Functional trait diversity across trophic levels determines herbivore impact on plant community biomass

Understanding the consequences of trophic interactions for ecosystem functioning is challenging, as contrasting effects of species and functional diversity can be expected across trophic levels. We experimentally manipulated functional identity and diversity of grassland insect herbivores and tested their impact on plant community biomass. Herbivore resource acquisition traits, i.e. mandible strength and the diversity of mandibular traits, had more important effects on plant biomass than body size. Higher herbivore functional diversity increased overall impact on plant biomass due to feeding niche complementarity. Higher plant functional diversity limited biomass pre‐emption by herbivores. The functional diversity within and across trophic levels therefore regulates the impact of functionally contrasting consumers on primary producers. By experimentally manipulating the functional diversity across trophic levels, our study illustrates how trait‐based approaches constitute a promising way to tackle existing links between trophic interactions and ecosystem functioning.     

Plant diversity induces shifts in the functional structure and diversity across trophic levels

Changes to primary producer diversity can cascade up to consumers and affect ecosystem processes. Although the effect of producer diversity on higher trophic groups have been studied, these studies often quantify taxonomy‐based measures of biodiversity, like species richness, which do not necessarily reflect the functioning of these communities. In this study, we assess how plant species richness affects the functional composition and diversity of higher trophic levels and discuss how this might affect ecosystem processes, such as herbivory, predation and decomposition. Based on six different consumer traits, we examined the functional composition of arthropod communities sampled in experimental plots that differed in plant species richness. The two components we focused on were functional variation in the consumer community structure (functional structure) and functional diversity, expressed as functional richness, evenness and divergence. We found a consistent positive effect of plant species richness on the functional richness of herbivores, carnivores, and omnivores, but not decomposers, and contrasting patterns for functional evenness and divergence. Increasing plant species richness shifted the omnivore community to more predatory and less mobile species, and the herbivore community to more specialized and smaller species. This was accompanied by a shift towards more species occurring in the vegetation than in the ground layer. Our study shows that plant species richness strongly affects the functional structure and diversity of aboveground arthropod communities. The observed shifts in body size (herbivores), specialization (herbivores), and feeding mode (omnivores) together with changes in the functional diversity may underlie previously observed increases in herbivory and predation in plant communities of higher diversity.     

On the prevalence and dynamics of inverted trophic pyramids and otherwise top‐heavy communities

Classically, biomass partitioning across trophic levels was thought to add up to a pyramidal distribution. Numerous exceptions have, however, been noted including complete pyramidal inversions. Elevated levels of biomass top‐heaviness (i.e. high consumer/resource biomass ratios) have been reported from Arctic tundra communities to Brazilian phytotelmata, and in species assemblages as diverse as those dominated by sharks and ants. We highlight two major pathways for creating top‐heaviness, via: (1) endogenous channels that enhance energy transfer across trophic boundaries within a community and (2) exogenous pathways that transfer energy into communities from across spatial and temporal boundaries. Consumer–resource models and allometric trophic network models combined with niche models reveal the nature of core mechanisms for promoting top‐heaviness. Outputs from these models suggest that top‐heavy communities can be stable, but they also reveal sources of instability. Humans are both increasing and decreasing top‐heaviness in nature with ecological consequences. Current and future research on the drivers of top‐heaviness can help elucidate fundamental mechanisms that shape the architecture of ecological communities and govern energy flux within and between communities. Questions emerging from the study of top‐heaviness also usefully draw attention to the incompleteness and inconsistency by which ecologists often establish definitional boundaries for communities.     

