Cumulative nitrogen enrichment alters the drivers of grassland overyielding.

Effects of plant diversity on grassland productivity, or overyielding, are found to be robust to nutrient enrichment. However, the impact of cumulative nitrogen (N) addition (total N added over time) on overyielding and its drivers are underexplored. Synthesizing data from 15 multi-year grassland biodiversity experiments with N addition, we found that N addition decreases complementarity effects and increases selection effects proportionately, resulting in no overall change in overyielding regardless of N addition rate. However, we observed a convex relationship between overyielding and cumulative N addition, driven by a shift from complementarity to selection effects. This shift suggests diminishing positive interactions and an increasing contribution of a few dominant species with increasing N accumulation. Recognizing the importance of cumulative N addition is vital for understanding its impacts on grassland overyielding, contributing essential insights for biodiversity conservation and ecosystem resilience in the face of increasing N deposition.

Effects of plant diversity on productivity strengthen over time due to trait-dependent shifts in species overyielding.

Plant diversity effects on community productivity often increase over time. Whether the strengthening of diversity effects is caused by temporal shifts in species-level overyielding (i.e., higher species-level productivity in diverse communities compared with monocultures) remains unclear. Here, using data from 65 grassland and forest biodiversity experiments, we show that the temporal strength of diversity effects at the community scale is underpinned by temporal changes in the species that yield. These temporal trends of species-level overyielding are shaped by plant ecological strategies, which can be quantitatively delimited by functional traits. In grasslands, the temporal strengthening of biodiversity effects on community productivity was associated with increasing biomass overyielding of resource-conservative species increasing over time, and with overyielding of species characterized by fast resource acquisition either decreasing or increasing. In forests, temporal trends in species overyielding differ when considering above- versus belowground resource acquisition strategies. Overyielding in stem growth decreased for species with high light capture capacity but increased for those with high soil resource acquisition capacity. Our results imply that a diversity of species with different, and potentially complementary, ecological strategies is beneficial for maintaining community productivity over time in both grassland and forest ecosystems.

Invasive earthworms can change understory plant community traits and reduce plant functional diversity.

Among the most important impacts of biological invasions on biodiversity is biotic homogenization, which may further compromise key ecosystem processes. However, the extent to which they homogenize functional diversity and shift dominant ecological strategies of invaded communities remains uncertain. Here, we investigated changes in plant communities in a northern North American forest in response to invasive earthworms, by examining the taxonomic and functional diversity of the plant community and soil ecosystem functions. We found that although plant taxonomic diversity did not change in response to invasive earthworms, they modified the dominance structure of plant functional groups. Invasive earthworms promoted the dominance of fast-growing plants at the expense of slow-growing ones. Moreover, earthworms decreased plant functional diversity, which coincided with changes in abiotic and biotic soil properties. Our study reveals that invasive earthworms erode multiple biodiversity facets of invaded forests, with potential cascading effects on ecosystem functioning.

Feedback loops drive ecological succession: towards a unified conceptual framework.

The core principle shared by most theories and models of succession is that, following a major disturbance, plant-environment feedback dynamics drive a directional change in the plant community. The most commonly studied feedback loops are those in which the regrowth of the plant community causes changes to the abiotic (e.g. soil nutrients) or biotic (e.g. dispersers) environment, which differentially affect species availability or performance. This, in turn, leads to shifts in the species composition of the plant community. However, there are many other PE feedback loops that potentially drive succession, each of which can be considered a model of succession. While plant-environment feedback loops in principle generate predictable successional trajectories, succession is generally observed to be highly variable. Factors contributing to this variability are the stochastic processes involved in feedback dynamics, such as individual mortality and seed dispersal, and extrinsic causes of succession, which are not affected by changes in the plant community but do affect species performance or availability. Both can lead to variation in the identity of dominant species within communities. This, in turn, leads to further contingencies if these species differ in their effect on their environment (priority effects). Predictability and variability are thus intrinsically linked features of ecological succession. We present a new conceptual framework of ecological succession that integrates the propositions discussed above. This framework defines seven general causes: landscape context, disturbance and land-use, biotic factors, abiotic factors, species availability, species performance, and the plant community. When involved in a feedback loop, these general causes drive succession and when not, they are extrinsic causes that create variability in successional trajectories and dynamics. The proposed framework provides a guide for linking these general causes into causal pathways that represent specific models of succession. Our framework represents a systematic approach to identifying the main feedback processes and causes of variation at different successional stages. It can be used for systematic comparisons among study sites and along environmental gradients, to conceptualise studies, and to guide the formulation of research questions and design of field studies. Mapping an extensive field study onto our conceptual framework revealed that the pathways representing the study's empirical outcomes and conceptual model had important differences, underlining the need to move beyond the conceptual models that currently dominate in specific fields and to find ways to examine the importance of and interactions among alternative causal pathways of succession. To further this aim, we argue for integrating long-term studies across environmental and anthropogenic gradients, combined with controlled experiments and dynamic modelling.

