Biodiversity and pollination benefits trade off against profit in an intensive farming system.

Agricultural expansion and intensification have boosted global food production but have come at the cost of environmental degradation and biodiversity loss. Biodiversity-friendly farming that boosts ecosystem services, such as pollination and natural pest control, is widely being advocated to maintain and improve agricultural productivity while safeguarding biodiversity. A vast body of evidence showing the agronomic benefits of enhanced ecosystem service delivery represent important incentives to adopt practices enhancing biodiversity. However, the costs of biodiversity-friendly management are rarely taken into account and may represent a major barrier impeding uptake by farmers. Whether and how biodiversity conservation, ecosystem service delivery, and farm profit can go hand in hand is unknown. Here, we quantify the ecological, agronomic, and net economic benefits of biodiversity-friendly farming in an intensive grassland-sunflower system in Southwest France. We found that reducing land-use intensity on agricultural grasslands drastically enhances flower availability and wild bee diversity, including rare species. Biodiversity-friendly management on grasslands furthermore resulted in an up to 17% higher revenue on neighboring sunflower fields through positive effects on pollination service delivery. However, the opportunity costs of reduced grassland forage yields consistently exceeded the economic benefits of enhanced sunflower pollination. Our results highlight that profitability is often a key constraint hampering adoption of biodiversity-based farming and uptake critically depends on society's willingness to pay for associated delivery of public goods such as biodiversity.

Global patterns in bumble bee pollen collection show phylogenetic conservation of diet.

Bumble bees (Bombus) are a group of eusocial bees with a strongly generalised feeding pattern, collecting pollen from many different botanical families. Though predominantly generalists, some bumble bee species seem to have restricted dietary choices. It is unclear whether restricted diets in bumble bees are inherent or a function of local conditions due to a lack of data for many species across different regions. The objective of this study was to determine whether bumble bee species displayed specific patterns of pollen collection, and whether patterns were influenced by phylogenetic relatedness or tongue length, a trait known to be associated with structuring floral visitation. Bumble bee pollen collection patterns were quantified from 4,132 pollen loads taken from 58 bumble bee species, representing 24% of the pollen-collecting diversity of this genus. Phylogenetic trait mapping showed a conserved pattern of dietary dissimilarity across species, but not for dietary breadth. Dietary dissimilarity was driven by collection of Fabaceae, with the most similar species collecting around 50%-60% of their diet from this botanical family. The proportion of the diet collected from Fabaceae also showed a conserved phylogenetic signal. Greater collection of Fabaceae was associated with longer tongue lengths, with shorter tongued species focusing on alternative botanical families. However, this result was largely driven by phylogenetic relatedness, not tongue length per se. These results demonstrate that, though generalists, bumble bees are still subject to dietary restrictions that constrain their foraging choices. These dietary constraints have implications for their persistence should their core resources decline in abundance.

Feeding specialization and longer generation time are associated with relatively larger brains in bees.

Despite their miniature brains, insects exhibit substantial variation in brain size. Although the functional significance of this variation is increasingly recognized, research on whether differences in insect brain sizes are mainly the result of constraints or selective pressures has hardly been performed. Here, we address this gap by combining prospective and retrospective phylogenetic-based analyses of brain size for a major insect group, bees (superfamily Apoidea). Using a brain dataset of 93 species from North America and Europe, we found that body size was the single best predictor of brain size in bees. However, the analyses also revealed that substantial variation in brain size remained even when adjusting for body size. We consequently asked whether such variation in relative brain size might be explained by adaptive hypotheses. We found that ecologically specialized species with single generations have larger brains-relative to their body size-than generalist or multi-generation species, but we did not find an effect of sociality on relative brain size. Phylogenetic reconstruction further supported the existence of different adaptive optima for relative brain size in lineages differing in feeding specialization and reproductive strategy. Our findings shed new light on the evolution of the insect brain, highlighting the importance of ecological pressures over social factors and suggesting that these pressures are different from those previously found to influence brain evolution in other taxa.

Effects of landscape complexity on pollinators are moderated by pollinators' association with mass-flowering crops.

Conserving and restoring semi-natural habitat, i.e. enhancing landscape complexity, is one of the main strategies to mitigate pollinator decline in agricultural landscapes. However, we still have limited understanding of how landscape complexity shapes pollinator communities in both crop and non-crop habitat, and whether pollinator responses to landscape complexity vary with their association with mass-flowering crops. Here, we surveyed pollinator communities on mass-flowering leek crops and in nearby semi-natural habitat in landscapes of varying complexity. Surveys were done before and during crop bloom and distinguished between pollinators that visit the crop frequently (dominant), occasionally (opportunistic), or not at all (non-crop). Forty-seven per cent of the species in the wider landscape were also observed on leek flowers. Crop pollinator richness increased with local pollinator community size and increasing landscape complexity, but relationships were stronger for opportunistic than for dominant crop pollinators. Relationships between pollinator richness in semi-natural habitats and landscape complexity differed between groups with the most pronounced positive effects on non-crop pollinators. Our results indicate that while dominant crop pollinators are core components of crop pollinator communities in all agricultural landscapes, opportunistic crop pollinators largely determine species-richness responses and complex landscapes are local hotspots for both biodiversity conservation and potential ecosystem service provision.

Insect pollination is at least as important for marketable crop yield as plant quality in a seed crop.

