Elevationally biased avian predation as a contributor to the spatial distribution of geometrid moth outbreaks in sub‐arctic mountain birch forest

1. Population dynamics and interactions that vary over a species' range are of particular importance in the context of latitudinal clines in biological diversity. Winter moth (Operophtera brumata) and autumnal moth (Epirrita autumnata) are two species of eruptive geometrids that vary widely in outbreak tendency over their range, which generally increases from south to north and with elevation. 2. The predation pressure on geometrid larvae and pupae over an elevational gradient was tested. The effects of background larval density and bird occupancy of monitoring nest boxes on predation rates were also tested. Predation on larvae was tested through exclusion treatments at 20 replicate stations over four elevations at one site, while pupae were set out to measure predation at two elevations at three sites. 3. Larval densities were reduced by bird predation at three lower elevations, but not at the highest elevation, and predation rates were 1.9 times higher at the lowest elevation than at the highest elevation. The rate of predation on larvae was not related to background larval density or nest box occupancy, although there were more eggs and chicks at the lowest elevation. There were no consistent differences in predation on pupae by elevation. 4. These results suggest that elevational variation in avian predation pressure on larvae may help to drive elevational differences in outbreak tendency, and that birds may play a more important role in geometrid population dynamics than the focus on invertebrate and soil predators of previous work would suggest.     

Modeling nitrogen flux by larval insect herbivores from a temperate hardwood forest

Herbivorous insects flux considerable amounts of nitrogen from the forest canopy to the soil in the form of frass. The amount of nitrogen fluxed varies depending on the characteristics of the herbivores, their food resources, and their physical environment. We used concepts from metabolic ecology and ecological stoichiometry to develop a general model of individual nitrogen flux via frass fall for moth and sawfly larvae from a temperate hardwood forest in northern Wisconsin, USA. We found that individual nitrogen flux (Q _N, mg N/day) was related to larval body mass (M _B, mg dry), short-term variation in environmental temperature (T, K), and larval nitrogen concentration (N _B, proportion dry mass) as Q _N = e^25.75 M _B^0.77 e^−0.83/kT N _B^−1.56, where k is Boltzmann’s constant (8.62 × 10⁻⁵ eV/K). We also found that larval nitrogen flux did not vary with the nitrogen concentration of food, and suggest that this was due to compensatory feeding by larvae living on low-quality leaves. With further work, models of individual N flux could be used to scale individual fluxes to population and community levels, and thus link the characteristics of insect herbivore communities with the flow of nitrogen through forested ecosystems.     

Large herbivores facilitate an insect herbivore by modifying plant community composition in a temperate grassland

Large herbivores often co‐occur and share plant resources with herbivorous insects in grassland ecosystems; yet, how they interact with each other remains poorly understood. We conducted a series of field experiments to investigate whether and how large domestic herbivores (sheep; Ovis aries) may affect the abundance of a common herbivorous insect (aphid; Hyalopterus pruni) in a temperate grassland of northeast China. Our exclosure experiment showed that 3 years (2010–2012) of sheep grazing had led to 86% higher aphid abundance compared with ungrazed sites. Mechanistically, this facilitative effect was driven by grazing altering the plant community, rather than by changes in food availability and predator abundance for aphids. Sheep significantly altered plant community by reducing the abundance of unpalatable forbs for the aphids. Our small‐scale forb removal experiment revealed an “associational plant defense” by forbs which protect the grass Phragmites australis from being attacked by the aphids. However, selective grazing on forbs by sheep indirectly disrupted such associational plant defense, making P. australis more susceptible to aphids, consequentially increasing the density of aphids. These findings provide a novel mechanistic explanation for the effects of large herbivores on herbivorous insects by linking selective grazing to plant community composition and the responses of insect populations in grassland ecosystems.     

Elevated CO2 reduces leaf damage by insect herbivores in a forest community

By altering foliage quality, exposure to elevated levels of atmospheric CO(2) potentially affects the amount of herbivore damage experienced by plants. Here, we quantified foliar carbon (C) and nitrogen (N) content, C : N ratio, phenolic levels, specific leaf area (SLA) and the amount of leaf tissue damaged by chewing insects for 12 hardwood tree species grown in plots exposed to elevated CO(2) (ambient plus 200 microl l(-1)) using free-air CO(2) enrichment (FACE) over 3 yr. The effects of elevated CO(2) varied considerably by year and across species. Elevated CO(2) decreased herbivore damage across 12 species in 1 yr but had no detectable effect in others. Decreased damage may have been related to lower average foliar N concentration and SLA and increased C : N ratio and phenolic content for some species under elevated compared with ambient CO(2). It remains unclear how these changes in leaf properties affect herbivory. Damage to the leaves of hardwood trees by herbivorous insects may be reduced in the future as the concentration of CO(2) continues to increase, perhaps altering the trophic structure of forest ecosystems.     

