Ecological network complexity scales with area.

Larger geographical areas contain more species-an observation raised to a law in ecology. Less explored is whether biodiversity changes are accompanied by a modification of interaction networks. We use data from 32 spatial interaction networks from different ecosystems to analyse how network structure changes with area. We find that basic community structure descriptors (number of species, links and links per species) increase with area following a power law. Yet, the distribution of links per species varies little with area, indicating that the fundamental organization of interactions within networks is conserved. Our null model analyses suggest that the spatial scaling of network structure is determined by factors beyond species richness and the number of links. We demonstrate that biodiversity-area relationships can be extended from species counts to higher levels of network complexity. Therefore, the consequences of anthropogenic habitat destruction may extend from species loss to wider simplification of natural communities.

Distribution and pollination services of wild bees and hoverflies along an altitudinal gradient in mountain hay meadows.

Extensively managed and flower-rich mountain hay meadows, hotspots of Europe's biodiversity, are subject to environmental and climatic gradients linked to altitude. While the shift of pollinators from bee- to fly-dominated communities with increasing elevation across vegetation zones is well established, the effect of highland altitudinal gradients on the community structure of pollinators within a specific habitat is poorly understood. We assessed wild bee and hoverfly communities, and their pollination service to three plant species common in mountain hay meadows, in eighteen extensively managed yellow oat grasslands (Trisetum flavescens) with an altitudinal gradient spanning approx. 300 m. Species richness and abundance of pollinators increased with elevation, but no shift between hoverflies and wild bees (mainly bumblebees) occurred. Seedset of the woodland cranesbill (Geranium sylvaticum) increased with hoverfly abundance, and seedset of the marsh thistle (Cirsium palustre) increased with wild bee abundance. Black rampion (Phyteuma nigrum) showed no significant response. The assignment of specific pollinator communities, and their response to altitude in highlands, to different plant species underlines the importance of wild bees and hoverflies as pollinators in extensive grassland systems.

Convergent evolution of specialized generalists: Implications for phylogenetic and functional diversity of carabid feeding groups.

Closely related species are often assumed to be functionally similar. Phylogenetic information is thus widely used to infer functional diversity and assembly of communities. In contrast, evolutionary processes generating functional similarity of phylogenetically distinct taxa are rarely addressed in this context.To investigate the impact of convergent evolution on functional diversity (FD) and phylogenetic diversity (PD), we reconstructed the phylogenetic structure of carabid trophic groups. We then analyzed the mandible shapes using geometric morphometrics to link specialization in functional morphology with feeding specialization among herbivores, generalist carnivores, and specialized consumers of Collembola.Our results show that carabid feeding groups are paraphyletic. Herbivory evolved at least twice and specialization to Collembola predation at least three times. Species within feeding groups share a remarkably similar mandible morphology, which evolved convergently. While specialized mandibles of herbivores and collembolan specialists represent an adaptation to their main food source, the particular mandible morphologies do not necessarily reflect the degree of food specialization within feeding groups. Only a few species with a specialized herbivorous mandible may occasionally feed on animals, but the range of specific food resources in generalist carnivore species is large, despite an almost identical mandible shape.Thus, convergent evolution in specialized feeding groups reverses the relationship between PD and functional similarity compared with generalist carnivores. We conclude that phylogenetic relationship is a poor proxy of FD in carabids. Moreover, the inconsistencies between relatedness, morphological adaptation, and ecological function require caution in the characterization of functional groups. Rather than assuming general relationships between PD and FD, we suggest integrating the analysis of evolutionary processes into functional community analyses.

The interplay of landscape composition and configuration: new pathways to manage functional biodiversity and agroecosystem services across Europe.

Managing agricultural landscapes to support biodiversity and ecosystem services is a key aim of a sustainable agriculture. However, how the spatial arrangement of crop fields and other habitats in landscapes impacts arthropods and their functions is poorly known. Synthesising data from 49 studies (1515 landscapes) across Europe, we examined effects of landscape composition (% habitats) and configuration (edge density) on arthropods in fields and their margins, pest control, pollination and yields. Configuration effects interacted with the proportions of crop and non-crop habitats, and species' dietary, dispersal and overwintering traits led to contrasting responses to landscape variables. Overall, however, in landscapes with high edge density, 70% of pollinator and 44% of natural enemy species reached highest abundances and pollination and pest control improved 1.7- and 1.4-fold respectively. Arable-dominated landscapes with high edge densities achieved high yields. This suggests that enhancing edge density in European agroecosystems can promote functional biodiversity and yield-enhancing ecosystem services.

Partitioning wild bee and hoverfly contributions to plant-pollinator network structure in fragmented habitats.

The risk of ecosystem function degradation with biodiversity loss has emerged as a major scientific concern in recent years. Possible relationships between taxonomic diversity and magnitude and stability of ecosystem processes build upon species' functional characteristics, which determine both susceptibility to environmental change and contribution to ecosystem properties. The functional diversity within communities thus provides a potential buffer against environmental disturbance, especially for properties emerging from interactions among species. In complex plant-pollinator networks, distantly related taxa spanning a great trait diversity shape network architecture. Here, we address the question of whether network properties are maintained after habitat loss by complementary contributions of phylogenetically distant pollinator taxa. We quantified contributions of wild bees and hoverflies to network structure (connectance, network specialization, specialization asymmetry) in 32 calcareous grassland fragments varying in size. Although hoverflies are often regarded less susceptible to environmental change than wild bees, species richness of both taxa was similarly affected by habitat loss. The associated loss of 80% of interactions resulted in small and tightly connected networks, which was more strongly attributed to wild bee loss than hoverfly loss. Networks in small fragments were less specialized due to equivalent losses of species and interactions in both pollinators and plants. Because wild bee and hoverfly loss contributed similarly to declining network specialization, we conclude that trait diversity among distantly related pollinators does not necessarily provide insurance against functional homogenization during community disassembly following habitat loss.