Assembly of functional diversity in an oceanic island flora.

Oceanic island floras are well known for their morphological peculiarities and exhibit striking examples of trait evolution1-3. These morphological shifts are commonly attributed to insularity and are thought to be shaped by the biogeographical processes and evolutionary histories of oceanic islands2,4. However, the mechanisms through which biogeography and evolution have shaped the distribution and diversity of plant functional traits remain unclear5. Here we describe the functional trait space of the native flora of an oceanic island (Tenerife, Canary Islands, Spain) using extensive field and laboratory measurements, and relate it to global trade-offs in ecological strategies. We find that the island trait space exhibits a remarkable functional richness but that most plants are concentrated around a functional hotspot dominated by shrubs with a conservative life-history strategy. By dividing the island flora into species groups associated with distinct biogeographical distributions and diversification histories, our results also suggest that colonization via long-distance dispersal and the interplay between inter-island dispersal and archipelago-level speciation processes drive functional divergence and trait space expansion. Contrary to our expectations, speciation via cladogenesis has led to functional convergence, and therefore only contributes marginally to functional diversity by densely packing trait space around shrubs. By combining biogeography, ecology and evolution, our approach opens new avenues for trait-based insights into how dispersal, speciation and persistence shape the assembly of entire native island floras.

Tree islands enhance biodiversity and functioning in oil palm landscapes.

In the United Nations Decade on Ecosystem Restoration1, large knowledge gaps persist on how to increase biodiversity and ecosystem functioning in cash crop-dominated tropical landscapes2. Here, we present findings from a large-scale, 5-year ecosystem restoration experiment in an oil palm landscape enriched with 52 tree islands, encompassing assessments of ten indicators of biodiversity and 19 indicators of ecosystem functioning. Overall, indicators of biodiversity and ecosystem functioning, as well as multidiversity and ecosystem multifunctionality, were higher in tree islands compared to conventionally managed oil palm. Larger tree islands led to larger gains in multidiversity through changes in vegetation structure. Furthermore, tree enrichment did not decrease landscape-scale oil palm yield. Our results demonstrate that enriching oil palm-dominated landscapes with tree islands is a promising ecological restoration strategy, yet should not replace the protection of remaining forests.

Extreme weather events threaten biodiversity and functions of river ecosystems: evidence from a meta-analysis.

Both gradual and extreme weather changes trigger complex ecological responses in river ecosystems. It is still unclear to what extent trend or event effects alter biodiversity and functioning in river ecosystems, adding considerable uncertainty to predictions of their future dynamics. Using a comprehensive database of 71 published studies, we show that event - but not trend - effects associated with extreme changes in water flow and temperature substantially reduce species richness. Furthermore, event effects - particularly those affecting hydrological dynamics - on biodiversity and primary productivity were twice as high as impacts due to gradual changes. The synthesis of the available evidence reveals that event effects induce regime shifts in river ecosystems, particularly affecting organisms such as invertebrates. Among extreme weather events, dryness associated with flow interruption caused the largest effects on biota and ecosystem functions in rivers. Effects on ecosystem functions (primary production, organic matter decomposition and respiration) were asymmetric, with only primary production exhibiting a negative response to extreme weather events. Our meta-analysis highlights the disproportionate impact of event effects on river biodiversity and ecosystem functions, with implications for the long-term conservation and management of river ecosystems. However, few studies were available from tropical areas, and our conclusions therefore remain largely limited to temperate river systems. Further efforts need to be directed to assemble evidence of extreme events on river biodiversity and functioning.

Strong floristic distinctiveness across Neotropical successional forests.

Forests that regrow naturally on abandoned fields are important for restoring biodiversity and ecosystem services, but can they also preserve the distinct regional tree floras? Using the floristic composition of 1215 early successional forests (≤20 years) in 75 human-modified landscapes across the Neotropic realm, we identified 14 distinct floristic groups, with a between-group dissimilarity of 0.97. Floristic groups were associated with location, bioregions, soil pH, temperature seasonality, and water availability. Hence, there is large continental-scale variation in the species composition of early successional forests, which is mainly associated with biogeographic and environmental factors but not with human disturbance indicators. This floristic distinctiveness is partially driven by regionally restricted species belonging to widespread genera. Early secondary forests contribute therefore to restoring and conserving the distinctiveness of bioregions across the Neotropical realm, and forest restoration initiatives should use local species to assure that these distinct floras are maintained.

