How collectively integrated are ecological communities?

Beyond abiotic conditions, do population dynamics mostly depend on a species' direct predators, preys and conspecifics? Or can indirect feedback that ripples across the whole community be equally important? Determining where ecological communities sit on the spectrum between these two characterizations requires a metric able to capture the difference between them. Here we show that the spectral radius of a community's interaction matrix provides such a metric, thus a measure of ecological collectivity, which is accessible from imperfect knowledge of biotic interactions and related to observable signatures. This measure of collectivity integrates existing approaches to complexity, interaction structure and indirect interactions. Our work thus provides an original perspective on the question of to what degree communities are more than loose collections of species or simple interaction motifs and explains when pragmatic reductionist approaches ought to suffice or fail when applied to ecological communities.

Decoupled responses of above- and below-ground beta-diversity to nitrogen enrichment in a typical steppe.

Increased atmospheric nitrogen (N) deposition affects biodiversity in terrestrial ecosystems. However, we do not know whether the effects of N on above-ground plant ß-diversity are coupled with changes occurring in the soil seed bank. We conducted a long-term N-addition experiment in a typical steppe and found that above-ground ß-diversity increased and then decreased with increasing N addition, whereas below-ground ß-diversity decreased linearly. This suggests decoupled dynamics of plant communities and their soil seed bank under N enrichment. Species substitution determined above- and below-ground ß-diversity change via an increasing role of deterministic processes with N addition. These effects were mostly driven by differential responses of the above-ground vegetation and the soil seed bank ß-diversities to N-induced changes in environmental heterogeneity, increased soil inorganic N concentrations and soil acidification. Our findings highlight the importance of considering above- and below-ground processes simultaneously for effectively conserving grassland ecosystems under N enrichment.

Partitioning the biodiversity effects on productivity into density and size components.

Plant density and size - two factors that represent plant survival and growth - are key determinants of yield but have rarely been analysed explicitly in the context of biodiversity-productivity relationships. Here, we derive equations to partition the net, complementarity and selection effects of biodiversity into additive components that reflect diversity-induced changes in plant density and size. Applications of the new method to empirical datasets reveal contrasting ways in which plant density and size regulate yield in species mixtures. In an annual plant diversity experiment, overyielding is largely explained by selection effects associated with increased size of highly productive plant species. In a tree diversity experiment, the cause of overyielding shifts from enhanced growth in tree size to reduced mortality by complementary use of canopy space during stand development. These results highlight the capability of the new method to resolve crucial, yet understudied, demographic links between biodiversity and productivity.

Spatial heterogeneity of biomass turnover has contrasting effects on synchrony and stability in trophic metacommunities.

Spatial heterogeneity is a fundamental feature of ecosystems, and ecologists have identified it as a factor promoting the stability of population dynamics. In particular, differences in interaction strengths and resource supply between patches generate an asymmetry of biomass turnover with a fast and a slow patch coupled by a mobile predator. Here, we demonstrate that asymmetry leads to opposite stability patterns in metacommunities receiving localized perturbations depending on the characteristics of the perturbed patch. Perturbing prey in the fast patch synchronizes the dynamics of prey biomass between the two patches and destabilizes predator dynamics by increasing the predator's temporal variability. Conversely, perturbing prey in the slow patch decreases the synchrony of the prey's dynamics and stabilizes predator dynamics. Our results have implications for conservation ecology and suggest reinforcing protection policies in fast patches to dampen the effects of perturbations and promote the stability of population dynamics at the regional scale.

Clarifying the effect of biodiversity on productivity in natural ecosystems with longitudinal data and methods for causal inference.

