Chengjin Chu

High‐dimensional coexistence based on individual variation: a synthesis of evidence

High biodiversity of forests is not predicted by traditional models, and evidence for trade-offs those models require is limited. High-dimensional regulation (e.g., N factors to regulate N species) has long been recognized as a possible alternative explanation, but it has not be been seriously pursued, because only a few limiting resources are evident for trees, and analysis of multiple interactions is challenging. We develop a hierarchical model that allows us to synthesize data from long-term, experimental, data sets with processes that control growth, maturation, fecundity, and survival. We allow for uncertainty at all stages and variation among 26 000 individuals and over time, including 268 000 tree years, for dozens of tree species. We estimate population-level parameters that apply at the species level and the interactions among latent states, i.e., the demographic rates for each individual, every year. The former show that the traditional trade-offs used to explain diversity are not present. Demographic rates overlap among species, and they do not show trends consistent with maintenance of diversity by simple mechanisms (negative correlations and limiting similarity). However, estimates of latent states at the level of individuals and years demonstrate that species partition environmental variation. Correlations between responses to variation in time are high for individuals of the same species, but not for individuals of different species. We demonstrate that these relationships are pervasive, providing strong evidence that high- dimensional regulation is critical for biodiversity regulation.

Regional Contingencies in the Relationship between Aboveground Biomass and Litter in the World's Grasslands

Based on regional-scale studies, aboveground production and litter decomposition are thought to positively covary, because they are driven by shared biotic and climatic factors. Until now we have been unable to test whether production and decomposition are generally coupled across climatically dissimilar regions, because we lacked replicated data collected within a single vegetation type across multiple regions, obfuscating the drivers and generality of the association between production and decomposition. Furthermore, our understanding of the relationships between production and decomposition rests heavily on separate meta-analyses of each response, because no studies have simultaneously measured production and the accumulation or decomposition of litter using consistent methods at globally relevant scales. Here, we use a multi-country grassland dataset collected using a standardized protocol to show that live plant biomass (an estimate of aboveground net primary production) and litter disappearance (represented by mass loss of aboveground litter) do not strongly covary. Live biomass and litter disappearance varied at different spatial scales. There was substantial variation in live biomass among continents, sites and plots whereas among continent differences accounted for most of the variation in litter disappearance rates. Although there were strong associations among aboveground biomass, litter disappearance and climatic factors in some regions (e.g. U.S. Great Plains), these relationships were inconsistent within and among the regions represented by this study. These results highlight the importance of replication among regions and continents when characterizing the correlations between ecosystem processes and interpreting their global-scale implications for carbon flux. We must exercise caution in parameterizing litter decomposition and aboveground production in future regional and global carbon models as their relationship is complex.

Grassland productivity limited by multiple nutrients

Terrestrial ecosystem productivity is widely accepted to be nutrient limited1. Although nitrogen (N) is deemed a key determinant of aboveground net primary production (ANPP)2,3, the prevalence of co-limitation by N and phosphorus (P) is increasingly recognized4–8. However, the extent to which terrestrial productivity is co-limited by nutrients other than N and P has remained unclear. Here, we report results from a standardized factorial nutrient addition experiment, in which we added N, P and potassium (K) combined with a selection of micronutrients (K+μ), alone or in concert, to 42 grassland sites spanning five continents, and monitored ANPP. Nutrient availability limited productivity at 31 of the 42 grassland sites. And pairwise combinations of N, P, and K+μ co-limited ANPP at 29 of the sites. Nitrogen limitation peaked in cool, high latitude sites. Our findings highlight the importance of less studied nutrients, such as K and micronutrients, for grassland productivity, and point to significant variations in the type and degree of nutrient limitation. We suggest that multiple-nutrient constraints must be considered when assessing the ecosystem-scale consequences of nutrient enrichment.

Abundance of introduced species at home predicts abundance away in herbaceous communities

Many ecosystems worldwide are dominated by introduced plant species, leading to loss of biodiversity and ecosystem function. A common but rarely tested assumption is that these plants are more abundant in introduced vs. native communities, because ecological or evolutionary-based shifts in populations underlie invasion success. Here, data for 26 herbaceous species at 39 sites, within eight countries, revealed that species abundances were similar at native (home) and introduced (away) sites - grass species were generally abundant home and away, while forbs were low in abundance, but more abundant at home. Sites with six or more of these species had similar community abundance hierarchies, suggesting that suites of introduced species are assembling similarly on different continents. Overall, we found that substantial changes to populations are not necessarily a pre-condition for invasion success and that increases in species abundance are unusual. Instead, abundance at home predicts abundance away, a potentially useful additional criterion for biosecurity programmes.

Species abundance at home predicts abundance away in grasslands

Many ecosystems worldwide are dominated by introduced plant species, leading to loss of biodiversity and ecosystem function. A common but rarely tested assumption is that these plants are more abundant in introduced vs. native communities, because ecological or evolutionary-based shifts in populations underlie invasion success. Here, data for 26 herbaceous species at 39 sites, within eight countries, revealed that species abundances were similar at native (home) and introduced (away) sites - grass species were generally abundant home and away, while forbs were low in abundance, but more abundant at home. Sites with six or more of these species had similar community abundance hierarchies, suggesting that suites of introduced species are assembling similarly on different continents. Overall, we found that substantial changes to populations are not necessarily a pre-condition for invasion success and that increases in species abundance are unusual. Instead, abundance at home predicts abundance away, a potentially useful additional criterion for biosecurity programmes.

