Deforestation-free land-use change and organic matter-centered management improve the C footprint of oil palm expansion.

In recent decades, mounting evidence has indicated that the expansion of oil palm (OP) plantations at the expense of tropical forest has had a far pernicious effect on ecosystem aspects. While various deforestation-free strategies have been proposed to enhance OP sustainability, field-based evidence still need to be consolidated, in particular with respect to savanna regions where OP expansion has recently occurred and that present large area with potential for OP cultivation. Here we show that the common management practice creating within the plantation the so-called management zones explained nearly five times more variability of soil biogeochemical properties than the savanna land-use change per se. We also found that clayey-soil savanna conversion into OP increased total ecosystem C stocks by 40 ± 13 Mg C ha-1 during a full OP cultivation cycle, which was due to the higher OP-derived C accumulated in the biomass and in the soil as compared to the loss of savanna-derived C. In addition, application of organic residues in specific management zones enhanced the accumulation of soil organic carbon by up to 1.9 Mg ha-1  year-1 over the full cycle. Within plantation, zones subjected to organic amendments sustained similar soil microbial activity as in neighboring savannas. Our findings represent an empirical proof-of-concept that the conversion of non-forested land in parallel with organic matter-oriented management strategies can enhance OP agroecosystems C sink capacity while promoting microbe-mediated soil functioning. Nonetheless, savannas are unique and threatened ecosystems that support a vast biodiversity. Therefore, we suggest to give priority attention to conservation of natural savannas and direct more research toward the impacts of the conversion and subsequent management of degraded savannas.

Perceptions from non-governmental actors on forest and landscape restoration, challenges and strategies for successful implementation across Asia, Africa and Latin America.

Forest and Landscape Restoration (FLR) has been defined as a planned process that aims to regain ecological functionality and enhance human well-being in degraded landscapes. Several governments and organizations worldwide rose to the challenge of halting degradation and restoring landscapes. Commitments are ambitious, thus a synthesis of current experiences with and strategies for implementation is important to inform future actions. To guide successful implementation, the Global Partnership on FLR put forward six principles, namely, the conservation and enhancement of ecosystems at landscape scales, the restoration of multiple functions, the engagement of multiple stakeholders, with allowances for context dependency and adaptive management. Non-governmental organizations, acting globally, regionally and (or) at national and local scales, play a fundamental role in supporting governments fulfill their commitments. Therefore, we gathered the perceptions of actors within non-governmental organizations engaged in FLR across countries in Asia, Africa and Latin America about what FLR is and their perceived challenges and strategies for implementation. We employed the six principles of FLR to organize and evaluate the responses. Results show that the principles of landscape scale, ecosystem conservation and enhancement, and multi stakeholder engagement are all considered by interviewees as core components of an FLR program. Yet several restoration projects shared by interviewees still required further evidence of a landscape vision, and the integration of actors beyond local communities and the environmental government sectors. Context dependency was evident in the clear incorporation of local natural resource governance norms, such as tribal and community management in project structure, yet few projects appeared to be designed by local actors. The principle of "adaptive management" was mostly missing from the responses, perhaps because most projects had not had sufficient time to learn from intervention outcomes. Key financial challenges for FLR implementation were the short duration and availability of funding, high-up front costs and few short-term returns. To overcome these challenges, promising strategies relate to the development of tangible economic returns for local actors engaged in productive restorative actions that are planned alongside conservation and ecological restoration actions in the landscape. The challenges of negotiating actions with a multitude of actors and the lack of supportive policies highlighted in the interviews require organizations to focus efforts on leveraging the enactment and enforcement of legislations that look beyond jurisdictional boundaries and support landscape management with clear, long term incentive mechanisms and cross-sectoral collaboration. In addition, implementation can be further supported with the scientifically robust sharing of results on how different FLR projects move forward in meeting the social and environmental objectives of a successful, integrative restoration of degraded landscapes.

Urban bumblebees are smaller and more phenotypically diverse than their rural counterparts.

