FOREST PRODUCTIVITY PREDICTS INVERTEBRATE BIOMASS AND OVENBIRD (SEIURUS AUROCAPILLUS) REPRODUCTION IN APPALACHIAN LANDSCAPES

Most nutrient enrichment studies in aquatic systems have focused on autotrophic food webs in systems where primary producers dominate the resource base. We tested the heterotrophic response to long-term nutrient enrichment in a forested, headwater stream. Our study design consisted of 2 years of pretreatment data in a reference and treatment stream and 2 years of continuous nitrogen (N) + phosphorus addition to the treatment stream. Studies were conducted with two leaf species that differed in initial C:N, Rhododendron maximum (rhododendron) and Acer rubrum (red maple). We determined the effects of nutrient addition on detrital resources (leaf breakdown rates, litter C:N and microbial activity) and tested whether nutrient enrichment affected macroinvertebrate consumers via increased biomass. Leaf breakdown rates were ca. 1.5 and 3x faster during the first and second years of enrichment, respectively, in the treatment stream for both leaf types. Microbial respiration rates of both leaf types were 3x higher with enrichment, and macroinvertebrate biomass associated with leaves increased ca. 2-3x with enrichment. The mass of N in macroinvertebrate biomass relative to leaves tended to increase with enrichment up to 6x for red maple and up to 44x for rhododendron leaves. Lower quality (higher C:N) rhododendron leaves exhibited greater changes in leaf nutrient content and macroinvertebrate response to nutrient enrichment than red maple leaves, suggesting a unique response by different leaf species to nutrient enrichment. Nutrient concentrations used in this study were moderate and equivalent to those in streams draining watersheds with altered land use. Thus, our results suggest that similarly moderate levels of enrichment may affect detrital resource quality and subsequently lead to altered energy and nutrient flow in detrital food webs.

Detritus processing, the breaking down of organic matter into smaller particles, is an essential operation in aquatic systems because it provides resources to filter feeders and accelerates nutrient release by microorganisms. Detrital foodweb dynamics are influenced by both consumption (top-down) and production (bottom-up) effects. We tested the effects of predators and detritivores on the abundance of microorganisms in an inquiline community in pitcher plants. We manipulated densities of mosquitoes (top predator) and midges (processing detritivore) in a factorial press experiment and measured the response (density) of bacteria, protozoa, and rotifer populations over several generations. We hypothesized that: 1) midges would have a positive effect on microorganisms by increasing nutrient availability (bottom-up effects), 2) mosquitoes would depress microorganism populations through consumption (top-down effects), 3) top-down and bottom-up effects would operate independently, and 4) would attenuate with trophic position. Mosquitoes (predators) had a negative effect on all measured populations. Midges (processing detritivores) had a positive effect on bacteria, but a negative effect on rotifers and some protozoan taxa. The increase in bacterial density probably was the result of nutrient enrichment from detritus processing, whereas the decrease in rotifers seems to have been the result of consumption by midges. Our study shows that the role of processing detritivores is complex and can enhance both bottom-up and top-down effects. Specifically, omnivory can complicate simple top-down and bottom-up predictions. Although they accelerate decomposition by microorganisms and, thereby, can increase resource availability, processing detritivores can also be important consumers in detrital food webs.

Spatial interpolation methods are widely used to estimate some ecological and environmental parameters that are difficult to measure. One of these parameters is forest site index, which is a demonstration of forest productivity. The aim of this study was to estimate forest site index in a beech forest ecosystem in Turkey. In this context, soil characteristics, stand parameters, and topographic features were measured in 70 temporary sample plots of beech forest stands. Forest site index of beech forest stands was predicted using different modeling techniques such as multiple regression analysis (MLR), multilayer perceptron (MLP), radial basis function (RBF), multiple regression kriging (MLRK), multilayer perceptron kriging (MLPK), and radial basis function kriging (RBFK). The results showed that the RBFK (R2 = 0.98) and MLRK (R2 = 0.96) outperformed the others to predict forest site index in the study area. The greatest improvement occurred when krigged residual used with MLR, which increase from 0.23 to 0.96. Thus, MLRK method significantly improved the prediction accuracy for site index. The models combined with krigged residuals were more successful than those used without krigged residuals. The results of this study suggest that the combined methods may help obtaining improved site index maps for forest management.