Biodiversity promotes ecosystem functioning despite environmental change.

Three decades of research have demonstrated that biodiversity can promote the functioning of ecosystems. Yet, it is unclear whether the positive effects of biodiversity on ecosystem functioning will persist under various types of global environmental change drivers. We conducted a meta-analysis of 46 factorial experiments manipulating both species richness and the environment to test how global change drivers (i.e. warming, drought, nutrient addition or CO2 enrichment) modulated the effect of biodiversity on multiple ecosystem functions across three taxonomic groups (microbes, phytoplankton and plants). We found that biodiversity increased ecosystem functioning in both ambient and manipulated environments, but often not to the same degree. In particular, biodiversity effects on ecosystem functioning were larger in stressful environments induced by global change drivers, indicating that high-diversity communities were more resistant to environmental change. Using a subset of studies, we also found that the positive effects of biodiversity were mainly driven by interspecific complementarity and that these effects increased over time in both ambient and manipulated environments. Our findings support biodiversity conservation as a key strategy for sustainable ecosystem management in the face of global environmental change.

Contrasts among cationic phytochemical landscapes in the southern United States.

Understanding the phytochemical landscapes of essential and nonessential chemical elements to plants provides an opportunity to better link biogeochemical cycles to trophic ecology. We investigated the formation and regulation of the cationic phytochemical landscapes of four key elements for biota: Ca, Mg, K, and Na. We collected aboveground tissues of plants in Atriplex, Helianthus, and Opuntia and adjacent soils from 51, 131, and 83 sites, respectively, across the southern United States. We determined the spatial variability of these cations in plants and soils. Also, we quantified the homeostasis coefficient for each cation and genus combination, by using mixed-effect models, with spatially correlated random effects. Additionally, using random forest models, we modeled the influence of bioclimatic, soil, and spatial variables on plant cationic concentrations. Sodium variability and spatial autocorrelation were considerably greater than for Ca, Mg, or K. Calcium, Mg, and K exhibited strongly homeostatic patterns, in striking contrast to non-homeostatic Na. Even so, climatic and soil variables explained a large proportion of plants' cationic concentrations. Essential elements (Ca, Mg, and K) appeared to be homeostatically regulated, which contrasted sharply with Na, a nonessential element for most plants. In addition, we provide evidence for the No-Escape-from-Sodium hypothesis in real-world ecosystems, indicating that plant Na concentrations tend to increase as substrate Na levels increase.

Multidimensional tropical forest recovery.

Tropical forests disappear rapidly because of deforestation, yet they have the potential to regrow naturally on abandoned lands. We analyze how 12 forest attributes recover during secondary succession and how their recovery is interrelated using 77 sites across the tropics. Tropical forests are highly resilient to low-intensity land use; after 20 years, forest attributes attain 78% (33 to 100%) of their old-growth values. Recovery to 90% of old-growth values is fastest for soil (<1 decade) and plant functioning (<2.5 decades), intermediate for structure and species diversity (2.5 to 6 decades), and slowest for biomass and species composition (>12 decades). Network analysis shows three independent clusters of attribute recovery, related to structure, species diversity, and species composition. Secondary forests should be embraced as a low-cost, natural solution for ecosystem restoration, climate change mitigation, and biodiversity conservation.

Functional recovery of secondary tropical forests.

One-third of all Neotropical forests are secondary forests that regrow naturally after agricultural use through secondary succession. We need to understand better how and why succession varies across environmental gradients and broad geographic scales. Here, we analyze functional recovery using community data on seven plant characteristics (traits) of 1,016 forest plots from 30 chronosequence sites across the Neotropics. By analyzing communities in terms of their traits, we enhance understanding of the mechanisms of succession, assess ecosystem recovery, and use these insights to propose successful forest restoration strategies. Wet and dry forests diverged markedly for several traits that increase growth rate in wet forests but come at the expense of reduced drought tolerance, delay, or avoidance, which is important in seasonally dry forests. Dry and wet forests showed different successional pathways for several traits. In dry forests, species turnover is driven by drought tolerance traits that are important early in succession and in wet forests by shade tolerance traits that are important later in succession. In both forests, deciduous and compound-leaved trees decreased with forest age, probably because microclimatic conditions became less hot and dry. Our results suggest that climatic water availability drives functional recovery by influencing the start and trajectory of succession, resulting in a convergence of community trait values with forest age when vegetation cover builds up. Within plots, the range in functional trait values increased with age. Based on the observed successional trait changes, we indicate the consequences for carbon and nutrient cycling and propose an ecologically sound strategy to improve forest restoration success.