With urbanization identified as being one of the key drivers of change in global land use, and the rapid expansion of urban areas world-wide, it is relevant to evaluate how novel ecological conditions in cities shape species functional traits, which are essential for how species interact with their environments and with each other. Despite the many comparative studies on organisms living in urban and non-urban areas, our knowledge on species responses to urban environments remains limited. For one, much of the ecological research has assumed that the environment changes in a linear fashion from the city core to the city edges, whereas in reality the environments within the cities are highly heterogeneous. Furthermore, studies on species responses to these highly variable ecosystems are often based on interspecific mean trait values, which ignore the potential for high levels of intraspecific variation among individuals in key functional traits. The current study investigated intraspecific functional trait differences for four functional traits associated with body size, mobility and resource selection among rural and urban populations of two common bumblebee species, Bombus pascuorum and Bombus lapidarius, in urban centres and adjacent rural areas in Switzerland. We document shifts in functional traits towards smaller individuals and higher multidimensional trait variation in urban populations compared to rural conspecifics of both species. This shows that urban individuals for both species are on average smaller sized but populations are distinctively different from rural population by increasing their trait richness and diversifying their trait combinations. In addition, we found bimodality in tongue length within urban B. pascuorum populations. Our results suggest that urban and rural populations possibly experience differential selection pressures resulting in trait differences across and among populations. We argue that variations in the respective foraging landscapes in cities leads to smaller sized but phenotypically more diverse populations, and drive functional trait divergence.

The effects of rainforest fragment area on the strength of plant-pathogen interactions.

Pathogenic interactions between fungi and plants facilitate plant species coexistence and tropical rainforest diversity. Such interactions, however, may be affected by forest fragmentation as fungi are susceptible to anthropogenic disturbance. To examine how fragmentation affects fungus-induced seed and seedling mortality, we sowed seeds of six plant species in soils collected from 21 forest fragments. We compared seedling establishment in unmanipulated soils to soils treated with fungicides. Fungicides increased germination of Toona ciliata seeds and decreased mortality of Syzygium rubicundum and Olea dioica seedlings. The fungus-induced mortality of one of these species, S. rubicundum, decreased with decreasing fragment size, indicating that its interactions with pathogenic fungi may weaken as fragments become smaller. We provide evidence that a potential diversity-maintaining plant-fungus interaction weakens in small forest fragments and suggest that such disruptions may have important long-term consequences for plant diversity. However, we emphasize the need for further research across rainforest plant communities to better understand the future of diversity in fragmented rainforest landscapes.

Non-bee insects are important contributors to global crop pollination.

Wild and managed bees are well documented as effective pollinators of global crops of economic importance. However, the contributions by pollinators other than bees have been little explored despite their potential to contribute to crop production and stability in the face of environmental change. Non-bee pollinators include flies, beetles, moths, butterflies, wasps, ants, birds, and bats, among others. Here we focus on non-bee insects and synthesize 39 field studies from five continents that directly measured the crop pollination services provided by non-bees, honey bees, and other bees to compare the relative contributions of these taxa. Non-bees performed 25-50% of the total number of flower visits. Although non-bees were less effective pollinators than bees per flower visit, they made more visits; thus these two factors compensated for each other, resulting in pollination services rendered by non-bees that were similar to those provided by bees. In the subset of studies that measured fruit set, fruit set increased with non-bee insect visits independently of bee visitation rates, indicating that non-bee insects provide a unique benefit that is not provided by bees. We also show that non-bee insects are not as reliant as bees on the presence of remnant natural or seminatural habitat in the surrounding landscape. These results strongly suggest that non-bee insect pollinators play a significant role in global crop production and respond differently than bees to landscape structure, probably making their crop pollination services more robust to changes in land use. Non-bee insects provide a valuable service and provide potential insurance against bee population declines.

Evaluating realized seed dispersal across fragmented tropical landscapes: a two-fold approach using parentage analysis and the neighbourhood model.

Despite the importance of seed dispersal for survival of plant species in fragmented landscapes, data on seed dispersal at landscape scales remain sparse. Effective seed dispersal among fragments determines recolonization and plant species persistence in such landscapes. We present the first large-scale (216-km2 ) direct estimates of realized seed dispersal of a high-value timber tree (Dysoxylum malabaricum) across an agro-forest landscape in the Western Ghats, India. Based upon an exhaustive inventory of adult trees and a sample of 488 seedlings all genotyped at 10 microsatellite loci, we estimated realized seed dispersal using parentage analysis and the neighbourhood model. Our estimates found that most realized seed dispersal was within 200 m, which is insufficient to effectively bridge the distances between forest patches. We conclude that using mobility of putative animal dispersers can be misleading when estimating tropical tree species vulnerability to habitat fragmentation. This raises serious concerns about the potential of many tropical trees to recolonize isolated forest patches where high-value tree species have already been removed.

