Leaf dynamics and insect herbivory in a Eucalyptus camaldulensis forest under moisture stress

Abstract The influence of soil moisture content on leaf dynamics and insect herbivory was examined between September 1991 and March 1992 in a river red gum (Eucalyptus camaldulensis) forest in southern central New South Wales. Long-term observations of leaves were made in trees standing either within intermittently flooded waterways or at an average of 37. 5m from the edge of the waterways. The mean soil moisture content was significantly (P≤0.05) greater in the waterways than in the non-flooded areas. Trees in the higher soil moisture regime produced significantly larger basal area increments and increased canopy leaf area. This increase in canopy leaf area was achieved, in part, through a significant increase in leaf longevity and mean leaf size. Although a greater number of leaves was initiated and abscissed per shoot from the non-flooded trees, more leaves were collected from litter traps beneath the denser canopies of the flooded trees. Consumption of foliage by insects on the trees subjected to flooding compared to the non-flooded trees was not significantly different. However, the relative impact of insect herbivory was significantly greater on the non-flooded trees. Leaf chewing was the most common form of damage by insects, particularly Chryso-melidae and Curculionidae. No species was present in outbreak during this study. Leaf survival decreased as the per cent area eaten per leaf increased. In addition, irrespective of the level of herbivory, leaf abscission tended to be higher in E. camaldulensis under moisture deficit. The influence of soil moisture content on the balance between river red gum growth and insect herbivory is discussed.     

Fire Increases Insect Herbivory in a Neotropical Savanna

Fire is an important agent of disturbance in many tropical ecosystems that can potentially influence plant consumers. Nevertheless, there are few reports on whether levels of plant damage change as a result of fire. Here we present the results of a 1‐yr study evaluating the effects of fire on rates of herbivory and damage by pathogens in leaves of cerrado (Brazilian savanna) tree species. Damage by leaf chewers was over two times greater in burned than in unburned trees. Levels of damage by leaf miners, leaf scrapers, galling insects, and leaf pathogens were relatively low and increased, remained the same, or even decreased as a result of fire. Nevertheless, in all three plant species studied, total herbivore damage was significantly greater in burned than in unburned trees given the preponderance of damage caused by leaf chewers compared with the other types of damage. Leaf chewers, mainly leaf‐cutter ants, caterpillars, and grasshoppers, completely ate over 50 percent of the >2000 leaves we marked in burned trees. That our results were consistent among different plant species with contrasting leaf phenologies suggests that the observed increase in herbivory is a general phenomenon in our study system. Because herbivore pressure is augmented dramatically in recently burned areas, herbivory may act synergistically with fire in influencing the structure of cerrado vegetation.     

Influence of boreal forest succession and dead wood qualities on saproxylic beetles

1 Saproxylic insects, a functional group dominated by beetles, are dependent on dead or moribund trees as habitat elements. 2 Although there are few studies of saproxylic insects from the North American boreal zone, European studies demonstrate that forest harvest can lead to a biologically significant decrease in saproxylic beetle diversity. 3 We studied saproxylic beetles in the North American boreal mixedwood forest using flight intercept traps established on naturally dead and girdled trembling aspen and spruce trees along a successional gradient of undisturbed stands from deciduous to coniferous overstory trees. 4 Composition and diversity of beetle assemblages differed among forest successional types. 5 Snag age class was an important determinant of composition for saproxylic beetle assemblages. 6 Multivariate regression analysis of these data indicated that saproxylic beetles are responding to changes in coarse woody debris, and not to the relative densities of canopy tree species, although these variables are strongly correlated. 7 Coarse woody debris management should be a primary concern in forest management plans seeking to conserve saproxylic organisms and the critical ecosystem functions (i.e. nutrient cycling) in which they participate.     