Contrasting effects of insect and molluscan herbivores on plant diversity in a long-term field experiment

The importance of invertebrate herbivores in regulating plant communities remains unclear, due to the absence of long-term exclusion experiments. An experiment in an English grassland involving long-term exclusions of insect and mollusc herbivores, along with rabbit fencing, showed strong, but opposing, effects of the invertebrate herbivores. Plant species richness declined and biomass increased following insect exclusion, due to increased dominance by a grass species, whereas mollusc exclusion led to increased herbs abundance. The two herbivores had a compensatory interaction: molluscs had no effects in the absence of insects and large insect effects depended on the absence of molluscs. The effects of invertebrate exclusion became apparent only after 8 years, and would have been seriously underestimated in shorter studies. Our results suggest that theorists and conservation managers need to shift from their historic focus on vertebrate herbivory, to a recognition that invertebrates can be equally important drivers of plant community structure.     

Bottom‐up vs. top‐down effects on terrestrial insect herbivores: a meta‐analysis

Primary consumers are under strong selection from resource (‘bottom‐up’) and consumer (‘top‐down’) controls, but the relative importance of these selective forces is unknown. We performed a meta‐analysis to compare the strength of top‐down and bottom‐up forces on consumer fitness, considering multiple predictors that can modulate these effects: diet breadth, feeding guild, habitat/environment, type of bottom‐up effects, type of top‐down effects and how consumer fitness effects are measured. We focused our analyses on the most diverse group of primary consumers, herbivorous insects, and found that in general top‐down forces were stronger than bottom‐up forces. Notably, chewing, sucking and gall‐making herbivores were more affected by top‐down than bottom‐up forces, top‐down forces were stronger than bottom‐up in both natural and controlled (cultivated) environments, and parasitoids and predators had equally strong top‐down effects on insect herbivores. Future studies should broaden the scope of focal consumers, particularly in understudied terrestrial systems, guilds, taxonomic groups and top‐down controls (e.g. pathogens), and test for more complex indirect community interactions. Our results demonstrate the surprising strength of forces exerted by natural enemies on herbivorous insects, and thus the necessity of using a tri‐trophic approach when studying insect‐plant interactions.     

The legacy of harvest and fire on ecosystem carbon storage in a north temperate forest

Forest harvesting and wildfire were widespread in the upper Great Lakes region of North America during the early 20th century. We examined how long this legacy of disturbance constrains forest carbon (C) storage rates by quantifying C pools and fluxes after harvest and fire in a mixed deciduous forest chronosequence in northern lower Michigan, USA. Study plots ranged in age from 6 to 68 years and were created following experimental clear‐cut harvesting and fire disturbance. Annual C storage was estimated biometrically from measurements of wood, leaf, fine root, and woody debris mass, mass losses to herbivory, soil C content, and soil respiration. Maximum annual C storage in stands that were disturbed by harvest and fire twice was 26% less than a reference stand receiving the same disturbance only once. The mechanism for this reduction in annual C storage was a long‐lasting decrease in site quality that endured over the 62‐year timeframe examined. However, during regrowth the harvested and burned forest rapidly became a net C sink, storing 0.53 Mg C ha⁻¹ yr⁻¹ after 6 years. Maximum net ecosystem production (1.35 Mg C ha⁻¹ yr⁻¹) and annual C increment (0.95 Mg C ha⁻¹ yr⁻¹) were recorded in the 24‐ and 50‐year‐old stands, respectively. Net primary production averaged 5.19 Mg C ha⁻¹ yr⁻¹ in experimental stands, increasing by < 10% from 6 to 50 years. Soil heterotrophic respiration was more variable across stand ages, ranging from 3.85 Mg C ha⁻¹ yr⁻¹ in the 6‐year‐old stand to 4.56 Mg C ha⁻¹ yr⁻¹ in the 68‐year‐old stand. These results suggest that harvesting and fire disturbances broadly distributed across the region decades ago caused changes in site quality and successional status that continue to limit forest C storage rates.     