Nitrogen addition enhances the bottom-up effects in the detrital food web

Biomass or growth? How to measure soil food webs to understand structure and function

Prey subsidies originating from detritus add nutrients and energy to arboreal communities. Measurement of this subsidy is required in the understanding of how food web dynamics respond to changes in surrounding environments. Shrub spiders are one of the key predators involved in food web coupling. We evaluate the effects of potential changes in prey availabilities during secondary succession on the contribution of subsidy from detrital food webs to shrub spiders and how different spider feeding guilds used the subsidy of prey from detrital food webs. We measured the relative importance of the subsidy for the spider feeding guilds, using the ratios of stable isotopes of C (δ(13)C), and N (δ(15)N) and C isotope discrimination (Δ(14)C). Diet age was calculated from Δ(14)C values, because old diet ages of spiders indicate that the spiders consume prey from detrital food sources. Dominant aerial prey (Diptera) had a distinctively old diet age compared with arboreal prey, which indicates that aerial prey were subsidized from detrital food webs. Sit-and-wait spiders tended to have an older diet age than active hunting spiders, which indicates that sit-and-wait spiders depended more on subsidies. Diet age varied only slightly for spiders in stands of different ages, indicating that rates at which spiders use grazing and detrital prey are probably determined more by foraging strategies and not by stand age. A dominance of sit-and-wait predators will lead to higher detrital subsidy inputs in shrub habitats. This study highlights the effect of shrub spider community structure (feeding guild composition) on the volume of the subsidy received from the detrital food web.

Site and climate conditions are the key determinants controlling dominant height growth and forest productivity, both independently and interactively. Secondary natural oak forests are a typical forest type in China, especially in Hunan Province, but little is known about the site index of this forest under the complex site and climate variables in the subtropics. Based on survey data of dominant trees and site variables from 101 plots in Hunan oak natural secondary forests and climate data obtained using spatial interpolation, we used the random forest method, correlation analysis, and the analysis of variance to determine the main site and climate factors affecting oak forest dominant height and proposed a modeling method of an oak natural secondary forest site index based on the random effect of site–climate interaction type. Of the site variables, elevation affected stand dominant height the most, followed by slope direction and position. Winter precipitation and summer mean maximum temperature had the greatest impact on stand dominant height. To develop the modeling method, we created 10 popular base models but found low performance (R2 ranged from 0.1731 to 0.2030). The optimal base model was Mitscherlich form M3 (R2 = 0.1940) based on parameter significance tests. Since site and climate factors affect the site index curve, the dominant site and climate factors were combined into site types and climate types, respectively, and a nonlinear mixed-effects approach was used to simulate different site types, climate types, site–climate interaction types, and their combinations as random effects. Site–climate interaction type as a random factor enhanced model (M3.4) performance and prediction accuracy (R2 from 0.1940 to 0.8220) compared to the optimum base model. After clustering the 62 site–climate interaction types into three, five, and eight groups using hierarchical clustering, a mixed-effects model with the random effects of eight groups improved model performance (R2 = 0.8265) and applicability. The modeling method developed in this study could be used to assess a regional secondary natural oak forest site index under complex site and climate variables to evaluate the forest productivity.

Recent studies have shown that organisms from the detritus food web subsidize generalist predators in aboveground food webs, but its significance in space and time is largely unknown. Here we report seasonal dynamics of aerial insects from grazing and detritus food webs in both forest and grassland habitats, and show how these patterns influence the dependence of web spiders on the detritus food web. Detrital insects were more abundant in spring, decreased in summer, and then increased slightly in autumn. This pattern was most conspicuous in Nematocera. Due to different seasonal activity patterns of grazing and detrital insects, the proportion of detrital insects was greater in spring and autumn. Detrital insects were relatively more abundant in the forest than in the grassland. Prey captured by web spiders generally reflected seasonal and spatial patterns of aerial insect abundance. In particular, Leucauge spiders reversed their dependence on the two food webs seasonally. Body size of spiders was negatively correlated with the proportion of detrital prey, suggesting that the detrital subsidy is essential for relatively small predators. This size effect probably resulted from interaction of the following two factors: 1) the maximum body size of prey that can be caught increased with spider body size, 2) larger body size classes of aerial insects included a higher proportion of insects from the grazing food web.