Effects of genetic variability and habitat of Qualea parviflora (Vochysiaceae) on herbivory by free-feeding and gall-forming insects

• Background and Aims Differences in the chemical and physical traits of plants caused by both genetic and habitat characteristics may influence attack by herbivores. Leaves of Qualea parviflora (Vochysiaceae), a common tree of different habitats in the Brazilian Neotropical savannas (cerrado), are susceptible to severe attack by herbivorous free-living and gall-forming insects. Attack by free-living and gall-forming insects within and between populations of Q. parviflora were examined and it was determined to what extent genetic variability (detected by RAPD markers), phenotypic characteristics of the plants and habit traits influence the number of free-living and gall-forming insect species and individuals attacking the plants, and the intensity of attack.• Methods On four occasions in 2000, leaves were sampled from ten individual trees in each of three types of vegetation in the cerrado: campo sujo, cerrado sensu stricto and cerradão at the Ecological Station of Pirapitinga (ESP), in Três Marias, north-western Minas Gerais, Brazil. Genetic variability was detected by RAPD markers, and concentrations of nutrients, phenols and tannins, sclerophylly and pre-dawn water potential of leaves were measured. Water and nutrient contents in the soil below each tree characterized the habitat. The free-living and gall-forming herbivorous insects were determined.• Key Results Of 69 RAPD markers analysed, 41 were polymorphic and were used for analyses of genetic variability of Q. parviflora. Most of the variability occurred within habitats, accounting for 97·65 % of the genetic variability. Plants in the cerrado sensu stricto and campo sujo were the most similar. There were no significant associations between genetic similarity and the chemical and physical traits of Q. parviflora, or with habitat, nor was there significant correlation between phenotypic and habitat traits. Increasing concentrations of tannins and sulphur, C : N ratio and sclerophylly correlated with increasing percentage of leaf area damaged by herbivores. Decreased sclerophylly, concentration of tannins and C : N ratio, and increased concentration of nutrients in leaves correlated with increased severity of attack by gall-forming insects.• Conclusions Nutrient concentration in the soil had more influence, indirectly, on free-feeding insects than did composition of Q. parviflora leaves. However, gall-forming insects are affected more by leaf quality, attacking fewer sclerophyllous leaves, with larger nutrient but smaller tannin concentrations.Key words: Cerrado, genetic variability, gall attack, herbivory, insect galls, plant quality, Qualea parviflora, RAPD, Vochysiaceae     

Gall-forming and free-feeding herbivory along vertical gradients in a lowland tropical rainforest: the importance of leaf sclerophylly

In contrast to most insect guilds, gall-forming insects are thought to reach highest diversity on sclerophyllous vegetation, such as Neotropical savannas and Mediterranean vegetation types. The water and nutrient stress endured by meristems of canopy trees in tall wet tropical rainforests may cause leaf sclerophylly. Hence, the upper canopies of such ecosystems may represent a suitable habitat for gall-forming insects. At the San Lorenzo Protected Area, Panama, we estimated free-feeding herbivory and gall densities within five sites in 2003 and 2004, by surveying leaves in vertical and horizontal transects. In each sample, we recorded leaf density (mature and young foliage), free-feeding herbivore damage and number of galls, including the presence of live larvae, parasitoids or fungi. We surveyed 43 994 leaves, including 231 plants and 73 tree and liana species. We collected 5014 galls from 17 host-plant species, including 32 gall species of which 59% were restricted to the canopy (overall infestation rates: 2.4% in 2003, 5.5% in 2004). In 2003, 16% of the galls were occupied by live larvae, against 5% in 2004. About 17–20% of leaves surveyed suffered from free-feeding herbivory. Leaf sclerophylly increased significantly with sampling height, while free-feeding herbivory decreased inversely. Conversely, the number of live galls collected in the canopy was 13–16 times higher than in the understorey, a pattern consistent across sites and years. Hence, the probability of gall survivorship increased with increasing leaf sclerophylly as death by fungi, parasitoids or accidental chewing were greater in the understorey. Increasing harsh ecophysiological conditions towards the upper canopy appear favourable to galls-forming population maintenance, in support of the hypothesis of harsh environment. Hence, gall diversity and abundance in the upper canopy of tall tropical forests are perhaps among the highest in the world.     

