Biodiversity in changing environments: An external-driver internal-topology framework to guide intervention.

Accompanying the climate crisis is the more enigmatic biodiversity crisis. Rapid reorganization of biodiversity due to global environmental change has defied prediction and tested the basic tenets of conservation and restoration. Conceptual and practical innovation is needed to support decision making in the face of these unprecedented shifts. Critical questions include: How can we generalize biodiversity change at the community level? When are systems able to reorganize and maintain integrity, and when does abiotic change result in collapse or restructuring? How does this understanding provide a template to guide when and how to intervene in conservation and restoration? To this end, we frame changes in community organization as the modulation of external abiotic drivers on the internal topology of species interactions, using plant-plant interactions in terrestrial communities as a starting point. We then explore how this framing can help translate available data on species abundance and trait distributions to corresponding decisions in management. Given the expectation that community response and reorganization are highly complex, the external-driver internal-topology (EDIT) framework offers a way to capture general patterns of biodiversity that can help guide resilience and adaptation in changing environments.

Extinction debt and functional traits mediate community saturation over large spatiotemporal scales.

Determining if ecological communities are saturated (have a limit to the number of species they can support) has important implications for understanding community assembly, species invasions, and climate change. However, previous studies have generally been limited to short time frames that overlook extinction debt and have not explicitly considered how functional trait diversity may mediate patterns of community saturation. Here, we combine data from biodiversity surveys with functional and phylogenetic data to explore if the colonisation events after the Great American Biotic Interchange (closure of the Panamanian Isthmus) resulted in increases in species richness of communities of the snake family Dipsadidae. We determined the number and the direction of dispersal events between Central and South America by estimating ancestral areas based on a Bayesian time-calibrated phylogenetic analysis. We then evaluated whether variation in community saturation was mediated by the functional similarity of six traits for the resident and colonizing snakes and/or local environmental conditions. We found that colonised communities did not support more species than those that were not colonised. Moreover, we did not find an association between the functional diversity across sites and whether they were colonised by members from the lineages dispersing across the Isthmus or not. Instead, variation in species richness was predicted best by covariates such as time since colonisation and local environment. Taken together, our results suggest that snake communities of the Dipsadidae across the neotropics are saturated. Moreover, our research highlights two important factors to consider in studies of community saturation: extinction debt and the functional differences and similarities in species' ecological roles.

Determinar si las comunidades ecológicas están saturadas (si tienen un límite en el número de especies que pueden albergar) tiene importantes implicaciones para entender el ensamblaje de comunidades, las invasiones de especies y el cambio climático. Sin embargo, los estudios previos en esta área se han limitado generalmente a marcos temporales cortos, ignorando el concepto de deuda de extinción y no considerando explícitamente cómo la diversidad de rasgos funcionales puede mediar en los patrones de saturación de las comunidades. En este trabajo combinamos datos publicados de muestreos de campo con datos funcionales y filogenéticos para explorar si los eventos de colonización después del Gran Intercambio Biótico Americano (ocurrido con el cierre del istmo de Panamá) resultaron en aumentos en la riqueza de especies de las comunidades de la familia de serpientes Dipsadidae. Determinamos el número y la dirección de los eventos de dispersión entre América Central y América del Sur mediante la estimación de áreas ancestrales basada en un análisis filogenético Bayesiano calibrado en el tiempo. Luego evaluamos si la variación en la saturación de las comunidades estaba mediada por la similitud funcional de seis rasgos para las serpientes residentes y colonizadoras y/o por las condiciones ambientales locales. Encontramos que las comunidades colonizadas no contienen más especies que aquellas que no fueron colonizadas. Además, no encontramos ninguna relación entre la diversidad funcional de los sitios considerados y el hecho de que estuvieran colonizados o no por miembros de los linajes que se dispersaron a través del Istmo. En cambio, la variación en la riqueza de especies se predijo mejor por covariantes como el tiempo transcurrido desde la colonización y el clima local. En conjunto, nuestros resultados sugieren que las comunidades de Dipsadidae a lo largo del neotrópico están saturadas. Además, nuestra investigación destaca dos factores importantes a considerar en los estudios de saturación de comunidades: la existencia de una deuda de extinción y las diferencias y similitudes funcionales en los papeles ecológicos de las especies.

