinning increased alpha (+3.5 species) and beta diversity (0.59). Over the course of a stand rotation in a single location, biomass harvesting generally led to less biodiversity. Cross-taxa responses to resource extraction were poorly predicted by alpha diversity: correlations in responses between taxonomic groups were highly variable (-0.2 to 0.4) with large uncertainties. In contrast, beta diversity patterns were highly consistent and predictable across taxa, where correlations in responses between taxonomic groups were all positive (0.05-0.4) with more narrow uncertainties. Beta diversity may, therefore, be a more reliable and information-rich indicator than alpha diversity in understanding animal community response to landscape change. Patterns in beta diversity were primarily driven by turnover instead of species loss or gain, indicating that wood extraction generates habitats that support different biological communities.

Conservación de la Diversidad Alfa y Beta en Paisajes de Producción Maderera Resumen La demanda internacional de madera y otros productos forestales sigue creciendo rápidamente mientras permanecen las incertidumbres sobre cómo responderán las comunidades animales a la intensificación de la extracción de recursos asociada con la producción de bioenergía leñosa. Examinamos los cambios en la diversidad alfa y beta de murciélagos, abejas, aves y reptiles en los paisajes de producción maderera en el sureste de los Estados Unidos, un punto caliente de biodiversidad y una de las fuentes principales de biomasa leñosa a nivel mundial. Muestreamos a lo largo de un gradiente espacial de usos de suelo forestales emparejados (representando la pre- y postcosecha) que nos permitió evaluar los cambios en las comunidades biológicas resultantes de varios tipos de recolección de biomasa. Las prácticas de corta rotación y de eliminación de residuos después de la tala estuvieron asociadas con la reducción de la diversidad alfa (−14.1 y −13.9 especies, respectivamente) y una diversidad beta más baja (es decir, diferencia de Jaccard) entre los pares de uso de suelo (0.46 y 0.50, respectivamente), mientras que el raleo de rotación media incrementó la diversidad alfa (+3.5 especies) y beta (0.59). Durante la duración de una rotación permanente en una sola ubicación, la cosecha de biomasa generalmente derivó en menos biodiversidad. La respuesta de los taxones a la extracción de recursos estuvo muy mal pronosticada por la diversidad alfa: la correlación de las respuestas entre los grupos taxonómicos fue altamente variable (−0.2 a 0.4) con muchas incertidumbres. Como contraste, los patrones de diversidad beta fueron fuertemente coherentes y predecibles en todos los taxones, mientras que la correlación de las respuestas entre los grupos taxonómicos siempre fue positiva (0.05 a 0.4) con incertidumbres más limitadas. Por lo tanto, la diversidad beta puede ser un indicador más confiable y rico en información que la diversidad alfa para entender las respuestas de la comunidad animal a los cambios en el paisaje. Los patrones de la diversidad beta estuvieron impulsados principalmente por la rotación en lugar de la pérdida o ganancia de especies, lo que indica que la extracción de madera genera hábitats que mantienen a diferentes comunidades biológicas.

The context dependency of pollinator interference: How environmental conditions and co-foraging species impact floral visitation.

Animals often change their behaviour in the presence of other species and the environmental context they experience, and these changes can substantially modify the course their populations follow. In the case of animals involved in mutualistic interactions, it is still unclear how to incorporate the effects of these behavioural changes into population dynamics. We propose a framework for using pollinator functional responses to examine the roles of pollinator-pollinator interactions and abiotic conditions in altering the times between floral visits of a focal pollinator. We then apply this framework to a unique foraging experiment with different models that allow resource availability and sublethal exposure to a neonicotinoid pesticide to modify how pollinators forage alone and with co-foragers. We found that all co-foragers interfere with the focal pollinator under at least one set of abiotic conditions; for most species, interference was strongest at higher levels of resource availability and with pesticide exposure. Overall our results highlight that density-dependent responses are often context-dependent themselves.

Plant-pollinator interaction niche broadens in response to severe drought perturbations.

