Making sense of multivariate community responses in global change experiments

Ecological communities are being impacted by global change worldwide. Experiments are a powerful tool to understand how global change will impact communities by comparing control and treatment replicates. Communities consist of multiple species, and their associated abundances make multivariate methods an effective approach to study community compositional differences between control and treated replicates. Dissimilarity metrics are a commonly employed multivariate measure of compositional differences; however, while highly informative, dissimilarity metrics do not elucidate the specific ways in which communities differ. Integrating two multivariate methods, dissimilarity metrics and rank abundance curves (RACs), have the potential to detect complex differences based on dissimilarity metrics and detail the how these differences came about through differences in richness, evenness, species ranks, or species identity. Here we use a database of 106 global change experiments located in herbaceous ecosystems and explore how patterns of ordinations based on dissimilarity metrics relate to RAC-based differences. We find that combining dissimilarity metrics alongside RAC-based measures clarifies how global change treatments are altering communities. We find that when there is no difference in community composition (no distance between centroids of control and treated replicates), there are rarely differences in species ranks or species identities and more often differences in richness or evenness alone. In contrast, when there are differences between centroids of control and treated replicates, this is most often associated with differences in ranks either alone or co-occurring with differences in richness, evenness, or species identities. We suggest that integrating these two multivariate measures of community composition results in a deeper understanding of how global change impacts communities

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

General Destabilizing Effects of Eutrophication on Grassland Productivity at Multiple Spatial Scales

Eutrophication is a widespread environmental change that usually reduces the stabilizing effect of plant diversity on productivity in local communities. Whether this effect is scale dependent remains to be elucidated. Here, we determine the relationship between plant diversity and temporal stability of productivity for 243 plant communities from 42 grasslands across the globe and quantify the effect of chronic fertilization on these relationships. Unfertilized local communities with more plant species exhibit greater asynchronous dynamics among species in response to natural environmental fluctuations, resulting in greater local stability (alpha stability). Moreover, neighborhood communities that have greater spatial variation in plant species composition within sites (higher beta diversity) have greater spatial asynchrony of productivity among communities, resulting in greater stability at the larger scale (gamma stability). Importantly, fertilization consistently weakens the contribution of plant diversity to both of these stabilizing mechanisms, thus diminishing the positive effect of biodiversity on stability at differing spatial scales. Our findings suggest that preserving grassland functional stability requires conservation of plant diversity within and among ecological communities

Global change effects on plant communities are magnified by time and the number of global change factors imposed

Global change drivers (GCDs) are expected to alter community structure and consequently, the services that ecosystems provide. Yet, few experimental investigations have examined effects of GCDs on plant community structure across multiple ecosystem types, and those that do exist present conflicting patterns. In an unprecedented global synthesis of over 100 experiments that manipulated factors linked to GCDs, we show that herbaceous plant community responses depend on experimental manipulation length and number of factors manipulated. We found that plant communities are fairly resistant to experimentally manipulated GCDs in the short term (<10 y). In contrast, long-term (≥10 y) experiments show increasing community divergence of treatments from control conditions. Surprisingly, these community responses occurred with similar frequency across the GCD types manipulated in our database. However, community responses were more common when 3 or more GCDs were simultaneously manipulated, suggesting the emergence of additive or synergistic effects of multiple drivers, particularly over long time periods. In half of the cases, GCD manipulations caused a difference in community composition without a corresponding species richness difference, indicating that species reordering or replacement is an important mechanism of community responses to GCDs and should be given greater consideration when examining consequences of GCDs for the biodiversity–ecosystem function relationship. Human activities are currently driving unparalleled global changes worldwide. Our analyses provide the most comprehensive evidence to date that these human activities may have widespread impacts on plant community composition globally, which will increase in frequency over time and be greater in areas where communities face multiple GCDs simultaneously

Global change effects on plant communities are magnified by time and the number of global change factors imposed

Global change drivers (GCDs) are expected to alter community structure and consequently, the services that ecosystems provide. Yet, few experimental investigations have examined effects of GCDs on plant community structure across multiple ecosystem types, and those that do exist present conflicting patterns. In an unprecedented global synthesis of over 100 experiments that manipulated factors linked to GCDs, we show that herbaceous plant community responses depend on experimental manipulation length and number of factors manipulated. We found that plant communities are fairly resistant to experimentally manipulated GCDs in the short term (\u3c10 y). In contrast, long-term (≥10 y) experiments show increasing community divergence of treatments from control conditions. Surprisingly, these community responses occurred with similar frequency across the GCD types manipulated in our database. However, community responses were more common when 3 or more GCDs were simultaneously manipulated, suggesting the emergence of additive or synergistic effects of multiple drivers, particularly over long time periods. In half of the cases, GCD manipulations caused a difference in community composition without a corresponding species richness difference, indicating that species reordering or replacement is an important mechanism of community responses to GCDs and should be given greater consideration when examining consequences of GCDs for the biodiversity.ecosystem function relationship. Human activities are currently driving unparalleled global changes worldwide. Our analyses provide the most comprehensive evidence to date that these human activities may have widespread impacts on plant community composition globally, which will increase in frequency over time and be greater in areas where communities face multiple GCDs simultaneously