Assessing invertebrate herbivory in human‐modified tropical forest canopies

1. Studies on the effects of human‐driven forest disturbance usually focus on either biodiversity or carbon dynamics but much less is known about ecosystem processes that span different trophic levels. Herbivory is a fundamental ecological process for ecosystem functioning, but it remains poorly quantified in human‐modified tropical rainforests.
2. Here, we present the results of the largest study to date on the impacts of human disturbances on herbivory. We quantified the incidence (percentage of leaves affected) and severity (the percentage of leaf area lost) of canopy insect herbivory caused by chewers, miners, and gall makers in leaves from 1,076 trees distributed across 20 undisturbed and human‐modified forest plots in the Amazon.
3. We found that chewers dominated herbivory incidence, yet were not a good predictor of the other forms of herbivory at either the stem or plot level. Chewing severity was higher in both logged and logged‐and‐burned primary forests when compared to undisturbed forests. We found no difference in herbivory severity between undisturbed primary forests and secondary forests. Despite evidence at the stem level, neither plot‐level incidence nor severity of the three forms of herbivory responded to disturbance.
4. Synthesis. Our large‐scale study of canopy herbivory confirms that chewers dominate the herbivory signal in tropical forests, but that their influence on leaf area lost cannot predict the incidence or severity of other forms. We found only limited evidence suggesting that human disturbance affects the severity of leaf herbivory, with higher values in logged and logged‐and‐burned forests than undisturbed and secondary forests. Additionally, we found no effect of human disturbance on the incidence of leaf herbivory.

     

Differential Responses of Herbivores and Herbivory to Management in Temperate European Beech

Forest management not only affects biodiversity but also might alter ecosystem processes mediated by the organisms, i.e. herbivory the removal of plant biomass by plant-eating insects and other arthropod groups. Aiming at revealing general relationships between forest management and herbivory we investigated aboveground arthropod herbivory in 105 plots dominated by European beech in three different regions in Germany in the sun-exposed canopy of mature beech trees and on beech saplings in the understorey. We separately assessed damage by different guilds of herbivores, i.e. chewing, sucking and scraping herbivores, gall-forming insects and mites, and leaf-mining insects. We asked whether herbivory differs among different forest management regimes (unmanaged, uneven-aged managed, even-aged managed) and among age-classes within even-aged forests. We further tested for consistency of relationships between regions, strata and herbivore guilds. On average, almost 80% of beech leaves showed herbivory damage, and about 6% of leaf area was consumed. Chewing damage was most common, whereas leaf sucking and scraping damage were very rare. Damage was generally greater in the canopy than in the understorey, in particular for chewing and scraping damage, and the occurrence of mines. There was little difference in herbivory among differently managed forests and the effects of management on damage differed among regions, strata and damage types. Covariates such as wood volume, tree density and plant diversity weakly influenced herbivory, and effects differed between herbivory types. We conclude that despite of the relatively low number of species attacking beech; arthropod herbivory on beech is generally high. We further conclude that responses of herbivory to forest management are multifaceted and environmental factors such as forest structure variables affecting in particular microclimatic conditions are more likely to explain the variability in herbivory among beech forest plots.     

森林冠层昆虫多样性研究方法

森林冠层是森林昆虫栖息、取食、避敌的重要生境,其中生活着十分丰富的昆虫物种。但由于乔木树体高大,冠层难于接近,在很大程度上限制了冠层昆虫多样性的研究。冠层昆虫类群结构的划分和冠层昆虫取样技术也是冠层昆虫多样性研究的重要基础。文章综述当前冠层接近和冠层昆虫抽样技术的最新进展,并评述冠层昆虫类群结构划分的方法。     

Seasonal patterns of herbivory,leaf traits and productivity consumption in dry and wet Patagonian forests

1. Endemic herbivory can influence forest ecosystem function, but how annual productivity consumption relates to seasonal resource utilisation by folivore guilds remains poorly understood. 2. Monthly changes in leaf damage and foliage traits were monitored in ‘dry’ and ‘wet’Nothofagus pumilio (Fagales: Nothofagaceae) deciduous forests in northern Patagonia, Argentina. Herbivore‐induced leaf abscission was assessed and foliar productivity consumption was measured in the canopy and in litterfall harvests. 3. Seasonal damage ranged from 8% to 32% in dry forest, but remained below 5% in wet forest although foliar quality was higher in the latter. In dry forest, dominant guilds were temporally separated; leaf miners consumed younger foliage in spring to early summer, whereas leaf tiers prevailed in late summer to autumn. In wet forest, damage created by external chewers was concentrated in early summer. 4. Insect damage induced premature leaf abscission, especially in dry forest. Although foliar production in wet forest doubled that in dry forest, the percentage of productivity lost to folivores was higher in dry (14–20%) than in wet (1.2–1.8%) forest. 5. The overall greater impact of herbivory in dry forest canopies countered the expectation that consumption would increase with plant productivity and nutritional quality. Lower temperatures and a shorter growing season are likely to constrain folivory in wet forest stands.     

