Proximity to wildflower strips did not boost crop pollination on small,diversified farms harboring diverse wild bees

The yield of many agricultural crops depends on pollination services provided by wild and managed bees, many of which are experiencing declines due to factors such as reductions in floral resources. Thus, improving pollinator habitat on farmlands using management strategies like planting wildflower strips is vital for wild bee conservation and sustainable crop pollination. Yet, few studies have examined whether and at what spatial scales wildflower strips enhance crop pollination and yields, and most research has been conducted in large-scale commercial agriculture. Therefore, we investigated the effects of wildflower strips on crop pollination on small, diversified farms (i.e., those growing a variety of crop species) where wild bee diversity and abundance is predicted to be comparatively high. Over three years, on four diversified farms in Montana USA, we tested the hypothesis that distance (20, 60, and 180 m) of crops from native perennial wildflower strips planted alongside crop fields affected wild bee visitation, pollination, and yields of squash and sunflower crop plants. We found that distance to wildflower strips did not affect bee visitation or pollination in crops. Squash yield was pollen-limited in the growing season prior to wildflower strip establishment, and in one of the two years after wildflower strip establishment, but proximity to wildflower strips did not influence the magnitude of pollen limitation. Sunflower seed production was not pollen-limited in any year. Our findings demonstrate that even on diverse farms with wildflower strips and a demonstrated high diversity of bees, some crops do not necessarily receive maximum pollination, regardless of distance from the wildflower strips. However, the value of wildflower strips for supporting wild bee diversity, and other ecological or economic benefits, needs consideration for a full understanding of this pollinator habitat management strategy.     

Pollinator sharing between mass-flowering oilseed rape and co-flowering wild plants: implications for wild plant pollination

Pollinating insects are not only important in wild plant pollination, but also in the production of a large number of crops. Oilseed rape production is increasing globally due to demands for biofuels which may have impacts on pollinating insects which visit the crop and on the pollination services delivered to co-flowering wild plants. In this study, we tested (1) the degree of pollinator sharing between oilseed rape and native wild plants in field margins and hedgerows and (2) the effects of oilseed rape on the quality of pollination service delivered to these wild plants. We found large overlap between flower visitors of wild plants and oilseed rape, but the composition of species overlap differed with respect to each wild plant species. Nearly all individual visitors caught on both the crop and foraging on wild species carried crop pollen, but more than half the insects also carried pollen from wild plants. However, very little oilseed rape pollen was deposited on wild plant stigmas. This shows that (1) oilseed rape overlaps in pollinator niche with most co-flowering wild plants, and (2) crop pollen deposition on wild plant stigmas is low which may indicate that it is unlikely to cause reductions in seed set of wild plants, although this was not measured here. Furthermore, wild plants in field margins and hedgerows are important sources of alternative forage for pollinating insects even when a crop is mass flowering, and we suggest maintenance and augmentation of field margins and hedgerows to provide alternative forage for pollinator conservation to continue provision of pollination services to both crops and wild plants.     

Moths pollinate four crops of Cucurbitaceae in Asia

Pollination ecology of many crops is not fully known, especially in tropical and subtropical regions. Non-bee pollinators may contribute substantially to crop yield, even if they do not receive much attention. Although moth pollination has fascinated ecologists and evolutionary biologists since Darwin, crop pollination by moths has not been well investigated and experimentally examined. Hence, we explored the pollination ecology of four cucurbit crops with crepuscular or nocturnal flowers. Lagenaria siceraria (Molina) Standl., Luffa acutangula (L.) Roxb., Trichosanthes anguina L., and Trichosanthes kirilowii Maxim. all display floral traits suggestive of moth pollination, such as opening around dusk or in evenings and secreting ample dilute nectar. We demonstrated that these crops’ flowers attracted a wide range of moth species, especially hawkmoths. The assemblage of flower-visiting moths varied according to location and season. Pollination treatments and pollen load analysis confirmed the pollination of the four crops by moths, especially hawkmoths. Our results provide evidence for the value as wild pollinators for the four crops, for which domesticated bees cannot provide reliable pollination services in practice. This study lends support to the proposal to pay more attention to the value of non-bee pollinators and to leave some areas unexploited in rural landscapes for the conservation of wild pollinators, including moths.     

