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.     

Land use change has stronger effects on functional diversity than taxonomic diversity in tropical Andean hummingbirds

Land use change modifies the environment at multiple spatial scales, and is a main driver of species declines and deterioration of ecosystem services. However, most of the research on the effects of land use change has focused on taxonomic diversity, while functional diversity, an important predictor of ecosystem services, is often neglected. We explored how local and landscape scale characteristics influence functional and taxonomic diversity of hummingbirds in the Andes Mountains in southern Ecuador. Data was collected in six landscapes along a land use gradient, from an almost intact landscape to one dominated by cattle pastures. We used point counts to sample hummingbirds from 2011 to 2012 to assessed how local factors (i.e., vegetation structure, flowering plants richness, nectar availability) and landscape factors (i.e., landscape heterogeneity, native vegetation cover) influenced taxonomic and functional diversity. Then, we analyzed environment – trait relationships (RLQ test) to explore how different hummingbird functional traits influenced species responses to these factors. Taxonomic and functional diversity of hummingbirds were positively associated with landscape heterogeneity but only functional diversity was positively related to native vegetation coverage. We found a weak response of taxonomic and functional diversity to land use change at the local scale. Environment‐trait associations showed that body mass of hummingbirds likely influenced species sensitivity to land use change. In conclusion, landscape heterogeneity created by land use change can positively influence hummingbird taxonomic and functional diversity; however, a reduction of native vegetation cover could decrease functional diversity. Given that functional diversity can mediate ecosystem services, the conservation of native vegetation cover could play a key role in the maintenance of hummingbird pollination services in the tropical Andes. Moreover, there are particular functional traits, such as body mass, that increase a species sensitivity to land use change.     

Abundance of common species,not species richness,drives delivery of a real‐world ecosystem service

Biodiversity‐ecosystem functioning experiments have established that species richness and composition are both important determinants of ecosystem function in an experimental context. Determining whether this result holds for real‐world ecosystem services has remained elusive, however, largely due to the lack of analytical methods appropriate for large‐scale, associational data. Here, we use a novel analytical approach, the Price equation, to partition the contribution to ecosystem services made by species richness, composition and abundance in four large‐scale data sets on crop pollination by native bees. We found that abundance fluctuations of dominant species drove ecosystem service delivery, whereas richness changes were relatively unimportant because they primarily involved rare species that contributed little to function. Thus, the mechanism behind our results was the skewed species‐abundance distribution. Our finding that a few common species, not species richness, drive ecosystem service delivery could have broad generality given the ubiquity of skewed species‐abundance distributions in nature.     

Global agricultural productivity is threatened by increasing pollinator dependence without a parallel increase in crop diversification

The global increase in the proportion of land cultivated with pollinator‐dependent crops implies increased reliance on pollination services. Yet agricultural practices themselves can profoundly affect pollinator supply and pollination. Extensive monocultures are associated with a limited pollinator supply and reduced pollination, whereas agricultural diversification can enhance both. Therefore, areas where agricultural diversity has increased, or at least been maintained, may better sustain high and more stable productivity of pollinator‐dependent crops. Given that >80% of all crops depend, to varying extents, on insect pollination, a global increase in agricultural pollinator dependence over recent decades might have led to a concomitant increase in agricultural diversification. We evaluated whether an increase in the area of pollinator‐dependent crops has indeed been associated with an increase in agricultural diversity, measured here as crop diversity, at the global, regional, and country scales for the period 1961–2016. Globally, results show a relatively weak and decelerating rise in agricultural diversity over time that was largely decoupled from the strong and continually increasing trend in agricultural dependency on pollinators. At regional and country levels, there was no consistent relationship between temporal changes in pollinator dependence and crop diversification. Instead, our results show heterogeneous responses in which increasing pollinator dependence for some countries and regions has been associated with either an increase or a decrease in agricultural diversity. Particularly worrisome is a rapid expansion of pollinator‐dependent oilseed crops in several countries of the Americas and Asia that has resulted in a decrease in agricultural diversity. In these regions, reliance on pollinators is increasing, yet agricultural practices that undermine pollination services are expanding. Our analysis has thereby identified world regions of particular concern where environmentally damaging practices associated with large‐scale, industrial agriculture threaten key ecosystem services that underlie productivity, in addition to other benefits provided by biodiversity.     