The sustainability of agriculture can be improved by integrating management of ecosystem services, such as insect pollination, into farming practices. However, large-scale adoption of ecosystem services-based practices in agriculture is lacking, possibly because growers undervalue the benefits of ecosystem services compared to those of conventional management practices. Here we show that, under representative real-world conditions, pollination and plant quality made similar contributions to marketable seed yield of hybrid leek (Allium porrum). Relative to the median, a 25% improvement of plant quality and pollination increased crop value by an estimated $18 007 and $17 174 ha-1 respectively. Across five crop lines, bumblebees delivered most pollination services, while other wild pollinator groups made less frequent but nevertheless substantial contributions. Honeybees actively managed for pollination services did not make significant contributions. Our results show that wild pollinators are an undervalued agricultural input and managing for enhancing pollinators makes sense economically in high-revenue insect-pollinated cropping systems.

A retrospective analysis of pollen host plant use by stable and declining bumble bee species.

Understanding population declines has been the objective of a wide range of ecological studies. When species have become rare such studies are complicated because particular behavior or life history traits may be the cause but also the result of the decline of a species. We approached this problem by studying species' characteristics on specimens that were collected before the onset of their decline and preserved in natural history museums. In northwestern Europe, some bumble bee species declined dramatically during the 20th century whereas other, ecologically similar, species maintained stable populations. A long-standing debate focuses on whether this is caused by declining species having stricter host plant preferences. We compared the composition of pollen loads of five bumble bee species with stable populations and five with declining populations using museum specimens collected before 1950 in Belgium, England, and The Netherlands. Prior to 1950, the number of plant taxa in pollen loads of declining species was almost one-third lower than that in stable species even though individuals of stable and declining species generally originated from the same areas. There were no systematic differences in the composition of pollen loads between stable and declining species, but the plant taxa preferred by declining species before 1950 had experienced a stronger decline in the 20th century than those preferred by stable species. In 2004 and 2005, we surveyed the areas where bumble bees had been caught in the past and compared the composition of past and present pollen loads of the stable, but not of the by now locally extinct declining species. The number of collected pollen taxa was similar, but the composition differed significantly between the two periods. Differences in composition reflected the major changes in land use in northwestern Europe but also the spread of the invasive plant species Impatiens glandulifera. The main question now is why declining species apparently were not able to switch to less preferred food plants when stable species were. This study shows that natural history collections can play an important role in improving our understanding of the ecological mechanisms driving species population change.

Arthropod assemblages are best predicted by plant species composition.

Insects and spiders comprise more than two-thirds of the Earth's total species diversity. There is wide concern, however, that the global diversity of arthropods may be declining even more rapidly than the diversity of vertebrates and plants. For adequate conservation planning, ecologists need to understand the driving factors for arthropod communities and devise methods, that provide reliable predictions when resources do not permit exhaustive ground surveys. Which factor most successfully predicts arthropod community structure is still a matter of debate, however. The purpose of this study was to identify the factor best predicting arthropod assemblage composition. We investigated the species composition of seven functionally different arthropod groups (epigeic spiders, grasshoppers, ground beetles, weevils, hoppers, hoverflies, and bees) at 47 sites in The Netherlands comprising a range of seminatural grassland types and one heathland type. We then compared the actual arthropod composition with predictions based on plant species composition, vegetation structure, environmental data, flower richness, and landscape composition. For this we used the recently published method of predictive co-correspondence analysis, and a predictive variant of canonical correspondence analysis, depending on the type of predictor data. Our results demonstrate that local plant species composition is the most effective predictor of arthropod assemblage composition, for all investigated groups. In predicting arthropod assemblages, plant community composition consistently outperforms both vegetation structure and environmental conditions (even when the two are combined), and also performs better than the surrounding landscape. These results run against a common expectation of vegetation structure as the decisive factor. Such expectations, however, have always been biased by the fact that until recently no methods existed that could use an entire (plant) species composition in the explanatory role. Although more recent experimental diversity work has reawakened interest in the role of plant species, these studies still have not used (or have not been able to use) entire species compositions. They only consider diversity measures, both for plant and insect assemblages, which may obscure relationships. The present study demonstrates that the species compositions of insect and plant communities are clearly linked.

First host record of Myopa pellucida Robineau-Desvoidy (Diptera: Conopidae) identified using DNA barcoding.

Thick-headed flies of the genus Myopa Fabricius are a common sight in early spring in Europe. Several species can be found flying among Salix catkins together with their supposed Andrena Fabricius hosts (Hymenoptera: Apidae) (John Smit observations). Despite the fact that some of these Myopa species are very common, little is known about their actual host-parasitoid associations. Only a few scattered host records can be found in the literature, most of which refer to M. testacea s.l.. Whether or not these records actually concern M. testacea (Linnaeus) is uncertain due to the confusion over the identity of several species within the M. testacea species group (Stuke Clements 2008). Stuke (2017), in his World Catalogue of Conopids, provides all recorded host associations and distinguishes between unambiguously accepted host records and doubtful records based on three criteria: 1) the conopid has been reared from the host or a larva has been identified by DNA barcoding; 2) there is no doubt concerning the identity of the conopid species concerned, and 3) there is no doubt concerning the identity of the host species. Because the vast majority of the host records in the literature do not fulfil one or more of these criteria they are regarded as doubtful (Stuke 2017). Thus, only three host associations for the genus Myopa can be regarded as confirmed: Andrena vaga Panzer, for both M. hirsuta Stuke Clements (Jentzsch 2009) and M. testacea (Erteld 1998, Fellendorf et al. 2004, De Meijere 1912), and the Nearctic Andrena regularis Malloch, for the Holarctic M. vicaria Walker (Miliczky Osgood 1995). Here we report on a fourth host association for the genus Myopa, since a second instar larva of M. pellucida Robineau-Desvoidy was found in the abdomen of an Andrena nitida (Müller) female and identified using DNA barcoding.