Data from: Herbivore size matters for productivity-richness relationships in African savannas

1.Productivity and herbivory often interact to shape plant community composition and species richness with levels of production mediating the impact of herbivory. Yet, differences in herbivore traits such as size, feeding guild, and dietary requirements may result in different impacts of diverse herbivore guilds across productivity gradients. 2.We used size-selective herbivore exclosures to separate the effects of herbivory by larger herbivores, such as elephant, Burchell's zebra, and blue wildebeest from those of medium/smaller herbivores, such as impala and warthog, on herbaceous plant communities. These exclosures were established along a 10-fold productivity gradient, ranging from 90-950 g m−2 of standing plant biomass in the Kruger National Park, South Africa. 3.Exclusion of all herbivores generally increased plant species richness at low productivity but decreased richness at high productivity. Exclusion of medium/smaller herbivores (e.g. impala, warthog) showed stronger effects on plant richness, particularly loss of forbs, at higher productivity rather than at lower productivity. In contrast, exclusion of larger herbivores had stronger effects on plant richness, typically with increasing forb richness, at low rather than high productivity. 4.The change in species richness appeared linked to changes in light availability following herbivore exclusion. Strong increases in shading led to declines in species richness while more moderate increases in shading led in increases in species richness, possibly due to amelioration of heat and water stress by modest increases in shading. 5.Increasing plant dominance, which likely alters multiple mechanisms of plant interactions, was correlated with declines in plant richness following herbivore exclusion. The impact of increasing dominance on plant richness operated independent of productivity, with the exclusion of impala appearing particularly important in driving this relationship. 6.Synthesis. We show that the impact of herbivore losses on plant diversity will be strongly situation dependent and will vary with the herbivores lost (e.g. larger vs. smaller, grazers vs. browsers), plant functional type (e.g. grasses vs. forbs), and environmental context (e.g. productivity). Although larger herbivores are often emphasized for their strong impacts on community dynamics and ecosystem processes, we show that smaller, abundant herbivores can exert strong top-down control on plant communities

Productivity_Spp_Comp_Burkepile_et_al_JoE

This file contains data for plant species composition, productivity, and light availability from a herbivore exclusion experiment conducted across a productivity gradient in Kruger National Park, South Afric

Data from: Fire frequency drives habitat selection by a diverse herbivore guild impacting top–down control of plant communities in an African savanna

In areas with diverse herbivore communities such as African savannas, the frequency of disturbance by fire may alter the top–down role of different herbivore species on plant community dynamics. In a seven year experiment in the Kruger National Park, South Africa, we examined the habitat use of nine common herbivore species across annually burned, triennially burned and unburned areas. We also used two types of exclosures (plus open access controls) to examine the impacts of different herbivores on plant community dynamics across fire disturbance regimes. Full exclosures excluded all herbivores > 0.5 kg (e.g. elephant, zebra, impala) while partial exclosures allowed access only to animals with shoulder heights ≤ 0.85 m (e.g. impala, steenbok). Annual burns attracted a diverse suite of herbivores, and exclusion of larger herbivores (e.g. elephant, zebra, wildebeest) increased plant abundance. When smaller species, mainly impala, were also excluded there were declines in plant diversity, likely mediated by a decline in open space available for colonization of uncommon plant species. Unburned areas attracted the least diverse suite of herbivores, dominated by impala. Here, herbivore exclusion, especially of impala, led to strong declines in plant richness and diversity. With no fire disturbance, herbivore exclusion led to competitive exclusion via increases in plant dominance and light limitation. In contrast, on triennial burns, herbivore exclusion had no effect on plant richness or diversity, potentially due to relatively little open space for colonization across exclosure treatments but also little competitive exclusion due to the intermediate fire disturbance. Further, the diverse suite of grazers and browsers on triennial burns may have had a compensating effect of on the diversity of grasses and forbs. Ultimately, our work shows that differential disturbance regimes can result in differential consumer pressure across a landscape and result in heterogeneous patterns in top–down control of community dynamics

Appendix A. Site and herbivore exclosure design information.

Site and herbivore exclosure design information

Appendix B. Plant community response metrics through time.

Plant community response metrics through time