Landscape perspectives on agricultural intensification and biodiversity – ecosystem service management

Understanding the negative and positive effects of agricultural land use for the conservation of biodiversity, and its relation to ecosystem services, needs a landscape perspective. Agriculture can contribute to the conservation of high‐diversity systems, which may provide important ecosystem services such as pollination and biological control via complementarity and sampling effects. Land‐use management is often focused on few species and local processes, but in dynamic, agricultural landscapes, only a diversity of insurance species may guarantee resilience (the capacity to reorganize after disturbance). Interacting species experience their surrounding landscape at different spatial scales, which influences trophic interactions. Structurally complex landscapes enhance local diversity in agroecosystems, which may compensate for local high‐intensity management. Organisms with high‐dispersal abilities appear to drive these biodiversity patterns and ecosystem services, because of their recolonization ability and larger resources experienced. Agri‐environment schemes (incentives for farmers to benefit the environment) need to broaden their perspective and to take the different responses to schemes in simple (high impact) and complex (low impact) agricultural landscapes into account. In simple landscapes, local allocation of habitat is more important than in complex landscapes, which are in total at risk. However, little knowledge of the relative importance of local and landscape management for biodiversity and its relation to ecosystem services make reliable recommendations difficult.     

Comparison of pollinators and natural enemies: a meta‐analysis of landscape and local effects on abundance and richness in crops

To manage agroecosystems for multiple ecosystem services, we need to know whether the management of one service has positive, negative, or no effects on other services. We do not yet have data on the interactions between pollination and pest‐control services. However, we do have data on the distributions of pollinators and natural enemies in agroecosystems. Therefore, we compared these two groups of ecosystem service providers, to see if the management of farms and agricultural landscapes might have similar effects on the abundance and richness of both. In a meta‐analysis, we compared 46 studies that sampled bees, predatory beetles, parasitic wasps, and spiders in fields, orchards, or vineyards of food crops. These studies used the proximity or proportion of non‐crop or natural habitats in the landscapes surrounding these crops (a measure of landscape complexity), or the proximity or diversity of non‐crop plants in the margins of these crops (a measure of local complexity), to explain the abundance or richness of these beneficial arthropods. Compositional complexity at both landscape and local scales had positive effects on both pollinators and natural enemies, but different effects on different taxa. Effects on bees and spiders were significantly positive, but effects on parasitoids and predatory beetles (mostly Carabidae and Staphylinidae) were inconclusive. Landscape complexity had significantly stronger effects on bees than it did on predatory beetles and significantly stronger effects in non‐woody rather than in woody crops. Effects on richness were significantly stronger than effects on abundance, but possibly only for spiders. This abundance‐richness difference might be caused by differences between generalists and specialists, or between arthropods that depend on non‐crop habitats (ecotone species and dispersers) and those that do not (cultural species). We call this the ‘specialist‐generalist’ or ‘cultural difference’ mechanism. If complexity has stronger effects on richness than abundance, it might have stronger effects on the stability than the magnitude of these arthropod‐mediated ecosystem services. We conclude that some pollinators and natural enemies seem to have compatible responses to complexity, and it might be possible to manage agroecosystems for the benefit of both. However, too few studies have compared the two, and so we cannot yet conclude that there are no negative interactions between pollinators and natural enemies, and no trade‐offs between pollination and pest‐control services. Therefore, we suggest a framework for future research to bridge these gaps in our knowledge.     

What does species richness tell us about functional trait diversity? Predictions and evidence for responses of species and functional trait diversity to land‐use change

In the conservation literature on land‐use change, it is often assumed that land‐use intensification drives species loss, driving a loss of functional trait diversity and ecosystem function. Modern research, however, does not support this cascade of loss for all natural systems. In this paper we explore the errors in this assumption and present a conceptual model taking a more mechanistic approach to the species–functional trait association in a context of land‐use change. We provide empirical support for our model's predictions demonstrating that the association of species and functional trait diversity follows various trajectories in response to land‐use change. The central premise of our model is that land‐use change impacts upon processes of community assembly, not species per se. From the model, it is clear that community context (i.e. type of disturbance, species pool size) will affect the response trajectory of the relationship between species and functional trait diversity in communities undergoing land‐use change. The maintenance of ecosystem function and of species diversity in the face of increasing land‐use change are complementary goals. The use of a more ecologically realistic model of responses of species and functional traits will improve our ability to make wise management decisions to achieve both aims in specific at‐risk systems.     

