Nathaniel Pope

Native wildflower plantings support wild bee abundance and diversity in agricultural landscapes across the United States.

Global trends in pollinator-dependent crops have raised awareness of the need to support managed and wild bee populations to ensure sustainable crop production. Provision of sufficient forage resources is a key element for promoting bee populations within human impacted landscapes, particularly those in agricultural lands where demand for pollination service is high and land use and management practices have reduced available flowering resources. Recent government incentives in North America and Europe support the planting of wildflowers to benefit pollinators; surprisingly, in North America there has been almost no rigorous testing of the performance of wildflower mixes, or their ability to support wild bee abundance and diversity. We tested different wildflower mixes in a spatially replicated, multiyear study in three regions of North America where production of pollinator-dependent crops is high: Florida, Michigan, and California. In each region, we quantified flowering among wildflower mixes composed of annual and perennial species, and with high and low relative diversity. We measured the abundance and species richness of wild bees, honey bees, and syrphid flies at each mix over two seasons. In each region, some but not all wildflower mixes provided significantly greater floral display area than unmanaged weedy control plots. Mixes also attracted greater abundance and richness of wild bees, although the identity of best mixes varied among regions. By partitioning floral display size from mix identity we show the importance of display size for attracting abundant and diverse wild bees. Season-long monitoring also revealed that designing mixes to provide continuous bloom throughout the growing season is critical to supporting the greatest pollinator species richness. Contrary to expectation, perennials bloomed in their first season, and complementarity in attraction of pollinators among annuals and perennials suggests that inclusion of functionally diverse species may provide the greatest benefit. Wildflower mixes may be particularly important for providing resources for some taxa, such as bumble bees, which are known to be in decline in several regions of North America. No mix consistently attained the full diversity that was planted. Further study is needed on how to achieve the desired floral display and diversity from seed mixes.

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.

VASCULAR PLANTS OF ADJACENT SERPENTINE AND GRANITE OUTCROPS ON THE DEER ISLES, MAINE, U.S.A.

Abstract We performed a comparative study of the vascular flora of a serpentine outcrop, Pine Hill, and that of a granite outcrop, Settlement Quarry, from Little Deer Isle and Deer Isle, respectively, Hancock County, Maine. We established four transects along a gradient from exposed to forested areas within each outcrop. Plants were recorded for presence and percent cover from circular plots along each transect. Soil and tissue samples were collected to examine soil-tissue elemental relations. One hundred thirty-two taxa were recorded from serpentine and 89 from granite. Fifty-seven taxa were shared by both sites. Species richness (α diversity) and diversity indices (Shannon-Weaver and Simpson) suggested significant differences between sites and within sites. Principle Component Analysis suggested substrates differed significantly between sites and between exposures within sites. Tissue analyses suggested intraspecific variation with respect to tissue elemental concentrations, especially in Achillea millefolium, Oenothera biennis, Prunus virginiana, Selaginella rupestris, Spiraea alba var. latifolia, and Vaccinium angustifolium. Serpentine populations of many taxa showed low tissue Ca∶Mg ratios (< 1) and high Ni concentrations. Two-way ANOVA showed significant substrate × species effects for several elements, including those that typically characterize serpentine substrates (Ca, Mg, Cr, Ni), suggesting significant genetic variation within species with respect to substrate. Finally, we compared our species list for Pine Hill with a plant survey done at Pine Hill and five additional serpentine sites of Maine in 1977 and provide a list of 285 vascular plant taxa from 62 families for serpentine in Maine.

ORNITHOCOPROPHILOUS PLANTS OF MOUNT DESERT ROCK, A REMOTE BIRD-NESTING ISLAND IN THE GULF OF MAINE, U.S.A.