Different initial responses of the canopy herbivory rate in mature oak trees to experimental soil and branch warming in a soil‐freezing area

Plants and insects comprise more than 50% of known species on earth, and their interactions are of major importance in most natural ecosystems. To understand the mechanisms by which global warming affects plant–insect interactions in the canopy of mature cool‐temperate forests with a freeze–thaw cycle, we examined changes in the herbivory rate and leaf traits in oak Quercus crispula. From 2007 to 2009, we experimentally increased the temperature of the surrounding soil and canopy branches of mature oak trees by approximately 5°C using electric heating cables. Soil warming decreased the rate of herbivory in the canopy, whereas branch warming had no effect. The magnitude of the effect of soil warming on canopy herbivory varied. For the first year, the decrease was 32%, but this doubled (63%) in the third year. Branch warming did not affect canopy leaf traits; however, soil warming decreased the leaf nutritional quality by decreasing N and increasing the carbon:nitrogen (CN) ratio for three years. Additionally, soil warming increased total phenolics in the third year. Stepwise multiple regression models showed that among the leaf traits that were changed by soil warming, N explained the variation in herbivory for the first and second years, whereas total phenolics explained it for the third year. Our experimental results demonstrate that soil warming drives the rate of herbivory in the canopy of mature oak trees, and the magnitude of the soil warming effect was gradually enhanced during the initial three years. This suggests the importance of belowground temperature elevation in predicting the effect of global warming on plant–insect interactions in a forest canopy.     

Nutrient supply and bird predation additively control insect herbivory and tree growth in two contrasting forest habitats

It has been suggested that bottom–up and top–down forces interactively control food web dynamics. While top–down effects would increase with resource availability to plants, bottom–up effects would be stronger under low predator abundance. These predictions, however, have rarely been tested at contrasting sites while keeping the dominant plant species unchanged. Furthermore, few studies have factorially manipulated both types of forces in forest communities. For two years, we evaluated the effects of fertiliser (NPK) addition and bird exclusion on tree growth, leaf traits, insect abundance, and folivory rates in a dry/warm and a wet/cold Nothofagus pumilio forest in Patagonia, Argentina. Overall, we found no interaction between nutrient supply and bird predation, although the strength of bottom–up and top–down forces differed markedly between forest sites. Treatment effects were generally weak in the wet forest, where tree growth rates and insect herbivory were low relative to the dry forest. In the dry forest, fertilisation increased sapling growth, insect abundance and folivory, whereas bird exclusion increased leaf damage and reduced tree growth. In the wet forest, fertilisation enhanced leaf nutrient contents and folivore abundance but not sapling growth, while bird exclusion had little impact on insects or trees. These results imply that factors other than nutrients and birds were important in controlling tree growth and folivore activity in the wet forest. While treatment effect sizes varied widely among feeding guilds, in general, nutrient effects on folivores were stronger than predator effects. We conclude that, within the time‐frame of this study, tree growth and herbivory were additively affected by soil nutrients and predator presence, as bird exclusion effects did not change with elevated folivore activity on fertilised trees. We also show that both top–down and bottom–up cascades were weaker in a forest site characterised by slow‐growing juvenile trees subjected to low folivore pressure.     