Biovera-Epi: A new database on species diversity, community composition and leaf functional traits of vascular epiphytes along gradients of elevation and forest-use intensity in Mexico.

BACKGROUND: This data paper describes a new, comprehensive database (BIOVERA-Epi) on species distributions and leaf functional traits of vascular epiphytes, a poorly studied plant group, along gradients of elevation and forest-use intensity in the central part of Veracruz State, Mexico. The distribution data include frequencies of 271 vascular epiphyte species belonging to 92 genera and 23 families across 120 20 m × 20 m forest plots at eight study sites along an elevational gradient from sea level to 3500 m a.s.l. In addition, BIOVERA-Epi provides information on 1595 measurements of nine morphological and chemical leaf traits from 474 individuals and 102 species. For morphological leaf traits, we provide data on each sampled leaf. For chemical leaf traits, we provide data at the species level per site and land-use type. We also provide complementary information for each of the sampled plots and host trees. BIOVERA-Epi contributes to an emerging body of synthetic epiphytes studies combining functional traits and community composition. NEW INFORMATION: BIOVERA-Epi includes data on species frequency and leaf traits from 120 forest plots distributed along an elevational gradient, including six different forest types and three levels of forest-use intensity. It will expand the breadth of studies on epiphyte diversity, conservation and functional plant ecology in the Neotropics and will contribute to future synthetic studies on the ecology and diversity of tropical epiphyte assemblages.

BIOVERA-Tree: tree diversity, community composition, forest structure and functional traits along gradients of forest-use intensity and elevation in Veracruz, Mexico.

BACKGROUND: Here, we describe BIOVERA-Tree, a database on tree diversity, community composition, forest structure and functional traits collected in 120 forest plots, distributed along an extensive elevational gradient in Veracruz State, Mexico. BIOVERA-Tree includes information on forest structure from three levels of forest-use intensity, namely old-growth, degraded and secondary forest, replicated across eight elevations from sea-level to near the tree line at 3500 m and on size and location of 4549 tree individuals with a diameter at breast height ≥ 5 cm belonging to 216 species, 154 genera and 80 families. We also report measurements of eight functional traits, namely wood density for 143 species, maximum height for 216 species and leaf traits including: specific leaf area, lamina density, leaf thickness, chlorophyll content and leaf area for 148 species and leaf dry matter content for 145 species. NEW INFORMATION: BIOVERA-Tree is a new database comprising data collected in a rigorous sampling design along forest-use intensity and elevational gradients, contributing to our understanding of how interactive effects of forest-use intensity and elevation affect tree diversity, community composition and functional traits in tropical forests.

Data Descriptor: Pacific Introduced Flora (PaciFLora).

BACKGROUND: The Pacific Region has the highest density of naturalised plant species worldwide, which makes it an important area for research on the ecology, evolution and biogeography of biological invasions. While different data sources on naturalised plant species exist for the Pacific, there is no taxonomically and spatially harmonised database available for different subsets of species and islands. A comprehensive, accessible database containing the distribution of naturalised vascular plant species in the Pacific will enable new basic and applied research for researchers and will be an important information source for practitioners working in the Region. NEW INFORMATION: Here, we present PacIFlora, an updated and taxonomically standardised list of naturalised species, their unified nativeness, cultivation and invasive status and their distribution across the Pacific Ocean, including harmonised location denoination. This list is based on the two largest databases on naturalised plants for the Region, specifically the Pacific Island Ecosystems at Risk (PIER) and the Global Naturalised Alien Flora (GloNAF) databases. We provide an outlook for how this database can contribute to numerous research questions and conservation efforts.

Successional syndromes of saplings in tropical secondary forests emerge from environment-dependent trait-demography relationships.

Identifying generalisable processes that underpin population dynamics is crucial for understanding successional patterns. While longitudinal or chronosequence data are powerful tools for doing so, the traditional focus on community-level shifts in taxonomic and functional composition rather than species-level trait-demography relationships has made generalisation difficult. Using joint species distribution models, we demonstrate how three traits-photosynthetic rate, adult stature, and seed mass-moderate recruitment and sapling mortality rates of 46 woody species during secondary succession. We show that the pioneer syndrome emerges from higher photosynthetic rates, shorter adult statures and lighter seeds that facilitate exploitation of light in younger secondary forests, while 'long-lived pioneer' and 'late successional' syndromes are associated with trait values that enable species to persist in the understory or reach the upper canopy in older secondary forests. Our study highlights the context dependency of trait-demography relationships, which drive successional shifts in sapling's species composition in secondary forests.