Abstract Plants growing on seabird-nesting islands are uniquely adapted to deal with guano-derived soils high in N and P. Such ornithocoprophilous plants found in isolated, oceanic settings provide useful models for ecological and evolutionary investigations. The current study explored the plants found on Mount Desert Rock (MDR), a small seabird-nesting, oceanic island 44 km south of Mount Desert Island (MDI), Hancock County, Maine, U.S.A. Twenty-seven species of vascular plants from ten families were recorded. Analyses of guano-derived soils from the rhizosphere of the three most abundant species from bird-nesting sites of MDR showed significantly higher (P < 0.05) NO3−, available P, extractable Cd, Cu, Pb, and Zn, and significantly lower Mn compared to soils from the rhizosphere of conspecifics on non-bird nesting coastal bluffs from nearby MDI. Bio-available Pb was several-fold higher in guano soils than for background levels for Maine. Leaf tissue elemental analyses from conspecifics on and off guano soils showed significant differences with respect to N, Ca, K, Mg, Fe, Mn, Zn, and Pb, although trends were not always consistent. Two-way ANOVA indicated a significant interaction between species and substrate for Ca, Mg, Zn, and Pb tissue accumulation, showing that for these four elements there is substantial differentiation among species found on and off of guano soil. A compilation of species lists from other important seabird-nesting islands in the region suggested an ornithocoprophilous flora for northeastern North America consisting of 168 species from 39 families, with Asteraceae (29 taxa; 17.3%), Poaceae (25 taxa; 14.9%), Polygonaceae (10 taxa; 5.95%), Caryophyllaceae (9 taxa; 5.4%), and Rosaceae (9 taxa; 5.4%) as the most species-rich families. The taxa were predominantly hermaphroditic (69%) and perennial (66%) species, native (60%) to eastern North America.

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.

Positive density-dependent reproduction regulated by local kinship and size in an understorey tropical tree

BACKGROUND AND AIMS Global pollinator declines and continued habitat fragmentation highlight the critical need to understand reproduction and gene flow across plant populations. Plant size, conspecific density and local kinship (i.e. neighbourhood genetic relatedness) have been proposed as important mechanisms influencing the reproductive success of flowering plants, but have rarely been simultaneously investigated. METHODS We conducted this study on a continuous population of the understorey tree Miconia affinis in the Forest Dynamics Plot on Barro Colorado Island in central Panama. We used spatial, reproductive and population genetic data to investigate the effects of tree size, conspecific neighbourhood density and local kinship on maternal and paternal reproductive success. We used a Bayesian framework to simultaneously model the effects of our explanatory variables on the mean and variance of maternal viable seed set and siring success. KEY RESULTS Our results reveal that large trees had lower proportions of viable seeds in their fruits but sired more seeds. We documented differential effects of neighbourhood density and local kinship on both maternal and paternal reproductive components. Trees in more dense neighbourhoods produced on average more viable seeds, although this positive density effect was influenced by variance-inflation with increasing local kinship. Neighbourhood density did not have significant effects on siring success. CONCLUSIONS This study is one of the first to reveal an interaction among tree size, conspecific density and local kinship as critical factors differentially influencing maternal and paternal reproductive success. We show that both maternal and paternal reproductive success should be evaluated to determine the population-level and individual traits most essential for plant reproduction. In addition to conserving large trees, we suggest the inclusion of small trees and the conservation of dense patches with low kinship as potential strategies for strengthening the reproductive status of tropical trees.

The role of elevation and soil chemistry in the distribution and ion accumulation of floral morphs of Streptanthus polygaloides Gray (Brassicaceae), a Californian nickel hyperaccumulator

Background: The flora of serpentine/ultramafic soils provides an excellent model system for the study of natural selection in plant populations. Streptanthus polygaloides is a nickel hyperaccumulator that is endemic to serpentine soils in the Sierra Nevada of California, and has four floral morphs (yellow, purple, yellow-to-purple and undulate). Aims: We investigate three hypotheses: (1) the purple morph occurs in colder, wetter climates than the yellow morph; (2) tissue–soil ionic relationships differ among morphs; and (3) morphs occur on soils with differing elemental concentrations. Methods: We queried herbarium records to investigate patterns of occurrence among the yellow and purple floral morphs, and analysed soil and tissue samples from wild populations of all four morphs. Results: The purple morph inhabited serpentine outcrops with colder temperatures and greater precipitation levels than the yellow morph. Concentrations of elements in leaf tissue and rhizosphere soil differed little among populations of the morphs, but showed substantial within-site variation. Conclusions: Our results suggest that a climatic gradient may be responsible for divergence in floral colour among populations of S. polygaloides. Because of the large within-site variation in soil and tissue elemental concentrations, plants appear to have a varied physiological response to edaphic factors, regardless of morph membership.