Causal effects of biodiversity on ecosystem functions can be estimated using experimental or observational designs - designs that pose a tradeoff between drawing credible causal inferences from correlations and drawing generalizable inferences. Here, we develop a design that reduces this tradeoff and revisits the question of how plant species diversity affects productivity. Our design leverages longitudinal data from 43 grasslands in 11 countries and approaches borrowed from fields outside of ecology to draw causal inferences from observational data. Contrary to many prior studies, we estimate that increases in plot-level species richness caused productivity to decline: a 10% increase in richness decreased productivity by 2.4%, 95% CI [-4.1, -0.74]. This contradiction stems from two sources. First, prior observational studies incompletely control for confounding factors. Second, most experiments plant fewer rare and non-native species than exist in nature. Although increases in native, dominant species increased productivity, increases in rare and non-native species decreased productivity, making the average effect negative in our study. By reducing the tradeoff between experimental and observational designs, our study demonstrates how observational studies can complement prior ecological experiments and inform future ones.

Will a large complex system be productive?

While the relationship between food web complexity and stability has been well documented, how complexity affects productivity remains elusive. In this study, we combine food web theory and a data set of 149 aquatic food webs to investigate the effect of complexity (i.e. species richness, connectance, and average interaction strength) on ecosystem productivity. We find that more complex ecosystems tend to be more productive, although different facets of complexity have contrasting effects. A higher species richness and/or average interaction strength increases productivity, whereas a higher connectance often decreases it. These patterns hold not only between realized complexity and productivity, but also characterize responses of productivity to simulated declines of complexity. Our model also predicts a negative association between productivity and stability along gradients of complexity. Empirical analyses support our predictions on positive complexity-productivity relationships and negative productivity-stability relationships. Our study provides a step forward towards reconciling ecosystem complexity, productivity and stability.

Opportunities to advance the synthesis of ecology and evolution.

Despite decades of research on the interactions between ecology and evolution, opportunities still remain to further integrate the two disciplines, especially when considering multispecies systems. Here, we discuss two such opportunities. First, the traditional emphasis on the distinction between evolutionary and ecological processes should be further relaxed as it is particularly unhelpful in the study of microbial communities, where the very notion of species is hard to define. Second, key processes of evolutionary theory such as adaptation should be exported to hierarchical levels higher than populations to make sense of biodiversity dynamics. Together, we argue that broadening our perspective of eco-evolutionary dynamics to be more inclusive of all biodiversity, both phylogenetically and hierarchically, will open up fertile new research directions and help us to address one of the major scientific challenges of our time, that is, to understand and predict changes in biodiversity in the face of rapid environmental change.

Consistent stabilizing effects of plant diversity across spatial scales and climatic gradients.

Biodiversity has widely been documented to enhance local community stability but whether such stabilizing effects of biodiversity extend to broader scales remains elusive. Here, we investigated the relationships between biodiversity and community stability in natural plant communities from quadrat (1 m2) to plot (400 m2) and regional (5-214 km2) scales and across broad climatic conditions, using an extensive plant community dataset from the National Ecological Observatory Network. We found that plant diversity provided consistent stabilizing effects on total community abundance across three nested spatial scales and climatic gradients. The strength of the stabilizing effects of biodiversity increased modestly with spatial scale and decreased as precipitation seasonality increased. Our findings illustrate the generality of diversity-stability theory across scales and climatic gradients, which provides a robust framework for understanding ecosystem responses to biodiversity and climate changes.

Human-nature connectedness as a pathway to sustainability: A global meta-analysis.

Internationally agreed sustainability goals are being missed. Here, we conduct global meta-analyses to assess how the extent to which humans see themselves as part of nature-known as human-nature connectedness (HNC)-can be used as a leverage point to reach sustainability. A meta-analysis of 147 correlational studies shows that individuals with high HNC had more pronature behaviours and were significantly healthier than those with low HNC. A meta-analysis of 59 experimental studies shows significant increases in HNC after manipulations involving contact with nature and mindfulness practices. Surprisingly, this same meta-analysis finds no significant effect of environmental education on HNC. Thus, HNC is positively linked to mind-sets that value sustainability and behaviours that enhance it. Further, we argue that HNC can be enhanced by targeted practices, and we identify those most likely to succeed. Our results suggest that enhancing HNC, via promotion of targeted practices, can improve sustainability and should be integrated into conservation policy.