Vertebrate predators have minimal cascading effects on plant production or seed predation in an intact grassland ecosystem

The strength of trophic cascades in terrestrial habitats has been the subject of considerable interest and debate. We conducted an 8-year experiment to determine how exclusion of vertebrate predators, ungulates alone (to control for ungulate exclusion from predator exclusion plots) or none of these animals influenced how strongly a three-species assemblage of rodent consumers affected plant productivity. We also examined whether predator exclusion influenced the magnitude of post-dispersal seed predation by mice. Both ungulates and rodents had strong direct effects on graminoid biomass. However, rodent impacts on plant biomass did not differ across plots with or without predators and/or ungulates. Deer mice removed more seeds from seed depots on predator exclusion plots, suggesting trait-mediated indirect effects of predators, but this short-term behavioural response did not translate into longer-term impacts on seed survival. These results suggest that vertebrate predators do not fundamentally influence primary production or seed survival in our system.     

Assessment of trends in predation pressure on insects across temperate forest microhabitats

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Responses of insect herbivores and herbivory to habitat fragmentation: a hierarchical meta‐analysis

Loss and fragmentation of natural habitats can lead to alterations of plant–animal interactions and ecosystems functioning. Insect herbivory, an important antagonistic interaction is expected to be influenced by habitat fragmentation through direct negative effects on herbivore community richness and indirect positive effects due to losses of natural enemies. Plant community changes with habitat fragmentation added to the indirect effects but with little predictable impact. Here, we evaluated habitat fragmentation effects on both herbivory and herbivore diversity, using novel hierarchical meta‐analyses. Across 89 studies, we found a negative effect of habitat fragmentation on abundance and species richness of herbivores, but only a non‐significant trend on herbivory. Reduced area and increased isolation of remaining fragments yielded the strongest effect on abundance and species richness, while specialist herbivores were the most vulnerable to habitat fragmentation. These fragmentation effects were more pronounced in studies with large spatial extent. The strong reduction in herbivore diversity, but not herbivory, indicates how important common generalist species can be in maintaining herbivory as a major ecosystem process.     

Impact of crimson clover dying mulch on two eggplant insect herbivores

Eggplant Solanum melongena L., is often colonized by two early season insect defoliators. The Colorado potato beetle (CPB), Leptinotarsa decemlineata (Say), and flea beetles Epitrix spp., emerge from their overwintering sites in early spring and seek out emerging host plants such as eggplant. During the 2009 and 2010 growing season, field studies were conducted to investigate the impact of inter‐planting eggplant into a crimson clover (CC), Trifolium incarnatum L., winter cover crop on populations of flea beetles, CPB and their associated arthropod predators. The experiment consisted also of two levels of insecticide usage such as an application of azadirachtins plus pyrethrins followed by several applications of spinosad or no insecticide sprays as subplot treatments. During both study years, significantly fewer (adults, larvae and egg masses) were found on eggplant inter‐planted into CC than in bare‐ground (BG) eggplant plots. Although flea beetle abundance was greater in BG eggplant during 2010, they appeared to be less influenced by the presence of CC than were CPB. Additionally, there was no apparent impact of insecticide treatment on CPB populations on eggplant inter‐planted into CC. However, there was a decline in CPB following treatments with insecticides in BG eggplant plots. This suggests that a winter cover crop such as CC can be used to help manage CPB in eggplant, however, using this tactic in tandem with insecticide sprays may not result in greater CPB management.     

Impacts of forest fragmentation on species composition and forest structure in the temperate landscape of southern Chile

Aim Few studies have explicitly examined the influence of spatial attributes of forest fragments when examining the impacts of fragmentation on woody species. The aim of this study was to assess the diverse impacts of fragmentation on forest habitats by integrating landscape‐level and species‐level approaches. Location The investigation was undertaken in temperate rain forests located in southern Chile. This ecosystem is characterized by high endemism and by intensive recent changes in land use. Method Measures of diversity, richness, species composition, forest structure and anthropogenic disturbances were related to spatial attributes of the landscape (size, shape, connectivity, isolation and interior forest area) of forest fragments using generalized linear models. A total of 63 sampling plots distributed in 51 forest fragments with different spatial attributes were sampled. Results Patch size was the most important attribute influencing different measures of species composition, stand structure and anthropogenic disturbances. The abundance of tree and shrub species associated with interior and edge habitats was significantly related to variation in patch size. Basal area, a measure of forest structure, significantly declined with decreasing patch size, suggesting that fragmentation is affecting successional processes in the remaining forests. Small patches also displayed a greater number of stumps, animal trails and cow pats, and lower values of canopy cover as a result of selective logging and livestock grazing in relatively accessible fragments. However, tree richness and β‐diversity of tree species were not significantly related to fragmentation. Main conclusions This study demonstrates that progressive fragmentation by logging and clearance is associated with dramatic changes in the structure and composition of the temperate forests in southern Chile. If this fragmentation process continues, the ability of the remnant forests to maintain their original biodiversity and ecological processes will be significantly reduced.     

