Traits reveal ecological strategies driving carrion insect community assembly

1. The succession of carrion-associated (necrophilous) insects on decomposing carrion is well documented. To exploit the changing nutritious and dynamic resources available throughout the carrion decomposition process, different species colonise and consume carrion in a predictable temporal sequence. The traits of these necrophilous insects should reflect their ecological strategies. Morphological traits of these insects, such as body size and wing size, however, have not previously been examined during active and advanced decomposition. 2. We used fourth-corner multivariate generalised linear models to identify insect community morphological trait patterns and to quantify their change through time on decomposing rabbit carcasses in grassland and woodland environments. 3. We found that larger-bodied species of flies and carrion-specialist beetles were associated with the early stages of decomposition. The morphological traits of ants, in contrast, showed no changes at carcasses through time and instead showed body size differences between grassland and woodland environments. 4. Our findings indicate that specialist flies and beetles that arrive early in the decomposition process possess traits that enable rapid discovery of carrion at a large scale. Generalist beetles and ants do not share this same trait and are instead adapted to locate and consume a wider variety of resources in their preferred habitat type at their local scale. 5. Our results provide insights into the morphological adaptations linked to the ecological strategies of distinct components of carrion insect communities. Further, our results offer insights into the community assembly dynamics that structure the communities of necrophilous insect species.     

Decomposition rate of carrion is dependent on composition not abundance of the assemblages of insect scavengers

Environmental factors and biodiversity affect ecosystem processes. As environmental change modifies also biodiversity it is unclear whether direct effects of environmental factors on ecosystem processes are more important than indirect effects mediated by changes in biodiversity. High-quality resources like carrion occur as heterogeneous pulses of energy and nutrients. Consequently, the distribution of scavenging insects is related to resource availability. Therefore, carrion decomposition represents a suitable process from which to unravel direct effects of environmental change from indirect biodiversity-related effects on ecosystem processes. During three field seasons in 2010 we exposed traps baited with small-mammal carrion at 21 sites along a temperature gradient to explore the insect carrion fauna and decomposition rate in the Bohemian Forest, Germany. The abundance component of beetle and fly assemblages decreased with decreasing temperature. Independently, the composition component of both taxa changed with temperature and season. The change in the composition component of beetles depicted a loss of larger species at higher temperatures. Decomposition rate did not change directly along the temperature gradient but was directly influenced by season. The composition component of beetles, and to a small extent of flies, but not their abundance component, directly affected carrion decomposition. Consequently, lower decomposition rates at lower temperatures can be explained by the absence of larger beetle species. Thus, we predict that future environmental change will modify carrion fauna composition and thereby indirectly decomposition rate. Moreover, reorganizations of the insect carrion composition will directly translate into modified decomposition rates, with potential consequences for nutrient availability and carbon storage.     

Species richness and composition patterns across trophic levels of true bugs (Heteroptera) in the agricultural landscape of the lower reach of the Tisza River Basin

River basins are among the most threatened ecosystems. The species diversity of several European river basins decreased seriously during the last decade due to loss of habitats and increasing land use pressure on the remaining habitats. We studied true bug assemblages in various land use types of grassland fragments and dikes as linear grassland habitats in the agricultural landscape of the lower reach of the Tisza River Basin. We tested the effects of the recorded variables of habitat quality, surrounding landscape and land use type on the abundance, species richness and composition of true bugs. Altogether, 5,389 adult Heteroptera individuals representing 149 species in 13 families were collected. The factors which influenced significantly the species richness of different trophic levels (i.e. herbivors, predators) and degrees of food specialization (i.e. generalist and specialist herbivors) were concordant. Contrary to this, the factors which influenced the abundance of the different feeding groups varied strongly. We emphasise the vegetation and land use types as primarily influential factors for insects. Excluding the grass-feeding species, the number of both generalist, specialist herbivorous and predaceous species were lower in agricultural swards, i.e. hay-meadows and pastures than in old field and dike habitats and their number increased with increasing vegetation diversity. Due to the high species richness and abundance observed in dike and old field habitats compared to agricultural swards, we emphasise their importance for conservation of insect diversity and we stress the negative effects of agricultural intensification on the remaining grasslands of the lower reach of the Tisza River Basin.     