Native bees buffer the negative impact of climate warming on honey bee pollination of watermelon crops

If climate change affects pollinator‐dependent crop production, this will have important implications for global food security because insect pollinators contribute to production for 75% of the leading global food crops. We investigate whether climate warming could result in indirect impacts upon crop pollination services via an overlooked mechanism, namely temperature‐induced shifts in the diurnal activity patterns of pollinators. Using a large data set on bee pollination of watermelon crops, we predict how pollination services might change under various climate change scenarios. Our results show that under the most extreme IPCC scenario (A1F1), pollination services by managed honey bees are expected to decline by 14.5%, whereas pollination services provided by most native, wild taxa are predicted to increase, resulting in an estimated aggregate change in pollination services of +4.5% by 2099. We demonstrate the importance of native biodiversity in buffering the impacts of climate change, because crop pollination services would decline more steeply without the native, wild pollinators. More generally, our study provides an important example of how biodiversity can stabilize ecosystem services against environmental change.     

Species Distribution Models for Crop Pollination: A Modelling Framework Applied to Great Britain

Insect pollination benefits over three quarters of the world''s major crops. There is growing concern that observed declines in pollinators may impact on production and revenues from animal pollinated crops. Knowing the distribution of pollinators is therefore crucial for estimating their availability to pollinate crops; however, in general, we have an incomplete knowledge of where these pollinators occur. We propose a method to predict geographical patterns of pollination service to crops, novel in two elements: the use of pollinator records rather than expert knowledge to predict pollinator occurrence, and the inclusion of the managed pollinator supply. We integrated a maximum entropy species distribution model (SDM) with an existing pollination service model (PSM) to derive the availability of pollinators for crop pollination. We used nation-wide records of wild and managed pollinators (honey bees) as well as agricultural data from Great Britain. We first calibrated the SDM on a representative sample of bee and hoverfly crop pollinator species, evaluating the effects of different settings on model performance and on its capacity to identify the most important predictors. The importance of the different predictors was better resolved by SDM derived from simpler functions, with consistent results for bees and hoverflies. We then used the species distributions from the calibrated model to predict pollination service of wild and managed pollinators, using field beans as a test case. The PSM allowed us to spatially characterize the contribution of wild and managed pollinators and also identify areas potentially vulnerable to low pollination service provision, which can help direct local scale interventions. This approach can be extended to investigate geographical mismatches between crop pollination demand and the availability of pollinators, resulting from environmental change or policy scenarios.     

Integrated crop pollination to buffer spatial and temporal variability in pollinator activity

Insect pollination improves the yield and quality of many crops, yet there is increasing evidence of insufficient insect pollinators limiting crop production. Effective Integrated Crop Pollination (ICP) involves adaptable, targeted and cost-effective management of crop pollination and encourages the use of both wild and managed pollinators where appropriate. In this study we investigate how the addition of honeybee hives affects the community of insects visiting oilseed rape, and if hive number and location affect pollinator foraging and oilseed rape pollination in order to provide evidence for effective ICP. We found that introducing hives increased overall flower visitor numbers and altered the pollinator community, which became dominated by honeybees. Furthermore a greater number of hives did not increase bee numbers significantly but did result in honeybees foraging further into fields. The timing of surveys and proximity to the field edge influenced different pollinators in different ways and represents an example of spatial and temporal complementarity. For example dipteran flower visitor numbers declined away from the field edge whereas honeybees peaked at intermediate distances into the field. Furthermore, no significant effects of survey round on wild bees overall was observed but honeybee numbers were relatively lower during peak flowering and dipteran abundance was greater in later survey rounds. Thus combining diverse wild pollinators and managed species for crop pollination buffers spatial and temporal variation in flower visitation. However we found no effect of insect pollination on seed set or yield of oilseed rape in our trial, highlighting the critical need to understand crop demand for insect pollination before investments are made in managing pollination services.     

Pollination crisis in the butterfly-pollinated wild carnation Dianthus carthusianorum?

