Modeling foundation species in food webs

Foundation species are basal species that play an important role in determining community composition by physically structuring ecosystems and modulating ecosystem processes. Foundation species largely operate via non-trophic interactions, presenting a challenge to incorporating them into food web models. Here, we used non-linear, bioenergetic predator-prey models to explore the role of foundation species and their non-trophic effects. We explored four types of models in which the foundation species reduced the metabolic rates of species in a specific trophic position. We examined the outcomes of each of these models for six metabolic rate “treatments” in which the foundation species altered the metabolic rates of associated species by one-tenth to ten times their allometric baseline metabolic rates. For each model simulation, we looked at how foundation species influenced food web structure during community assembly and the subsequent change in food web structure when the foundation species was removed. When a foundation species lowered the metabolic rate of only basal species, the resultant webs were complex, species-rich, and robust to foundation species removals. On the other hand, when a foundation species lowered the metabolic rate of only consumer species, all species, or no species, the resultant webs were species-poor and the subsequent removal of the foundation species resulted in the further loss of species and complexity. This suggests that in nature we should look for foundation species to predominantly facilitate basal species.Organismic and Evolutionary Biolog

Species richness and phylogenetic diversity of native and non-native species respond differently to area and environmental factors

© 2018 John Wiley & Sons Ltd Aim: To test whether native and non-native species have similar diversity–area relationships (species–area relationships [SARs] and phylogenetic diversity–area relationships [PDARs]) and whether they respond similarly to environmental variables. Location: United States. Methods: Using lists of native and non-native species as well as environmental variables for \u3e250 US national parks, we compared SARs and PDARs of native and non-native species to test whether they respond similarly to environmental conditions. We then used multiple regressions involving climate, land cover and anthropogenic variables to further explore underlying predictors of diversity for plants and birds in US national parks. Results: Native and non-native species had different slopes for SARs and PDARs, with significantly higher slopes for native species. Corroborating this pattern, multiple regressions showed that native and non-native diversity of plants and birds responded differently to a greater number of environmental variables than expected by chance. For native species richness, park area and longitude were the most important variables while the number of park visitors, temperature and the percentage of natural area were among the most important ones for non-native species richness. Interestingly, the most important predictor of native and non-native plant phylogenetic diversity, temperature, had positive effects on non-native plants but negative effects on natives. Main conclusions: SARs, PDARs and multiple regressions all suggest that native and non-native plants and birds responded differently to environmental factors that influence their diversity. The agreement between diversity–area relationships and multiple regressions with environmental variables suggests that SARs and PDARs can be both used as quick proxies of overall responses of species to environmental conditions. However, more importantly, our results suggest that global change will have different effects on native and non-native species, making it inappropriate to apply the large body of knowledge on native species to understand patterns of community assembly of non-native species

Organic-Matter Loading Determines Regime Shifts and Alternative States in an Aquatic Ecosystem

Slow changes in underlying state variables can lead to “tipping points,” rapid transitions between alternative states (“regime shifts”) in a wide range of complex systems. Tipping points and regime shifts routinely are documented retrospectively in long time series of observational data. Experimental induction of tipping points and regime shifts is rare, but could lead to new methods for detecting impending tipping points and forestalling regime shifts. By using controlled additions of detrital organic matter (dried, ground arthropod prey), we experimentally induced a shift from aerobic to anaerobic states in a miniature aquatic ecosystem: the self-contained pools that form in leaves of the carnivorous northern pitcher plant, Sarracenia purpurea. In unfed controls, the concentration of dissolved oxygen ($[O_2]$) in all replicates exhibited regular diurnal cycles associated with daytime photosynthesis and nocturnal plant respiration. In low prey-addition treatments, the regular diurnal cycles of $[O_2]$ were disrupted, but a regime shift was not detected. In high prey-addition treatments, the variance of the $[O_2]$ time series increased until the system tipped from an aerobic to an anaerobic state. In these treatments, replicate $[O_2]$ time series predictably crossed a tipping point at ∼45 h as $[O_2]$ was decoupled from diurnal cycles of photosynthesis and respiration. Increasing organic-matter loading led to predictable changes in $[O_2]$ dynamics, with high loading consistently driving the system past a well-defined tipping point. The Sarracenia microecosystem functions as a tractable experimental system in which to explore the forecasting and management of tipping points and alternative regimes.Organismic and Evolutionary Biolog

Identifying foundation species in North American forests using long‐term data on ant assemblage structure