Although belowground food webs have received much attention, studies concerning microarthropods in nondetrital food webs are scarce. Because adult oribatid mites often number between 250,000–500,000/m2 in coniferous forests, microarthropods are a potential food resource for macroarthropod and vertebrate predators of the forest floor. Although the contribution of microarthropods to aboveground food webs has received little attention, sufficient data concerning macroarthropods and vertebrate predators were available at the Savannah River Site (SRS, Aiken, South Carolina) to construct a food web model of the various trophic interactions. To supplement this analysis, literature of microarthropod predation by arthropods and vertebrates was reviewed. This information was incorporated with the existing data to produce a model for taxa occurring in coniferous forests at the SRS. Because of the diversity and natural history of microarthropod predators, both vertebrate and invertebrate, the resulting web is quite connected and includes transfers to many trophic levels. The diets of arthropods and vertebrates are variable; yet feeding patterns reflect the relative abundance of prey at a place and time. Also, many predators feed on members of their own group. These factors suggest that belowground transfers are deserved of more attention in these and other forest food webs where substantial numbers of detritus feeding invertebrates inhabit the soil/litter interface and are available as prey items. Moreover, this model can be generalized to describe the dynamics of arthropod and vertebrate communities in other coniferous forests. The functioning of terrestrial ecosystems is dependent upon the interrelationships between aboveground and belowground food webs, and transfers of biotic components of the decomposer subsystem to aboveground consumers connect the two subsystems. It is hoped that those consumers traditionally associated with foliage-based food webs be reconsidered, as they may be gaining a proportion of their nutrition from organisms in the detrital pathway.

Ecological research demonstrates how intraspecific phenotypic variation can have consequences for community dynamics. However, the effects of animal personalities (i.e., intraspecific behavioral variation) on ecological processes remains relatively understudied. Using Red-backed Salamanders (Plethodon cinereus) and the detrital food web as a model system, we conducted a laboratory mesocosm experiment to explore whether or not the personality type of a top-level predator could affect the community structure within a complex, terrestrial food web. We used behavioral assays to investigate the repeatability of salamander behaviors, and classified individuals as either active or inactive. We then subjected laboratory mesocosms to one of four treatments for 3 mo: one active salamander, one inactive salamander, control (no salamander), and a pre-experimental reference. Our results indicate that the effect of P. cinereus on the detrital food web might be behaviorally mediated, with only the most active salamanders affecting community structure. Specifically, mesocosms housing active salamanders contained less diverse invertebrate communities than all other treatments. This difference was primarily driven by springtails, which were more abundant within mesocosms housing active salamanders. We also found that salamander personality was associated with cover board use inside mesocosms, with inactive salamanders exhibiting a more philopatric use of cover objects than active individuals. Leaf-litter degradation did not differ between treatments, indicating that the ecological effects of salamanders were too weak to influence basal resources within the detrital food web. Our study prompts further questions regarding the potential for animal personalities to influence ecological processes within terrestrial communities.

The trophic cascade effects of predators on detrital food webs have crucial impacts on ecosystem functioning. However, due to the complexity of detrital food webs, no consistent pattern of cascading effects has emerged. Furthermore, global changes, such as increased nitrogen (N) deposition, likely have significant impacts on detrital trophic interactions but are not well understood. Here, we evaluate how increased N deposition influences trophic cascades using a simplified detrital food web (spider–Collembola) that has previously exhibited trophic cascades in an experimental context. We tested the impact of N addition on the direct and cascading effects of spiders in detritus food web. We manipulated the abundance of spiders with sit‐and‐wait or actively hunting foraging strategies in microcosms. Under ambient N conditions (control treatments), we did not detect trophic cascade effects of spiders on litter decomposition. In contrast, N addition enhanced the negative trophic cascade effects of spiders on decomposition by directly reducing Collembola density. Our findings suggest that understanding how increased N deposition affects trophic interactions and ecosystem functioning is important for predicting the impact of global change on soil ecosystems.