Adding landscape genetics and individual traits to the ecosystem function paradigm reveals the importance of species functional breadth

Significance Pollinators mediate reproductive processes between plants; however, little is known about how plant and pollinator traits impact pollen dispersal and resulting genetic connectivity. Our study quantifies pollen-dispersal distances and pollination effectiveness for individual pollinator species. We demonstrate that seed set is positively driven by pollinator body size, but long-distance pollen dispersal is mediated by both large-bodied and small-bodied bees. We also reveal that individual plant and population attributes impact pollen-dispersal distances and seed production, respectively. Thus, we show that plant and pollinator traits mediate pollination function and that the entire pollinator community, large and small, plays an important role in the maintenance of genetic connectivity. Animal pollination mediates both reproduction and gene flow for the majority of plant species across the globe. However, past functional studies have focused largely on seed production; although useful, this focus on seed set does not provide information regarding species-specific contributions to pollen-mediated gene flow. Here we quantify pollen dispersal for individual pollinator species across more than 690 ha of tropical forest. Specifically, we examine visitation, seed production, and pollen-dispersal ability for the entire pollinator community of a common tropical tree using a series of individual-based pollinator-exclusion experiments followed by molecular-based fractional paternity analyses. We investigate the effects of pollinator body size, plant size (as a proxy of floral display), local plant density, and local plant kinship on seed production and pollen-dispersal distance. Our results show that while large-bodied pollinators set more seeds per visit, small-bodied bees visited flowers more frequently and were responsible for more than 49% of all long-distance (beyond 1 km) pollen-dispersal events. Thus, despite their size, small-bodied bees play a critical role in facilitating long-distance pollen-mediated gene flow. We also found that both plant size and local plant kinship negatively impact pollen dispersal and seed production. By incorporating genetic and trait-based data into the quantification of pollination services, we highlight the diversity in ecological function mediated by pollinators, the influential role that plant and population attributes play in driving service provision, and the unexpected importance of small-bodied pollinators in the recruitment of plant genetic diversity.

Elevation, Not Deforestation, Promotes Genetic Differentiation in a Pioneer Tropical Tree.

The regeneration of disturbed forest is an essential part of tropical forest ecology, both with respect to natural disturbance regimes and large-scale human-mediated logging, grazing, and agriculture. Pioneer tree species are critical for facilitating the transition from deforested land to secondary forest because they stabilize terrain and enhance connectivity between forest fragments by increasing matrix permeability and initiating disperser community assembly. Despite the ecological importance of early successional species, little is known about their ability to maintain gene flow across deforested landscapes. Utilizing highly polymorphic microsatellite markers, we examined patterns of genetic diversity and differentiation for the pioneer understory tree Miconia affinis across the Isthmus of Panama. Furthermore, we investigated the impact of geographic distance, forest cover, and elevation on genetic differentiation among populations using circuit theory and regression modeling within a landscape genetics framework. We report marked differences in historical and contemporary migration rates and moderately high levels of genetic differentiation in M. affinis populations across the Isthmus of Panama. Genetic differentiation increased significantly with elevation and geographic distance among populations; however, we did not find that forest cover enhanced or reduced genetic differentiation in the study region. Overall, our results reveal strong dispersal for M. affinis across human-altered landscapes, highlighting the potential use of this species for reforestation in tropical regions. Additionally, this study demonstrates the importance of considering topography when designing programs aimed at conserving genetic diversity within degraded tropical landscapes.

Diversity and soil-tissue elemental relations of vascular plants of Callahan Mine, Brooksville, Maine, U.S.A

Abstract Metal-contaminated soils provide numerous stressors to plant life, resulting in unique plant communities worldwide. The current study focuses on the vascular plants of Callahan Mine in Brooksville, ME, USA, a Superfund site contaminated with Cu, Zn, Pb, and other pollutants. One hundred and fifty-five taxa belonging to 50 families were identified, with the Asteraceae (21%), Poaceae (11%), and Rosaceae (9%) as the most species-rich families. Ninety-six species encountered at the Mine were native to North America (62%), including 11 taxa (7%) with rarity status in at least one New England state. Fifty-one species were non-native (33%), including nine taxa (6%) considered invasive in at least one New England state. We characterized how the plant community changed across different habitats at the Mine, from disturbed and exposed (waste rock piles, tailings pond) to inundated and relatively undisturbed (wetland, shore), and documented concurrent shifts in the ionic content of the soils across the habitats. We found substantial differences in both the plant community and soil chemical features among habitats. Habitats separated out along a single axis of an ordination of the plant community, with wetland and shore habitats at one extreme and tailings pond and waste rock-pile habitats at the other. The first principal component axis of the 21 soil variables was significantly predicted by the ordination of the plant community, indicating a gradient of increasing organic matter, Fe, Mg, Mn, total N, Na, and K roughly parallel to the gradient of increasing wetland vegetation. None of the plant species tested accumulated substantial concentrations of metals in their leaf tissue except Salix bebbiana and Populus balsamifera, which accumulated 1070 ppm and 969 ppm Zn in dry leaf tissue, respectively—approximately one-third of the concentration considered as hyperaccumulation for Zn.