Although invasive plants are a major source of terrestrial ecosystem degradation worldwide, it remains unclear which trophic levels above the base of the food web are most vulnerable to plant invasions. We performed a meta-analysis of 38 independent studies from 32 papers to examine how invasive plants alter major groupings of primary and secondary consumers in three globally distributed ecosystems: wetlands, woodlands and grasslands. Within each ecosystem we examined if green (grazing) food webs are more sensitive to plant invasions compared to brown (detrital) food webs. Invasive plants have strong negative effects on primary consumers (detritivores, bacterivores, fungivores, and/or herbivores) in woodlands and wetlands, which become less abundant in both green and brown food webs in woodlands and green webs in wetlands. Plant invasions increased abundances of secondary consumers (predators and/or parasitoids) only in woodland brown food webs and green webs in wetlands. Effects of invasive plants on grazing and detrital food webs clearly differed between ecosystems. Overall, invasive plants had the most pronounced effects on the trophic structure of wetlands and woodlands, but caused no detectable changes to grassland trophic structure.
1. Functional traits are useful for characterizing variation in community and ecosystem dynamics. Most advances in trait-based ecology to date centre on plant functional traits, although there is an increasing recognition that animal traits are also key contributors to processes operating at the community or ecosystem scale.2. Terrestrial invertebrates are incredibly diverse and ubiquitous animals with important roles in nutrient cycling. Despite their widespread influence on ecosystem processes, we currently lack a synthetic understanding of how invertebrate functional traits affect terrestrial nutrient cycling.3. We present a meta-analysis of 511 paired observations from 122 papers that examined how invertebrate functional traits affected litter decomposition rates, nitrogen pools and litter C:N ratios. Based on the available data, we specifically assessed the effects of feeding mode (bioturbation, detritus shredding, detritus grazing, leaf chewing, leaf piercing, ambush predators, active hunting predators) and body size (macro-and micro-invertebrates) on nutrient cycling.4. The effects of invertebrates on terrestrial nutrient cycling varied according to functional trait. The inclusion of both macro-(≥2 mm) and micro-invertebrates (<2 mm) increased litter decomposition by 20% and 19%, respectively. All detritivorous feeding modes enhanced litter decomposition rates, with bioturbators, detritus shredders and detritus grazers increasing decomposition by 28%, 22% and 15%, respectively. Neither herbivore feeding mode (e.g. leaf chewers and leaf piercers) nor predator hunting mode (ambush and active hunting) affected decomposition. We also revealed that bioturbators and detritus grazers increased soil nitrogen availability by 99% and 70%, respectively, and that leaf-chewing herbivores had a weak effect on litterfall stoichiometry via reducing C:N ratios by 11%.5. Although functional traits might be useful predictors of ecosystem processes, our findings suggest context-dependent effects of invertebrate traits on terrestrial nutrient cycling. Detritivore functional traits (i.e. bioturbators, detritus shredders and detritus grazers) are more consistent with increased rates of nutrient cycling, whereas our currently characterized predator and herbivore traits are less predictive. Future research is needed to identify, standardize and deliberately study the impacts of invertebrate functional traits on nutrient cycling in hopes of revealing the key functional traits governing ecosystem functioning worldwide.
Pulsed fluxes of organisms across ecosystem boundaries can exert top‐down and bottom‐up effects in recipient food webs, through both direct effects on the subsidized trophic levels and indirect effects on other components of the system. While previous theoretical and empirical studies demonstrate the influence of allochthonous subsidies on bottom‐up and top‐down processes, understanding how these forces act in conjunction is still limited, particularly when an allochthonous resource can simultaneously subsidize multiple trophic levels. Using the Lake Mývatn region in Iceland as an example system of allochthony and its potential effects on multiple trophic levels, we analyzed a mathematical model to evaluate how pulsed subsidies of aquatic insects affect the dynamics of a soil–plant–arthropod food web. We found that the relative balance of top‐down and bottom‐up effects on a given food web compartment was determined by trophic position, subsidy magnitude, and top predators’ ability to exploit the subsidy. For intermediate trophic levels (e.g., detritivores and herbivores), we found that the subsidy could either alleviate or intensify top‐down pressure from the predator. For some parameter combinations, alleviation and intensification occurred sequentially during and after the resource pulse. The total effect of the subsidy on detritivores and herbivores, including top‐down and bottom‐up processes, was determined by the rate at which predator consumption saturated with increasing size of the allochthonous subsidy, with greater saturation leading to increased bottom‐up effects. Our findings illustrate how resource pulses to multiple trophic levels can influence food web dynamics by changing the relative strength of bottom‐up and top‐down effects, with bottom‐up predominating top‐down effects in most scenarios in this subarctic system.
Animals can be important vectors of nutrient transfer within and across landscapes, with important implications for ecosystem productivity and composition. While it is presumed large ungulates are agents of nutrient dispersal via movement and activity, research analyzing their net effects on landscapes remains scarce. We present an individual‐based model that investigates how caribou affect the distribution of nutrients on a landscape through consumption only, as well through the cumulative effects of consumption and nutrient deposition (i.e. fecal waste and carcass deposition). We explored these dynamics in simulations that altered the context of environments, either initially containing heterogeneous or homogeneous nutrient distributions, animal densities and sociality of caribou behavior. In the consumption‐only simulations, caribou density and sociality created different patterns of heterogeneity at both the landscape and local scale depending on the initial landscape conditions. In these simulations, caribou populations crashed at high densities because the lack of animal deposition resulted in low resources across the landscape. This was not the case when considering the cumulative effects of consumption and deposition, indicating the return of nutrients from animals may be important for population stability. Additionally, in simulations considering the cumulative effects of caribou, increasing caribou density increased landscape heterogeneity irrespective of the initial condition (i.e. heterogeneous and homogeneous landscapes), and maintained or increased local heterogeneity in heterogeneous and homogenous landscape, respectively. Importantly, in all simulations the net impact of caribou at the individual patch level was extremely variable, suggesting that animal inputs are highly varied throughout the landscape. Our results indicate the movement of large ungulates such as caribou can increase the heterogeneity of available nutrients within a landscape and provide an important feedback for population stability. Thus, the loss of large ungulates from natural ecosystems via anthropogenic activity is likely to result in less heterogeneous natural landscapes.
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