Kevin R. Wilcox

Contrasting above‐ and belowground sensitivity of three Great Plains grasslands to altered rainfall regimes

Intensification of the global hydrological cycle with atmospheric warming is expected to increase interannual variation in precipitation amount and the frequency of extreme precipitation events. Although studies in grasslands have shown sensitivity of aboveground net primary productivity (ANPP) to both precipitation amount and event size, we lack equivalent knowledge for responses of belowground net primary productivity (BNPP) and NPP. We conducted a 2-year experiment in three US Great Plains grasslands--the C4-dominated shortgrass prairie (SGP; low ANPP) and tallgrass prairie (TGP; high ANPP), and the C3-dominated northern mixed grass prairie (NMP; intermediate ANPP)--to test three predictions: (i) both ANPP and BNPP responses to increased precipitation amount would vary inversely with mean annual precipitation (MAP) and site productivity; (ii) increased numbers of extreme rainfall events during high-rainfall years would affect high and low MAP sites differently; and (iii) responses belowground would mirror those aboveground. We increased growing season precipitation by as much as 50% by augmenting natural rainfall via (i) many (11-13) small or (ii) fewer (3-5) large watering events, with the latter coinciding with naturally occurring large storms. Both ANPP and BNPP increased with water addition in the two C4 grasslands, with greater ANPP sensitivity in TGP, but greater BNPP and NPP sensitivity in SGP. ANPP and BNPP did not respond to any rainfall manipulations in the C3 -dominated NMP. Consistent with previous studies, fewer larger (extreme) rainfall events increased ANPP relative to many small events in SGP, but event size had no effect in TGP. Neither system responded consistently above- and belowground to event size; consequently, total NPP was insensitive to event size. The diversity of responses observed in these three grassland types underscores the challenge of predicting responses relevant to C cycling to forecast changes in precipitation regimes even within relatively homogeneous biomes such as grasslands.

Plant community response to loss of large herbivores differs between North American and South African savanna grasslands

Herbivory and fire shape plant community structure in grass-dominated ecosystems, but these disturbance regimes are being altered around the world. To assess the consequences of such alterations, we excluded large herbivores for seven years from mesic savanna grasslands sites burned at different frequencies in North America (Konza Prairie Biological Station, Kansas, USA) and South Africa (Kruger National Park). We hypothesized that the removal of a single grass-feeding herbivore from Konza would decrease plant community richness and shift community composition due to increased dominance by grasses. Similarly, we expected grass dominance to increase at Kruger when removing large herbivores, but because large herbivores are more diverse, targeting both grasses and forbs, at this study site, the changes due to herbivore removal would be muted. After seven years of large-herbivore exclusion, richness strongly decreased and community composition changed at Konza, whereas little change was evident at Kruger. We found that this divergence in response was largely due to differences in the traits and numbers of dominant grasses between the study sites rather than the predicted differences in herbivore assemblages. Thus, the diversity of large herbivores lost may be less important in determining plant community dynamics than the functional traits of the grasses that dominate mesic, disturbance-maintained savanna grasslands.

Stoichiometric homeostasis predicts plant species dominance, temporal stability, and responses to global change

Why some species are consistently more abundant than others, and predicting how species will respond to global change, are fundamental questions in ecology. Long-term observations indicate that plant species with high stoichiometric homeostasis for nitrogen (HN), i.e., the ability to decouple foliar N levels from variation in soil N availability, were more common and stable through time than low-HN species in a central U.S. grassland. However, with nine years of nitrogen addition, species with high H(N) decreased in abundance, while those with low H(N) increased in abundance. In contrast, in climate change experiments simulating a range of forecast hydrologic changes, e.g., extreme drought (two years), increased rainfall variability (14 years), and chronic increases in rainfall (21 years), plant species with the highest H(N) were least responsive to changes in soil water availability. These results suggest that H(N) may be predictive of plant species success and stability, and how plant species and ecosystems will respond to global-change-driven alterations in resource availability.