Habitat connectivity and local conditions shape taxonomic and functional diversity of arthropods on green roofs.

Increasing development of urban environments creates high pressure on green spaces with potential negative impacts on biodiversity and ecosystem services. There is growing evidence that green roofs - rooftops covered with vegetation - can contribute mitigate the loss of urban green spaces by providing new habitats for numerous arthropod species. Whether green roofs can contribute to enhance taxonomic and functional diversity and increase connectivity across urbanized areas remains, however, largely unknown. Furthermore, only limited information is available on how environmental conditions shape green roof arthropod communities. We investigated the community composition of arthropods (Apidae, Curculionidae, Araneae and Carabidae) on 40 green roofs and 40 green sites at ground level in the city of Zurich, Switzerland. We assessed how the site's environmental variables (such as area, height, vegetation, substrate and connectivity among sites) affect species richness and functional diversity using generalized linear models. We used an extension of co-inertia analysis (RLQ) and fourth-corner analysis to highlight the mechanism underlying community assemblages across taxonomic groups on green roof and ground communities. Species richness was higher at ground-level sites, while no difference in functional diversity was found between green roofs and ground sites. Green roof arthropod diversity increased with higher connectivity and plant species richness, irrespective of substrate depth, height and area of green roofs. The species trait analysis reviewed the mechanisms related to the environmental predictors that shape the species assemblages of the different taxa at ground and roof sites. Our study shows the important contribution of green roofs in maintaining high functional diversity of arthropod communities across different taxonomic groups, despite their lower species richness compared with ground sites. Species communities on green roofs revealed to be characterized by specific trait assemblages. The study also provides details on the environmental conditions that influence arthropod diversity and gives new perspectives on how the design of green roofs can be improved to increase their ecological value. Furthermore, the study highlights the importance of integrating green roofs in planning policies which aim to enhance urban habitat connectivity.

Interactive effects among ecosystem services and management practices on crop production: pollination in coffee agroforestry systems.

Crop productivity is improved by ecosystem services, including pollination, but this should be set in the context of trade-offs among multiple management practices. We investigated the impact of pollination services on coffee production, considering variation in fertilization, irrigation, shade cover, and environmental variables such as rainfall (which stimulates coffee flowering across all plantations), soil pH, and nitrogen availability. After accounting for management interventions, bee abundance improved coffee production (number of berries harvested). Some management interventions, such as irrigation, used once to trigger asynchronous flowering, dramatically increased bee abundance at coffee trees. Others, such as the extent and type of tree cover, revealed interacting effects on pollination and, ultimately, crop production. The effects of management interventions, notably irrigation and addition of lime, had, however, far more substantial positive effects on coffee production than tree cover. These results suggest that pollination services matter, but managing the asynchrony of flowering was a more effective tool for securing good pollination than maintaining high shade tree densities as pollinator habitat. Complex interactions across farm and landscape scales, including both management practices and environmental conditions, shape pollination outcomes. Effective production systems therefore require the integrated consideration of management practices in the context of the surrounding habitat structure. This paper points toward a more strategic use of ecosystem services in agricultural systems, where ecosystem services are shaped by the coupling of management interventions and environmental variables.

Ten principles for a landscape approach to reconciling agriculture, conservation, and other competing land uses.

"Landscape approaches" seek to provide tools and concepts for allocating and managing land to achieve social, economic, and environmental objectives in areas where agriculture, mining, and other productive land uses compete with environmental and biodiversity goals. Here we synthesize the current consensus on landscape approaches. This is based on published literature and a consensus-building process to define good practice and is validated by a survey of practitioners. We find the landscape approach has been refined in response to increasing societal concerns about environment and development tradeoffs. Notably, there has been a shift from conservation-orientated perspectives toward increasing integration of poverty alleviation goals. We provide 10 summary principles to support implementation of a landscape approach as it is currently interpreted. These principles emphasize adaptive management, stakeholder involvement, and multiple objectives. Various constraints are recognized, with institutional and governance concerns identified as the most severe obstacles to implementation. We discuss how these principles differ from more traditional sectoral and project-based approaches. Although no panacea, we see few alternatives that are likely to address landscape challenges more effectively than an approach circumscribed by the principles outlined here.