Apex predators and plant resources are both critical for maintaining diversity in biotic communities, but the indirect (‘cascading’) effects of top‐down and bottom‐up forces on diversity at different trophic levels are not well resolved in terrestrial systems. Manipulations of predators or resources can cause direct changes of diversity at one trophic level, which in turn can affect diversity at other trophic levels. The indirect diversity effects of resource and consumer variation should be strongest in aquatic systems, moderate in terrestrial systems, and weakest in decomposer food webs. We measured effects of top predators and plant resources on the diversity of endophytic animals in an understorey shrubPiper cenocladum(Piperaceae). Predators and resource availability had significant direct and indirect effects on the diversity of the endophytic animal community, but the effects were not interactive, nor were they consistent between living vs. detrital food webs. The addition of fourth trophic level beetle predators increased diversity of consumers supported by living plant tissue, whereas balanced plant resources (light and nutrients) increased the diversity of primary through tertiary consumers in the detrital resources food web. These results support the hypotheses that top‐down and bottom‐up diversity cascades occur in terrestrial systems, and that diversity is affected by different factors in living vs. detrital food webs.

Impacts of Disturbance on Detritus Food Webs in Agro-Ecosystems of Contrasting Tillage and Weed Management Practices

Nearly all ecosystems are contaminated with highly toxic methylmercury (MeHg), but the specific sources and pathways leading to the uptake of MeHg within and among food webs are not well understood. In this study, we report stable mercury (Hg) isotope compositions in food webs in a river and an adjacent forest in northern California and demonstrate the utility of Hg isotopes for studying MeHg sources and cross-habitat transfers. We observed large differences in both δ(202)Hg (mass-dependent fractionation) and Δ(199)Hg (mass-independent fractionation) within both food webs. The majority of isotopic variation within each food web could be accounted for by differing proportions of inorganic Hg [Hg(II)] and MeHg along food chains. We estimated mean isotope values of Hg(II) and MeHg in each habitat and found a large difference in δ(202)Hg between Hg(II) and MeHg (∼2.7‰) in the forest but not in the river (∼0.25‰). This is consistent with in situ Hg(II) methylation in the study river but suggests Hg(II) methylation may not be important in the forest. In fact, the similarity in δ(202)Hg between MeHg in forest food webs and Hg(II) in precipitation suggests that MeHg in forest food webs may be derived from atmospheric sources (e.g., rainfall, fog). Utilizing contrasting δ(202)Hg values between MeHg in river food webs (-1.0‰) and MeHg in forest food webs (+0.7‰), we estimate with a two-source mixing model that ∼55% of MeHg in two riparian spiders is derived from riverine sources while ∼45% of MeHg originates from terrestrial sources. Thus, stable Hg isotopes can provide new information on subtle differences in sources of MeHg and trace MeHg transfers within and among food webs in natural ecosystems.

The effects of soil and topographic variables on forest site index were determined for two mesophytic tree species, northern red oak (Quercus rubraL.) and yellow-poplar (Liriodendron tulipiferaL.) in the Southern Appalachian Mountains of North Carolina. Stand variables included soil solum thickness, soil A-horizon thickness, elevation, aspect, slope gradient, and landform index. Landform index is a recently devised environmental variable that has been used to quantify the influence of topography surrounding a stand on productivity. Regression analysis indicated that among the variables only landform index had a significant () relationship with site index and explained 46 percent of the variation for northern red oak and 56 percent for yellow-poplar. Plot data from this study were also used to validate a previously developed prediction equation for estimating yellow-poplar site index and results indicated that unbiased estimates would be within 2.5 m. Results from this study suggest that landform accounts for variation in site index of mesophytic species in mountainous terrain that is not explained by conventional stand variables associated with soil and topography.