Land use intensification in grasslands: higher trophic levels are more negatively affected than lower trophic levels

Increasing land use intensity and human influence are leading to a reduction in plant and animal species diversity. However, little is known about how these changes may affect higher trophic levels, apart from simply reducing species numbers. Here we investigated, over 3 years, the influence of different land practices on a tritrophic system in grassland habitats. The system consisted of the host plant Plantago lanceolata L. (Plantaginaceae), two monophagous weevils, Mecinus labilis Herbst and Mecinus pascuorum Gyllenhal (Coleoptera: Curculionidae), and their parasitoid Mesopolobus incultus Walker (Hymenoptera: Pteromalidae). At over 70 sites across three geographic regions in Germany, we measured plant species diversity and vegetation structure, as well as abundance of P. lanceolata, the two weevils, and the parasitoid. Land use intensity (fertilization) and type (mowing vs. grazing) negatively affected not only plant species richness but also the occurrence of the two specialized herbivores and their parasitoid. In contrast, land use had a mostly positive effect on host plant size, vegetation structure, and parasitization rate. This study reveals that intensification of land use influences higher trophic organisms even without affecting the availability of the host plant. The observed relationships between land use, vegetation complexity, and the tritrophic system are not restricted locally; rather they are measureable along a broad range of environmental conditions and years throughout Germany. Our findings may have important implications for the conservation of insect species of nutrient‐poor grasslands.     

Species richness and trophic diversity increase decomposition in a co-evolved food web

Ecological communities show great variation in species richness, composition and food web structure across similar and diverse ecosystems. Knowledge of how this biodiversity relates to ecosystem functioning is important for understanding the maintenance of diversity and the potential effects of species losses and gains on ecosystems. While research often focuses on how variation in species richness influences ecosystem processes, assessing species richness in a food web context can provide further insight into the relationship between diversity and ecosystem functioning and elucidate potential mechanisms underpinning this relationship. Here, we assessed how species richness and trophic diversity affect decomposition rates in a complete aquatic food web: the five trophic level web that occurs within water-filled leaves of the northern pitcher plant, Sarracenia purpurea. We identified a trophic cascade in which top-predators--larvae of the pitcher-plant mosquito--indirectly increased bacterial decomposition by preying on bactivorous protozoa. Our data also revealed a facultative relationship in which larvae of the pitcher-plant midge increased bacterial decomposition by shredding detritus. These important interactions occur only in food webs with high trophic diversity, which in turn only occur in food webs with high species richness. We show that species richness and trophic diversity underlie strong linkages between food web structure and dynamics that influence ecosystem functioning. The importance of trophic diversity and species interactions in determining how biodiversity relates to ecosystem functioning suggests that simply focusing on species richness does not give a complete picture as to how ecosystems may change with the loss or gain of species.     

Traits across trophic levels interact to influence parasitoid establishment in biological control releases

A central goal in ecology is to predict what governs a species’ ability to establish in a new environment. One mechanism driving establishment success is individual species’ traits, but the role of trait combinations among interacting species across different trophic levels is less clear. Deliberate or accidental species additions to existing communities provide opportunities to study larger scale patterns of establishment success. Biological control introductions are especially valuable because they contain data on both the successfully established and unestablished species. Here, we used a recent dataset of importation biological control introductions to explore how life‐history traits of 132 parasitoid species and their herbivorous hosts interact to affect parasitoid establishment. We find that of five parasitoid and herbivore traits investigated, one parasitoid trait—host range—weakly predicts parasitoid establishment; parasitoids with higher levels of phylogenetic specialization have higher establishment success, though the effect is marginal. In addition, parasitoids are more likely to establish when their herbivore host has had a shorter residence time. Interestingly, we do not corroborate earlier findings that gregarious parasitoids and endoparasitoids are more likely to establish. Most importantly, we find that life‐history traits of the parasitoid species and their hosts can interact to influence establishment. Specifically, parasitoids with broader host ranges are more likely to establish when the herbivore they have been released to control is also more of a generalist. These results provide insight into how multiple species’ traits and their interactions, both within and across trophic levels, can influence establishment of species of higher trophic levels.     