Knowledge of pollination services provided by flower visitors is a prerequisite for understanding (co)evolutionary processes between plants and their pollinators, for evaluating the degree of specialization in the pollination system, and for assessing threats from a potential pollination crisis. This study examined pollination efficiency and visitation frequency of pollinators--key traits of pollinator-mediated fecundity--in a natural population of the wild carnation Dianthus carthusianorum. The five lepidopteran pollinator species observed differed in pollination efficiency and visitation frequency. Pollinator importance, the product of pollination efficiency and visitation frequency, was determined by the pollinator's visitation frequency. Pollination of D. carthusianorum depended essentially on only two of the five recorded pollinator species. Seed set was pollen-limited and followed a saturating dose-response function with a threshold of c. 50 deposited pollen grains for fruit development. Our results confirm that D. carthusianorum is specialized to lepidopteran pollinators, but is not particularly adapted to the two main pollinator species identified. The local persistence of D. carthusianorum is likely to be at risk as its reproduction depends essentially on only two of the locally abundant, but generally vulnerable, butterfly species.     

Bumble bee abundance and richness improves honey bee pollination behaviour in sweet cherry

Pollination service in agricultural crops increases significantly with pollinator diversity and wild pollinator abundance. Differences in the foraging behaviour of pollinating insects are one of the reasons why pollinator diversity and abundance enhances crop pollination. Here, we focused on the foraging behaviour of honey bees and bumble bees in sweet cherry orchards. In addition, we studied the influence of bee diversity and abundance on the foraging behaviour of honey bees and bumble bees. Honey bees were found to visit fewer flowers than bumble bees. Bumble bees also showed a higher probability of changing trees between rows than honey bees. Both visitation rate and probability of row changes of honey bees increased with bumble bee diversity and with bumble bee abundance. We also found that the probability of row changes of honey bees increased with increasing bumble bee abundance. These effects of bumble bee richness and abundance on the pollination behaviour of honey bees can improve the pollination performance of honey bees in crops that depend on cross pollination. Our results highlight the higher pollination performance of bumble bees and the facilitative effect of wild pollinators to crop pollination.     

Effects of landscape metrics on Apis and non-Apis pollinators and seed set in common buckwheat

Crop pollination is an essential ecosystem service, which, when provided by wild pollinator species, can be negatively affected by human activities at a landscape scale. Various pollinator species can be affected by landscape factors and/or act at different spatial scales. We investigated whether different landscape factors at different spatial scales are correlated with pollinator abundances and pollination success. We examined common buckwheat, which is a self-incompatible crop species, pollinated by managed honeybees, native honeybees, and wild non-honeybee insects. To determine the effects of landscape factors at different spatial scales, we sampled flower-visiting insects and recorded seed sets. Results from the sampled flower visitors indicated that the abundances of managed honeybees, native honeybees, and wild non-honeybee insects were significantly related to the distance of managed hives, the area of forest cover within a 3-km radius, and the area of forest and grassland cover within a 100-m radius, respectively. Additionally, the latter two influential factors of the landscape, but not the distance of managed hives, showed significant positive correlations with the seed set. These results suggest that considering landscape factors at different spatial scales associated with the characteristics of managed and wild pollinators would help to ensure crop yields.     

Economic gain,stability of pollination and bee diversity decrease from southern to northern Europe

Bees are in decline potentially leading to reduced pollination and hence production of insect-pollinated crops in many countries. It is however still unclear whether the consequences of pollinator shortages differ among countries with different environmental and societal conditions. Here, we calculated economic gains attributed to insect (particularly bee) pollination (EVIP) as well as their contribution to the total value of crop production (vulnerability), and analyzed their temporal trends and inter-annual variability from 1991 to 2009 for each country of the European Union (EU). To understand which factors drive country-specific differences in pollinator dependency and stability of insect-dependent crop yields, we further asked whether EVIP, vulnerability and stability of yields were influenced by a country's climate, the number of wild bee species and/or managed honeybee hives per country, and (agricultural) gross domestic product (GDP).Across Europe, crop pollination by insects accounted for 14.6 [±3.3] billion EUR annually (EVIP), which equaled 12 (±0.8)% of the total economic value of annual crop production. Gains strongly varied among countries. Both EVIP and vulnerability increased (and the inter-annual variation of vulnerability decreased) significantly from the colder northern to the warmer Mediterranean EU countries, in parallel with increases in the number of wild bee species. Across years, economic importance of pollination increased in all but three EU countries. Apples were the most important insect-pollinated crop in the EU, accounting for 16% of the EU's total EVIP. Our results show that whereas dependency on insect pollination increased from the colder north to the warmer south, variation in economic gain from insect pollination decreased, indicating that Mediterranean countries had more stable yields of pollinator-dependent crops across years and thus more reliable gains from pollination services.     