Effect of soil macroaggregates crushing on C mineralization in a tropical deciduous forest ecosystem

Soil aggregates strongly influence C dynamics by affecting microbial activity. Our study tested the effect of soil crushing on C mineralization by laboratory incubation experiments of soil samples from a tropical deciduous forest ecosystem in Western Mexico. Soil samples were taken in January (dry season) and in October (rainy season). For each sampling date, the incubation experiment had a two factorial design with litter and macroaggregates (>250 μm) crushing as the main factors, both with two levels (with and without). At both sampling dates, the soil samples with intact macroaggregates had significantly higher C mineralization than the soil samples in which macroaggregates were crushed. The pH of leached solution was higher in the crushed soil samples than in uncrushed soil samples. The reduction of C mineralization caused by the disruption of soil aggregates is explained by the disturbance of environmental conditions within macroaggregates. The effect of macroaggregates crushing also reduced the differences of C mineralization between both seasons. We concluded that macroaggregates promote microbial activity by reducing the impact of variations in soil chemical and physical environmental conditions.     

Concordant population dynamics of Lepidoptera herbivores in a forest ecosystem

It is frequently assumed that population fluctuations are largely independent within a community of trophically‐similar species, but this need not be so. If population fluctuations are partly synchronized or concordant, this will produce interannual variability in the community's aggregate abundance and generate temporal variance in ecosystem structure. We studied the community of Lepidoptera inhabiting northern hardwood forests in New Hampshire, USA, to evaluate the hypothesis that fluctuations in consumer communities can arise from concordant dynamics of constituent populations. Interannual comparisons of moth abundances for >75 species sampled at three sites over four years revealed that concordant dynamics contribute strongly to interannual variability in the abundance of consumers. A conspicuous decline in community abundance from 2004 to 2005 was the result of predominantly negative population growth rates of the component species, while an increase in community abundance from 2006 to 2007 was the result of predominantly positive population growth rates. Population dynamics most strongly linked species that feed in the early season (perhaps due to shared responses to climatic effects), but not species that might share natural enemies or host plants. The observed concordant dynamics introduced conspicuous temporal variation in the abundance of primary consumers relative to plants and secondary consumers, thereby altering the forest's trophic structure. Such variance in the aggregate abundance of forest primary consumers could generate time‐lagged fluctuations in abundances of secondary consumers and will generally have important consequences for ecosystem properties and processes that are nonlinear functions of consumer abundance, such as plant community structure and nutrient cycling.     

The significance of unripe seeds and animal tissues in the protein nutrition of herbivores

Many herbivorous animals selectively eat flowers and unripe fruit or seeds. Some preferentially eat new tissues growing from germinating seeds. This behaviour enables access to otherwise limited or unavailable amino acids that are necessary to sustain successful production and growth of young. For the same reason the diet of breeding females and neonates of many presumed strictly herbivorous animals is supplemented with animal protein. However, because these foods are often only eaten for limited periods, and make up only a small proportion of the total diet, they are usually dismissed as unimportant to the animals' nutrition. It is suggested that actively looking for such feeding may well reveal it to be far more common and important to the successful breeding of herbivores than has been thought in the past.     

Effects of pepper (Capsicum chinense) genotypic diversity on insect herbivores

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Tree seedling richness,but not neighborhood composition,influences insect herbivory in a temperate deciduous forest community