Effects of trophic skewing of species richness on ecosystem functioning in a diverse marine community

Widespread overharvesting of top consumers of the world's ecosystems has "skewed" food webs, in terms of biomass and species richness, towards a generally greater domination at lower trophic levels. This skewing is exacerbated in locations where exotic species are predominantly low-trophic level consumers such as benthic macrophytes, detritivores, and filter feeders. However, in some systems where numerous exotic predators have been added, sometimes purposefully as in many freshwater systems, food webs are skewed in the opposite direction toward consumer dominance. Little is known about how such modifications to food web topology, e.g., changes in the ratio of predator to prey species richness, affect ecosystem functioning. We experimentally measured the effects of trophic skew on production in an estuarine food web by manipulating ratios of species richness across three trophic levels in experimental mesocosms. After 24 days, increasing macroalgal richness promoted both plant biomass and grazer abundance, although the positive effect on plant biomass disappeared in the presence of grazers. The strongest trophic cascade on the experimentally stocked macroalgae emerged in communities with a greater ratio of prey to predator richness (bottom-rich food webs), while stronger cascades on the accumulation of naturally colonizing algae (primarily microalgae with some early successional macroalgae that recruited and grew in the mesocosms) generally emerged in communities with greater predator to prey richness (the more top-rich food webs). These results suggest that trophic skewing of species richness and overall changes in food web topology can influence marine community structure and food web dynamics in complex ways, emphasizing the need for multitrophic approaches to understand the consequences of marine extinctions and invasions.     

Analysis of trophic interactions reveals highly plastic response to climate change in a tri-trophic High-Arctic ecosystem

As a response to current climate changes, individual species have changed various biological traits, illustrating an inherent phenotypic plasticity. However, as species are embedded in an ecological network characterised by multiple consumer–resource interactions, ecological mismatches are likely to arise when interacting species do not respond homogeneously. The approach of biological networks analysis calls for the use of structural equation modelling (SEM), a multidimensional analytical setup that has proven particularly useful for analysing multiple interactions across trophic levels. Here we apply SEM to a long-term dataset from a High-Arctic ecosystem to analyse how phenological responses across three trophic levels are coupled to snowmelt patterns and how changes may cascade through consumer–resource interactions. Specifically, the model included the effect of snowmelt on a High-Arctic tri-trophic system of flowers, insects and waders (Charadriiformes), with latent factors representing phenology (timing of life history events) and performance (abundance or reproduction success) for each trophic level. The effects derived from the model demonstrated that the time of snowmelt directly affected plant and arthropod phenology as well as the performance of all included trophic levels. Additionally, timing of snowmelt appeared to indirectly influence wader phenology as well as plant, arthropod and wader performance through effects on adjacent trophic levels and lagged effects. The results from the tri-trophic community presented here emphasise that effects of climate on species in consumer–resource systems may propagate through trophic levels.     

Plant diversity effects on arthropods and arthropod-dependent ecosystem functions in a biodiversity experiment

Biodiversity-ecosystem function experiments test how species diversity influences fundamental ecosystem processes. Historically, arthropod driven functions, such as herbivory and pest-control, have been thought to be influenced by direct and indirect associations among species. Although a number of studies have evaluated how plant diversity affects arthropod communities and arthropod-mediated ecosystem processes, it remains unclear whether diversity effects on arthropods are sufficiently consistent over time such that observed responses can be adequately predicted by classical hypotheses based on associational effects. By combining existing results from a long-term grassland biodiversity experiment (Jena Experiment) with new analyses, we evaluate the consistency of consumer responses within and across taxonomic, trophic, and trait-based (i.e. vertical stratification) groupings, and we consider which changes in arthropod community composition are associated with changes in consumer-mediated ecosystem functions.Overall, higher plant species richness supported more diverse and complex arthropod communities and this pattern was consistent across multiple years. Vegetation-associated arthropods responded more strongly to changes in plant species richness than ground-dwelling arthropods. Additionally, increases in plant species richness were associated with shifts in the species-abundance distributions for many, but not all taxa. For example, highly specialized consumers showed a decrease in dominance and an increase in the number of rare species with increasing plant species richness. Most ecosystem processes investigated responded to increases in plant species richness in the same way as the trophic group mediating the process, e.g. both herbivory and herbivore diversity increase with increasing plant species richness. In the Jena Experiment and other studies, inconsistencies between predictions based on classic hypotheses of associational effects and observed relationships between plant species richness and arthropod diversity likely reflect the influence of multi-trophic community dynamics and species functional trait distributions. Future research should focus on testing a broader array of mechanisms to unravel the biological processes underlying the biodiversity-ecosystem functioning relationships.     