Plant population differentiation and climate change: responses of grassland species along an elevational gradient.

Mountain ecosystems are particularly susceptible to climate change. Characterizing intraspecific variation of alpine plants along elevational gradients is crucial for estimating their vulnerability to predicted changes. Environmental conditions vary with elevation, which might influence plastic responses and affect selection pressures that lead to local adaptation. Thus, local adaptation and phenotypic plasticity among low and high elevation plant populations in response to climate, soil and other factors associated with elevational gradients might underlie different responses of these populations to climate warming. Using a transplant experiment along an elevational gradient, we investigated reproductive phenology, growth and reproduction of the nutrient-poor grassland species Ranunculus bulbosus, Trifolium montanum and Briza media. Seeds were collected from low and high elevation source populations across the Swiss Alps and grown in nine common gardens at three different elevations with two different soil depths. Despite genetic differentiation in some traits, the results revealed no indication of local adaptation to the elevation of population origin. Reproductive phenology was advanced at lower elevation in low and high elevation populations of all three species. Growth and reproduction of T. montanum and B. media were hardly affected by garden elevation and soil depth. In R. bulbosus, however, growth decreased and reproductive investment increased at higher elevation. Furthermore, soil depth influenced growth and reproduction of low elevation R. bulbosus populations. We found no evidence for local adaptation to elevation of origin and hardly any differences in the responses of low and high elevation populations. However, the consistent advanced reproductive phenology observed in all three species shows that they have the potential to plastically respond to environmental variation. We conclude that populations might not be forced to migrate to higher elevations as a consequence of climate warming, as plasticity will buffer the detrimental effects of climate change in the three investigated nutrient-poor grassland species.

Landscape genetics as a tool for conservation planning: predicting the effects of landscape change on gene flow.

For conservation managers, it is important to know whether landscape changes lead to increasing or decreasing gene flow. Although the discipline of landscape genetics assesses the influence of landscape elements on gene flow, no studies have yet used landscape-genetic models to predict gene flow resulting from landscape change. A species that has already been severely affected by landscape change is the large marsh grasshopper (Stethophyma grossum), which inhabits moist areas in fragmented agricultural landscapes in Switzerland. From transects drawn between all population pairs within maximum dispersal distance (< 3 km), we calculated several measures of landscape composition as well as some measures of habitat configuration. Additionally, a complete sampling of all populations in our study area allowed incorporating measures of population topology. These measures together with the landscape metrics formed the predictor variables in linear models with gene flow as response variable (F(ST) and mean pairwise assignment probability). With a modified leave-one-out cross-validation approach, we selected the model with the highest predictive accuracy. With this model, we predicted gene flow under several landscape-change scenarios, which simulated construction, rezoning or restoration projects, and the establishment of a new population. For some landscape-change scenarios, significant increase or decrease in gene flow was predicted, while for others little change was forecast. Furthermore, we found that the measures of population topology strongly increase model fit in landscape genetic analysis. This study demonstrates the use of predictive landscape-genetic models in conservation and landscape planning.

Remotely sensed evidence of tropical peatland conversion to oil palm.

Rising global demands for food and biofuels are driving forest clearance in the tropics. Oil-palm expansion contributes to biodiversity declines and carbon emissions in Southeast Asia. However, the magnitudes of these impacts remain largely unquantified until now. We produce a 250-m spatial resolution map of closed canopy oil-palm plantations in the lowlands of Peninsular Malaysia (2 million ha), Borneo (2.4 million ha), and Sumatra (3.9 million ha). We demonstrate that 6% (or ≈880,000 ha) of tropical peatlands in the region had been converted to oil-palm plantations by the early 2000s. Conversion of peatswamp forests to oil palm led to biodiversity declines of 1% in Borneo (equivalent to four species of forest-dwelling birds), 3.4% in Sumatra (16 species), and 12.1% in Peninsular Malaysia (46 species). This land-use change also contributed to the loss of ≈140 million Mg of aboveground biomass carbon, and annual emissions of ≈4.6 million Mg of belowground carbon from peat oxidation. Additionally, the loss of peatswamp forests implies the loss of carbon sequestration service through peat accumulation, which amounts to ≈660,000 Mg of carbon annually. By 2010, 2.3 million ha of peatswamp forests were clear-felled, and currently occur as degraded lands. Reforestation of these clearings could enhance biodiversity by up to ≈20%, whereas oil-palm establishment would exacerbate species losses by up to ≈12%. To safeguard the region's biodiversity and carbon stocks, conservation and reforestation efforts should target Central Kalimantan, Riau, and West Kalimantan, which retain three-quarters (3.9 million ha) of the remaining peatswamp forests in Southeast Asia.