Asymmetric responses of primary productivity to precipitation extremes: A synthesis of grassland precipitation manipulation experiments

Climatic changes are altering Earths hydrological cycle, resulting in altered precipitation amounts, increased interannual variability of precipitation, and more frequent extreme precipitation events. These trends will likely continue into the future, having substantial impacts on net primary productivity (NPP) and associated ecosystem services such as food production and carbon sequestration. Frequently, experimental manipulations of precipitation have linked altered precipitation regimes to changes in NPP. Yet, findings have been diverse and substantial uncertainty still surrounds generalities describing patterns of ecosystem sensitivity to altered precipitation. Additionally, we do not know whether previously observed correlations between NPP and precipitation remain accurate when precipitation changes become extreme. We synthesized results from 83 case studies of experimental precipitation manipulations in grasslands worldwide. We used meta-analytical techniques to search for generalities and asymmetries of aboveground NPP (ANPP) and belowground NPP (BNPP) responses to both the direction and magnitude of precipitation change. Sensitivity (i.e., productivity response standardized by the amount of precipitation change) of BNPP was similar under precipitation additions and reductions, but ANPP was more sensitive to precipitation additions than reductions; this was especially evident in drier ecosystems. Additionally, overall relationships between the magnitude of productivity responses and the magnitude of precipitation change were saturating in form. The saturating form of this relationship was likely driven by ANPP responses to very extreme precipitation increases, although there were limited studies imposing extreme precipitation change, and there was considerable variation among experiments. This highlights the importance of incorporating gradients of manipulations, ranging from extreme drought to extreme precipitation increases into future climate change experiments. Additionally, policy and land management decisions related to global change scenarios should consider how ANPP and BNPP responses may differ, and that ecosystem responses to extreme events might not be predicted from relationships found under moderate environmental changes.

Does ecosystem sensitivity to precipitation at the site‐level conform to regional‐scale predictions?

Central to understanding global C cycle dynamics is the functional relationship between precipitation and net primary production (NPP). At large spatial (regional) scales, the responsiveness of aboveground NPP (ANPP) to interannual variation in annual precipitation (AP; ANPPsens) is inversely related to site-level ANPP, coinciding with turnover of plant communities along precipitation gradients. Within ecosystems experiencing chronic alterations in water availability, plant community change will also occur with unknown consequences for ANPPsens. To examine the role plant community shifts may play in determining alterations in site-level ANPPPsens, we experimentally increased precipitation by approximately 35% for two decades in a native Central U.S. grassland. Consistent with regional models, ANPPsens decreased initially as water availability and ANPP increased. However, ANPPsens shifted back to ambient levels when mesic species increased in abundance in the plant community. Similarly, in grassland sites with distinct mesic and xeric plant communities and corresponding 50% differences in ANPP, ANPPsens did not differ over almost three decades. We conclude that responses in ANPPsens to chronic alterations in water availability within an ecosystem may not conform to regional AP-ANPP patterns, despite expected changes in ANPP and plant communities. The result is unanticipated functional resistance to climate change at the site scale.

A framework for quantifying the magnitude and variability of community responses to global change drivers

A major challenge in global change ecology is to predict the trajectory and magnitude of community change in response to global change drivers (GCDs). Here, we present a new framework that not only increases the predictive power of individual studies, but also allows for synthesis across GCD studies and ecosystems. First, we suggest that by quantifying community dissimilarity of replicates both among and within treatments, we can infer both the magnitude and predictability of community change, respectively. Second, we demonstrate the utility of integrating rank abundance curves with measures of community dissimilarity to understand the species-level dynamics driving community changes and propose a series of testable hypotheses linking changes in rank abundance curves with shifts in community dissimilarity. Finally, we review six case studies that demonstrate how our new conceptual framework can be applied. Overall, we present a new framework for holistically predicting community responses to GCDs that has broad applicability in this era of unprecedented global change and novel environmental conditions.

