Human activities are transforming grassland biomass via changing climate, elemental nutrients, and herbivory. Theory predicts that food-limited herbivores will consume any additional biomass ...stimulated by nutrient inputs ('consumer-controlled'). Alternatively, nutrient supply is predicted to increase biomass where herbivores alter community composition or are limited by factors other than food ('resource-controlled'). Using an experiment replicated in 58 grasslands spanning six continents, we show that nutrient addition and vertebrate herbivore exclusion each caused sustained increases in aboveground live biomass over a decade, but consumer control was weak. However, at sites with high vertebrate grazing intensity or domestic livestock, herbivores consumed the additional fertilization-induced biomass, supporting the consumer-controlled prediction. Herbivores most effectively reduced the additional live biomass at sites with low precipitation or high ambient soil nitrogen. Overall, these experimental results suggest that grassland biomass will outstrip wild herbivore control as human activities increase elemental nutrient supply, with widespread consequences for grazing and fire risk.
Gastric acidity is likely a key factor shaping the diversity and composition of microbial communities found in the vertebrate gut. We conducted a systematic review to test the hypothesis that a key ...role of the vertebrate stomach is to maintain the gut microbial community by filtering out novel microbial taxa before they pass into the intestines. We propose that species feeding either on carrion or on organisms that are close phylogenetic relatives should require the most restrictive filter (measured as high stomach acidity) as protection from foreign microbes. Conversely, species feeding on a lower trophic level or on food that is distantly related to them (e.g. herbivores) should require the least restrictive filter, as the risk of pathogen exposure is lower. Comparisons of stomach acidity across trophic groups in mammal and bird taxa show that scavengers and carnivores have significantly higher stomach acidities compared to herbivores or carnivores feeding on phylogenetically distant prey such as insects or fish. In addition, we find when stomach acidity varies within species either naturally (with age) or in treatments such as bariatric surgery, the effects on gut bacterial pathogens and communities are in line with our hypothesis that the stomach acts as an ecological filter. Together these results highlight the importance of including measurements of gastric pH when investigating gut microbial dynamics within and across species.
Background Herbivores modify the structure and function of tundra ecosystems. Understanding their impacts is necessary to assess the responses of these ecosystems to ongoing environmental changes. ...However, the effects of herbivores on plants and ecosystem structure and function vary across the Arctic. Strong spatial variation in herbivore effects implies that the results of individual studies on herbivory depend on local conditions, i.e., their ecological context. An important first step in assessing whether generalizable conclusions can be produced is to identify the existing studies and assess how well they cover the underlying environmental conditions across the Arctic. This systematic map aims to identify the ecological contexts in which herbivore impacts on vegetation have been studied in the Arctic. Specifically, the primary question of the systematic map was: "What evidence exists on the effects of herbivores on Arctic vegetation?". Methods We used a published systematic map protocol to identify studies addressing the effects of herbivores on Arctic vegetation. We conducted searches for relevant literature in online databases, search engines and specialist websites. Literature was screened to identify eligible studies, defined as reporting primary data on herbivore impacts on Arctic plants and plant communities. We extracted information on variables that describe the ecological context of the studies, from the studies themselves and from geospatial data. We synthesized the findings narratively and created a Shiny App where the coded data are searchable and variables can be visually explored. Review findings We identified 309 relevant articles with 662 studies (representing different ecological contexts or datasets within the same article). These studies addressed vertebrate herbivory seven times more often than invertebrate herbivory. Geographically, the largest cluster of studies was in Northern Fennoscandia. Warmer and wetter parts of the Arctic had the largest representation, as did coastal areas and areas where the increase in temperature has been moderate. In contrast, studies spanned the full range of ecological context variables describing Arctic vertebrate herbivore diversity and human population density and impact. Conclusions The current evidence base might not be sufficient to understand the effects of herbivores on Arctic vegetation throughout the region, as we identified clear biases in the distribution of herbivore studies in the Arctic and a limited evidence base on invertebrate herbivory. In particular, the overrepresentation of studies in areas with moderate increases in temperature prevents robust generalizations about the effects of herbivores under different climatic scenarios. Keywords: Browsing, Grazing, Grubbing, Defoliation, Tundra, Invertebrate, Vertebrate, Forest-tundra, Plant-herbivore interaction
Understanding and predicting large-scale ecological responses to global environmental change requires comparative studies across geographic scales with coordinated efforts and standardized ...methodologies. We designed, applied, and assessed standardized protocols to measure tundra herbivory at three spatial scales: plot, site (habitat), and study area (landscape). The plot- and site-level protocols were tested in the field during summers 2014–2015 at 11 sites, nine of them consisting of warming experimental plots included in the International Tundra Experiment (ITEX). The study area protocols were assessed during 2014–2018 at 24 study areas across the Arctic. Our protocols provide comparable and easy to implement methods for assessing the intensity of invertebrate herbivory within ITEX plots and for characterizing vertebrate herbivore communities at larger spatial scales. We discuss methodological constraints and make recommendations for how these protocols can be used and how sampling effort can be optimized to obtain comparable estimates of herbivory, both at ITEX sites and at large landscape scales. The application of these protocols across the tundra biome will allow characterizing and comparing herbivore communities across tundra sites and at ecologically relevant spatial scales, providing an important step towards a better understanding of tundra ecosystem responses to large-scale environmental change.