Components of ecosystem evaporation in a temperate coniferous rainforest, with canopy transpiration scaled using sapwood density

Here we develop and test a method to scale sap velocity measurements from individual trees to canopy transpiration (E(c)) in a low-productivity, old-growth rainforest dominated by the conifer Dacrydium cupressinum. Further, E(c) as a component of the ecosystem water balance is quantified in relation to forest floor evaporation rates and measurements of ecosystem evaporation using eddy covariance (E(eco)) in conditions when the canopy was dry and partly wet. Thermal dissipation probes were used to measure sap velocity of individual trees, and scaled to transpiration at the canopy level by dividing trees into classes based on sapwood density and canopy position (sheltered or exposed). When compared with ecosystem eddy covariance measurements, E(c) accounted for 51% of E(eco) on dry days, and 22% of E(eco) on wet days. Low transpiration rates, and significant contributions to E(eco) from wet canopy evaporation and understorey transpiration (35%) and forest floor evaporation (25%), were attributable to the unique characteristics of the forest: in particular, high rainfall, low leaf area index, low stomatal conductance and low productivity associated with severe nutrient limitation.     

Potential effects of elevated carbon dioxide on leaf-feeding forest insects

The elevated concentration of atmospheric CO2 may result in a decline of leaf nutritional quality (especially N) and an increase in some kinds of defensive secondary components (such as phenolics). The changes in the phytochemistry of trees, combined with the effect of elevated CO2 per se, have a potential negative influence on insect herbivores. Here, we review the effect of elevated CO2 on the performance of leaf-feeding forest insects at individual-level and commu-nity-level. The elevated CO2 per se have little influence on the metabolism of insects. Over half of the tree-insect experimental systems show that the performance of individual insect become poorer under high-CO2 grown trees; but the others show that the insects have just little or no response to the treatments. The direction and magnitude of the changes in the performance of insects could be mediated by various factors. The effects of treatment are strongly species-dependent. The magni-tude of changes in the phytochemistry, the sensitivity and adaptive capacity of insects to the poorer leaf quality, the differences in plant growth conditions and experimental methods, and the mediated effects of other environmental factors (such as soil nutrient availability, light, temperature, O3) were all closely related to the final performance of insects. However, the larvae's consumption usually increased under enriched CO2 treatment, which was widely thought to be a compensa-tory response to poorer plant quality. The experiments on forest community-level found identically a reduction in herbivory, which was contrary to the results from small-scale experiments. The changes in insect popula-tion and the actual response of consumption by leaf-feeding forest insects under CO2 enrichment remain unclear, and more field-based experiments need to be conducted.     

The influence of a neotropical herbivore (<Emphasis Type="Italic">Lamponius portoricensis</Emphasis>) on nutrient cycling and soil processes

The role of phytophagous insects in ecosystem nutrient cycling remains poorly understood. By altering the flow of litterfall nutrients from the canopy to the forest floor, herbivores may influence key ecosystem processes. We manipulated levels of herbivory in a lower montane tropical rainforest of Puerto Rico using the common herbivore, Lamponius portoricensis (Phasmatidea), on a prevalent understory plant, Piper glabrescens (Piperaceae), and measured the effects on nutrient input to the forest floor and on rates of litter decomposition. Four treatment levels of herbivory generated a full range of leaf area removal, from plants experiencing no herbivory to plants that were completely defoliated (>4,000 cm² leaf area removed during the 76-day study duration). A significant (P<0.05) positive regression was found between all measures of herbivory (total leaf area removed, greenfall production, and frass-related inputs) and the concentration of NO ₃ ⁻ in ion exchange resin bags located in the litter layer. No significant relationship was found between any of the herbivory components and resin bag concentrations of NH ₄ ⁺ or PO ₄ ⁻ . Rates of litter decay were significantly affected by frass-related herbivore inputs. A marginally significant negative relationship was also found between the litter mass remaining at 47 days and total leaf area removed. This study demonstrated a modest, but direct relationship between herbivory and both litter decomposition and NO ₃ ⁻ transfer to the forest floor. These results suggest that insect herbivores can influence forest floor nutrient dynamics and thus merit further consideration in discussions on ecosystem nutrient dynamics.     