Grassland harvesting alters ant community trophic structure: An isotopic study in tallgrass prairies

Disturbances have long been recognized as important forces for structuring natural communities but their effects on trophic structure are not well understood, particularly in terrestrial systems. This is in part because quantifying trophic linkages is a challenge, especially for small organisms with cryptic feeding behaviors such as insects, and often relies on conducting labor‐intensive feeding trials or extensive observations in the field. In this study, we used stable isotopes of carbon and nitrogen to examine how disturbance (annual biomass harvesting) in tallgrass prairies affected the trophic position, trophic range, and niche space of ants, a widespread grassland consumer. We hypothesized that biomass harvest would remove important food and nesting resources of insects thus affecting ant feeding relationships and trophic structure. We found shifts in the feeding relationships inferred by isotopic signatures with harvest. In particular, these shifts suggest that ants within harvest sites utilized resources at lower trophic levels (possibly plant‐based resources or herbivores), expanded trophic breadth, and occupied different niche spaces. Shifts in resource use following harvest could be due to harvest‐mediated changes in both the plant and arthropod communities that might affect the strength of competition or alter plant nitrogen availability. Because shifts in resource use alter the flow of nutrients across the food web, disturbance effects on ants could have ecosystem‐level consequences through nutrient cycling.     

Changes of community composition at multiple trophic levels due to hunting in Nigerian tropical forests

Hunting in tropical forests decimates large mammals, and this may have direct and indirect effects on other trophic levels and lead to trophic cascades. We compared replicated sites of hunted and protected forests in southeastern Nigeria, with respect to community composition of primates, other mammals, birds, plant seedlings, and mature trees. We make predictions regarding the community composition at the different trophic levels. In forests where large primates are rare, we hypothesize that their ecological role will not be fully compensated for by small frugivores. We apply multivariate methods to assess changes in community composition of mammals, birds, and seedlings, controlling for any differences between sites in the other groups, including mature trees. Medium and large (4–180 kg) primates were much rarer in hunted sites, while porcupine and rock hyrax increased in abundance with hunting. In contrast, the community composition of birds was similar in both types of forests. Seedling communities were significantly related to the community composition of mammals, and thus strongly affected by hunting. In protected forests primate dispersed plant seedling species dominated, whereas in hunted forests the seedling community was shifted towards one dominated by abiotically dispersed species. This was probably both a consequence of reduced seed dispersal by primates, and increased seed predation by rodents and hyrax. Hence we found no evidence for buffering effects on tree regeneration through functional compensation by non‐hunted animals (such as birds). Our results highlight how seedling communities are changed by the complex plant–animal intera ctions, triggered by the loss of seed dispersers. The results predict a rarity of primate‐dispersed trees in future tropical forest canopies; a forest less diverse in timber and non‐timber resources.     

Relationship between freshwater gastropods and plant communities reflecting various trophic levels

Plant communities were examined in ponds in Brittany (north-west France) and then classified into six types reflecting different trophic levels: oligotrophic, oligodystrophic, mesotrophic, meso-dystrophic, meso-eutrophic an eutrophic. In 45 of these ponds, aquatic snails were sampled in order to determine the relationship between the gastropod species richness and the water trophic levels as indicated by plant community types. The second aim of this study was to determine whether some gastropod species were characteristic of a particular trophic level. The number of plant communities in the ponds was also taken into consideration.A trophic gradient was found along the F1 axis on the principal plane of the correspondence analyses. The species richnesses low or zero and especially the lymneid, Lymnaea glabra were close to the oligo-dystrophic and oligotrophic communities. In contrast, the highest numbers of snail species (5 and above) were found in the most eutrophic ponds where Hippeutis complanatus, Planorbis planorbis, Lymnaea stagnalis and Planorbarius corneus were particularly common. The latter species inhabited the ponds including on average the greatest number of macrophyte communities but no significant differences were found between snail species. The ponds which contained the greatest numbers of plant communities included the richest gastropod communities (7 and more) but also the poorest ones (0 or 1 species). Relationships between freshwater snails, macrophytes and trophic levels are discussed.Laboratoire de Zoologie et d'Ecophysiologie     