Seasonal complementary in pollinators of soft-fruit crops

Understanding the relative contributions of wild and managed pollinators, and the functional contributions made by a diverse pollinator community, is essential to the maintenance of yields in the 75% of our crops that benefit from insect pollination. We describe a field study and pollinator exclusion experiments conducted on two soft-fruit crops in a system with both wild and managed pollinators. We test whether fruit quality and quantity is limited by pollination, and whether different pollinating insects respond differently to varying weather conditions. Both strawberries and raspberries produced fewer marketable fruits when insects were excluded, demonstrating dependence on insect pollinators. Raspberries had a short flowering season which coincided with peak abundance of bees, and attracted many bees per flower. In contrast, strawberries had a much longer flowering season and appeared to be much less attractive to pollinators, so that ensuring adequate pollination is likely to be more challenging. The proportion of high-quality strawberries was positively related to pollinator abundance, suggesting that yield was limited by inadequate pollination on some farms. The relative abundance of different pollinator taxa visiting strawberries changed markedly through the season, demonstrating seasonal complementarity. Insect visitors responded differently to changing weather conditions suggesting that diversity can reduce the risk of pollination service shortfalls. For example, flies visited the crop flowers in poor weather and at the end of the flowering season when other pollinators were scarce, and so may provide a unique functional contribution. Understanding how differences between pollinator groups can enhance pollination services to crops strengthens the case for multiple species management. We provide evidence for the link between increased diversity and function in real crop systems, highlighting the risks of replacing all pollinators with managed alternatives.     

Characterization of wild bee communities in apple and blueberry orchards

1. Wild bees provide invaluable ecosystem services in agricultural landscapes such as pollination. However, in recent decades, pollinator biodiversity, especially in wild bees, is declining on a global scale, with potentially far‐reaching consequences for crop production. Thus, there is an urgent need to determine whether wild bees are present in agricultural systems, such as fruit orchards.
2. In the present study, we examined the wild bee fauna at species and community levels during the period of bee activity (May to August) in apple and high‐bush blueberry orchards in New England.
3. Bee communities are crop‐specific and dominated by very few species, which fluctuate according to crop and season. The blueberry associated bee fauna was more diverse. In apple, communities were phylogenetically clustered at the genus level and dominated by solitary ground nesting bees within the genus Andrena. Species fluctuated widely in presence and abundance throughout the season, leading to differences in community composition and functional trait structure.
4. The results obtained in the present study show that apple and blueberry harbour a distinct and diverse bee fauna that performs vital pollination services in orchards. Our results provide essential baseline data for wild bees in blueberry and apple orchards and this can be used to improve management and conservation strategies for wild bee preservation in these crops.

     

Valuing insect pollination services with cost of replacement

Value estimates of ecosystem goods and services are useful to justify the allocation of resources towards conservation, but inconclusive estimates risk unsustainable resource allocations. Here we present replacement costs as a more accurate value estimate of insect pollination as an ecosystem service, although this method could also be applied to other services. The importance of insect pollination to agriculture is unequivocal. However, whether this service is largely provided by wild pollinators (genuine ecosystem service) or managed pollinators (commercial service), and which of these requires immediate action amidst reports of pollinator decline, remains contested. If crop pollination is used to argue for biodiversity conservation, clear distinction should be made between values of managed- and wild pollination services. Current methods either under-estimate or over-estimate the pollination service value, and make use of criticised general insect and managed pollinator dependence factors. We apply the theoretical concept of ascribing a value to a service by calculating the cost to replace it, as a novel way of valuing wild and managed pollination services. Adjusted insect and managed pollinator dependence factors were used to estimate the cost of replacing insect- and managed pollination services for the Western Cape deciduous fruit industry of South Africa. Using pollen dusting and hand pollination as suitable replacements, we value pollination services significantly higher than current market prices for commercial pollination, although lower than traditional proportional estimates. The complexity associated with inclusive value estimation of pollination services required several defendable assumptions, but made estimates more inclusive than previous attempts. Consequently this study provides the basis for continued improvement in context specific pollination service value estimates.     