Drivers of bacterial and fungal root endophyte communities: understanding the relative influence of host plant, environment, and space.

Bacterial and fungal root endophytes can impact the fitness of their host plants, but the relative importance of drivers for root endophyte communities is not well known. Host plant species, the composition and density of the surrounding plants, space, and abiotic drivers could significantly affect bacterial and fungal root endophyte communities. We investigated their influence in endophyte communities of alpine plants across a harsh high mountain landscape using high-throughput sequencing. There was less compositional overlap between fungal than bacterial root endophyte communities, with four 'cosmopolitan' bacterial OTUs found in every root sampled, but no fungal OTUs found across all samples. We found that host plant species, which included nine species from three families, explained the greatest variation in root endophyte composition for both bacterial and fungal communities. We detected similar levels of variation explained by plant neighborhood, space, and abiotic drivers on both communities, but the plant neighborhood explained less variation in fungal endophytes than expected. Overall, these findings suggest a more cosmopolitan distribution of bacterial OTUs compared to fungal OTUs, a structuring role of the plant host species for both communities, and largely similar effects of the plant neighborhood, abiotic drivers, and space on both communities.

Precipitation legacies amplify ecosystem nitrogen losses from nitric oxide emissions in a Pinyon-Juniper dryland.

Climate change is increasing the variability of precipitation, altering the frequency of soil drying-wetting events and the distribution of seasonal precipitation. These changes in precipitation can alter nitrogen (N) cycling and stimulate nitric oxide (NO) emissions (an air pollutant at high concentrations), which may vary according to legacies of past precipitation and represent a pathway for ecosystem N loss. To identify whether precipitation legacies affect NO emissions, we excluded or added precipitation during the winter growing season in a Pinyon-Juniper dryland and measured in situ NO emissions following experimental wetting. We found that the legacy of both excluding and adding winter precipitation increased NO emissions early in the following summer; cumulative NO emissions from the winter precipitation exclusion plots (2750 ± 972 µg N-NO m-2 ) and winter water addition plots (2449 ± 408 µg N-NO m-2 ) were higher than control plots (1506 ± 397 µg N-NO m-2 ). The increase in NO emissions with previous precipitation exclusion was associated with inorganic N accumulation, while the increase in NO emissions with previous water addition was associated with an upward trend in microbial biomass. Precipitation legacies can accelerate soil NO emissions and may amplify ecosystem N loss in response to more variable precipitation.

Beta diversity as a driver of forest biomass across spatial scales.

Despite the importance of biodiversity-ecosystem functioning (BEF) relationships in ecology and conservation, relatively little is known about how BEF relationships change across spatial scales. Theory predicts that change in BEF relationships with increasing spatial scale will depend on variation in species composition across space (ß-diversity), but empirical evidence for this is limited. Moreover, studies have not quantified the direct and indirect role the environment plays in costructuring ecosystem functioning across spatial scales. We used 14 temperate-forest plots 1.4 ha in size containing 18,323 trees to quantify scale-dependence between aboveground tree biomass and three components of tree-species diversity-α-diversity (average local diversity), γ-diversity (total diversity), and ß-diversity. Using structural-equation models, we quantified the direct effects of each diversity component and the environment (soil nutrients and topography), as well as indirect effects of the environment, on tree biomass across 11 spatial extents ranging from 400 to 14,400 m2 . Our results show that the relationship between ß-diversity and tree biomass strengthened with increasing spatial extent. Moreover, ß-diversity appeared to be a stronger predictor of biomass than α-diversity and γ-diversity at intermediate to large spatial extents. The environment had strong direct and indirect effects on biomass, but, in contrast to diversity, these effects did not strengthen with increasing spatial extent. This study provides some of the first empirical evidence that ß-diversity underpins the scaling of BEF relationships in naturally complex ecosystems.