Functional diversity enhances dryland forest productivity under long-term climate change.

Short-term experimental studies provided evidence that plant diversity increases ecosystem resilience and resistance to drought events, suggesting diversity to serve as a nature-based solution to address climate change. However, it remains unclear whether the effects of diversity are momentary or still hold over the long term in natural forests to ensure that the sustainability of carbon sinks. By analyzing 57 years of inventory data from dryland forests in Canada, we show that productivity of dryland forests decreased at an average rate of 1.3% per decade, in concert with the temporally increasing temperature and decreasing water availability. Increasing functional trait diversity from its minimum (monocultures) to maximum value increased productivity by 13%. Our results demonstrate the potential role of tree functional trait diversity in alleviating climate change impacts on dryland forests. While recognizing that nature-based climate mitigation (e.g., planting trees) can only be partial solutions, their long-term (decadal) efficacy can be improved by enhancing functional trait diversity across the forest community.

Diverse pollen nutrition can improve the development of solitary bees but does not mitigate negative pesticide impacts.

Floral resource loss and pesticide exposure are major threats to bees in intensively managed agroecosystems, but interactions among these drivers remain poorly understood. Altered composition and lowered diversity of pollen nutrition may reinforce negative pesticide impacts on bees. Here we investigated the development and survival of the solitary bee Osmia bicornis provisioned with three different pollen types, as well as a mixture of these types representing a higher pollen diversity. We exposed bees of each nutritional treatment to five pesticides at different concentrations in the laboratory. Two field-realistic concentrations of three nicotinic acetylcholine receptor (nAChR) modulating insecticides (thiacloprid, sulfoxaflor and flupyradifurone), as well as of two fungicides (azoxystrobin and tebuconazole) were examined. We further measured the expression of two detoxification genes (CYP9BU1, CYP9BU2) under exposure to thiacloprid across different nutrition treatments as a potential mechanistic pathway driving pesticide-nutrition interactions. We found that more diverse pollen nutrition reduced development time, enhanced pollen efficacy (cocoon weight divided by consumed pollen weight) and pollen consumption, and increased weight of O. bicornis after larval development (cocoon weight). Contrary to fungicides, high field-realistic concentrations of all three insecticides negatively affected O. bicornis by extending development times. Moreover, sulfoxaflor and flupyradifurone also reduced pollen efficacy and cocoon weight, and sulfoxaflor reduced pollen consumption and increased mortality. The expression of detoxification genes differed across pollen nutrition types, but was not enhanced after exposure to thiacloprid. Our findings highlight that lowered diversity of pollen nutrition and high field-realistic exposure to nAChR modulating insecticides negatively affected the development of O. bicornis, but we found no mitigation of negative pesticide impacts through increased pollen diversity. These results have important implications for risk assessment for bee pollinators, indicating that negative effects of nAChR modulating insecticides to developing solitary bees are currently underestimated.

Extensive contemporary pollen-mediated gene flow in two herb species, Ranunculus bulbosus and Trifolium montanum, along an altitudinal gradient in a meadow landscape.

BACKGROUND AND AIMS: Genetic connectivity between plant populations allows for exchange and dispersal of adaptive genes, which can facilitate plant population persistence particularly in rapidly changing environments. METHODS: Patterns of historic gene flow, flowering phenology and contemporary pollen flow were investigated in two common herbs, Ranunculus bulbosus and Trifolium montanum, along an altitudinal gradient of 1200-1800 m a.s.l. over a distance of 1 km among five alpine meadows in Switzerland. KEY RESULTS: Historic gene flow was extensive, as revealed by Fst values of 0·01 and 0·007 in R. bulbosus and T. montanum, respectively, by similar levels of allelic richness among meadows and by the grouping of all individuals into one genetic cluster. Our data suggest contemporary pollen flow is not limited across altitudes in either species but is more pronounced in T. montanum, as indicated by the differential decay of among-sibships correlated paternity with increasing spatial distance. Flowering phenology among meadows was not a barrier to pollen flow in T. montanum, as the large overlap between meadow pairs was consistent with the extensive pollen flow. The smaller flowering overlap among R. bulbosus meadows might explain the slightly more limited pollen flow detected. CONCLUSIONS: High levels of pollen flow among altitudes in both R. bulbosus and T. montanum should facilitate exchange of genes which may enhance adaptive responses to rapid climate change.