Does habitat loss affect the communities of plants and insects equally in plant–pollinator interactions? Preliminary findings

Habitat loss is a major threat to biodiversity and ecosystem function. As habitats are lost, one factor affecting their community structures is the niche-width demand of species, which ranges from specialist to generalist. This study focused on specialist and generalist species in plant–pollinator interactions and tested the hypothesis that plant and pollinator communities become more generalized as habitat loss increases. The study was made in seven selected sites in southern Ontario, Canada, at the level of landscape that is characterized by distributed forests within intensively managed agricultural fields. We quantified both the degree of habitat loss and the degree of specialization/generalization for each of the plant and insect communities using a sampling method of hexagonal transects. Regression analysis indicated a significant relationship between the increase of habitat loss and the shift to generalization in insect, but not in plant, communities. Our results suggest that, in plant–pollinator interactions, insect communities are more sensitive and/or quicker than plant communities to respond to the effects of habitat loss.     

The effect of forest age and habitat structure on the ground‐dwelling ant assemblages of lowland poplar plantations

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Influence of surrounding vegetation on insect herbivory: A matter of spatial scale and herbivore specialisation

The diversity of surrounding vegetation is thought to modify the interactions between a focal plant and its herbivores, disrupting (associational resistance) or enhancing (associational susceptibility) host plant location and colonisation. We compared the effects of host plant concentration on herbivory by generalist and specialist insects feeding on oak seedlings by increasing local concentration of seedlings. We also assessed the effects of the composition and structure of surrounding vegetation, both at stand and local levels. The damage caused by generalist leaf-feeding insects depended on the structure of plant communities at stand level, and increased with tree cover. By contrast, infestation by specialist leaf miners was affected by local understorey vegetation surrounding oak seedlings, and decreased with increasing shrub cover and stratification diversity. Leaf mine abundance was higher at higher oak seedling density, giving support to the host concentration hypothesis. However, the abundance of these specialist herbivores was also negatively correlated with damage caused by the generalist external leaf-feeders, suggesting competitive interactions.     

Tracking the effects of one century of habitat loss and fragmentation on calcareous grassland butterfly communities

Habitat loss and fragmentation are known to reduce patch sizes and increase their isolation, consequently leading to modifications in species richness and community structure. Calcareous grasslands are among the richest ecosystems in Europe for insect species. About 10% (1,150 ha) of the total area of a calcareous ridge region (Calestienne, Belgium) and its butterfly community was analysed over a timeframe of about 100 years. Since 1905 to present day (2005), the Calestienne region has undergone both calcareous grassland loss and fragmentation: not only did calcareous grassland size decrease and isolation increase, but also, the number of calcareous grassland patches within the landscape increased until 1965, and subsequently decreased, clearly reflecting the effects of fragmentation. These processes have had a profound effect on the butterfly community: extinction and rarefaction affected significantly more often specialist species, which means that generalist species are more and more overrepresented. This ecological drift, i.e. the replacement of specialists by generalists in species assemblages is likely to be a general effect of habitat loss and fragmentation on natural communities.     

A preliminary investigation of temporal patterns in semiarid ant communities: Variation with habitat type

Abstract This study reports on preliminary findings of habitat‐contingent temporal variability in ant assemblages in Purnululu National Park in northern Australia's semiarid tropics, by sampling at the end of the dry season (October 2004) and the end of the wet season (April 2005). Six grids of 15 pitfall traps were established in each of the spinifex, sandplain and gorge habitats. Community composition was dominated by behaviourally dominant ants (Iridomyrmex spp.) and climate specialists (Melophorus and Meranoplus spp.). Ant activity was higher in the wet season sampling period, with greater species richness and abundance. Interestingly, temporal variation in ant assemblage richness, abundance and composition varied markedly with habitat type. While there were large differences between sampling periods for the spinifex and sandplain habitat, this was not the case in the gorges. These temporal changes in ant assemblages are postulated to be linked with major environmental differences between the two sampling periods, driven by seasonal climatic conditions. It is likely that these changes influenced the ant assemblages through species differences in physiological tolerance levels, ecological requirements and competitive ability. This study demonstrates the need, in highly seasonal environments, to consider the temporal context of studies in relation to habitat type, particularly when undertaking biodiversity surveys and monitoring.     