Insect herbivores can serve as important regulators of plant dynamics, but their impacts in temperate forest understories have received minimal attention at local scales. Here, we test several related hypotheses about the influence of plant neighborhood composition on insect leaf damage in southwestern Pennsylvania, USA. Using data on seedlings and adult trees sampled at 36 sites over an approximately 900 ha area, we tested for the effects of total plant density, rarefied species richness (i.e., resource concentration and dietary‐mixing hypotheses), conspecific density (i.e., Janzen–Connell hypothesis), and heterospecific density (i.e., herd‐immunity hypothesis), on the proportion of leaf tissue removed from 290 seedlings of 20 species. We also tested for the effects of generic‐ and familial‐level neighborhoods. Our results showed that the proportion of leaf tissue removed ranged from zero to just under 50% across individuals, but was generally quite low (<2%). Using linear mixed models, we found a significant negative relationship between insect damage and rarefied species richness, but no relationship with neighborhood density or composition. In addition, leaf damage had no significant effect on subsequent seedling growth or survival, likely due to the low levels of damage experienced by most individuals. Our results provide some support for the resource concentration hypothesis, but suggest a limited role for insect herbivores in driving local‐scale seedling dynamics in temperate forest understories.     

Quantifying resilience of multiple ecosystem services and biodiversity in a temperate forest landscape

Resilience is increasingly being considered as a new paradigm of forest management among scientists, practitioners, and policymakers. However, metrics of resilience to environmental change are lacking. Faced with novel disturbances, forests may be able to sustain existing ecosystem services and biodiversity by exhibiting resilience, or alternatively these attributes may undergo either a linear or nonlinear decline. Here we provide a novel quantitative approach for assessing forest resilience that focuses on three components of resilience, namely resistance, recovery, and net change, using a spatially explicit model of forest dynamics. Under the pulse set scenarios, we explored the resilience of nine ecosystem services and four biodiversity measures following a one‐off disturbance applied to an increasing percentage of forest area. Under the pulse + press set scenarios, the six disturbance intensities explored during the pulse set were followed by a continuous disturbance. We detected thresholds in net change under pulse + press scenarios for the majority of the ecosystem services and biodiversity measures, which started to decline sharply when disturbance affected >40% of the landscape. Thresholds in net change were not observed under the pulse scenarios, with the exception of timber volume and ground flora species richness. Thresholds were most pronounced for aboveground biomass, timber volume with respect to the ecosystem services, and ectomycorrhizal fungi and ground flora species richness with respect to the biodiversity measures. Synthesis and applications. The approach presented here illustrates how the multidimensionality of stability research in ecology can be addressed and how forest resilience can be estimated in practice. Managers should adopt specific management actions to support each of the three components of resilience separately, as these may respond differently to disturbance. In addition, management interventions aiming to deliver resilience should incorporate an assessment of both pulse and press disturbances to ensure detection of threshold responses to disturbance, so that appropriate management interventions can be identified.     

A review of predation as a limiting factor for bird populations in mesopredator‐rich landscapes: a case study of the UK

The impact of increasing vertebrate predator numbers on bird populations is widely debated among the general public, game managers and conservationists across Europe. However, there are few systematic reviews of whether predation limits the population sizes of European bird species. Views on the impacts of predation are particularly polarised in the UK, probably because the UK has a globally exceptional culture of intensive, high‐yield gamebird management where predator removal is the norm. In addition, most apex predators have been exterminated or much depleted in numbers, contributing to a widely held perception that the UK has high numbers of mesopredators. This has resulted in many high‐quality studies of mesopredator impacts over several decades. Here we present results from a systematic review of predator trends and abundance, and assess whether predation limits the population sizes of 90 bird species in the UK. Our results confirm that the generalist predators Red Fox (Vulpes vulpes) and Crows (Corvus corone and C. cornix) occur at high densities in the UK compared with other European countries. In addition, some avian and mammalian predators have increased numerically in the UK during recent decades. Despite these high and increasing densities of predators, we found little evidence that predation limits populations of pigeons, woodpeckers and passerines, whereas evidence suggests that ground‐nesting seabirds, waders and gamebirds can be limited by predation. Using life‐history characteristics of prey species, we found that mainly long‐lived species with high adult survival and late onset of breeding were limited by predation. Single‐brooded species were also more likely to be limited by predation than multi‐brooded species. Predators that depredate prey species during all life stages (i.e. from nest to adult stages) limited prey numbers more than predators that depredated only specific life stages (e.g. solely during the nest phase). The Red Fox and non‐native mammals (e.g. the American Mink Neovison vison) were frequently identified as numerically limiting their prey species. Our review has identified predator–prey interactions that are particularly likely to result in population declines of prey species. In the short term, traditional predator‐management techniques (e.g. lethal control or fencing to reduce predation by a small number of predator species) could be used to protect these vulnerable species. However, as these techniques are costly and time‐consuming, we advocate that future research should identify land‐use practices and landscape configurations that would reduce predator numbers and predation rates.