Landscape composition has limited impact on local genetic structure in mountain clover, Trifolium montanum L.

Semi-dry grasslands in the European Alps have been increasingly fragmented over the last 150 years. Few studies have investigated the implications of landscape configuration for genetic structure and gene flow among remnant habitat patches. Conservation management of semi-dry grassland plants rarely accounts for possible effects of major landscape elements, such as forest patches, as barriers to gene flow and dispersal via seed and pollen, despite their potential importance for biodiversity conservation. Using 1416 individuals from 61 sampling sites across 2 valleys in South-Eastern Switzerland and Amplified fragment length polymorphism (AFLP) fingerprints, we applied a spatial strip and a circle approach to determine the impact of different landscape elements on genetic differentiation in the semi-dry grassland herb Trifolium montanum (mountain clover). Overall genetic differentiation among sampling sites was low (overall F ST = 0.044). Forest area had no effect on gene flow at the landscape scale, but area of semi-dry grassland, the potential habitat of T. montanum, road area, and altitude influenced genetic differentiation among sampling sites. The observed pattern of genetic differentiation suggests that a future increase in forest area, due to land use abandonment, at least in the short term, are unlikely to directly impact patterns of genetic variation in T. montanum.

Spatially explicit scenario analysis for reconciling agricultural expansion, forest protection, and carbon conservation in Indonesia.

Palm oil is the world's most important vegetable oil in terms of production quantity. Indonesia, the world's largest palm-oil producer, plans to double its production by 2020, with unclear implications for the other national priorities of food (rice) production, forest and biodiversity protection, and carbon conservation. We modeled the outcomes of alternative development scenarios and show that every single-priority scenario had substantial tradeoffs associated with other priorities. The exception was a hybrid approach wherein expansion targeted degraded and agricultural lands that are most productive for oil palm, least suitable for food cultivation, and contain the lowest carbon stocks. This approach avoided any loss in forest or biodiversity and substantially ameliorated the impacts of oil-palm expansion on carbon stocks (limiting net loss to 191.6 million tons) and annual food production capacity (loss of 1.9 million tons). Our results suggest that the environmental and land-use tradeoffs associated with oil-palm expansion can be largely avoided through the implementation of a properly planned and spatially explicit development strategy.

Incentives and barriers to private finance for forest and landscape restoration.

Increased private finance can accelerate forest and landscape restoration globally. Here we conduct semi-structured interviews with asset managers, corporations and restoration finance experts to examine incentives and barriers to private restoration finance. Next, we assess what type of restoration projects and regions appeal to different private funders and how current financial barriers can be overcome. We show that market incentives for corporations include meeting net-emission-reduction commitments, impact and sustainable branding opportunities, and promotion of sustainability in supply chains. Conversely, asset managers face stronger barriers to investing in restoration as it is deemed a high-risk, unknown investment with low profitability. We find that investment finance biases towards restoration projects in low-risk areas and corporate finance towards areas with business presence. Both private finance types tend to omit projects focusing on natural regeneration. Through expanded and diversified markets for restoration benefits, strong public policy support and new financial instruments, private finance for restoration can be scaled for a wider variety of restoration projects in more diverse geographical contexts.

Eight microsatellite markers for the bulbous buttercup Ranunculus bulbosus (Ranunculaceae).

PREMISE OF THE STUDY: Pollen dispersal is a key biological process enabling plant populations to maintain genetic connectivity. Direct estimates of pollen dispersal using paternity assignment or correlated paternity estimates require highly variable genetic markers, of which microsatellites are the markers of choice. • METHODS AND RESULTS: Eight species-specific microsatellites have been developed for Ranunculus bulbosus, combining classical enrichment methods with 454 sequencing. These markers have been used in paternity analysis as well as in pollen-pool analyses and proven to be highly polymorphic (seven to 63 alleles in the largest population studied). An excess of homozygotes in six loci indicate the presence of null alleles. • CONCLUSIONS: These markers are the first microsatellites isolated and tested on R. bulbosus and provide a useful tool for population genetic studies in this common grassland herb.