A bioenergetic framework for the temperature dependence of trophic interactions

Changing temperature can substantially shift ecological communities by altering the strength and stability of trophic interactions. Because many ecological rates are constrained by temperature, new approaches are required to understand how simultaneous changes in multiple rates alter the relative performance of species and their trophic interactions. We develop an energetic approach to identify the relationship between biomass fluxes and standing biomass across trophic levels. Our approach links ecological rates and trophic dynamics to measure temperature‐dependent changes to the strength of trophic interactions and determine how these changes alter food web stability. It accomplishes this by using biomass as a common energetic currency and isolating three temperature‐dependent processes that are common to all consumer–resource interactions: biomass accumulation of the resource, resource consumption and consumer mortality. Using this framework, we clarify when and how temperature alters consumer to resource biomass ratios, equilibrium resilience, consumer variability, extinction risk and transient vs. equilibrium dynamics. Finally, we characterise key asymmetries in species responses to temperature that produce these distinct dynamic behaviours and identify when they are likely to emerge. Overall, our framework provides a mechanistic and more unified understanding of the temperature dependence of trophic dynamics in terms of ecological rates, biomass ratios and stability.     

Divergent trophic responses to biogeographic and environmental gradients

Following environmental changes, communities disassemble and reassemble in seemingly unpredictable ways. Whether species respond to such changes individualistically or collectively (e.g. as functional groups) is still unclear. To address this question, we used an extensive new dataset for the lake communities in the Azores' archipelago to test whether: 1) individual species respond concordantly within trophic groups; 2) trophic groups respond concordantly to biogeographic and environmental gradients. Spatial concordance in individual species distributions within trophic groups was always greater than expected by chance. In contrast, trophic groups varied non‐concordantly along biogeographic and environmental gradients revealing idiosyncratic responses to them. Whether communities respond individualistically to environmental gradients thus depends on the functional resolution of the data. Our study challenges the view that modelling environmental change effects on biodiversity always requires an individualist approach. Instead, it finds support for the longstanding idea that communities might be modelled as a cohort if the functional resolution is appropriate.     

Integration of ecosystem engineering and trophic effects of herbivores

Herbivores affect vegetation in a variety of ways, involving both trophic and ecosystem engineering interactions, but the study of these different interaction types has rarely been integrated. The aim of this study was to investigate both the trophic and engineering effects of herbivores on plant communities in the Negev desert, Israel, and to promote an integrative approach to the study of herbivore effects in ecosystems. First, we summarise previous studies of the Indian crested porcupine (Hystrix indica), which show that in digging for food, porcupines excavate soil pits, which accumulate resources and seeds resulting in marked changes in plant species richness, density and biomass. By contrast, their trophic effect, via consumption of bulbs, has little impact on populations of perennial plants. Second, we present an empirical study of the trophic and ecosystem engineering effects of harvester ants (Messor spp.). An exclusion experiment, using barriers to restrict ant access, failed to reveal any significant effect of seed collection by harvester ants on plant species incidence (proportional occurrence in samples) or abundance (number of individuals). However, we show that vegetation on nest mounds of M. ebeninus differs in plant density, species richness and biomass from that on undisturbed soil. An analysis of incidence and abundance responses of individual plant species suggests that the observed differences in vegetation resulted from multiple interacting mechanisms.
The case studies highlight that many interactions between herbivores and plant communities can occur simultaneously, and that ecosystem engineering and trophic processes can be closely associated, resulting from single actions of herbivores. We propose a conceptual framework that classifies the range of possible trophic and engineering interactions between herbivores and plant communities with respect to the level of association between trophic and engineering effects. The framework is presented as an aid to the design and interpretation of studies of interactions between herbivores and plant communities, and promotes integrative research into the roles of herbivores in ecosystems.     