The composition of plant-pollinator interactions-i.e., who interacts with whom in diverse communities-is highly dynamic, and we have a very limited understanding of how interaction identities change in response to perturbations in nature. One prediction from niche and diet theory is that resource niches will broaden to compensate for resource reductions driven by perturbations, yet this has not been empirically tested in plant-pollinator systems in response to real-world perturbations in the field. Here, we use a long-term dataset of floral visitation to Ipomopsis aggregata, a montane perennial herb, to test whether the breadth of its floral visitation niche (i.e., flower visitor richness) changed in response to naturally occurring drought perturbations. Fewer floral resources are available in drought years, which could drive pollinators to expand their foraging niches, thereby expanding plants' floral visitation niches. We compared two drought years to three non-drought years to analyze changes in niche breadth and community composition of floral visitors to I. aggregata, predicting broadened niche breadth and distinct visitor community composition in drought years compared to non-drought years. We found statistically significant increases in niche breadth in drought years as compared to non-drought conditions, but no statistically distinguishable changes in community composition of flower visitors. Our findings suggest that plants' floral visitation niches may exhibit considerable plasticity in response to disturbance. This may have widespread consequences for community-level stability as well as functional consequences if increased niche overlap affects pollination services.

Persistent effects of management history on honeybee colony virus abundances.

Infectious diseases are a major threat to both managed and wild pollinators. One key question is how the movement or transplantation of honeybee colonies under different management regimes affects honeybee disease epidemiology. We opportunistically examined any persistent effect of colony management history following relocation by characterising the virus abundances of honeybee colonies from three management histories, representing different management histories: feral, low-intensity management, and high-intensity "industrial" management. The colonies had been maintained for one year under the same approximate 'common garden' condition. Colonies in this observational study differed in their virus abundances according to management history, with the feral population history showing qualitatively different viral abundance patterns compared to colonies from the two managed population management histories; for example, higher abundance of sacbrood virus but lower abundances of various paralysis viruses. Colonies from the high-intensity management history exhibited higher viral abundances for all viruses than colonies from the low-intensity management history. Our results provide evidence that management history has persistent impacts on honeybee disease epidemiology, suggesting that apicultural intensification could be majorly impacting on pollinator health, justifying much more substantial investigation.

Agricultural intensification drives changes in hybrid network robustness by modifying network structure.

Within ecological communities, species engage in myriad interaction types, yet empirical examples of hybrid species interaction networks composed of multiple types of interactions are still scarce. A key knowledge gap is understanding how the structure and stability of such hybrid networks are affected by anthropogenic disturbance. Using 15,169 interaction observations, we constructed 16 hybrid herbivore-plant-pollinator networks along an agricultural intensification gradient to explore changes in network structure and robustness to local extinctions. We found that agricultural intensification led to declines in modularity but increases in nestedness and connectance. Notably, network connectance, a structural feature typically thought to increase robustness, caused declines in hybrid network robustness, but the directionality of changes in robustness along the gradient depended on the order of local species extinctions. Our results not only demonstrate the impacts of anthropogenic disturbance on hybrid network structure, but they also provide unexpected insights into the structure-stability relationship of hybrid networks.

Bat community response to intensification of biomass production for bioenergy across the southeastern United States.

Human demand for food, fiber, and space is accelerating the rate of change of land cover and land use. Much of the world now consists of a matrix of natural forests, managed forests, agricultural cropland, and urbanized plots. Expansion of domestic energy production efforts in the United States is one driver predicted to influence future land-use and land management practices across large spatial scales. Favorable growing conditions make the southeastern United States an ideal location for producing a large portion of the country's renewable bioenergy. We investigated patterns of bat occurrence in two bioenergy feedstocks commonly grown in this region (corn, Zea mays, and pine, Pinus taeda and P. elliottii). We also evaluated potential impacts of the three major pathways of woody biomass extraction (residue removal following clearcut harvest, short-rotation energy plantations, and mid-rotation forest thinning) to bat occurrence through a priori land-use contrasts. We acoustically sampled bat vocalizations at 84 sites in the Southeastern Plains and Southern Coastal Plains of the southeastern United States across three years. We found that mid-rotation thinning resulted in positive effects on bat occurrence, and potential conversion of unmanaged (reference) forest to managed forest for timber and/or bioenergy harvest resulted in negative effects on bat occurrence when effects were averaged across all species. The effects of short-rotation energy plantations, removal of logging residues from plantation clearcuts, and corn were equivocal for all bat species examined. Our results suggest that accelerated production of biomass for energy production through either corn or intensively managed pine forests is not likely to have an adverse effect on bat communities, so long as existing older unmanaged forests are not converted to managed bioenergy or timber plantations. Beyond bioenergy crop production, mid-rotation thinning of even-aged pine stands intended for timber production, increases to the duration of plantation rotations to promote older forest stands, arranging forest stands and crop fields to maximize edge habitat, and maintaining unmanaged forests could benefit bat communities by augmenting roosting and foraging opportunities.