Managed bumble bees increase flower visitation but not fruit weight in polytunnel strawberry crops

Animal-mediated pollination is essential for the production and quality of fruits and seeds of many crops consumed by humans. However, crop pollination services might be compromised when wild pollinators are scarce. Managed pollinators are commonly used in crops to supplement such services with the assumption that they will enhance crop yield. However, information on the spatiotemporal pollinator-dependence of crops is still limited. We assessed the contribution of commercial bumble bee colonies compared to the available pollinator community on strawberry (‘Fortuna’ variety) flower visitation and strawberry quality across a landscape gradient of agricultural intensification (i.e. polytunnel berry crop cover). We used colonies of bumble bees in winter and in spring, i.e. when few and most wild pollinators are in their flight period, respectively. The placement of colonies increased visits of bumble bees to strawberry flowers, especially in winter. The use of bumble bee colonies did not affect flower visitation by other insects, mainly honey bees, hoverflies and other Diptera. Flower visitation by both honey bees and wild insects did not vary between seasons and was unrelated to the landscape gradient of berry crop cover. Strawberries were of the highest quality (i.e. weight) when insect-mediated pollination was allowed, and their quality was positively related to wild flower visitors in winter but not in spring. However, increased visits to strawberry flowers by managed bumble bees and honey bees had no effect on strawberry weight. Our results suggest that the pollination services producing high quality strawberry fruits are provided by the flower visitor community present in the study region without the need to use managed bumble bees.     

Climate‐driven spatial mismatches between British orchards and their pollinators: increased risks of pollination deficits

Understanding how climate change can affect crop‐pollinator systems helps predict potential geographical mismatches between a crop and its pollinators, and therefore identify areas vulnerable to loss of pollination services. We examined the distribution of orchard species (apples, pears, plums and other top fruits) and their pollinators in Great Britain, for present and future climatic conditions projected for 2050 under the SRES A1B Emissions Scenario. We used a relative index of pollinator availability as a proxy for pollination service. At present, there is a large spatial overlap between orchards and their pollinators, but predictions for 2050 revealed that the most suitable areas for orchards corresponded to low pollinator availability. However, we found that pollinator availability may persist in areas currently used for fruit production, which are predicted to provide suboptimal environmental suitability for orchard species in the future. Our results may be used to identify mitigation options to safeguard orchard production against the risk of pollination failure in Great Britain over the next 50 years; for instance, choosing fruit tree varieties that are adapted to future climatic conditions, or boosting wild pollinators through improving landscape resources. Our approach can be readily applied to other regions and crop systems, and expanded to include different climatic scenarios.     

On the inconsistency of pollinator species traits for predicting either response to land‐use change or functional contribution

The response and effect trait framework, if supported empirically, would provide for powerful and general predictions about how biodiversity loss leads to loss in ecosystem function. This framework proposes that species traits will explain how different species respond to disturbance (i.e. response traits) as well as their contribution to ecosystem function (i.e. effect traits). However, predictive response and effect traits remain elusive for most systems. Here, we use data on crop pollination services provided by native, wild bees to explore the role of six commonly used species traits in determining both species’ response to land‐use change and the subsequent effect on crop pollination. Analyses were conducted in parallel for three crop systems (watermelon, cranberry, and blueberry) located within the same geographical region (mid‐Atlantic USA). Bee species traits did not strongly predict species’ response to land‐use change, and the few traits that were weakly predictive were not consistent across crops. Similarly, no trait predicted species’ overall functional contribution in any of the three crop systems, although body size was a good predictor of per capita efficiency in two systems. Overall we were unable to make generalizable predictions regarding species responses to land‐use change and its effect on the delivery of crop pollination services. Pollinator traits may be useful for understanding ecological processes in some systems, but thus far the promise of traits‐based ecology has yet to be fulfilled for pollination ecology.     