Loss of a large grazer impacts savanna grassland plant communities similarly in North America and South Africa

Abstract Large herbivore grazing is a widespread disturbance in mesic savanna grasslands which increases herbaceous plant community richness and diversity. However, humans are modifying the impacts of grazing on these ecosystems by removing grazers. A more general understanding of how grazer loss will impact these ecosystems is hampered by differences in the diversity of large herbivore assemblages among savanna grasslands, which can affect the way that grazing influences plant communities. To avoid this we used two unique enclosures each containing a single, functionally similar large herbivore species. Specifically, we studied a bison (Bos bison) enclosure at Konza Prairie Biological Station, USA and an African buffalo (Syncerus caffer) enclosure in Kruger National Park, South Africa. Within these enclosures we erected exclosures in annually burned and unburned sites to determine how grazer loss would impact herbaceous plant communities, while controlling for potential fire-grazing interactions. At both sites, removal of the only grazer decreased grass and forb richness, evenness and diversity, over time. However, in Kruger these changes only occurred with burning. At both sites, changes in plant communities were driven by increased dominance with herbivore exclusion. At Konza, this was caused by increased abundance of one grass species, Andropogon gerardii, while at Kruger, three grasses, Themeda triandra, Panicum coloratum, and Digitaria eriantha increased in abundance.

Dual mechanisms regulate ecosystem stability under decade-long warming and hay harvest

Past global change studies have identified changes in species diversity as a major mechanism regulating temporal stability of production, measured as the ratio of the mean to the standard deviation of community biomass. However, the dominant plant functional group can also strongly determine the temporal stability. Here, in a grassland ecosystem subject to 15 years of experimental warming and hay harvest, we reveal that warming increases while hay harvest decreases temporal stability. This corresponds with the biomass of the dominant C4 functional group being higher under warming and lower under hay harvest. As a secondary mechanism, biodiversity also explains part of the variation in temporal stability of production. Structural equation modelling further shows that warming and hay harvest regulate temporal stability through influencing both temporal mean and variation of production. Our findings demonstrate the joint roles that dominant plant functional group and biodiversity play in regulating the temporal stability of an ecosystem under global change.

Warming Effects on Ecosystem Carbon Fluxes Are Modulated by Plant Functional Types

Despite the importance of future carbon (C) pools for policy and land management decisions under various climate change scenarios, predictions of these pools under altered climate vary considerably. Chronic warming will likely impact both ecosystem C fluxes and the abundance and distribution of plant functional types (PFTs) within systems, potentially interacting to create novel patterns of C exchange. Here, we report results from a 3-year warming experiment using open top chambers (OTC) on the Tibetan Plateau meadow grassland. Warming significantly increased C uptake through gross primary productivity (GPP) but not ecosystem respiration (ER), resulting in a 31.0% reduction in net ecosystem exchange (NEE) in warmed plots. The OTC-induced changes in ecosystem C fluxes were not fully explained by the corresponding changes in soil temperature and moisture. Warming treatments significantly increased the biomass of graminoids and legumes by 12.9 and 27.6%. These functional shifts were correlated with enhanced local GPP, but not ER, resulting in more negative NEE in plots with larger increases in graminoid and legume biomass. This may be due to a link between greater legume abundance and higher levels of total inorganic nitrogen, which can potentially drive higher GPP, but not higher ER. Overall, our results indicate that C-climate feedbacks might be closely mediated by climate-induced changes in PFTs. This highlights the need to consider the impacts of changes in PFTs when predicting future responses of C pools under altered climate scenarios.

Asynchrony among local communities stabilises ecosystem function of metacommunities

Abstract Temporal stability of ecosystem functioning increases the predictability and reliability of ecosystem services, and understanding the drivers of stability across spatial scales is important for land management and policy decisions. We used species‐level abundance data from 62 plant communities across five continents to assess mechanisms of temporal stability across spatial scales. We assessed how asynchrony (i.e. different units responding dissimilarly through time) of species and local communities stabilised metacommunity ecosystem function. Asynchrony of species increased stability of local communities, and asynchrony among local communities enhanced metacommunity stability by a wide range of magnitudes (1–315%); this range was positively correlated with the size of the metacommunity. Additionally, asynchronous responses among local communities were linked with species’ populations fluctuating asynchronously across space, perhaps stemming from physical and/or competitive differences among local communities. Accordingly, we suggest spatial heterogeneity should be a major focus for maintaining the stability of ecosystem services at larger spatial scales.