The root microlandscape of arbuscular mycorrhizal fungi.

Understanding the drivers of assemblages of arbuscular mycorrhizal fungi (AMF) is essential to leverage the benefits of AMF for plant growth and health. Arbuscular mycorrhizal fungi are heterogeneously distributed in space even at small scale. We review the role of plant distribution in driving AMF assemblages (the passenger hypothesis), using a transposition of the conceptual framework of landscape ecology. Because rooting systems correspond to habitat patches with limited carrying capacity that differ in quality due to host-preference effects, we suggest considering plant communities as mosaics of AMF microhabitats. We review how predictions from landscape ecology apply to plant community effects on AMF, and the existing evidence that tests these predictions. Although many studies have been conducted on the effect of plant compositional heterogeneity on AMF assemblages, they mostly focused on the effect of plant richness, while only a few investigated the effect of configurational heterogeneity, plant connectivity or plant community temporal dynamics. We propose key predictions and future prospects to fill these gaps. Considering plant communities as landscapes extends the passenger hypothesis by including a spatially explicit dimension and its associated ecological processes and may help understand and manipulate AMF assemblages at small spatial scales.

Landscape effects on the thermotolerance of carabid beetles and the role of behavioral thermoregulation.

Physiological thermotolerance and behavioral thermoregulation are central to seasonal cold adaptation in ectothermic organisms. For species with enhanced mobility, behavioral responses may be of greater importance in the cold stress response. Employing the carabid beetles as a study organism, the current study compared physiological thermotolerance and behavioral thermoregulation in carabid species inhabiting cereal fields in different landscape contexts, from fine grain heterogeneous "complex" landscapes to homogenous "simple" landscapes. Physiological thermotolerance was determined via measurement of the CTmin and chill coma temperature. Behavioral responses to cold temperature exposure were determined employing a purpose built arena, and thoracic temperature measured to estimate the efficacy of the behavior as a form of behavioral thermoregulation. Results revealed an influence of landscape composition on the cold tolerance of carabid beetles, although species differed in their sensitivity to landscape intensification. A reduced effect of landscape on the thermotolerance of larger carabid beetles was observed, thought to be the consequence of greater mobility preventing local acclimation to microclimatic variation along the landscape intensification gradient. Investigation into behavioral thermoregulation of the 3 largest species revealed burrowing behavior to be the main behavioral response to cold stress, acting to significantly raise carabid body temperature. This finding highlights the importance of behavioral thermoregulation as a strategy to evade cold stress. The use of behavioral thermoregulation may negate the need to invest in physiological thermotolerance, further offering explanation for the lack of landscape effect on the physiological thermotolerance of larger carabids.

Multi-scale approach to biodiversity proxies of biological control service in European farmlands.

Intensive agriculture has profoundly altered biodiversity and trophic relationships in agricultural landscapes, leading to the deterioration of many ecosystem services such as pollination or biological control. Information on which spatio-temporal factors are simultaneously affecting crop pests and their natural enemies is required to improve conservation biological control practices. We conducted a study in 80 winter wheat crop fields distributed in three regions of North-western Europe (Brittany, Hauts-de-France and Wallonia), along intra-regional gradients of landscape complexity. Five taxa of major crop pests (aphids and slugs) and natural enemies (spiders, carabids, and parasitoids) were sampled three times a year, for two consecutive years. We analysed the influence of regional (meteorology), landscape (structure in both the years n and n-1) and local factors (hedge or grass strip field boundaries, and distance to boundary) on the abundance and species richness of crop-dwelling organisms, as proxies of the service/disservice they provide. Firstly, there was higher biocontrol potential in areas with mild winter climatic conditions. Secondly, natural enemy communities were less diverse and had lower abundances in landscapes with high crop and wooded continuities (sum of interconnected crop or wood surfaces), contrary to slugs and aphids. Finally, field boundaries with grass strips were more favourable to spiders and carabids than boundaries formed by hedges, while the opposite was found for crop pests, with the latter being less abundant towards the centre of the fields. We also revealed temporal modulation-and sometimes reversion-of the impact of local elements on crop biodiversity. To some extent, these results cause controversy because they show that hedgerows and woodlots should not be the unique cornerstones of agro-ecological landscape design strategies. We point out that combining woody and grassy habitats to take full advantage of the features and ecosystem services they both provide (biological pest control, windbreak effect, soil stabilization) may promote sustainable agricultural ecosystems. It may be possible to both reduce pest pressure and promote natural enemies by accounting for taxa-specific antagonistic responses to multi-scale environmental characteristics.

Increasing crop heterogeneity enhances multitrophic diversity across agricultural regions.