多样化松林中昆虫群落多样性特征

马尾松和湿地松是我国南方的2种主要松树。通过对6种不同林分结构下的马尾松林和湿地松林内昆虫群落调查与多样性指数分析,表明2种松树内的昆虫种类和数量无显著差异,混交林中的昆虫群落的种类和数量比纯林多,尤其以捕食天敌类群的种类和数量更为明显。整个昆虫群落和植食类群多样性指数以湿地松林内较大,而天敌(捕食类群和寄生类群)多样性指数则以马尾松林较高。从不同林分结构下昆虫多样性的比较来看,混交林内昆虫群落多样性指数波动较小,明显地高于纯林。但不同林分结构下昆虫多样性随水平分布和垂直分层格局而变化,松树北面和东面各样地之间的昆虫群落多样性指数差异显著,而南、西面之间差异较小;树冠层各样地之间的差异达极显著水平,而枯枝落叶层和树干层之间差异不显著。由此,还进一步讨论了混交林中昆虫群落稳定性问题。     

Phytophagous insects enhance nitrogen flux in a desert creosotebush community

Summary We tested the hypothesis that herbivorous insects on desert shrubs contribute to short-term nitrogen cycling, and increase rates of nitrogen flux from nutrient rich plants. Creosotebush (Larrea tridentata) shrubs were treated with different combinations of fertilizer and water augmentations, resulting in different levels of foliage production and foliar nitrogen contents. Foliage arthropod populations, and nitrogen in canopy dry throughfall, wet throughfall and stemflow were measured to assess nitrogen flux rates relative to arthropod abundances on manipulated and unmanipulated shrubs over a one-month period during peak productivity. Numbers and biomass of foliage arthropods were significantly higher on fertilized shrubs. Sap-sucking phytophagous insects accounted for the greatest numbers of foliage arthropods, but leaf-chewing phytophagous insects represented the greatest biomass of foliage arthropods. Measured amounts of bulk frass (from leaf-chewing insects) were not significantly different among the various treatments. Amounts of nitrogen from dry and wet throughfall and stemflow were significantly greater under fertilized shrubs due to fine frass input from sap-sucking insects. Increased numbers and biomass of phytophagous insects on fertilized shrubs increased canopy to soil nitrogen flux due to increased levels of herbivory and excrement. Nitrogen excreted by foliage arthropods accounted for about 20% of the total one month canopy to soil nitrogen flux, while leaf litter accounted for about 80%.     

Tree size and herbivory determine below-canopy grass quality and species composition in savannahs

Large single-standing trees are rapidly declining in savannahs, ecosystems supporting a high diversity of large herbivorous mammals. Savannah trees are important as they support both a unique flora and fauna. The herbaceous layer in particular responds to the structural and functional properties of a tree. As shrubland expands stem thickening occurs and large trees are replaced by smaller trees. Here we examine whether small trees are as effective in providing advantages for grasses growing beneath their crowns as large trees are. The role of herbivory in this positive tree-grass interaction is also investigated. We assessed soil and grass nutrient content, structural properties, and herbaceous species composition beneath trees of three size classes and under two grazing regimes in a South African savannah. We found that grass leaf content (N and P) beneath the crowns of particularly large (ca. 3.5 m) and very large trees (ca. 9 m) was as much as 40% greater than the same grass species not growing under a tree canopy, whereas nutrient contents of grasses did not differ beneath small trees (<2.3 m). Moderate herbivory enhanced these effects slightly. Grass species composition differed beneath and beyond the tree canopy but not between tree size classes. As large trees significantly improve the grass nutrient quality for grazers in contrast to smaller trees, the decline of the former should be halted. The presence of trees further increases grass species diversity and patchiness by favouring shade-tolerant species. Both grazing wildlife and livestock will benefit from the presence of large trees because of their structural and functional importance for savannahs.     

Diversity of pselaphine beetles (Coleoptera: Staphylinidae: Pselaphinae) in eastern Thailand