Foundation species are locally abundant and uniquely control associated biodiversity, whereas dominant species are locally abundant but are thought to be replaceable in ecological systems. It is important to distinguish foundation from dominant species to direct conservation efforts. Long‐term studies that remove abundant species while measuring community dynamics have the potential to (1) aid in the identification of foundation vs. dominant species and, (2) once a foundation species is identified, determine how long its effects persist within a community after its loss. Long‐term data on ant assemblages within two canopy‐manipulation experiments—the Harvard Forest Hemlock Removal Experiment (HF‐HeRE) and the Black Rock Future of Oak Forests Experiment (BRF‐FOFE)—provide insights into how ant assemblages change and reassemble following the loss of Tsuga canadensis or Quercus spp. Previous research documented foundation species effects on ants in the HF‐HeRE for up to four years after T. canadensis loss. Six additional years of data at HF‐HeRE presented for the first time here show that removal of T. canadensis resulted in taxonomic and some measures of functional shifts in ant assemblages that persisted for ten years, further supporting the hypothesis that T. canadensis is a foundation species at Harvard Forest. In contrast, ant assemblages at BRF‐FOFE varied little regardless of whether oaks or other tree species were removed from the canopy, suggesting that Quercusspecies do not act as foundation species at Black Rock Forest. Deer and moose exclosures within each experiment also allowed for comparisons between effects on ants of foundation or dominant tree species relative to effects of large herbivores. At HF‐HeRE, effects of T. canadensis were stronger than effects of large herbivores on taxonomic and functional diversity of ant assemblages. At BRF‐FOFE, in contrast, effects of Quercus species were weaker than effects of large herbivores on ant taxonomic diversity and some measures of ant functional diversity. These findings illustrate the importance of distinguishing between the roles of irreplaceable foundation species and replaceable dominant ones in forested ecosystems along with other drivers of biodiversity (e.g., herbivory)

Predicting food-web structure with metacommunity models

Synthesis Metacommunity theory aims to elucidate the relative influence of local and regional-scale processes in generating diversity patterns across the landscape. Metacommunity research has focused largely on assemblages of competing organisms within a single trophic level. Here, we test the ability of metacommunity models to predict the network structure of the aquatic food web found in the leaves of the northern pitcher plant Sarracenia purpurea. The species-sorting and patch-dynamics models most accurately reproduced nine food web properties, suggesting that local-scale interactions play an important role in structuring Sarracenia food webs. Our approach can be applied to any well-resolved food web for which data are available from multiple locations. The metacommunity framework explores the relative influence of local and regional-scale processes in generating diversity patterns across the landscape. Metacommunity models and empirical studies have focused mostly on assemblages of competing organisms within a single trophic level. Studies of multi-trophic metacommunities are predominantly restricted to simplified trophic motifs and rarely consider entire food webs. We tested the ability of the patch-dynamics, species-sorting, mass-effects, and neutral metacommunity models, as well as three hybrid models, to reproduce empirical patterns of food web structure and composition in the complex aquatic food web found in the northern pitcher plant Sarracenia purpurea. We used empirical data to determine regional species pools and estimate dispersal probabilities, simulated local food-web dynamics, dispersed species from regional pools into local food webs at rates based on the assumptions of each metacommunity model, and tested their relative fits to empirical data on food-web structure. The species-sorting and patch-dynamics models most accurately reproduced nine food web properties, suggesting that local-scale interactions were important in structuring Sarracenia food webs. However, differences in dispersal abilities were also important in models that accurately reproduced empirical food web properties. Although the models were tested using pitcher-plant food webs, the approach we have developed can be applied to any well-resolved food web for which data are available from multiple locations. © 2012 The Authors. Oikos © 2012 Nordic Society Oikos

Response of macroarthropod assemblages to the loss of hemlock (Tsuga canadensis), a foundation species

In eastern North American forests, eastern hemlock (Tsuga canadensis) is a foundation species. As hemlock is lost from forests due to the invasive hemlock woolly adelgid (Adelges tsugae) and preemptive salvage logging, the structure of assemblages of species associated with hemlock is expected to change. We manipulated hemlock canopy structure at hectare scales to investigate the effects of hemlock death on assemblages of ants, beetles, and spiders in a New England forest. Relative to reference hemlock stands, both in situ death of hemlock and logging and removal of hemlock altered composition and diversity of beetles and spiders, and logging increased the species richness and evenness of ant assemblages. Species composition of ant assemblages in disturbed habitats was non-random relative to the regional species pool, but we found no evidence that interspecific competition shaped the structure of ant, beetle, or spider assemblages, in either manipulated or intact forest stands. Environmental filtering by hemlock appears to maintain low levels of species richness and evenness in forest stands, suggesting that the loss of hemlock due to the hemlock woolly adelgid or human activities will not likely lead to extirpations of ant, beetle, or spider species at local scales