Agricultural landscape homogenization has detrimental effects on biodiversity and key ecosystem services. Increasing agricultural landscape heterogeneity by increasing seminatural cover can help to mitigate biodiversity loss. However, the amount of seminatural cover is generally low and difficult to increase in many intensively managed agricultural landscapes. We hypothesized that increasing the heterogeneity of the crop mosaic itself (hereafter "crop heterogeneity") can also have positive effects on biodiversity. In 8 contrasting regions of Europe and North America, we selected 435 landscapes along independent gradients of crop diversity and mean field size. Within each landscape, we selected 3 sampling sites in 1, 2, or 3 crop types. We sampled 7 taxa (plants, bees, butterflies, hoverflies, carabids, spiders, and birds) and calculated a synthetic index of multitrophic diversity at the landscape level. Increasing crop heterogeneity was more beneficial for multitrophic diversity than increasing seminatural cover. For instance, the effect of decreasing mean field size from 5 to 2.8 ha was as strong as the effect of increasing seminatural cover from 0.5 to 11%. Decreasing mean field size benefited multitrophic diversity even in the absence of seminatural vegetation between fields. Increasing the number of crop types sampled had a positive effect on landscape-level multitrophic diversity. However, the effect of increasing crop diversity in the landscape surrounding fields sampled depended on the amount of seminatural cover. Our study provides large-scale, multitrophic, cross-regional evidence that increasing crop heterogeneity can be an effective way to increase biodiversity in agricultural landscapes without taking land out of agricultural production.

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.

Response of wild bee diversity, abundance, and functional traits to vineyard inter-row management intensity and landscape diversity across Europe.

Agricultural intensification is a major driver of wild bee decline. Vineyards may be inhabited by plant and animal species, especially when the inter-row space is vegetated with spontaneous vegetation or cover crops. Wild bees depend on floral resources and suitable nesting sites which may be found in vineyard inter-rows or in viticultural landscapes. Inter-row vegetation is managed by mulching, tillage, and/or herbicide application and results in habitat degradation when applied intensively. Here, we hypothesize that lower vegetation management intensities, higher floral resources, and landscape diversity affect wild bee diversity and abundance dependent on their functional traits. We sampled wild bees semi-quantitatively in 63 vineyards representing different vegetation management intensities across Europe in 2016. A proxy for floral resource availability was based on visual flower cover estimations. Management intensity was assessed by vegetation cover (%) twice a year per vineyard. The Shannon Landscape Diversity Index was used as a proxy for landscape diversity within a 750 m radius around each vineyard center point. Wild bee communities were clustered by country. At the country level, between 20 and 64 wild bee species were identified. Increased floral resource availability and extensive vegetation management both affected wild bee diversity and abundance in vineyards strongly positively. Increased landscape diversity had a small positive effect on wild bee diversity but compensated for the negative effect of low floral resource availability by increasing eusocial bee abundance. We conclude that wild bee diversity and abundance in vineyards is efficiently promoted by increasing floral resources and reducing vegetation management frequency. High landscape diversity further compensates for low floral resources in vineyards and increases pollinating insect abundance in viticulture landscapes.

Land use, settlement, and plant diversity in Iron Age Northwest France.

Various studies using pollen stratigraphies have demonstrated significant correlations between Holocene plant diversity, climate, and human activities. Studies that have analyzed longer Holocene timescales tend to discuss cultural data very superficially. This is remarkable because detailed insights into past human activities may be key to gain an understanding of the observed trends in biodiversity. This study aims to reconstruct and explain spatio-temporal trends in past plant diversity (alpha, temporal, and spatial beta diversity) by integrating data on vegetation dynamics, human subsistence economy, and land-use patterns. The landscape of Northwest France during the greater part of the Iron Age and the start of the Roman period (600 BC-AD 100) is selected as a case study. In total, 30 high-quality pollen-stratigraphical sequences allow for the reconstruction of the main long-term trends in plant diversity and more generally of the changing fabric of the landscape. Additionally, increasingly detailed images of the Iron Age rural landscape are available because of a steep increase in archaeological data (aerial photography, surveys, and excavations). These different types of data are integrated and used as input for a wider discussion on the relation between human activities and plant diversity. In general, the taxonomic richness increases steadily during the period under study. Some spatio-temporal differences are observed. The increasing richness values correspond with the growing impact of human activities on the landscape. Archaeologically documented land-use changes on smaller timescales are less clearly reflected in the richness values and vegetation dynamics, which might result from the (large-scale) research design.

Author Correction: Soil biota in vineyards are more influenced by plants and soil quality than by tillage intensity or the surrounding landscape.

A correction to this article has been published and is linked from the HTML and PDF versions of this paper. The error has been fixed in the paper.

Landscape configurational heterogeneity by small-scale agriculture, not crop diversity, maintains pollinators and plant reproduction in western Europe.