Pselaphine beetles (Coleoptera: Staphylinidae: Pselaphinae) are cosmopolitan, species‐rich, and yet poorly studied, particularly in the tropics. We sampled beetles in three types of primary forest and two types of disturbed forest habitats in eastern Thailand to assess the utility of pselaphine beetles as bioindicators of forest disturbance. We simultaneously measured leaf litter mass, soil moisture, soil acidity and canopy cover at each site to infer which environmental factors affect pselaphine beetle diversity and abundance. At each site, pselaphine beetles were extracted from ten 1 m² samples of leaf litter and soil with Tullgren funnels. We sampled 1867 adult beetles representing six supertribes, 51 genera and 114 morphospecies; 7% of the genera and 92% of the species were undescribed. Forest types differed significantly in species richness, abundance, diversity and evenness. Primary forest had greater numbers of species and individuals, and higher diversity indices (H′). Teak plantation and secondary forest had substantially fewer individuals and species of pselaphine beetles. Species composition differed between primary and degraded forests. Canopy cover, soil moisture, and leaf litter mass positively correlated with beetle species richness and abundance. Leaf litter mass and soil moisture were the two most important factors affecting the diversity of pselaphine beetle assemblages. Among the 114 morphospecies collected, 43 morphospecies were specific to two or three habitats and 64 morphospecies were found only in a single habitat. Thus pselaphine beetles appear to have rather narrow habitat requirements and their presence/absence was correlated with environmental differences. These traits make pselaphine beetles a suitable bioindicator taxon for assessing forest litter diversity and monitoring habitat change.     

Tree Species Composition and Harvest Intensity Affect Herbivore Density and Leaf Damage on Beech,Fagus sylvatica,in Different Landscape Contexts

Most forests are exposed to anthropogenic management activities that affect tree species composition and natural ecosystem processes. Changes in ecosystem processes such as herbivory depend on management intensity, and on regional environmental conditions and species pools. Whereas influences of specific forest management measures have already been addressed for different herbivore taxa on a local scale, studies considering effects of different aspects of forest management across different regions are rare. We assessed the influence of tree species composition and intensity of harvesting activities on arthropod herbivores and herbivore-related damage to beech trees, Fagus sylvatica, in 48 forest plots in three regions of Germany. We found that herbivore abundance and damage to beech trees differed between regions and that – despite the regional differences - density of tree-associated arthropod taxa and herbivore damage were consistently affected by tree species composition and harvest intensity. Specifically, overall herbivore damage to beech trees increased with increasing dominance of beech trees – suggesting the action of associational resistance processes – and decreased with harvest intensity. The density of leaf chewers and mines was positively related to leaf damage, and several arthropod groups responded to beech dominance and harvest intensity. The distribution of damage patterns was consistent with a vertical shift of herbivores to higher crown layers during the season and with higher beech dominance. By linking quantitative data on arthropod herbivore abundance and herbivory with tree species composition and harvesting activity in a wide variety of beech forests, our study helps to better understand the influence of forest management on interactions between a naturally dominant deciduous forest tree and arthropod herbivores.     

Interannual changes in folivory and bird insectivory along a natural productivity gradient in northern Patagonian forests

Trophic regulation models suggest that the magnitude of herbivory and predation (top-down forces) should vary predictably with habitat productivity. Theory also indicates that temporal abiotic variation and within-trophic level heterogeneity both affect trophic dynamics, but few studies addressed how these factors interact over broad-scale environmental gradients. Here we document herbivory from leaf-feeding insects along a natural rainfall/productivity gradient in Nothofagus pumilio forests of northern Patagonia, Argentina, and evaluate the impact of insectivorous birds on foliar damage experienced by tree saplings at each end of the gradient. The study ran over three years (1997–2000) comprising a severe drought (1998–1999), which allowed us to test how climatic events alter top-down forces. Foliar damage tended to increase towards the xeric, least productive forests. However, we found a predictable change of insect guild prevalence across the forest gradient. Leaf miners accounted for the greater damage recorded in xeric sites, whereas leaf chewers dominated in the more humid and productive forests. Interannual folivory patterns depended strongly on the feeding guild and forest site. Whereas leaf-miner damage decreased during the drought in xeric sites, chewer damage increased after the drought in the wettest site. Excluding birds did not affect leaf damage from miners, but generally increased chewer herbivory on hydric and xeric forest saplings. Indirect effects elicited by bird exclusion became most significant after the drought, when total folivory levels were higher. Thus, interannual abiotic heterogeneity markedly influenced the amount of folivory and strength of top-down control observed across the forest gradient. Moreover, our results suggest that spatial turnovers between major feeding guilds may need be considered to predict the dynamics of insect herbivory along environmental gradients.