Time series data on arthropod populations are critical for understanding the magnitude, direction, and drivers of change. However, most arthropod monitoring programs are short-lived and restricted in ...taxonomic resolution. Monitoring data from the Arctic are especially underrepresented, yet critical to uncovering and understanding some of the earliest biological responses to rapid environmental change. Clear imprints of climate on the behavior and life history of some Arctic arthropods have been demonstrated, but a synthesis of population-level abundance changes across taxa is lacking. We utilized 24 y of abundance data from Zackenberg in High-Arctic Greenland to assess trends in abundance and diversity and identify potential climatic drivers of abundance changes. Unlike findings from temperate systems, we found a nonlinear pattern, with total arthropod abundance gradually declining during 1996 to 2014, followed by a sharp increase. Family-level diversity showed the opposite pattern, suggesting increasing dominance of a small number of taxa. Total abundance masked more complicated trajectories of family-level abundance, which also frequently varied among habitats. Contrary to expectation in this extreme polar environment, winter and fall conditions and positive density-dependent feedbacks were more common determinants of arthropod dynamics than summer temperature. Together, these data highlight the complexity of characterizing climate change responses even in relatively simple Arctic food webs. Our results underscore the need for data reporting beyond overall trends in biomass or abundance and for including basic research on life history and ecology to achieve a more nuanced understanding of the sensitivity of Arctic and other arthropods to global changes.
Rapid climate change in the Arctic is altering microbial structure and function, with important consequences for the global ecosystem. Emerging evidence suggests organisms in higher trophic levels ...may also influence microbial communities, but whether warming alters these effects is unclear. Wolf spiders are dominant Arctic predators whose densities are expected to increase with warming. These predators have temperature-dependent effects on decomposition via their consumption of fungal-feeding detritivores, suggesting they may indirectly affect the microbial structure as well. To address this, we used a fully factorial mesocosm experiment to test the effects of wolf spider density and warming on litter microbial structure in Arctic tundra. We deployed replicate litter bags at the surface and belowground in the organic soil profile and analyzed the litter for bacterial and fungal community structure, mass loss, and nutrient characteristics after 2 and 14 months. We found there were significant interactive effects of wolf spider density and warming on fungal but not bacterial communities. Specifically, higher wolf spider densities caused greater fungal diversity under ambient temperature but lower fungal diversity under warming at the soil surface. We also observed interactive treatment effects on fungal composition belowground. Wolf spider density influenced surface bacterial composition, but the effects did not change with warming. These findings suggest a widespread predator can have indirect, cascading effects on litter microbes and that effects on fungi specifically shift under future expected levels of warming. Overall, our study highlights that trophic interactions may play important, albeit overlooked, roles in driving microbial responses to warming in Arctic terrestrial ecosystems.IMPORTANCEThe Arctic contains nearly half of the global pool of soil organic carbon and is one of the fastest warming regions on the planet. Accelerated decomposition of soil organic carbon due to warming could cause positive feedbacks to climate change through increased greenhouse gas emissions; thus, changes in ecological dynamics in this region are of global relevance. Microbial structure is an important driver of decomposition and is affected by both abiotic and biotic conditions. Yet how activities of soil-dwelling organisms in higher trophic levels influence microbial structure and function is unclear. In this study, we demonstrate that predicted changes in abundances of a dominant predator and warming interactively affect the structure of litter-dwelling fungal communities in the Arctic. These findings suggest predators may have widespread, indirect cascading effects on microbial communities, which could influence ecosystem responses to future climate change.
Arctic terrestrial herbivores influence tundra carbon and nutrient dynamics through their consumption of resources, waste production, and habitat‐modifying behaviors. The strength of these effects is ...likely to change spatially and temporally as climate change drives shifts in herbivore abundance, distribution, and activity timing. Here, we review how herbivores influence tundra carbon and nutrient dynamics through their consumptive and nonconsumptive effects. We also present evidence for herbivore responses to climate change and discuss how these responses may alter the spatial and temporal distribution of herbivore impacts. Several current knowledge gaps limit our understanding of the changing functional roles of herbivores; these include limited characterization of the spatial and temporal variability in herbivore impacts and of how herbivore activities influence the cycling of elements beyond carbon. We conclude by highlighting approaches that will promote better understanding of herbivore effects on tundra ecosystems, including their integration into existing biogeochemical models, new applications of remote sensing techniques, and the continued use of distributed experiments.
Here we review how herbivores influence tundra carbon and nutrient dynamics through their consumptive and non‐consumptive effects. We also present evidence for herbivore responses to climate change and discuss how these responses may alter the spatial and temporal distribution of herbivore impacts. We conclude by highlighting several approaches that will promote better understanding of herbivore effects on tundra ecosystems, including their integration into existing biogeochemical models, new applications of remote sensing techniques, and the continued use of distributed experiments.
The Arctic is experiencing some of the fastest rates of warming on the planet. Although many studies have documented responses to such warming by individual species, the idiosyncratic nature of these ...findings has prevented us from extrapolating them to community-level predictions. Here, we leverage the availability of a long-term dataset from Zackenberg, Greenland (593 700 specimens collected between 1996 and 2014), to investigate how climate parameters influence the abundance of different arthropod groups and overall community composition. We find that variation in mean seasonal temperatures, winter duration and winter freeze–thaw events is correlated with taxon-specific and habitat-dependent changes in arthropod abundances. In addition, we find that arthropod communities have exhibited compositional changes consistent with the expected effects of recent shifts towards warmer active seasons and fewer freeze–thaw events in NE Greenland. Changes in community composition are up to five times more extreme in drier than wet habitats, with herbivores and parasitoids generally increasing in abundance, while the opposite is true for surface detritivores. These results suggest that species interactions and food web dynamics are changing in the Arctic, with potential implications for key ecosystem processes such as decomposition, nutrient cycling and primary productivity.
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