Species Traits Predict Assemblage Dynamics at Ephemeral Resource Patches Created by Carrion

Carrion is an ephemeral and spatially patchy resource that supports a diverse subset of species linked to nutrient cycling and the decomposition process. A number of studies have separately documented changes in the diversity of plants, arthropods and vertebrates at individual carcasses, but there are few studies that have examined how functional traits of different groups of organisms underpin their responses to carrion patches. We used a carrion addition experiment to compare changes in composition and functional traits of insect and plant assemblages at carcasses compared with control sites. We found that significant changes in insect assemblage evenness and heterogeneity was associated with species’ dispersal traits, and that plant assemblage responses to subsequent soil nitrogen changes was most apparent among graminoids and exotic species. Beetles at carcasses were twice as large as their counterparts at control sites during the first week of carrion decomposition, and also had higher wing loadings. Plants with high specific leaf area responded faster to the carcass addition, and twice as many species recolonised the centre of carcasses in exotic-dominated grassland compared with carcasses in native-dominated grassland. These results provide an example of how traits of opportunist species enable them to exploit patchy and dynamic resources. This increases our understanding of how carcasses can drive biodiversity dynamics, and has implications for the way carrion might be managed in ecosystems, such as appropriate consideration of spatial and temporal continuity in carrion resources to promote heterogeneity in nutrient cycling and species diversity within landscapes.     

Increasing habitat connectivity in agricultural landscapes as a weed management strategy reconciling ecology and agronomy

While studies have explored how habitat amount drives weed assemblages in agroecosystems, knowledge remains limited of the effects of habitat connectivity. The response-effect trait framework provides insights into the mechanisms underpinning the relationship between landscape structure and the taxonomic diversity and abundance of weed assemblages. This study evaluated how habitat connectivity and habitat amount affect weed diversity and abundance in winter cereal fields, and whether these effects are driven by the functional composition of weed assemblages. We sampled weeds in 27 winter cereal fields. We measured habitat connectivity and habitat amount provided by wooded, grassland and cropland elements. We selected five traits related to the dispersal, establishment, and competitive abilities of weed species likely to respond to landscape structure: seed number per plant, type of reproduction, seed dry mass, plant vegetative height and seed germination rate. The functional composition of weed assemblages was assessed using community weighted mean trait values. Weed diversity and abundance were used as proxies of weed management. The taxonomic approach did not reveal any effect of landscape structure on weed diversity and abundance. Only the grassland elements that contributed to habitat connectivity, and to a lesser extent to habitat amount, drove the functional composition of weed assemblages. High habitat amount favoured species with many seeds, while high habitat connectivity favoured species with fewer seeds, a higher ability to reproduce vegetatively and higher seed germination rates. In turn, higher seed germination rates increased weed evenness and reduced weed abundance. Some of these relationships were influenced by the presence of rare species. Overall, high connectivity provided by grassland elements increases weed evenness and reduces weed abundance by shaping weed functional composition. Our study suggests that land-use planning policies that enhance the connectivity provided by grassland elements could be considered as a weed management strategy reconciling ecology and agronomy.     