Agricultural intensification is one of the main causes for the current biodiversity crisis. While reversing habitat loss on agricultural land is challenging, increasing the farmland configurational heterogeneity (higher field border density) and farmland compositional heterogeneity (higher crop diversity) has been proposed to counteract some habitat loss. Here, we tested whether increased farmland configurational and compositional heterogeneity promote wild pollinators and plant reproduction in 229 landscapes located in four major western European agricultural regions. High-field border density consistently increased wild bee abundance and seed set of radish (Raphanus sativus), probably through enhanced connectivity. In particular, we demonstrate the importance of crop-crop borders for pollinator movement as an additional experiment showed higher transfer of a pollen analogue along crop-crop borders than across fields or along semi-natural crop borders. By contrast, high crop diversity reduced bee abundance, probably due to an increase of crop types with particularly intensive management. This highlights the importance of crop identity when higher crop diversity is promoted. Our results show that small-scale agricultural systems can boost pollinators and plant reproduction. Agri-environmental policies should therefore aim to halt and reverse the current trend of increasing field sizes and to reduce the amount of crop types with particularly intensive management.

The effect of landscape complexity and microclimate on the thermal tolerance of a pest insect.

Landscape changes are known to exacerbate the impacts of climate change. As such, understanding the combined effect of climate and landscape on agroecosystems is vital if we are to maintain the function of agroecosystems. This study aimed to elucidate the effects of agricultural landscape complexity on the microclimate and thermal tolerance of an aphid pest to better understand how landscape and climate may interact to affect the thermal tolerance of pest species within the context of global climate change. Meteorological data were measured at the landscape level, and cereal aphids (Sitobion avenae, Metopolophium dirhodum and Rhopalosiphum padi) sampled, from contrasting landscapes (simple and complex) in winter 2013/2014 and spring 2014 in cereal fields of Brittany, France. Aphids were returned to the laboratory and the effect of landscape of origin on aphid cold tolerance (as determined by CTmin ) was investigated. Results revealed that local landscape complexity significantly affected microclimate, with simple homogenous landscapes being on average warmer, but with greater temperature variation. Landscape complexity was shown to impact aphid cold tolerance, with aphids from complex landscapes being more cold tolerant than those from simple landscapes in both winter and spring, but with differences among species. This study highlights that future changes to land use could have implications for the thermal tolerance and adaptability of insects. Furthermore, not all insect species respond in a similar way to microhabitat and microclimate, which could disrupt important predator-prey relationships and the ecosystem service they provide.

Soil biota in vineyards are more influenced by plants and soil quality than by tillage intensity or the surrounding landscape.

Tillage is known for its adverse effects on soil biota, at least in arable agroecosystems. However, in vineyards effects might differ as tillage is often performed during dry periods or only in every other inter-row allowing species to re-colonise disturbed areas. We examined the response of earthworms (lumbricids), springtails (collembola) and litter decomposition to periodically mechanically disturbed (PMD) and permanently green covered (PGC) vineyard inter-rows and assessed whether site effects are altered by the surrounding landscape. In commercial vineyards in Austria we sampled earthworms by handsorting, springtails by soil coring and pitfall trapping and installed litter decomposition bags. Earthworm species diversity increased with plant biomass under PMD but not under PGC; earthworm density was unaffected by tillage but increased with plant biomass mainly at high soil quality (soil fertility index). Springtail species diversity was unaffected by tillage; springtail densities (mainly larger species) were reduced under PGC. Litter decomposition was little affected by investigated parameters. Landscape heterogeneity affected the functional diversity of surface springtails, but did not influence soil-dwelling springtails, earthworms or litter decomposition. We conclude that effects on soil biota of periodical tillage in vineyards need not necessarily be detrimental and will be modified by plant biomass and soil quality.

Why is there no impact of the host species on the cold tolerance of a generalist parasitoid?

For generalist parasitoids such as those belonging to the Genus Aphidius, the choice of host species can have profound implications for the emerging parasitoid. Host species is known to affect a variety of life history traits. However, the impact of the host on thermal tolerance has never been studied. Physiological thermal tolerance, enabling survival at unfavourable temperatures, is not a fixed trait and may be influenced by a number of external factors including characteristics of the stress, of the individual exposed to the stress, and of the biological and physical environment. As such, the choice of host species is likely to also have implications for the thermal tolerance of the emerging parasitoid. The current study aimed to investigate the effect of cereal aphid host species (Sitobion avenae, Rhopalosiphum padi and Metopolophium dirhodum) on adult thermal tolerance, in addition to sex and size, of the aphid parasitoids Aphidius avenae, Aphidius matricariae and Aphidius rhopalosiphi. Results revealed no effect of host species on the cold tolerance of the emerging parasitoid, as determined by CTmin and Chill Coma, for all parasitoid species. Host species significantly affected the size of the emerging parasitoid for A. rhopalosiphi only, with individuals emerging from R. padi being significantly larger than those emerging from S. avenae, although this did not correspond to a difference in thermal tolerance. Furthermore, a significant difference in the size of male and female parasitoids was observed for A. avenae and A. matricariae, although, once again this did not correspond to a difference in cold tolerance. It is suggested that potential behavioural thermoregulation via host manipulation may act to influence the thermal environment experienced by the wasp and thus wasp thermal tolerance and, in doing so, may negate physiological thermal tolerance or any impact of the aphid host.