Biotic homogenization destabilizes ecosystem functioning by decreasing spatial asynchrony.

Our planet is facing significant changes of biodiversity across spatial scales. Although the negative effects of local biodiversity (α diversity) loss on ecosystem stability are well documented, the consequences of biodiversity changes at larger spatial scales, in particular biotic homogenization, that is, reduced species turnover across space (ß diversity), remain poorly known. Using data from 39 grassland biodiversity experiments, we examine the effects of ß diversity on the stability of simulated landscapes while controlling for potentially confounding biotic and abiotic factors. Our results show that higher ß diversity generates more asynchronous dynamics among local communities and thereby contributes to the stability of ecosystem productivity at larger spatial scales. We further quantify the relative contributions of α and ß diversity to ecosystem stability and find a relatively stronger effect of α diversity, possibly due to the limited spatial scale of our experiments. The stabilizing effects of both α and ß diversity lead to a positive diversity-stability relationship at the landscape scale. Our findings demonstrate the destabilizing effect of biotic homogenization and suggest that biodiversity should be conserved at multiple spatial scales to maintain the stability of ecosystem functions and services.

Evaluating forest resilience to global threats using functional response traits and network properties.

Ecosystem functions provided by forests are threatened by direct and indirect effects of global change drivers such as climate warming land-use change, biological invasions, and shifting natural disturbance regimes. To develop resilience-based forest management, new tools and methods are needed to quantitatively estimate forest resilience to management and future natural disturbances. We propose a multidimensional evaluation of ecological resilience based on species functional response traits (e.g., functional response diversity and functional redundancy) and network properties of forested patches (e.g., connectivity, modularity, and centrality). Using a fragmented rural landscape in temperate south-eastern Canada as a reference landscape, we apply our multidimensional approach to evaluate two alternative management strategies at three levels of intensity: (1) functional enrichment of current forest patches and (2) multi-species plantations in previously non-forested patches. Within each management strategy, planted species are selected to maximize functional diversity, drought tolerance, or pest resistance. We further compare how ecological resilience under these alternative management strategies responds to three simulated disturbances: drought, pest outbreak, and timber harvesting. We found that both management strategies enhance resilience at the landscape scale by increasing functional response diversity and connectivity. Specifically, when the less functionally diverse patches are prioritized for management, functional enrichment is more effective than the establishment of new multi-species plantations in increasing resilience. In addition, randomly allocated multi-species plantations increased connectivity more than those allocated in riparian areas. Our results show that across various management strategies, planting species to enhance biodiversity led to the highest increase in functional response diversity while planting pest-resistant species led to the highest increase in landscape connectivity. Planting biodiversity-enhancing species (i.e., species that maximize functional diversity) mitigated drought effects equally well as planting with drought-tolerant species. Our multidimensional approach facilitates the characterization at the landscape scale of forest resilience to disturbances using both functional diversity and network properties while accounting for the importance of response traits to future disturbances. The simulation approach we used can be applied to forest landscapes across different biomes for the evaluation and comparison of forest management initiatives to enhance resilience.

Successional, spatial, and seasonal changes in seed rain in the Atlantic forest of southern Bahia, Brazil.

Seed arrival is a limiting factor for the regeneration of diverse tropical forests and may be an important mechanism that drives patterns of tree species' distribution. Here we quantify spatial and seasonal variation in seed rain of secondary forests in southern Bahia, Brazil. We also examine whether secondary forest age enhances seed dispersal and whether seed rain density and diversity in secondary forests decay with distance from mature forest. Across a chronosequence of 15 pairs of mature and secondary forests, 105 seed traps were installed and monitored for one year. We tested the effects of secondary forest age, distance from mature forest, and seasonality on monthly seed rain density, diversity, seed dispersal mode, and diaspore size. We found that secondary forest age had strong, positive effects on the diversity of seed rain, which was generally higher during the wet season. Moreover, contrasting patterns among diversity indices revealed that seeds of rare species occurred more often in 40 yr old secondary forests and mature forests. While the proportion of biotically and abiotically dispersed seeds did not change significantly with distance from mature forest across all forest age classes, we found that biotically dispersed seeds contributed disproportionately more to seed rain diversity. Our results emphasize the importance of biotic dispersal to enhance diversity during secondary succession and suggest that changes in secondary forest structure have the potential to enhance the diversity of tropical secondary forests, principally by increasing dispersal of rare species.

Author Correction: Diversity-dependent temporal divergence of ecosystem functioning in experimental ecosystems.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.