Multispecies forest plantations outyield monocultures across a broad range of conditions.

Multispecies tree planting has long been applied in forestry and landscape restoration in the hope of providing better timber production and ecosystem services; however, a systematic assessment of its effectiveness is lacking. We compiled a global dataset of matched single-species and multispecies plantations to evaluate the impact of multispecies planting on stand growth. Average tree height, diameter at breast height, and aboveground biomass were 5.4, 6.8, and 25.5% higher, respectively, in multispecies stands compared with single-species stands. These positive effects were mainly the result of interspecific complementarity and were modulated by differences in leaf morphology and leaf life span, stand age, planting density, and temperature. Our results have implications for designing afforestation and reforestation strategies and bridging experimental studies of biodiversity-ecosystem functioning relationships with real-world practices.

Habitat percolation transition undermines sustainability in social-ecological agricultural systems.

Steady increases in human population size and resource consumption are driving rampant agricultural expansion and intensification. Habitat loss caused by agriculture puts the integrity of ecosystems at risk and threatens the persistence of human societies that rely on ecosystem services. We develop a spatially explicit model describing the coupled dynamics of an agricultural landscape and human population size to assess the effect of different land-use management strategies, defined by agricultural clustering and intensification, on the sustainability of the social-ecological system. We show how agricultural expansion can cause natural habitats to undergo a percolation transition leading to abrupt habitat fragmentation that feedbacks on human's decision making, aggravating landscape degradation. We found that agricultural intensification to spare land from conversion is a successful strategy only in highly natural landscapes, and that clustering agricultural land is the most effective measure to preserve large connected natural fragments, prevent severe fragmentation and thus, enhance sustainability.

The hidden role of multi-trophic interactions in driving diversity-productivity relationships.

Resource-use complementarity of producer species is often invoked to explain the generally positive diversity-productivity relationships. Additionally, multi-trophic interactions that link processes across trophic levels have received increasing attention as a possible key driver. Given that both are integral to natural ecosystems, their interactive effect should be evident but has remained hidden. We address this issue by analysing diversity-productivity relationships in a simulation experiment of producer communities nested within complex food-webs, manipulating resource-use complementarity and multi-trophic animal richness. We show that these two mechanisms interactively create diverse communities of complementary producer species. This shapes diversity-productivity relationships such that their joint contribution generally exceeds their individual effects. Specifically, multi-trophic interactions in animal-rich ecosystems facilitate producer coexistence by preventing competitive exclusion despite overlaps in resource-use, which increases the realised complementarity. The interdependence of food-webs and producer complementarity in creating biodiversity-productivity relationships highlights the importance to adopt a multi-trophic perspective on biodiversity-ecosystem functioning relationships.

Grand challenges in biodiversity-ecosystem functioning research in the era of science-policy platforms require explicit consideration of feedbacks.

Feedbacks are an essential feature of resilient socio-economic systems, yet the feedbacks between biodiversity, ecosystem services and human wellbeing are not fully accounted for in global policy efforts that consider future scenarios for human activities and their consequences for nature. Failure to integrate feedbacks in our knowledge frameworks exacerbates uncertainty in future projections and potentially prevents us from realizing the full benefits of actions we can take to enhance sustainability. We identify six scientific research challenges that, if addressed, could allow future policy, conservation and monitoring efforts to quantitatively account for ecosystem and societal consequences of biodiversity change. Placing feedbacks prominently in our frameworks would lead to (i) coordinated observation of biodiversity change, ecosystem functions and human actions, (ii) joint experiment and observation programmes, (iii) more effective use of emerging technologies in biodiversity science and policy, and (iv) a more inclusive and integrated global community of biodiversity observers. To meet these challenges, we outline a five-point action plan for collaboration and connection among scientists and policymakers that emphasizes diversity, inclusion and open access. Efforts to protect biodiversity require the best possible scientific understanding of human activities, biodiversity trends, ecosystem functions and-critically-the feedbacks among them.