Phytophagous insect fauna tracks host plant responses to exotic grass invasion

The high dependence of herbivorous insects on their host plants implies that plant invaders can affect these insects directly, by not providing a suitable habitat, or indirectly, by altering host plant availability. In this study, we sampled Asteraceae flower heads in cerrado remnants with varying levels of exotic grass invasion to evaluate whether invasive grasses have a direct effect on herbivore richness independent of the current disturbance level and host plant richness. By classifying herbivores according to the degree of host plant specialization, we also investigated whether invasive grasses reduce the uniqueness of the herbivorous assemblages. Herbivorous insect richness showed a unimodal relationship with invasive grass cover that was significantly explained only by way of the variation in host plant richness. The same result was found for polyphagous and oligophagous insects, but monophages showed a significant negative response to the intensity of the grass invasion that was independent of host plant richness. Our findings lend support to the hypothesis that the aggregate effect of invasive plants on herbivores tends to mirror the effects of invasive plants on host plants. In addition, exotic plants affect specialist insects differently from generalist insects; thus exotic plants affect not only the size but also the structural profile of herbivorous insect assemblages.     

Fire effect on ground-foraging ant assemblages in northeastern Argentina

Fire is an important component of many natural ecosystems affecting plant communities and arthropods by mortality during combustion and/or indirectly through the modification of the habitat. The Iberá Natural Reserve (INR) is one of the most diverse ecosystems in northern Argentina; it is dominated by grasslands commonly affected by disturbances, such as grazing and fire. The objective of this work was to study the response of ground-foraging ant assemblages, particular species, and functional groups to an extended fire of high intensity in four natural INR habitats with >5 years of cattle exclusion (strict conservation area). A total of 12,798 ant workers of 67 species were captured in 39 sampling stations. The ant fauna was less abundant in burned sites only a few days after the fire; 6 months later, no effect was detected. Richness and abundance of ants differed among unburned habitats. However, fire effect on species richness and composition remained unclear. The rapid recovery of the ant fauna made these insects poor indicators of long-term fire-promoted changes on biodiversity in open habitats dominated by grassland, though some ant species showed a high level of habitat fidelity mainly in unburned habitats. These results agree with those from other areas of the world, indicating that ants are particularly unreliable biodiversity indicators, with the exception of severe disturbance with long-term habitat restoration. Management decisions at the INR should be oriented to preserve the closed savanna, one of the most diverse and threatened habitat of Argentina.     

Phytochemical defences and performance of specialist and generalist herbivores: a meta-analysis

1. Phytochemical coevolution theory, a long-standing paradigm in plant–insect interactions, predicts that specialist herbivores are less negatively affected by the allelochemicals of their host plants than are generalist herbivores. Although this theory is prevalent in plant–insect science, it is not always supported by empirical studies measuring the performance of specialist and generalist insects in response to allelochemicals. 2. The present study aimed to investigate: (i) whether there a difference between specialist and generalist performance in response to allelochemicals and (ii) whether the effect of allelochemicals on specialists and generalists depend upon allelochemical class or insect order. 3. A meta-analysis was conducted incorporating 76 effect sizes drawn from studies that directly compared the performance of specialist and generalist insects in response to treatment and control diets. Most of the effect sizes were related to the performance metric growth, the insect order Lepidoptera, and the allelochemical class nitrogen-containing compounds. 4. As predicted by phytochemical coevolution theory, specialist insects responded less negatively to allelochemicals of their hosts than generalist insects in terms of growth. There were no significant differences in terms of fecundity or survival, or among allelochemical classes or insect orders. 5. These results support the prediction of phytochemical coevolution theory that specialist insects respond less negatively to allelochemicals of their hosts than generalists, although only in terms of growth.     

Large and least isolated fragments preserve habitat specialist spiders best in dry sandy grasslands in Hungary

The role of fragment size, isolation and habitat diversity in the conservation of spider assemblages living in fragmented landscape were studied in dry sandy grasslands (East Hungary, Nyírség). Spiders were collected using pitfall traps at eight dry grassland fragments from 2001 to 2009 from March to October fortnightly. We tested the rules of island biogeography, which suggest that the species richness increases with the size and decreases with the isolation of fragments. The habitat diversity is an important factor for species richness, since large areas usually have more habitats; therefore, the number of species may be higher in these areas. During the 9-year study period, altogether 10,544 individuals belonging to 106 species were collected. Contradicting the classical theory, we found a significant negative relationship between the total number of spider species and the grassland size, while the ratio of sandy grassland specialist spider species increased with fragment size. The relationship between the ratio of generalist species and the fragment size was not significant. The overall species richness and the isolation of studied grasslands did not show a significant relationship. The ratio of sandy grassland specialist species decreased, while the ratio of generalist species increased with the increasing of isolation. The habitat diversity did not show any effect on spider species richness. We concluded that to conserve the habitat specialist species it is recommended to preserve the large and least isolated grassland fragments, furthermore to increase the size of small fragments with the restoration of the adjacent croplands.     