Clustering analysis of large-scale phenotypic data in the model filamentous fungus Neurospora crassa.

BACKGROUND: With 9730 protein-coding genes and a nearly complete gene knockout strain collection, Neurospora crassa is a major model organism for filamentous fungi. Despite this abundance of information, the phenotypes of these gene knockout mutants have not been categorized to determine whether there are broad correlations between phenotype and any genetic features. RESULTS: Here, we analyze data for 10 different growth or developmental phenotypes that have been obtained for 1168 N. crassa knockout mutants. Of these mutants, 265 (23%) are in the normal range, while 903 (77%) possess at least one mutant phenotype. With the exception of unclassified functions, the distribution of functional categories for genes in the mutant dataset mirrors that of the N. crassa genome. In contrast, most genes do not possess a yeast ortholog, suggesting that our analysis will reveal functions that are not conserved in Saccharomyces cerevisiae. To leverage the phenotypic data to identify pathways, we used weighted Partitioning Around Medoids (PAM) approach with 40 clusters. We found that genes encoding metabolic, transmembrane and protein phosphorylation-related genes are concentrated in subsets of clusters. Results from K-Means clustering of transcriptomic datasets showed that most phenotypic clusters contain multiple expression profiles, suggesting that co-expression is not generally observed for genes with shared phenotypes. Analysis of yeast orthologs of genes that co-clustered in MAPK signaling cascades revealed potential networks of interacting proteins in N. crassa. CONCLUSIONS: Our results demonstrate that clustering analysis of phenotypes is a promising tool for generating new hypotheses regarding involvement of genes in cellular pathways in N. crassa. Furthermore, information about gene clusters identified in N. crassa should be applicable to other filamentous fungi, including saprobes and pathogens.

Belowground impacts of alpine woody encroachment are determined by plant traits, local climate, and soil conditions.

Global climate and land use change are causing woody plant encroachment in arctic, alpine, and arid/semi-arid ecosystems around the world, yet our understanding of the belowground impacts of this phenomenon is limited. We conducted a globally distributed field study of 13 alpine sites across four continents undergoing woody plant encroachment and sampled soils from both woody encroached and nearby herbaceous plant community types. We found that woody plant encroachment influenced soil microbial richness and community composition across sites based on multiple factors including woody plant traits, site level climate, and abiotic soil conditions. In particular, root symbiont type was a key determinant of belowground effects, as Nitrogen-fixing woody plants had higher soil fungal richness, while Ecto/Ericoid mycorrhizal species had higher soil bacterial richness and symbiont types had distinct soil microbial community composition. Woody plant leaf traits indirectly influenced soil microbes through their impact on soil abiotic conditions, primarily soil pH and C:N ratios. Finally, site-level climate affected the overall magnitude and direction of woody plant influence, as soil fungal and bacterial richness were either higher or lower in woody encroached versus herbaceous soils depending on mean annual temperature and precipitation. All together, these results document global impacts of woody plant encroachment on soil microbial communities, but highlight that multiple biotic and abiotic pathways must be considered to scale up globally from site- and species-level patterns. Considering both the aboveground and belowground effects of woody encroachment will be critical to predict future changes in alpine ecosystem structure and function and subsequent feedbacks to the global climate system.

Publisher Correction: Open Science principles for accelerating trait-based science across the Tree of Life.

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

Climate and plant community diversity in space and time.