作物驯化的遗传学研究及其在大豆育种中的应用

驯化遗传学是进化生物学研究的一个重要分支, 对作物驯化历程进行深入探索, 可以更好地认识人类早期农耕文明的产生和发展, 了解物种的形成乃至生命的起源和进化, 对指导人们利用野生资源来改良现有的栽培作物和培育新的有价值的品种也具有重要的意义。本文综述了作物驯化的遗传学研究内容和进展, 以及重要作物大豆驯化起源的研究概况, 探讨了目前作物驯化研究对大豆驯化遗传学及野生大豆资源综合利用的指导意义。     

作物驯化的遗传学研究及其在大豆育种中的应用

驯化遗传学是进化生物学研究的一个重要分支,对作物驯化历程进行深入探索,可以更好地认识人类早期农耕文明的产生和发展,了解物种的形成乃至生命的起源和进化,对指导人们利用野生资源来改良现有的栽培作物和培育新的有价值的品种也具有重要的意义。本文综述了作物驯化的遗传学研究内容和进展,以及重要作物大豆驯化起源的研究概况,探讨了目前作物驯化研究对大豆驯化遗传学及野生大豆资源综合利用的指导意义。     

Pollinators and pollination of oilseed rape crops (Brassica napus L.) in Ireland: ecological and economic incentives for pollinator conservation

Pollinators are beneficial for many wild and crop plants. As a mass-flowering crop, oilseed rape has received much focus in terms of its pollination requirements but despite a threefold increase in area of cultivation of this crop in Ireland over the past 5 years, little is known about its pollination here. We surveyed the flower visiting insects found in commercial winter oilseed rape fields and evaluated the importance of different pollinator groups, investigated the contribution of insect pollination to oilseed rape seed production, and estimated the economic value of insect pollination to the crop at a national level. Our data showed that winter oilseed rape is visited by a wide variety of insect species, including the honeybee, bumblebees, solitary bees, and hoverflies. The honeybee, Eristalis hoverflies and bumblebees (especially Bombus sensu stricto and B. lapidarius) were the best pollinators of winter oilseed rape based on the number of pollen grains they carry, visitation rates per flower and their relative abundance per field. Exclusion of pollinators resulted in a 27 % decrease in the number of seeds produced, and a 30 % decrease in seed weight per pod in winter crops, with comparable values from a spring oilseed rape field also. The economic value of insect pollination to winter oilseed rape was estimated as €2.6 million per annum, while the contribution to spring oilseed rape was €1.3 million, resulting in an overall value of €3.9 million per annum. We can suggest the appropriate conservation and management of both honeybees and wild pollinators in agricultural areas to ensure continued provision of pollination services to oilseed rape, as a decrease in insect numbers has the potential to negatively influence crop yields.     

Pollination by the locally endangered island flying fox (Pteropus hypomelanus) enhances fruit production of the economically important durian (Durio zibethinus)

Fruit bats provide valuable pollination services to humans through a unique coevolutionary relationship with chiropterophilous plants. However, chiropterophily in the Old World and the pollination roles of large bats, such as flying foxes (Pteropus spp., Acerodon spp., Desmalopex spp.), are still poorly understood and require further elucidation. Efforts to protect these bats have been hampered by a lack of basic quantitative information on their role as ecosystem service providers. Here, we investigate the role of the locally endangered island flying fox Pteropus hypomelanus in the pollination ecology of durian (Durio zibethinus), an economically important crop in Southeast Asia. On Tioman Island, Peninsular Malaysia, we deployed 19 stations of paired infrared camera and video traps across varying heights at four individual flowering trees in a durian orchard. We detected at least nine species of animal visitors, but only bats had mutualistic interactions with durian flowers. There was a clear vertical stratification in the feeding niches of flying foxes and nectar bats, with flying foxes feeding at greater heights in the trees. Flying foxes had a positive effect on mature fruit set and therefore serve as important pollinators for durian trees. As such, semi‐wild durian trees—particularly tall ones—may be dependent on flying foxes for enhancing reproductive success. Our study is the first to quantify the role of flying foxes in durian pollination, demonstrating that these giant fruit bats may have far more important ecological, evolutionary, and economic roles than previously thought. This has important implications and can aid efforts to promote flying fox conservation, especially in Southeast Asian countries.