Complementary ecosystem services provided by pest predators and pollinators increase quantity and quality of coffee yields

Wild animals substantially support crop production by providing ecosystem services, such as pollination and natural pest control. However, the strengths of synergies between ecosystem services and their dependencies on land-use management are largely unknown. Here, we took an experimental approach to test the impact of land-use intensification on both individual and combined pollination and pest control services in coffee production systems at Mount Kilimanjaro. We established a full-factorial pollinator and vertebrate exclosure experiment along a land-use gradient from traditional homegardens (agroforestry systems), shaded coffee plantations to sun coffee plantations (total sample size = 180 coffee bushes). The exclusion of vertebrates led to a reduction in fruit set of ca 9%. Pollinators did not affect fruit set, but significantly increased fruit weight of coffee by an average of 7.4%. We found no significant decline of these ecosystem services along the land-use gradient. Pest control and pollination service were thus complementary, contributing to coffee production by affecting the quantity and quality of a major tropical cash crop across different coffee production systems at Mount Kilimanjaro.     

Natural hazard threats to pollinators and pollination

Natural hazards are naturally occurring physical events that can impact human welfare both directly and indirectly, via shocks to ecosystems and the services they provide. Animal‐mediated pollination is critical for sustaining agricultural economies and biodiversity, yet stands to lose both from present exposure to natural hazards, and future climate‐driven shifts in their distribution, frequency, and intensity. In contrast to the depth of knowledge available for anthropogenic‐related threats, our understanding of how naturally occurring extreme events impact pollinators and pollination has not yet been synthesized. We performed a systematic review and meta‐analysis to examine the potential impacts of natural hazards on pollinators and pollination in natural and cultivated systems. From a total of 117 studies (74% of which were observational), we found evidence of community and population‐level impacts to plants and pollinators from seven hazard types, including climatological (extreme heat, fire, drought), hydrological (flooding), meteorological (hurricanes), and geophysical (volcanic activity, tsunamis). Plant and pollinator response depended on the type of natural hazard and level of biological organization observed; 19% of cases reported no significant impact, whereas the majority of hazards held consistent negative impacts. However, the effects of fire were mixed, but taxa specific; meta‐analysis revealed that bee abundance and species richness tended to increase in response to fire, differing significantly from the mainly negative response of Lepidoptera. Building from this synthesis, we highlight important future directions for pollination‐focused natural hazard research, including the need to: (a) advance climate change research beyond static “mean‐level” changes by better incorporating “shock” events; (b) identify impacts at higher levels of organization, including ecological networks and co‐evolutionary history; and (c) address the notable gap in crop pollination services research—particularly in developing regions of the world. We conclude by discussing implications for safeguarding pollination services in the face of global climate change.     

Are ecosystem services stabilized by differences among species? A test using crop pollination

Biological diversity could enhance ecosystem service provision by increasing the mean level of services provided, and/or by providing more consistent (stable) services over space and time. Ecological theory predicts that when an ecosystem service is provided by many species, it will be stabilized against disturbance by a variety of 'stabilizing mechanisms.' However, few studies have investigated whether stabilizing mechanisms occur in real landscapes affected by human disturbance. We used two datasets on crop pollination by wild native bees to screen for and differentiate among three stabilizing mechanisms: density compensation (negative co-variance among species' abundances); response diversity (differential response to environmental variables among species); and cross-scale resilience (response to the same environmental variable at different scales by different species). In both datasets, we found response diversity and cross-scale resilience, but not density compensation. We conclude that stabilizing mechanisms may contribute to the stability of pollination services in our study areas, emphasizing the insurance value of seemingly 'redundant' species. Furthermore, the absence of density compensation that we found at the landscape scale contrasts with findings of previous small-scale experimental and modelling work, suggesting that we should not assume that density compensation will stabilize ecosystem services in real landscapes.     