Climate strongly shapes plant diversity over large spatial scales, with relatively warm and wet (benign, productive) regions supporting greater numbers of species. Unresolved aspects of this relationship include what causes it, whether it permeates to community diversity at smaller spatial scales, whether it is accompanied by patterns in functional and phylogenetic diversity as some hypotheses predict, and whether it is paralleled by climate-driven changes in diversity over time. Here, studies of Californian plants are reviewed and new analyses are conducted to synthesize climate-diversity relationships in space and time. Across spatial scales and organizational levels, plant diversity is maximized in more productive (wetter) climates, and these consistent spatial relationships are mirrored in losses of taxonomic, functional, and phylogenetic diversity over time during a recent climatic drying trend. These results support the tolerance and climatic niche conservatism hypotheses for climate-diversity relationships, and suggest there is some predictability to future changes in diversity in water-limited climates.

Open Science principles for accelerating trait-based science across the Tree of Life.

Synthesizing trait observations and knowledge across the Tree of Life remains a grand challenge for biodiversity science. Species traits are widely used in ecological and evolutionary science, and new data and methods have proliferated rapidly. Yet accessing and integrating disparate data sources remains a considerable challenge, slowing progress toward a global synthesis to integrate trait data across organisms. Trait science needs a vision for achieving global integration across all organisms. Here, we outline how the adoption of key Open Science principles-open data, open source and open methods-is transforming trait science, increasing transparency, democratizing access and accelerating global synthesis. To enhance widespread adoption of these principles, we introduce the Open Traits Network (OTN), a global, decentralized community welcoming all researchers and institutions pursuing the collaborative goal of standardizing and integrating trait data across organisms. We demonstrate how adherence to Open Science principles is key to the OTN community and outline five activities that can accelerate the synthesis of trait data across the Tree of Life, thereby facilitating rapid advances to address scientific inquiries and environmental issues. Lessons learned along the path to a global synthesis of trait data will provide a framework for addressing similarly complex data science and informatics challenges.

Accurate forest projections require long-term wood decay experiments because plant trait effects change through time.

Whether global change will drive changing forests from net carbon (C) sinks to sources relates to how quickly deadwood decomposes. Because complete wood mineralization takes years, most experiments focus on how traits, environments and decomposer communities interact as wood decay begins. Few experiments last long enough to test whether drivers change with decay rates through time, with unknown consequences for scaling short-term results up to long-term forest ecosystem projections. Using a 7 year experiment that captured complete mineralization among 21 temperate tree species, we demonstrate that trait effects fade with advancing decay. However, wood density and vessel diameter, which may influence permeability, control how decay rates change through time. Denser wood loses mass more slowly at first but more quickly with advancing decay, which resolves ambiguity about the after-life consequences of this key plant functional trait by demonstrating that its effect on decay depends on experiment duration and sampling frequency. Only long-term data and a time-varying model yielded accurate predictions of both mass loss in a concurrent experiment and naturally recruited deadwood structure in a 32-year-old forest plot. Given the importance of forests in the carbon cycle, and the pivotal role for wood decay, accurate ecosystem projections are critical and they require experiments that go beyond enumerating potential mechanisms by identifying the temporal scale for their effects.

Cascading effects of mammalian herbivores on ground-dwelling arthropods: Variable responses across arthropod groups, habitats and years.