Quantitative and qualitative assessment of pollen DNA metabarcoding using constructed species mixtures.

Pollen DNA metabarcoding-marker-based genetic identification of potentially mixed-species pollen samples-has applications across a variety of fields. While basic species-level pollen identification using standard DNA barcode markers is established, the extent to which metabarcoding (a) correctly assigns species identities to mixes (qualitative matching) and (b) generates sequence reads proportionally to their relative abundance in a sample (quantitative matching) is unclear, as these have not been assessed relative to known standards. We tested the quantitative and qualitative robustness of metabarcoding in constructed pollen mixtures varying in species richness (1-9 species), taxonomic relatedness (within genera to across class) and rarity (5%-100% of grains), using Illumina MiSeq with the markers rbcL and ITS2. Qualitatively, species composition determinations were largely correct, but false positives and negatives occurred. False negatives were typically driven by lack of a barcode gap or rarity in a sample. Species richness and taxonomic relatedness, however, did not strongly impact correct determinations. False positives were likely driven by contamination, chimeric sequences and/or misidentification by the bioinformatics pipeline. Quantitatively, the proportion of reads for each species was only weakly correlated with its relative abundance, in contrast to suggestions from some other studies. Quantitative mismatches are not correctable by consistent scaling factors, but instead are context-dependent on the other species present in a sample. Together, our results show that metabarcoding is largely robust for determining pollen presence/absence but that sequence reads should not be used to infer relative abundance of pollen grains.

Generalisation and specialisation in hoverfly (Syrphidae) grassland pollen transport networks revealed by DNA metabarcoding.

Pollination by insects is a key ecosystem service and important to wider ecosystem function. Most species-level pollination networks studied have a generalised structure, with plants having several potential pollinators, and pollinators in turn visiting a number of different plant species. This is in apparent contrast to a plant's need for efficient conspecific pollen transfer. The aim of this study was to investigate the structure of pollen transport networks at three levels of biological hierarchy: community, species and individual. We did this using hoverflies in the genus Eristalis, a key group of non-Hymenopteran pollinators. We constructed pollen transport networks using DNA metabarcoding to identify pollen. We captured hoverflies in conservation grasslands in west Wales, UK, removed external pollen loads, sequenced the pollen DNA on the Illumina MiSeq platform using the standard plant barcode rbcL, and matched sequences using a pre-existing plant DNA barcode reference library. We found that Eristalis hoverflies transport pollen from 65 plant taxa, more than previously appreciated. Networks were generalised at the site and species level, suggesting some degree of functional redundancy, and were more generalised in late summer compared to early summer. In contrast, pollen transport at the individual level showed some degree of specialisation. Hoverflies defined as "single-plant visitors" varied from 40% of those captured in early summer to 24% in late summer. Individual hoverflies became more generalised in late summer, possibly in response to an increase in floral resources. Rubus fruticosus agg. and Succisa pratensis were key plant species for hoverflies at our sites Our results contribute to resolving the apparent paradox of how generalised pollinator networks can provide efficient pollination to plant species. Generalised hoverfly pollen transport networks may result from a varied range of short-term specialised feeding bouts by individual insects. The generalisation and functional redundancy of Eristalis pollen transport networks may increase the stability of the pollination service they deliver.

Reproductive modification in forest plantations: impacts on biodiversity and society.