Contrasting effects of land‐use changes on herbivory and pollination networks

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The challenge of accurately documenting bee species richness in agroecosystems: bee diversity in eastern apple orchards

Bees are important pollinators of agricultural crops, and bee diversity has been shown to be closely associated with pollination, a valuable ecosystem service. Higher functional diversity and species richness of bees have been shown to lead to higher crop yield. Bees simultaneously represent a mega‐diverse taxon that is extremely challenging to sample thoroughly and an important group to understand because of pollination services. We sampled bees visiting apple blossoms in 28 orchards over 6 years. We used species rarefaction analyses to test for the completeness of sampling and the relationship between species richness and sampling effort, orchard size, and percent agriculture in the surrounding landscape. We performed more than 190 h of sampling, collecting 11,219 specimens representing 104 species. Despite the sampling intensity, we captured <75% of expected species richness at more than half of the sites. For most of these, the variation in bee community composition between years was greater than among sites. Species richness was influenced by percent agriculture, orchard size, and sampling effort, but we found no factors explaining the difference between observed and expected species richness. Competition between honeybees and wild bees did not appear to be a factor, as we found no correlation between honeybee and wild bee abundance. Our study shows that the pollinator fauna of agroecosystems can be diverse and challenging to thoroughly sample. We demonstrate that there is high temporal variation in community composition and that sites vary widely in the sampling effort required to fully describe their diversity. In order to maximize pollination services provided by wild bee species, we must first accurately estimate species richness. For researchers interested in providing this estimate, we recommend multiyear studies and rarefaction analyses to quantify the gap between observed and expected species richness.     

Mismatched outcomes for biodiversity and ecosystem services: testing the responses of crop pollinators and wild bee biodiversity to habitat enhancement

Supporting ecosystem services and conserving biodiversity may be compatible goals, but there is concern that service‐focused interventions mostly benefit a few common species. We use a spatially replicated, multiyear experiment in four agricultural settings to test if enhancing habitat adjacent to crops increases wild bee diversity and abundance on and off crops. We found that enhanced field edges harbored more taxonomically and functionally abundant, diverse, and compositionally different bee communities compared to control edges. Enhancements did not increase the abundance or diversity of bees visiting crops, indicating that the supply of pollination services was unchanged following enhancement. We find that actions to promote crop pollination improve multiple dimensions of biodiversity, underscoring their conservation value, but these benefits may not be spilling over to crops. More work is needed to identify the conditions that promote effective co‐management of biodiversity and ecosystem services.     

传粉动物多样性的保护与农业景观传粉服务的提升

传粉动物为许多植物尤其是作物提供了重要的传粉服务, 在保障全球粮食安全和人类福祉、缓冲气候变化对作物产量的影响等方面都发挥着重要的作用。然而来自全球土地利用变化、化学农药使用、外来物种入侵及气候变化等的威胁, 导致传粉动物的多样性下降并造成了依赖动物传粉的作物产量和品质的下降。针对这一情况, 作者提出了农业景观传粉动物多样性保护和利用的3种主要途径: (1)改善生产管理, 例如减少化学农药的使用、适当地采取有机种植; (2)促进景观多样性, 包括创建适宜野生传粉者的半自然生境、保护高价值的自然生境、作物多样化、合理配置资源和生境的空间分布; (3)加强对本土传粉动物的保护和开发利用。文章最后提出, 为进一步提升传粉服务, 还需加强对传粉者的生物学特征、传粉服务的需求与供给现状、影响传粉动物多样性和传粉服务的农作措施和景观因素等方面的研究。     

Pollinator‐mediated interactions between cultivated papaya and co‐flowering plant species

Many modern crop varieties rely on animal pollination to set fruit and seeds. Intensive crop plantations usually do not provide suitable habitats for pollinators so crop yield may depend on the surrounding vegetation to maintain pollination services. However, little is known about the effect of pollinator‐mediated interactions among co‐flowering plants on crop yield or the underlying mechanisms. Plant reproductive success is complex, involving several pre‐ and post‐pollination events; however, the current literature has mainly focused on pre‐pollination events in natural plant communities. We assessed pollinator sharing and the contribution to pollinator diet in a community of wild and cultivated plants that co‐flower with a focal papaya plantation. In addition, we assessed heterospecific pollen transfer to the stigmatic loads of papaya and its effect on fruit and seed production. We found that papaya shared at least one pollinator species with the majority of the co‐flowering plants. Despite this, heterospecific pollen transfer in cultivated papaya was low in open‐pollinated flowers. Hand‐pollination experiments suggest that heterospecific pollen transfer has no negative effect on fruit production or weight, but does reduce seed production. These results suggest that co‐flowering plants offer valuable floral resources to pollinators that are shared with cultivated papaya with little or no cost in terms of heterospecific pollen transfer. Although HP reduced seed production, a reduced number of seeds per se are not negative, given that from an agronomic perspective the number of seeds does not affect the monetary value of the papaya fruit.     