Flower visitor communities are similar on remnant and reconstructed tallgrass prairies despite forb community differences

One common goal of habitat restoration and reconstruction is to reinstate the biodiversity found at intact reference sites. However, few researchers have examined whether these practices reinstate communities of flower‐visiting insects. This is unfortunate, as anthropogenically mediated declines in flower visitors, including bees (the primary pollinators for most terrestrial ecosystems), beetles, flies, and butterflies, have been reported worldwide. Biodiversity declines may be especially severe in North America's tallgrass prairie, a once‐vast grassland that has experienced severe destruction and degradation due to agricultural conversion. As such, we assessed the structure of forb and flower‐visiting insect communities as a whole and two subsets of the flower visitor community—bees and phytophagous beetles—across five tallgrass prairie remnants and five reconstructed prairies (former crop fields) in Kansas from 2013 to 2015. Remnant prairies had significantly higher forb diversity and differed significantly in forb composition, compared to reconstructed prairies. Despite the dissimilarities in forb community structure, there were no differences in flower visitor diversity or abundance between remnants and reconstructed prairies. However, when considered separately, bee communities exhibited significantly greater variability in composition on reconstructed prairies, likely due to the abundance of generalist bee species visiting non‐native legumes at two reconstructed prairies. Our work provides evidence that prairie habitat reconstruction is a valuable tool for reestablishing flower‐visiting insect communities and also emphasizes the considerable role that non‐native species may play in structuring grassland plant–bee interactions.     

Measuring the efficiency of fen restoration on carabid beetles and vascular plants: a case study from north‐eastern Germany

The aim of the study was to assess the effects of fen rewetting on carabid beetle and vascular plant assemblages within riverine fens along the river Peene in north‐eastern Germany. Drained (silage grassland), rewetted (restored formerly drained silage grassland), and near‐natural (fairly pristine) stands were compared. Eighty‐four beetle species (7,267 individuals) and 135 plant species were recorded. The richness of vascular plant species and the number of endangered species were highest on near‐natural fens. Fourteen years of rewetting did not increase plant species numbers compared with drained fens. For carabid beetles, however, species richness and the number of stenotopic species were highest on rewetted fens. Rewetting caused the replacement of generalist carabids by wetland specialists, but did not provide suitable habitat for specialist fen carabids or for plant species of oligo‐ or mesotrophic fen communities. Therefore, raising the water table on fens with nutrient‐rich, degraded peat was not sufficient for restoring species assemblages of intact fens, although water level was the most important environmental factor separating species assemblages. Our study illustrated that insects and plants may respond differentially to restoration, stressing the need to consider different taxa when assessing the efficiency of fen restoration. Furthermore, species assemblages of intact fens could not be restored within 14 years, highlighting the importance of conserving pristine habitat.     

Changes in the composition of British butterfly assemblages over two decades

Changes in the abundance and distribution of individual species have been widely documented in Britain and other countries in recent decades, but little has been done to determine changes in community composition over broad geographic areas. Here, we studied species turnover in 51 butterfly assemblages in Britain since 1976, examining extinction and colonisation events together with variation in the abundances of the species. We showed that the species turnover that occurred over 20 years in Britain was associated with colonisation and extinction events and also with variability in the abundance of the species. These changes in community composition differed according to the habitat requirements of the species and their previous distributions, being more evident for habitat specialists and for southerly distributed species. Colonising species often became abundant components of the communities they joined, although this was more evident for generalist than for specialist species. The abundance of species following their arrival, increased with time since colonisation. Species turnover associated with southerly species expanding northwards is consistent with being a response to climate change. The results suggest that climate- and habitat-driven changes in the identity and abundance of species within communities are widespread, and probably ubiquitous. Similar changes are likely to be occurring in other groups of organisms that are similarly undertaking major range shifts associated with climate change.     