Large mammalian herbivores are well known to shape the structure and function of ecosystems world-wide, and these effects can in turn cascade through systems to indirectly influence other animal species. A wealth of studies has explored the effects of large mammals on arthropods, but to date they have reported such widely varying results that generalizations have been elusive. Three factors are likely drivers of this variability: the widely varying life-history characteristics of different arthropod groups, the highly variable landscapes that mammalian herbivores commonly inhabit and temporal variation in environmental conditions. Here, we use an 18-year-old exclosure experiment stratified across three distinct coastal prairie habitats in northern California to address the effects of a reintroduced mammalian herbivore, tule elk (Cervus canadensis nannodes) on the composition, richness and abundance of ground-dwelling arthropods over two years with very different precipitation regimes. We found that elk shifted the composition of arthropod communities, increasing the abundance of ants, beetles, spiders and mites, decreasing the abundance of woodlice and bristletails in some but not all habitats types, and having no effect on the abundance of bugs, crickets and springtails. Elk also increased richness and changed the composition of ant genera and beetle morpho-species. Interestingly, the effects of elk on arthropod composition, richness and abundance varied little between years, despite very different precipitation levels, biomass accumulation and thatch height. Elk reduced shrub cover, above-ground herbaceous biomass and thatch height and increased soil compaction, and these changes predicted the abundance and richness of arthropods, although taxonomic groups varied in their responses, presumably due to differences in environmental requirements. Synthesis. Our research highlights the importance of using long-term experiments to assess the cascading effects of large herbivores on the composition of grounddwelling arthropod communities and to identify the mechanisms that indirectly shape arthropod responses to herbivores among variable habitats and years in order to develop a greater understanding of the variable responses of arthropods to large mammalian herbivores.

Soil Microbial Networks Shift Across a High-Elevation Successional Gradient.

While it is well established that microbial composition and diversity shift along environmental gradients, how interactions among microbes change is poorly understood. Here, we tested how community structure and species interactions among diverse groups of soil microbes (bacteria, fungi, non-fungal eukaryotes) change across a fundamental ecological gradient, succession. Our study system is a high-elevation alpine ecosystem that exhibits variability in successional stage due to topography and harsh environmental conditions. We used hierarchical Bayesian joint distribution modeling to remove the influence of environmental covariates on species distributions and generated interaction networks using the residual species-to-species variance-covariance matrix. We hypothesized that as ecological succession proceeds, diversity will increase, species composition will change, and soil microbial networks will become more complex. As expected, we found that diversity of most taxonomic groups increased over succession, and species composition changed considerably. Interestingly, and contrary to our hypothesis, interaction networks became less complex over succession (fewer interactions per taxon). Interactions between photosynthetic microbes and any other organism became less frequent over the gradient, whereas interactions between plants or soil microfauna and any other organism were more abundant in late succession. Results demonstrate that patterns in diversity and composition do not necessarily relate to patterns in network complexity and suggest that network analyses provide new insight into the ecology of highly diverse, microscopic communities.

Plant functional trait change across a warming tundra biome.

The tundra is warming more rapidly than any other biome on Earth, and the potential ramifications are far-reaching because of global feedback effects between vegetation and climate. A better understanding of how environmental factors shape plant structure and function is crucial for predicting the consequences of environmental change for ecosystem functioning. Here we explore the biome-wide relationships between temperature, moisture and seven key plant functional traits both across space and over three decades of warming at 117 tundra locations. Spatial temperature-trait relationships were generally strong but soil moisture had a marked influence on the strength and direction of these relationships, highlighting the potentially important influence of changes in water availability on future trait shifts in tundra plant communities. Community height increased with warming across all sites over the past three decades, but other traits lagged far behind predicted rates of change. Our findings highlight the challenge of using space-for-time substitution to predict the functional consequences of future warming and suggest that functions that are tied closely to plant height will experience the most rapid change. They also reveal the strength with which environmental factors shape biotic communities at the coldest extremes of the planet and will help to improve projections of functional changes in tundra ecosystems with climate warming.

Plant diversity and density predict belowground diversity and function in an early successional alpine ecosystem.