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.     

Utilisation of plant functional diversity in wildflower strips for the delivery of multiple agroecosystem services

Increased plant diversity in cropping systems can play an important role in agriculture by enhancing arthropod‐mediated ecosystem services, including biological control and pollination. However, there is limited research investigating the concurrent influence of plant functional diversity within cultivated systems on different arthropod functional groups, the provision of multiple ecosystem services, and crop yield. During a field experiment, repeated over 2 years, we measured the effect of increasing plant functional diversity on community structure of arthropod visitors, the abundance of multiple pests and induced crop damage, and fruit production in two varieties of tomato. Plant resources (floral and extra‐floral nectar and pollen) were included within experimental plots in four levels, with each level increasing the plant functional group richness, based on floral morphology and availability of resources, in a replacement series. The presence of sown flower mixtures in experimental plots was associated with increased abundance and diversity of natural enemy functional groups and an enhanced abundance of bees (Hymenoptera: Apiformes). However, we only detected relatively small variability in arthropod visitors among types of mixtures, and increased abundance of natural enemies did not translate into stronger pest suppression or reduced crop damage. Lepidoptera pest damage was significantly higher in plots adjacent to wildflower strips, an ecosystem disservice, but a significantly higher crop productivity was recorded from these plots. Our results provide evidence that inclusion of non‐crop plant resources in agroecosystems can improve the conservation of beneficial arthropods and may lead to increased crop productivity.     

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.     

Trait‐based approaches to analyze links between the drivers of change and ecosystem services: Synthesizing existing evidence and future challenges

Understanding the responses of biodiversity to drivers of change and the effects of biodiversity on ecosystem properties and ecosystem services is a key challenge in the context of global environmental change. We performed a systematic review and meta‐analysis of the scientific literature linking direct drivers of change and ecosystem services via functional traits of three taxonomic groups (vegetation, invertebrates, and vertebrates) to: (1) uncover trends and research biases in this field; and (2) synthesize existing empirical evidence. Our results show the existence of important biases in published studies related to ecosystem types, taxonomic groups, direct drivers of change, ecosystem services, geographical range, and the spatial scale of analysis. We found multiple evidence of links between drivers and services mediated by functional traits, particularly between land‐use changes and regulating services in vegetation and invertebrates. Seventy‐five functional traits were recorded in our sample. However, few of these functional traits were repeatedly found to be associated with both the species responses to direct drivers of change (response traits) and the species effects on the provision of ecosystem services (effect traits). Our results highlight the existence of potential “key functional traits,” understood as those that have the capacity to influence the provision of multiple ecosystem services, while responding to specific drivers of change, across a variety of systems and organisms. Identifying “key functional traits” would help to develop robust indicator systems to monitor changes in biodiversity and their effects on ecosystem functioning and ecosystem services supply.     

Land use intensification alters ecosystem multifunctionality via loss of biodiversity and changes to functional composition

Global change, especially land‐use intensification, affects human well‐being by impacting the delivery of multiple ecosystem services (multifunctionality). However, whether biodiversity loss is a major component of global change effects on multifunctionality in real‐world ecosystems, as in experimental ones, remains unclear. Therefore, we assessed biodiversity, functional composition and 14 ecosystem services on 150 agricultural grasslands differing in land‐use intensity. We also introduce five multifunctionality measures in which ecosystem services were weighted according to realistic land‐use objectives. We found that indirect land‐use effects, i.e. those mediated by biodiversity loss and by changes to functional composition, were as strong as direct effects on average. Their strength varied with land‐use objectives and regional context. Biodiversity loss explained indirect effects in a region of intermediate productivity and was most damaging when land‐use objectives favoured supporting and cultural services. In contrast, functional composition shifts, towards fast‐growing plant species, strongly increased provisioning services in more inherently unproductive grasslands.