Molecular characterization of trophic ecology within an island radiation of insect herbivores (Curculionidae: Entiminae: Cratopus)

The phytophagous beetle family Curculionidae is the most species‐rich insect family known, with much of this diversity having been attributed to both co‐evolution with food plants and host shifts at key points within the early evolutionary history of the group. Less well understood is the extent to which patterns of host use vary within or among related species, largely because of the technical difficulties associated with quantifying this. Here we develop a recently characterized molecular approach to quantify diet within and between two closely related species of weevil occurring primarily within dry forests on the island of Mauritius. Our aim is to quantify dietary variation across populations and assess adaptive and nonadaptive explanations for this and to characterize the nature of a trophic shift within an ecologically distinct population within one of the species. We find that our study species are polyphagous, consuming a much wider range of plants than would be suggested by the literature. Our data suggest that local diet variation is largely explained by food availability, and locally specialist populations consume food plants that are not phylogenetically novel, but do appear to represent a novel preference. Our results demonstrate the power of molecular methods to unambiguously quantify dietary variation across populations of insect herbivores, providing a valuable approach to understanding trophic interactions within and among local plant and insect herbivore communities.     

Assessment of the biocontrol effects of three aromatic plants on multiple trophic levels of the arthropod community in an agroforestry ecosystem

1. Studies have shown that plant diversity plays a major role in influencing arthropod community composition. However, the effects of increasing plant species diversity on arthropod abundance at multiple trophic levels in the presence of aromatic plants have not been well documented. 2. To explore the potential of using aromatic plants to biocontrol arthropods at multiple trophic levels, three aromatic plant species – French marigold (Tagetes patula L.), Ageratum (Ageratum houstonianum Mill.) and Catnip (Nepeta cataria L.) – were introduced into an apple orchard to increase ground plant species composition. 3. The aromatic plants influenced the structure of arthropod communities at multiple trophic levels, particularly the herbivores in the tree canopy and predators in ground covers. Aromatic plants negatively influenced total arthropod community abundance. Compared with the control treatment, the total arthropod community abundance in the treated areas declined 24.99–33.84% and 14.35–24.65% in the tree canopy and ground covers, respectively. 4. Aromatic plants negatively influenced herbivore abundance, both overall and relative to the total community. By contrast, aromatic plants positively influenced predator abundance, both overall and relative to the total community, in the treatments containing both ageratum and catnip. However, aromatic plants had no effect on species richness at each trophic level or on parasitoid abundance. 5. These results suggest that increasing ground plant species diversity by introducing aromatic plants into apple orchards may considerably affect arthropod community composition, and that aromatic plants are potentially effective for the biocontrol of herbivore pests in agroforestry ecosystems.     

Species identity and negative density dependence lead to convergence in designed plant mixtures of twelve species

Competition effects on community development are difficult to quantify in species-rich plant communities due to the complexity of possible interactions. We used multispecies mixtures to investigate how species identity and competitive interactions influence the development of plant communities. Given the same set of species with differing initial abundance in various communities, we tested whether communities would become more similar (converge) or dissimilar (diverge) over time depending on the relative importance of species identity and competition. Twenty-four experimental communities were established by planting seedlings of twelve wetland species at different relative abundances and absolute densities. The development of the communities was monitored over three years, and yearly changes in biomass were modelled as a linear function of the species biomass at the start of each period. After three years, a clear dominance structure had developed, with four species making up 80% of the aboveground biomass. In all three years, community dynamics was driven by differences in relative growth rates among the species (i.e. an effect of species identity). However, in the second and third years negative density dependence was also important, with changes in the relative abundance of the most abundant species being negatively related to their biomass at the start of the period. Multiple species interactions – though generally weaker than effects of species identity and intraspecific competition – became increasingly important and also contributed to the dominance pattern. It is concluded that species identity and negative density dependence of the dominant species were the most important factors causing the experimental plant communities to converge. We suggest that model systems composed of several species offer a useful method for investigating the influence of functional traits upon community dynamics.     