Despite decades of interest, few studies have provided evidence supporting theoretical expectations for coupled relationships between aboveground and belowground diversity and ecosystem functioning in non-manipulated natural ecosystems. We characterized plant species richness and density, soil bacterial, fungal and eukaryotic species richness and phylogenetic diversity (using 16S, ITS, and 18S gene sequencing), and ecosystem function (levels of soil C and N, and rates of microbial enzyme activities) along a natural gradient in plant richness and density in high-elevation, C-deficient soils to examine the coupling between above- and belowground systems. Overall, we observed a strong positive relationship between aboveground (plant richness and density) and belowground (bacteria, fungi, and non-fungal eukaryotes) richness. In addition to the correlations between plants and soil communities, C and N pools, and rates of enzyme activities increased as plant and soil communities became richer and more diverse. Our results suggest that the theoretically expected positive correlation between above- and belowground communities does exist in natural systems, but may be undetectable in late successional ecosystems due to the buildup of legacy organic matter that results in extremely complex belowground communities. In contrast, microbial communities in early successional systems, such as the system described here, are more directly dependent on contemporary inputs from plants and therefore are strongly correlated with plant diversity and density.

Landscape Physiognomy Influences Abundance of the Lone Star Tick, Amblyomma americanum (Ixodida: Ixodidae), in Ozark Forests.

The lone star tick, Amblyomma americanum Linnaeus (Ixodida: Ixodidae), is emerging as an important human disease vector in the United States. While some recent studies have modeled broad-scale (regional or county-level) distribution patterns of A. americanum, less is known about how local-scale habitat characteristics drive A. americanum abundance. Such local-scale information is vital to identify targets for tick population control measures within land management units. We investigated how habitat features predict host-seeking A. americanum adult and nymph abundance within a 12-ha oak-hickory forest plot in the Missouri Ozarks. We trapped ticks using CO2-baited traps at 40 evenly spaced locations for three 24-h periods during the summer of 2015, and we measured biotic and abiotic variables surrounding each location. Of 2,008 A. americanum captured, 1,009 were nymphs, and 999 were adults. We observed spatial heterogeneity in local tick abundance (min = 0 ticks, max = 112 ticks, mean = 16.7 ticks per trap night). Using generalized linear mixed models, we found that both nymphs and adults had greater abundance in valleys as well as on northern-facing aspects. Moreover, nymph abundance was negatively related to temperature variance, while adult abundance had a negative relationship with elevation. These results demonstrate that managers in this region may be able to predict local tick abundance through simple physiognomic factors and use these parameters for targeted management action.

Integrating species traits into species pools.

Despite decades of research on the species-pool concept and the recent explosion of interest in trait-based frameworks in ecology and biogeography, surprisingly little is known about how spatial and temporal changes in species-pool functional diversity (SPFD) influence biodiversity and the processes underlying community assembly. Current trait-based frameworks focus primarily on community assembly from a static regional species pool, without considering how spatial or temporal variation in SPFD alters the relative importance of deterministic and stochastic assembly processes. Likewise, species-pool concepts primarily focus on how the number of species in the species pool influences local biodiversity. However, species pools with similar richness can vary substantially in functional-trait diversity, which can strongly influence community assembly and biodiversity responses to environmental change. Here, we integrate recent advances in community ecology, trait-based ecology, and biogeography to provide a more comprehensive framework that explicitly considers how variation in SPFD, among regions and within regions through time, influences the relative importance of community assembly processes and patterns of biodiversity. First, we provide a brief overview of the primary ecological and evolutionary processes that create differences in SPFD among regions and within regions through time. We then illustrate how SPFD may influence fundamental processes of local community assembly (dispersal, ecological drift, niche selection). Higher SPFD may increase the relative importance of deterministic community assembly when greater functional diversity in the species pool increases niche selection across environmental gradients. In contrast, lower SPFD may increase the relative importance of stochastic community assembly when high functional redundancy in the species pool increases the influence of dispersal history or ecological drift. Next, we outline experimental and observational approaches for testing the influence of SPFD on assembly processes and biodiversity. Finally, we highlight applications of this framework for restoration and conservation. This species-pool functional diversity framework has the potential to advance our understanding of how local- and regional-scale processes jointly influence patterns of biodiversity across biogeographic regions, changes in biodiversity within regions over time, and restoration outcomes and conservation efforts in ecosystems altered by environmental change.

Seed banks of native forbs, but not exotic grasses, increase during extreme drought.