Past and potential future effects of habitat fragmentation on structure and stability of plant-pollinator and host-parasitoid networks.

Habitat fragmentation is a primary threat to biodiversity, but how it affects the structure and stability of ecological networks is poorly understood. Here, we studied plant-pollinator and host-parasitoid networks on 32 calcareous grassland fragments covering a size gradient of several orders of magnitude and with amounts of additional habitat availability in the surrounding landscape that varied independent of fragment size. We find that additive and interactive effects of habitat fragmentation at local (fragment size) and landscape scales (1,750 m radius) directly shape species communities by altering the number of interacting species and, indirectly, their body size composition. These, in turn, affect plant-pollinator, but not host-parasitoid, network structure: the nestedness and modularity of plant-pollinator networks increase with pollinator body size. Moreover, pollinator richness increases modularity. In contrast, the modularity of host-parasitoid networks decreases with host richness, whereas neither parasitoid richness nor body size affects network structure. Simulating species coextinctions also reveals that the structure-stability relationship depends on species' sensitivity to coextinctions and their capacity for adaptive partner switches, which differ between mutualistic and antagonistic interaction partners. While plant-pollinator communities may cope with future habitat fragmentation by responding to species loss with opportunistic partner switches, past effects of fragmentation on the current structure of host-parasitoid networks may strongly affect their robustness to coextinctions under future habitat fragmentation.

Wild pollinators enhance oilseed rape yield in small-holder farming systems in China.

BACKGROUND: Insect pollinators play an important role in crop pollination, but the relative contribution of wild pollinators and honey bees to pollination is currently under debate. There is virtually no information available on the strength of pollination services and the identity of pollination service providers from Asian smallholder farming systems, where fields are small, and variation among fields is high. We established 18 winter oilseed rape (Brassica napus L.) fields along a large geographical gradient in Jiangxi province in China. In each field, oilseed rape plants were grown in closed cages that excluded pollinators and open cages that allowed pollinator access. The pollinator community was sampled by pan traps for the entire oilseed rape blooming period. RESULTS: Oilseed rape plants from which insect pollinators were excluded had on average 38% lower seed set, 17% lower fruit set and 12% lower yield per plant, but the seeds were 17% heavier, and the caged plants had 28% more flowers and 18% higher aboveground vegetative biomass than plants with pollinator access. Oilseed rape plants thus compensate for pollination deficit by producing heavier seeds and more flowers. Regression analysis indicated that local abundance and diversity of wild pollinators were positively associated with seed set and yield/straw ratio, while honey bee abundance was not related to yield parameters. CONCLUSIONS: Wild pollinator abundance and diversity contribute to oilseed rape yield by enhancing plant resource allocation to seeds rather than to above-ground biomass. This study highlights the importance of the conservation of wild pollinators to support oilseed rape production in small-holder farming systems in China.

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.

Delivery of crop pollination services is an insufficient argument for wild pollinator conservation.

There is compelling evidence that more diverse ecosystems deliver greater benefits to people, and these ecosystem services have become a key argument for biodiversity conservation. However, it is unclear how much biodiversity is needed to deliver ecosystem services in a cost-effective way. Here we show that, while the contribution of wild bees to crop production is significant, service delivery is restricted to a limited subset of all known bee species. Across crops, years and biogeographical regions, crop-visiting wild bee communities are dominated by a small number of common species, and threatened species are rarely observed on crops. Dominant crop pollinators persist under agricultural expansion and many are easily enhanced by simple conservation measures, suggesting that cost-effective management strategies to promote crop pollination should target a different set of species than management strategies to promote threatened bees. Conserving the biological diversity of bees therefore requires more than just ecosystem-service-based arguments.

Hover flies are efficient pollinators of oilseed rape.

Understanding the consequences of declining diversity and abundance of pollinators for crops and floral biodiversity is a major challenge for current conservation ecology. However, most studies on this issue focus on bees, while other invertebrate taxa are largely ignored. We investigated the pollination efficiency of the globally abundant hover fly Episyrphus balteatus on the common crop, oilseed rape (Brassica napus). The study was conducted over a period of 2 consecutive years by means of enclosure experiments at an agricultural site located in Central Hesse (Germany). E. balteatus significantly increased both seed set and yield. This effect was very constant in the 2 years, despite considerable interannual differences in total seed numbers and seed mass. It highlights the important role of hover flies as pollinators of arable crops under varying environmental conditions. In contrast to bees, the effect of E. balteatus was lower at high pollinator densities than at low pollinator densities. This suggests adverse effects of density-dependent factors on pollination efficiency at high densities. Thus, models ignoring the modulating effect of biotic interactions by generally assuming a simple positive relationship between pollinator density and pollination efficiency might not apply to a vital component of the pollinator community.