Geographic Variation in Network Structure of a Nearctic Aquatic Food Web

Aim: The network structure of food webs plays an important role in the maintenance of diversity and ecosystem functioning in ecological communities. Previous research has found that ecosystem size, resource availability, assembly history and biotic interactions can potentially drive food web structure. However, the relative influence of climatic variables that drive broad-scale biogeographic patterns of species richness and composition has not been explored for food web structure. In this study, we assess the influence of broad-scale climatic variables in addition to known drivers of food web structure on replicate observations of a single aquatic food web, sampled from the leaves of the pitcher plant (Sarracenia purpurea), at different geographic sites across a broad latitudinal and climatic range. Location: Using standardized sampling methods, we conducted an extensive ‘snapshot’ survey of 780 replicated aquatic food webs collected from the leaves of the pitcher plant S. purpurea at 39 sites from northern Florida to Newfoundland and westward to eastern British Columbia. Methods: We examined correlations of 15 measures of food web structure at the pitcher and site scales with geographic variation in temperature and precipitation, concentrations of nutrients from atmospheric nitrogen deposition, resource availability, ecosystem size and the abundance of the pitcher plant mosquito (Wyeomyia smithii), a potential keystone species. Results: At the scale of a single pitcher plant leaf, linkage density, species richness, measures of chain length and the proportion of omnivores in a web all increased with pitcher volume. Linkage density and species richness were greater at high-latitude sites, which experience low mean temperatures and precipitation and high annual variation in both of these variables. At the site scale, variation in 8 of the 15 food web metrics decreased at higher latitudes, and variation in measures of chain length increased with the abundance of mosquitoes. Main conclusions: Ecosystem size and climatic variables related to latitude were most strongly correlated with network structure of the Sarracenia food web. However, in spite of large sample sizes, thorough standardized sampling and the large geographic extent of the survey, even the best-fitting models explained less than 40% of the variation in food web structure. In contrast to biogeographic patterns of species richness, food web structure was largely independent of broad-scale climatic variables. The large proportion of unexplained variance in our analyses suggests that stochastic assembly may be an important determinant of local food web structure.Organismic and Evolutionary Biolog

Response of Macroarthropod Assemblages to the Loss of Hemlock (Tsuga canadensis), a Foundation Species

In eastern North American forests, eastern hemlock (Tsuga canadensis) is a foundation species. As hemlock is lost from forests due to the invasive hemlock woolly adelgid (Adelges tsugae) and pre-emptive salvage logging, the structure of assemblages of species associated with hemlock is expected to change. We manipulated hemlock canopy structure at hectare scales to investigate the effects of hemlock death on assemblages of ants, beetles, and spiders in a New England forest. Relative to reference hemlock stands, both in situ death of hemlock and logging and removal of hemlock altered composition and diversity of beetles and spiders, and logging increased the species richness and evenness of ant assemblages. Species composition of ant assemblages in disturbed habitats was non-random relative to the regional species pool, but we found no evidence that interspecific competition shaped the structure of ant, beetle, or spider assemblages, in either manipulated or intact forest stands. Environmental filtering by hemlock appears to maintain low levels of species richness and evenness in forest stands, suggesting that the loss of hemlock due to the hemlock woolly adelgid or human activities will not likely lead to extirpations of ant, beetle, or spider species at local scales.Organismic and Evolutionary BiologyOther Research Uni

Response of macroarthropod assemblages to the loss of hemlock (Tsuga canadensis), a foundation species

In eastern North American forests, eastern hemlock (Tsuga canadensis) is a foundation species. As hemlock is lost from forests due to the invasive hemlock woolly adelgid (Adelges tsugae) and preemptive salvage logging, the structure of assemblages of species associated with hemlock is expected to change. We manipulated hemlock canopy structure at hectare scales to investigate the effects of hemlock death on assemblages of ants, beetles, and spiders in a New England forest. Relative to reference hemlock stands, both in situ death of hemlock and logging and removal of hemlock altered composition and diversity of beetles and spiders, and logging increased the species richness and evenness of ant assemblages. Species composition of ant assemblages in disturbed habitats was non-random relative to the regional species pool, but we found no evidence that interspecific competition shaped the structure of ant, beetle, or spider assemblages, in either manipulated or intact forest stands. Environmental filtering by hemlock appears to maintain low levels of species richness and evenness in forest stands, suggesting that the loss of hemlock due to the hemlock woolly adelgid or human activities will not likely lead to extirpations of ant, beetle, or spider species at local scales