Close association between grasshopper and plant communities in suburban secondary grasslands and the indicator value of grasshoppers for conservation

Semi-natural grasslands in Japan have decreased due to management abandonment and urbanization over the last 100 years, but they remain in suburban areas in addition to rural areas. Because suburban grasslands have various land-use histories and disturbance regimes, plant and herbivorous insect communities are likely to differ among grassland types. To identify grasslands with high conservation value, we conducted a comprehensive survey of grasshoppers and plants in 150 grasslands with 5 grassland types differing in land-use history and current management in northern Chiba prefecture, Japan. We then analyzed the association of the distributions of grasshopper and plant species compositions. Our results showed that grasshoppers were classified into habitat specialists and generalists. Three out of four habitat specialists were almost exclusively found in semi-natural grasslands and vacant lots, while habitat generalists were commonly observed at the cropland margins. This habitat specialist–generalist distribution gradient corresponded well to that found in plant communities, which was probably due to current disturbance regimes. We suggest that vacant lots as well as semi-natural grasslands have high conservation value for grassland organisms of various taxa in suburban areas, and grasshoppers are candidate indicator species for monitoring grassland environments.

     

The relative importance of lethal and non-lethal effects of fish on insect colonisation of ponds

1. We hypothesised that adult insects actively monitor potential habitats for the presence of fish by means of chemical cues and avoid sites that pose significant risks. This was examined by quantifying colonisation of insects in outdoor pools with no fish (controls), fish (direct predation effect) or caged fish (chemical predator cues).
2. A significant direct effect of predation was found, but no indirect effect (avoidance of chemical cue pools), on the total biomass of colonising insects. However, predatory insects avoided fish-cue pools, thus releasing non-predatory insects from predation. This resulted in significantly greater biomass of non-predatory insects in fish-cue pools than control pools.
3. Fish reduced the number of species of colonising insects in pools through predation. This negative influence of fish implies that caution is necessary when stocking wetlands and ponds with fish if the goal is to maximise biodiversity.
4. Our data suggest that although predatory aquatic insects may use chemical signals to assess the quality of potential habitats with respect to predation risk, direct predation is the main method by which fish affect insect assemblages in ponds. Because fish and invertebrate predators may both have strong effects on prey mortality, behavioural adjustment by insects to the actual predator regime within a habitat should be more important than avoiding colonisation of habitats with fish.     

Habitat complementarity and butterfly traits are essential considerations when mitigating the effects of exotic plantation forestry

Habitat loss threatens insect diversity globally. However, complementary vegetation types in remaining habitat increases opportunities for species survival. We assess the extent to which indigenous forest patches moderate the impact of exotic commercial afforestation on grassland butterflies. Butterflies were sampled in grassland along uncorrelated gradients of landscape-scale indigenous forest and plantation cover, while controlling for variation in local vegetation composition. We separately assessed responses by butterfly groups differing in habitat preference, larval diet, and mobility. There was no effect of landscape- or local-scale variables on species richness, but there was a strong interactive effect of forest and plantation cover on butterfly assemblage structure. The effect varied according to species traits. When forest cover was high, assemblages did not differ at different levels of plantation cover. However, plantation cover significantly influenced assemblage structure when forest cover was low. Grassland with limited forest cover in the protected area supported unique assemblages with high frequency of less mobile, specialized species with herbaceous larval host plants, whereas grassland with low forest cover near plantations had a prevalence of mobile, generalist species. A positive association between forest cover and butterflies with woody larval host plants suggests that indigenous forest patches improved the suitability of fragmented grassland for a subset of butterflies, emphasising the value of natural heterogeneity in transformed areas. However, certain butterfly traits associated with large, open grassland were under-represented in grassland between plantations, underscoring the importance of open areas in the broader landscape to conserve the full diversity of species.