Large niche differences emerge at the recruitment stage to stabilize grassland coexistence

Niche differences and average fitness differences jointly determine coexistence. However, little empirical information about the magnitude of these two mechanisms is available. Using multispecies population models fit to long-term demographic data for common, co-occurring species in five grassland and shrubland plant communities in western North America, we estimated the strength of stabilizing niche differences and average fitness differences. In all five communities, both pairwise and full community comparisons showed evidence for strong stabilizing mechanisms and relatively small average fitness differences. For a total of 17 species pairs, a measure of niche differences based on simulations of invasion growth rates ranged from 0.59 to 0.93 with a mean of 0.81, where 0 indicates complete niche overlap and 1 indicates zero niche overlap. A corresponding measure of average fitness differences ranged from 1.02 to 2.54 with a mean of 1.53, where 1 indicates identical fitness and a value of 2 indicates a fo...

The Relationship between Species Richness and Evenness in Plant Communities along a Successional Gradient: A Study from Sub-Alpine Meadows of the Eastern Qinghai-Tibetan Plateau, China

The relationship between species richness and evenness across communities remains an unsettled issue in ecology from both theoretical and empirical perspectives. As a result, we do not know the mechanisms that could generate a relationship between species richness and evenness, and how this responds to spatial scale. Here we examine the relationship between species richness(S) and evenness (Pielou’s J′ evenness) using a chronosequence of successional sub-alpine meadow communities in the eastern Qinghai-Tibetan Plateau. These meadows range from natural community (never farmed), to those that have been protected from agricultural exploitation for periods ranging from 1 to 10 years. A total of 30 sampling quadrats with size of 0.5 m×0.5 m were laid out along two transects at each meadow. Using correlation analyses we found a consistent negative correlation between S and J′ in these communities along the successional gradient at the sampling scale of 0.5 m×0.5 m. We also explored the relationship between S and J′ at different sampling scales (from 0.5 m×0.5 m to10 m×10 m) using properly measured ramet-mapped data of a10 m×10 m quadrat in the natural community. We found that S was negatively corrected with J′ at the scales of 0.5 m×0.5 m to 2 m×2 m, but such a relationships disappeared at relative larger scales (≥2 m×4 m). When fitting different species abundance models combined with trait-specific methods, we found that niche preemption may be the determining mechanism of species evenness along the succession gradient. Considering all results together, we can conclude that such niche differentiation and spatial scale effects may help to explain the maintenance of high species richness in sub-alpine meadow communities.

Assessing facilitative responses to a nurse shrub at the community level: the example of Potentilla fruticosa in a sub-alpine grassland of northwest China

Biotic interaction studies have revealed a large discrepancy among experiments in target responses to the effects of neighbours, which may in part be due to both high species-specificity of plant responses and low number of target species used in experiments. Our aim was to assess facilitative responses at the community level and the role of both functional groups and ecological attributes of target species. In a sub-alpine grassland on the eastern Tibet plateau, we assessed growth responses of all species in the community to removal of a dominant shrub. We also measured changes in the main environmental variables. Species responses were analysed by functional group and in relation to their mean regional altitudinal distribution. All significant interactions were positive and affected one-third of the total species richness of the community. All functional groups were facilitated but forbs were less strongly facilitated than in the two other groups. High-alpine species were less strongly facilitated than low-sub-alpine species, but the strength of this relationship was weaker than that reported in previous work. There was evidence of a decrease in extreme temperatures below the canopy of the shrub but no variations in soil moisture. We conclude that the highly stressful conditions induced by the dry continental climate of the eastern Tibet plateau are a main driver of the exclusive dominance of positive interactions. Assessing interactive responses at the community level is likely to provide a useful tool to better understand the role of biotic interactions in community responses to environmental changes.

The effect of environmental heterogeneity on species richness depends on community position along the environmental gradient

Environmental heterogeneity is among the most important factors governing community structure. Besides the widespread evidence supporting positive relationships between richness and environmental heterogeneity, negative and unimodal relationships have also been reported. However, few studies have attempted to test the role of the heterogeneity on species richness after removing the confounding effect of resource availability or environmental severity. Here we constructed an individual-based spatially explicit model incorporating a long-recognized tradeoff between competitive ability and stress-tolerance ability of species. We explored the impact of the level of resource availability (i.e. the position of the community along a gradient of environmental severity) on the heterogeneity-diversity relationship (HDR). The results indicate that the shape of HDR depends on the community position along the environmental gradient: at either end of the gradient of environmental severity, a positive HDR occurred, whereas at the intermediate levels of the gradient, a unimodal HDR emerged. Our exploration demonstrates that resource availability/environmental severity should be considered as a potential factor influencing the shape of the HDR. Our theoretical predictions represent hypotheses in need of further empirical study.

Comment on "Worldwide evidence of a unimodal relationship between productivity and plant species richness"

Fraser et al. (Reports, 17 July 2015, p. 302) report a unimodal relationship between productivity and species richness at regional and global scales, which they contrast with the results of Adler et al. (Reports, 23 September 2011, p. 1750). However, both data sets, when analyzed correctly, show clearly and consistently that productivity is a poor predictor of local species richness.