In situ warming strengthens trophic cascades in a coastal food web

Global warming may affect most organisms and their interactions. Theory and simple mesocosm experiments suggest that consumer top–down control over primary producer biomass should strengthen with warming, since consumer respiration increases faster with warming than plant photosynthesis. However, these predictions have so far not been tested on natural communities that have experienced warming over many generations. Natural systems display a higher diversity, heterogeneity and complexity than mesocosms, which could alter predicted effects of warming. Here we used an artificially heated part of the northern Baltic Sea (the Forsmark Biotest basin) to test how warming influences trophic interactions in a shallow coastal food web with four trophic levels: omnivorous fish, invertivorous fish, herbivorous invertebrates, and filamentous macroalgae. Monitoring of fish assemblages over six years showed that small invertivorous fish (gobiids, sticklebacks and minnows) were much less abundant in the heated basin than in unheated references areas. Stomach content analyses of the dominating omnivorous fish – Eurasian perch Perca fluviatilis – revealed a strikingly different diet within and outside the Biotest basin; gammarid crustaceans were the dominating prey at heated sites, whereas invertivorous fish (e.g. gobiids) dominated at unheated sites. A 45‐day cage experiment showed that fish exclusion did not affect the biomass of algal herbivores (gastropods and gammarids), but reduced algal biomass in heated sites (but not unheated). This suggests that warming induced a trophic cascade from fish to algae, and that this effect was mediated by predator‐induced changes in herbivore behavior, rather than number. Overall, our study suggests that warming has effectively compressed the food chain from four to three trophic levels (algae, gammarids and perch), which have benefitted the primary producers by reducing grazing pressure. Consequently, warming appears to have restructured this coastal food web through a combination of direct (physiological) and indirect (species interactions) effects.     

Landscape complexity differentially benefits generalized fourth,over specialized third,trophic level natural enemies

The differential loss of higher trophic levels in the face of natural habitat loss can result in the disruption of important trophic interactions, such as biological control. Natural enemies of herbivorous pests in cropping systems often benefit from the presence of natural habitats in surrounding landscapes, as they provide key resources such as alternative hosts. However, any benefits from a biological control perspective may be dampened if this also enhances enemies at the fourth trophic level. Remarkably, studies of the influence of landscape structure on diversity and interactions of fourth trophic‐level natural enemies are largely lacking. We carried out a large‐scale sampling study to investigate the effects of landscape complexity (i.e. the proportion of non‐crop habitat in the landscapes surrounding focal study areas) on the parasitoid communities of aphids in wheat and on an abundant extra‐field plant, stinging nettle. Primary parasitoid communities (3rd trophic level) attacking the cereal aphid, Sitobion avenae, had little overlap with the communities attacking the nettle aphid, Microlophium carnosum, while secondary parasitoids (4th trophic level) showed high levels of species overlap across these two aphids (25 vs 73% shared species respectively), resulting in significantly higher linkage density and lower specialization for secondary than primary parasitoid webs. In wheat, parasitoid diversity was not related to landscape complexity for either primary or secondary parasitoids. Rates of primary parasitism were generally low, while secondary parasitism rates were high (37–94%) and increased significantly with increasing landscape complexity, although this pattern was driven by a single secondary parasitoid species. Overall, our results demonstrate that extra‐field habitats and landscape complexity can differentially benefit fourth, over third, trophic level natural enemies, and thereby, could dampen biological control. Our results further suggest that fourth trophic‐level enemies may play an important, yet understudied, role in linking insect population dynamics across habitat types.