1000 I. 1000 II. 1001 III. 1014 IV. 1015 V. 1016 1016 References 1016 SUMMARY: Genetic engineering (GE) can be used to improve forest plantation productivity and tolerance of biotic and abiotic stresses. However, gene flow from GE forest plantations is a large source of ecological, social and legal controversy. The use of genetic technologies to mitigate or prevent gene flow has been discussed widely and should be technically feasible in a variety of plantation taxa. However, potential ecological effects of such modifications, and their social acceptability, are not well understood. Focusing on Eucalyptus, Pinus, Populus and Pseudotsuga - genera that represent diverse modes of pollination and seed dispersal - we conducted in-depth reviews of ecological processes associated with reproductive tissues. We also explored potential impacts of various forms of reproductive modification at stand and landscape levels, and means for mitigating impacts. We found little research on potential reactions by the public and other stakeholders to reproductive modification in forest plantations. However, there is considerable research on related areas that suggest key dimensions of concern and support. We provide detailed suggestions for research to understand the biological and social dimensions of containment technologies, and consider the role of regulatory and market restrictions that obstruct necessary ecological and genetic research.

Foraging traits modulate stingless bee community disassembly under forest loss.

Anthropogenic land use change is an important driver of impacts to biological communities and the ecosystem services they provide. Pollination is one ecosystem service that may be threatened by community disassembly. Relatively little is known about changes in bee community composition in the tropics, where pollination limitation is most severe and land use change is rapid. Understanding how anthropogenic changes alter community composition and functioning has been hampered by high variability in responses of individual species. Trait-based approaches, however, are emerging as a potential method for understanding responses of ecologically similar species to global change. We studied how communities of tropical, eusocial stingless bees (Apidae: Meliponini) disassemble when forest is lost. These bees are vital tropical pollinators that exhibit high trait diversity, but are under considerable threat from human activities. We compared functional traits of stingless bee species found in pastures surrounded by differing amounts of forest in an extensively deforested landscape in southern Costa Rica. Our results suggest that foraging traits modulate competitive interactions that underlie community disassembly patterns. In contrast to both theoretical predictions and temperate bee communities, we found that stingless bee species with the widest diet breadths were less likely to persist in sites with less forest. These wide-diet-breadth species also tend to be solitary foragers, and are competitively subordinate to group-foraging stingless bee species. Thus, displacement by dominant, group-foraging species may make subordinate species more dependent on the larger or more diversified resource pool that natural habitats offer. We also found that traits that may reduce reliance on trees-nesting in the ground or inside nests of other species-correlated with persistence in highly deforested landscapes. The functional trait perspective we employed enabled capturing community processes in analyses and suggests that land use change may disassemble bee communities via different mechanisms in temperate and tropical areas. Our results further suggest that community processes, such as competition, can be important regulators of community disassembly under land use change. A better understanding of community disassembly processes is critical for conserving and restoring pollinator communities and the ecosystem services and functions they provide.

Experimental species removals impact the architecture of pollination networks.

Mutualistic networks are key for the creation and maintenance of biodiversity, yet are threatened by global environmental change. Most simulation models assume that network structure remains static after species losses, despite theoretical and empirical reasons to expect dynamic responses. We assessed the effects of experimental single bumblebee species removals on the structure of entire flower visitation networks. We hypothesized that network structure would change following processes linking interspecific competition with dietary niche breadth. We found that single pollinator species losses impact pollination network structure: resource complementarity decreased, while resource overlap increased. Despite marginally increased connectance, fewer plant species were visited after species removals. These changes may have negative functional impacts, as complementarity is important for maintaining biodiversity-ecological functioning relationships and visitation of rare plant species is critical for maintaining diverse plant communities.

Climatic Drivers of Diarrheagenic Escherichia coli Incidence: A Systematic Review and Meta-analysis.