Extreme droughts such as the one that affected California in 2012-2015 have been linked to severe ecological consequences in perennial-dominated communities such as forests. In annual communities, drought impacts are difficult to assess because many species persist through facultative multiyear seed dormancy, which leads to the development of seed banks. Impacts of extreme drought on the abundance and composition of the seed banks of whole communities are little known. In 80 heterogeneous grassland plots where cover is dominated by ~15 species of exotic annual grasses and diversity is dominated by ~70 species of native annual forbs, we grew out seeds from soil cores collected early in the California drought (2012) and later in the multiyear drought (2014), and analyzed drought-associated changes in the seed bank. Over the course of the study we identified more than 22,000 seedlings to species. We found that seeds of exotic annual grasses declined sharply in abundance during the drought while seeds of native annual forbs increased, a pattern that resembled but was even stronger than the changes in aboveground cover of these groups. Consistent with the expectation that low specific leaf area (SLA) is an indicator of drought tolerance, we found that the community-weighted mean SLA of annual forbs declined both in the seed bank and in the aboveground community, as low-SLA forbs increased disproportionately. In this system, seed dormancy reinforces the indirect benefits of extreme drought to the native forb community.

Mapping local and global variability in plant trait distributions.

Our ability to understand and predict the response of ecosystems to a changing environment depends on quantifying vegetation functional diversity. However, representing this diversity at the global scale is challenging. Typically, in Earth system models, characterization of plant diversity has been limited to grouping related species into plant functional types (PFTs), with all trait variation in a PFT collapsed into a single mean value that is applied globally. Using the largest global plant trait database and state of the art Bayesian modeling, we created fine-grained global maps of plant trait distributions that can be applied to Earth system models. Focusing on a set of plant traits closely coupled to photosynthesis and foliar respiration-specific leaf area (SLA) and dry mass-based concentrations of leaf nitrogen ([Formula: see text]) and phosphorus ([Formula: see text]), we characterize how traits vary within and among over 50,000 [Formula: see text]-km cells across the entire vegetated land surface. We do this in several ways-without defining the PFT of each grid cell and using 4 or 14 PFTs; each model's predictions are evaluated against out-of-sample data. This endeavor advances prior trait mapping by generating global maps that preserve variability across scales by using modern Bayesian spatial statistical modeling in combination with a database over three times larger than that in previous analyses. Our maps reveal that the most diverse grid cells possess trait variability close to the range of global PFT means.

Functional composition drives ecosystem function through multiple mechanisms in a broadleaved subtropical forest.

Understanding the role of biodiversity (B) in maintaining ecosystem function (EF) is a foundational scientific goal with applications for resource management and conservation. Two main hypotheses have emerged that address B-EF relationships: niche complementarity (NC) and the mass-ratio (MR) effect. We tested the relative importance of these hypotheses in a subtropical old-growth forest on the island nation of Taiwan for two EFs: aboveground biomass (ABG) and coarse woody productivity (CWP). Functional dispersion (FDis) of eight plant functional traits was used to evaluate complementarity of resource use. Under the NC hypothesis, EF will be positively correlated with FDis. Under the MR hypothesis, EF will be negatively correlated with FDis and will be significantly influenced by community-weighted mean (CWM) trait values. We used path analysis to assess how these two processes (NC and MR) directly influence EF and may contribute indirectly to EF via their influence on canopy packing (stem density). Our results indicate that decreasing functional diversity and a significant influence of CWM traits were linked to increasing AGB for all eight traits in this forest supporting the MR hypothesis. Interestingly, CWP was primarily influenced by NC and MR indirectly via their influence on canopy packing. Maximum height explained more of the variation in both AGB and CWP than any of the other plant functional traits. Together, our results suggest that multiple mechanisms operate simultaneously to influence EF, and understanding their relative importance will help to elucidate the role of biodiversity in maintaining ecosystem function.