Effects of Grazer Exclusion on Carbon Cycling in Created Freshwater Wetlands

Wetland ecosystems play a significant role in the global carbon cycle, and yet are increasingly threatened by human development and climate change. The continued loss of intact freshwater wetlands heightens the need for effective wetland creation and restoration. However, wetland structure and function are controlled by interacting abiotic and biotic factors, complicating efforts to replace ecosystem services associated with natural wetlands and making ecologically-driven management imperative. Increasing waterfowl populations pose a threat to the development and persistence of created wetlands, largely through intensive grazing that can shift vegetation community structure or limit desired plant establishment. This study capitalized on a long-term herbivore exclusion experiment to evaluate how herbivore management impacts carbon cycling and storage in a created wetland in Western New York, USA. Vegetation, above- and belowground biomass, soil carbon, carbon gas fluxes and decomposition rates were evaluated in control plots with free access by large grazers and in plots where grazers had been excluded for four years. Waterfowl were the dominant herbivore at the site. Grazing reduced peak growing season aboveground biomass by over 55%, and during the summer, gross primary productivity doubled in grazer exclusion plots. The shift in plant productivity led to a 34% increase in soil carbon after exclusion of grazers for five growing seasons, but no change in belowground biomass. Our results suggest that grazers may inhibit the development of soil carbon pools during the first decade following wetland creation, reducing the carbon sequestration potential and precluding functional equivalence with natural wetlands.

Short-term plant-community responses to large mammalian herbivore exclusion in a rewilded Javan savanna

Grassy biomes such as savannas are maintained by an interacting suite of ecosystem processes from herbivory to rainfall to fire. Many studies have examined the impacts of large mammalian herbivores on herbaceous plant communities, but few of these studies have been conducted in humid, fertile savannas. We present the findings of a short-term experiment that investigated the effects of herbivory in a fertile, humid, and semi-managed savanna. We erected large-herbivore exclosures in Alas Purwo National Park, Java, Indonesia where rainfall is high and fire is suppressed to test how herbivores impact plant community development across the growing season. Where large mammalian herbivores were excluded, herbaceous plant communities contained more non-grasses and were less similar; diverging in their composition as the growing season progressed. Effects of herbivore exclusion on plant species richness, evenness, and biomass per quadrat were generally weak. Notably, however, two weedy plant species (one native, Imperata cylindrica and one introduced, Senna cf. tora) appeared to benefit most from herbivore release. Our results suggest that heavy grazing pressure by native large mammalian herbivores controlled the composition of the herbaceous plant community. Moreover, exclusion of large mammalian herbivores led to divergence in the plant species composition of exclosures; compositional dissimilarity between herbivore-exclusion plots was higher than between plots exposed to large mammalian herbivores. Our findings suggest that, at this high-rainfall site, large mammalian herbivores constrained the developmental trajectory of plant communities across the growing season.

The rich get richer and the poor get poorer: commonness and rarity of arctic tundra plants diverge under warming and herbivore exclusion

While most species are rare, our understanding of how rare species persist remains limited. Consequently, little is also known about how the commonness and rarity of co-occurring species might be differentially impacted by direct and indirect effects of climate change. We report results of a 15-year field experiment investigating effects on commonness and rarity of 14 arctic tundra plant taxa to warming and exclusion of large herbivores, factors demonstrated to have important effects on plant community composition in many biomes. Across all taxa, pooled commonness was reduced by experimental warming, and more strongly under herbivore exclusion than under herbivory. However, taxon-specific analyses revealed that although warming elicited variable effects on commonness, herbivore exclusion disproportionately reduced the commonness of rare taxa. Over the course of the experiment, we also observed trends in commonness and rarity under all treatments through time. Sitewide commonness increased for two common taxa, the deciduous shrubs Betula nana and Salix glauca, and declined in six other taxa, all of which were rare. Across experimental treatments, rates of increase and decline in commonness (i.e., temporal trends over the duration of the experiment) were strongly related to baseline commonness of taxa early in the experiment. Hence, commonness itself may be a strong predictor of plant species responses to climate change in the arctic tundra biome, but large herbivores may mediate such responses in rare taxa, perhaps facilitating their persistence.

Nutrients and herbivores impact grassland stability across multiple spatial scales through different pathways

Nutrients and herbivores have independent effects on the temporal stability of aboveground biomass in grasslands; however, their joint effects may not be additive and may also depend on spatial scales. In an experiment adding nutrients and excluding herbivores in 34 globally distributed grasslands, we found that nutrients and herbivores mainly had additive effects. Nutrient addition consistently reduced stability at the local and larger spatial scales (aggregated local communities), while herbivore exclusion weakly reduced stability at these scales. Moreover, nutrient addition reduced stability primarily by causing changes in local community composition over time and by reducing local species richness and evenness. In contrast, herbivore exclusion weakly reduced stability at the larger scale mainly by decreasing asynchronous dynamics among local communities, but also by weakly decreasing local species richness. Our findings indicate disentangling the influences of processes operating at different spatial scales may improve conservation and management in stabilizing grassland biomass.