BACKGROUND: Positive associations have been noted between temperature and diarrhea incidence, but considerable uncertainty surrounds quantitative estimates of this relationship because of pathogen-specific factors and a scarcity of data on the influence of meteorological factors on the risk of disease. Quantifying these relationships is important for disease prevention and climate change adaptation. METHODS: To address these issues, we performed a systematic literature review of studies in which at least 1 full year of data on the monthly incidence of diarrheagenic Escherichia coli were reported. We characterized seasonal patterns of disease incidence from 28 studies. In addition, using monthly time- and location-specific weather data for 18 studies, we performed univariate Poisson models on individual studies and a meta-analysis, using a generalized estimating equation, on the entire data set. RESULTS: We found an 8% increase in the incidence of diarrheagenic E. coli (95% confidence interval, 5%-11%; P < .0001) for each 1°C increase in mean monthly temperature. We found a modest positive association between 1-month-lagged mean rainfall and incidence of diarrheagenic E. coli, which was not statistically significant when we controlled for temperature. CONCLUSIONS: These results suggest that increases in ambient temperature correspond to an elevated incidence of diarrheagenic E. coli and underscore the need to redouble efforts to prevent the transmission of these pathogens in the face of increasing global temperatures.

Niche partitioning due to adaptive foraging reverses effects of nestedness and connectance on pollination network stability.

Much research debates whether properties of ecological networks such as nestedness and connectance stabilise biological communities while ignoring key behavioural aspects of organisms within these networks. Here, we computationally assess how adaptive foraging (AF) behaviour interacts with network architecture to determine the stability of plant-pollinator networks. We find that AF reverses negative effects of nestedness and positive effects of connectance on the stability of the networks by partitioning the niches among species within guilds. This behaviour enables generalist pollinators to preferentially forage on the most specialised of their plant partners which increases the pollination services to specialist plants and cedes the resources of generalist plants to specialist pollinators. We corroborate these behavioural preferences with intensive field observations of bee foraging. Our results show that incorporating key organismal behaviours with well-known biological mechanisms such as consumer-resource interactions into the analysis of ecological networks may greatly improve our understanding of complex ecosystems.

Pollinator specialization: from the individual to the community.

1190 I. 1190 II. 1191 III. 1191 IV. 1193 V. 1193 1194 References 1194 SUMMARY: Most spermatophytes need conspecific pollen in order to produce seed. This need for specialization seems to conflict with the generalized nature of most plant-pollinator interactions. Specialization and generalization are dynamic - not fixed - and exist simultaneously in multiple states at different levels of biological hierarchy. Over the short term, specialization ensures conspecific pollen transfer, whereas over the long term, generalization improves system-level robustness. The balance between specialization and generalization at different scales is critical for different kinds of ecological functioning and is an important factor in plant speciation and the evolution of plant mating systems. Community context, including diversity and interaction network structure at different levels of aggregation, is a key driver of specialization dynamics.

Heterospecific pollen deposition in Delphinium barbeyi: linking stigmatic pollen loads to reproductive output in the field.

BACKGROUND AND AIMS: Most pollinators are generalists and therefore are likely to transfer heterospecific pollen among co-flowering plants. Most work on the impacts of heterospecific pollen deposition on plant fecundity has utilized hand-pollination experiments in greenhouse settings, and we continue to know very little about the reproductive effects of heterospecific pollen in field settings. METHODS: We explored how patterns of naturally deposited heterospecific pollen relate to the reproductive output of Delphinium barbeyi, a common subalpine perennial herb in the Rocky Mountains (USA). We assessed a wide range of naturally occurring heterospecific pollen proportions and pollen load sizes, and linked stigmatic pollen deposition directly to seed set in individual carpels in the field. KEY RESULTS: We found that heterospecific pollen deposition in D. barbeyi is common, but typically found at low levels across stigmas collected in our sites. Neither conspecific nor heterospecific pollen deposition was related to carpel abortion. By contrast, we saw a significant positive relationship between conspecific pollen amount and viable seed production, as well as a significant negative interaction between the effects of conspecific pollen and heterospecific pollen amount, whereby the effect of conspecific pollen on viable seed production became weaker with greater heterospecific deposition on stigmas. CONCLUSIONS: To our knowledge, this is the first demonstration of a relationship between heterospecific pollen and seed production in a field setting. In addition, it is the first report of an interaction between conspecific and heterospecific pollen quantities on seed production. These findings, taken with the results from other studies, suggest that greenhouse hand-pollination studies and field studies should be more tightly integrated in future work to better understand how heterospecific pollen transfer can be detrimental for plant reproduction.