Potential Host Manipulation by the Aphid Parasitoid Aphidius avenae to Enhance Cold Tolerance.

During parasitoid development, the immature parasitoid is confined to the host species. As a result, any potential to modify the physiology or behaviour of the host could play an important role in parasitoid fitness. The potential for host manipulation by the aphid parasitoid Aphidius avenae to increase cold thermotolerance was investigated using the aphid host species Metopolophium dirhodum and Sitobion avenae. Aphids were parasitized at L3/L4 instar stage (5 d old) and allowed to develop into pre-reproductive adults (10 d old) containing a 5 d old parasitoid larva. A control group was created of non-parasitized pre-reproductive adults (10 d old). The inherent physiological thermotolerance (LT50) and potential behavioural thermoregulation (behaviour in a declining temperature regime) of parasitized and non-parasitized aphids were investigated. Results revealed no effect of parasitism on the physiological thermotolerance of S. avenae and M. dirhodum. Significant differences in the behaviour of parasitized and non-parasitized aphids were observed, in addition to differences between host species, and such behaviours are discussed in view of the potential for host manipulation.

Could behaviour and not physiological thermal tolerance determine winter survival of aphids in cereal fields?

Traits of physiological thermotolerance are commonly measured in the laboratory as predictors of the field success of ectotherms at unfavourable temperatures (e.g. during harsh winters, heatwaves, or under conditions of predicted global warming). Due to being more complicated to measure, behavioural thermoregulation is less commonly studied, although both physiology and behaviour interact to explain the survival of ectotherms. The aphids Metopolophium dirhodum, Rhopalosiphum padi and Sitobion avenae are commercially important pests of temperate cereal crops. Although coexisting, these species markedly differ in winter success, with R. padi being the most abundant species during cold winters, followed by S. avenae and lastly M. dirhodum. To better understand the thermal physiology and behavioural factors contributing to differential winter success, the lethal temperature (physiological thermotolerance) and the behaviour of aphids in a declining temperature regime (behavioural thermotolerance) of these three species were investigated. Physiological thermotolerance significantly differed between the three species, with R. padi consistently the least cold tolerant and S. avenae the most cold tolerant. However, although the least cold tolerant of the study species, significantly more R. padi remained attached to the host plant at extreme sub-zero temperatures than S. avenae and M. dirhodum. Given the success of anholocyclic R. padi in harsh winters compared to its anholocyclic counterparts, this study illustrates that behavioural differences could be more important than physiological thermotolerance in explaining resistance to extreme temperatures. Furthermore it highlights that there is a danger to studying physiological thermotolerance in isolation when ascertaining risks of ectotherm invasions, the establishment potential of exotic species in glasshouses, or predicting species impacts under climate change scenarios.

Functional landscape heterogeneity and animal biodiversity in agricultural landscapes.

Biodiversity in agricultural landscapes can be increased with conversion of some production lands into 'more-natural'- unmanaged or extensively managed - lands. However, it remains unknown to what extent biodiversity can be enhanced by altering landscape pattern without reducing agricultural production. We propose a framework for this problem, considering separately compositional heterogeneity (the number and proportions of different cover types) and configurational heterogeneity (the spatial arrangement of cover types). Cover type classification and mapping is based on species requirements, such as feeding and nesting, resulting in measures of 'functional landscape heterogeneity'. We then identify three important questions: does biodiversity increase with (1) increasing heterogeneity of the more-natural areas, (2) increasing compositional heterogeneity of production cover types and (3) increasing configurational heterogeneity of production cover types? We discuss approaches for addressing these questions. Such studies should have high priority because biodiversity protection globally depends increasingly on maintaining biodiversity in human-dominated landscapes.

A hierarchical matrix model to assess the impact of habitat fragmentation on population dynamics: an elasticity analysis.

To better understand the role of habitat quality and boundaries on population dynamics at the landscape scale, we develop a model combining a spatially implicit approach, a spatial population Leslie-type model and an implicit model of habitat fragmentation. An original approach of elasticity permits to identify which types of element and boundary influence the most population viability according to the wood fragmentation degree. The studied species is a corridor forest insect sensitive to fragmentation (Abax parallelepipedus, Coleoptera, Carabidae). We show that a single large patch of wood is better than several small patches for the population viability.