Multiple resource limitations explain biomass-precipitation relationships in grasslands

Interannual variability in grassland primary production is strongly driven by precipitation, nutrient availability and herbivory, but there is no general consensus on the mechanisms linking these variables. If grassland biomass is limited by the single most limiting resource at a given time, then we expect that nutrient addition will not affect biomass production at arid sites. We conducted a distributed experiment manipulating nutrients and herbivores at 44 grassland sites in 8 regions around the world, spanning a broad range in aridity. We estimated the effects of 5-11 years of nutrient addition and herbivore exclusion treatments on precipitation sensitivity of biomass (proportional change in biomass relative to proportional change in rainfall among years), and the biomass in the driest year (to measure treatment effects when water was most limiting) at each site. Grazer exclusion did not interact with nutrients to influence driest year biomass or sensitivity. Nutrient addition increased driest year biomass by 74% and sensitivity by 0.12 (proportional units), and that effect did not change across the range of aridity spanned by our sites. Grazer exclusion did not interact with nutrients to influence sensitivity or driest year biomass. At almost half of our sites, the previous year's rainfall explained as much variation in biomass as current year precipitation. Overall, our distributed fertilization experiment detected co-limitation between nutrients and water governing grasslands, with biomass sensitivity to precipitation being limited by nutrient availability irrespective of site aridity and herbivory. Our findings refute the classical ideas that grassland plant performance is limited by the single most limiting resource at a site. This suggests that nutrient eutrophication will destabilize grassland ecosystems through increased sensitivity to precipitation variation.

Rapid relative increase of crustose coralline algae following herbivore exclusion in a reef of El Salvador

The Eastern Tropical Pacific (ETP) is one of the most isolated and least studied regions in the world. This particularly applies to the coast of El Salvador, where the only reef between Guatemala and Nicaragua, called Los Cóbanos reef, is located. There is very little published information about the reef’s biodiversity, and to our knowledge, no research on its ecology and responses to anthropogenic impacts, such as overfishing, has been conducted. The present study, therefore, described the benthic community of Los Cóbanos reef, El Salvador, using the Line-Point-Intercept-Transect method and investigated changes in the benthic community following the exclusion of piscine macroherbivores over a period of seven weeks. Results showed high benthic algae cover (up to 98%), dominated by turf and green algae, and low coral cover (0–4%). Porites lobata was the only hermatypic coral species found during the surveys. Surprisingly, crustose coralline algae (CCA) showed a remarkable total cover increase by 58%, while turf algae cover decreased by 82%, in experimental plots after seven weeks of piscine macroherbivore exclusion. These findings apparently contradict the results of most previous similar studies. While it was not possible to ascertain the exact mechanisms leading to these drastic community changes, the most likely explanation is grazing on turf by small grazing macroherbivores that had access to the cages during the experiment and clearing of CCA initially covered by epiphytes and sediments. A higher CCA cover would promote the succesful settlement by corals and prevent further erosion of the reef framework. Therefore it is crucial to better understand algal dynamics, herbivory, and implications of overfishing at Los Cóbanos to avoid further reef deterioration. This could be achieved through video surveys of the fish community, night-time observations of the macroinvertebrate community, exclusion experiments that also keep out herbivorous macroinvertebrates, and/or experimental assessments of turf algae/CCA interactions.

Herbivory and warming interact in opposing patterns of covariation between arctic shrub species at large and local scales

A major challenge in predicting species’ distributional responses to climate change involves resolving interactions between abiotic and biotic factors in structuring ecological communities. This challenge reflects the classical conceptualization of species’ regional distributions as simultaneously constrained by climatic conditions, while by necessity emerging from local biotic interactions. A ubiquitous pattern in nature illustrates this dichotomy: potentially competing species covary positively at large scales but negatively at local scales. Recent theory poses a resolution to this conundrum by predicting roles of both abiotic and biotic factors in covariation of species at both scales, but empirical tests have lagged such developments. We conducted a 15-y warming and herbivore-exclusion experiment to investigate drivers of opposing patterns of covariation between two codominant arctic shrub species at large and local scales. Climatic conditions and biotic exploitation mediated both positive covariation between these species at the landscape scale and negative covariation between them locally. Furthermore, covariation between the two species conferred resilience in ecosystem carbon uptake. This study thus lends empirical support to developing theoretical solutions to a long-standing ecological puzzle, while highlighting its relevance to understanding community compositional responses to climate change.