Mycorrhizal feedbacks influence global forest structure and diversity.

One mechanism proposed to explain high species diversity in tropical systems is strong negative conspecific density dependence (CDD), which reduces recruitment of juveniles in proximity to conspecific adult plants. Although evidence shows that plant-specific soil pathogens can drive negative CDD, trees also form key mutualisms with mycorrhizal fungi, which may counteract these effects. Across 43 large-scale forest plots worldwide, we tested whether ectomycorrhizal tree species exhibit weaker negative CDD than arbuscular mycorrhizal tree species. We further tested for conmycorrhizal density dependence (CMDD) to test for benefit from shared mutualists. We found that the strength of CDD varies systematically with mycorrhizal type, with ectomycorrhizal tree species exhibiting higher sapling densities with increasing adult densities than arbuscular mycorrhizal tree species. Moreover, we found evidence of positive CMDD for tree species of both mycorrhizal types. Collectively, these findings indicate that mycorrhizal interactions likely play a foundational role in global forest diversity patterns and structure.

Climate sensitive size-dependent survival in tropical trees.

Survival rates of large trees determine forest biomass dynamics. Survival rates of small trees have been linked to mechanisms that maintain biodiversity across tropical forests. How species survival rates change with size offers insight into the links between biodiversity and ecosystem function across tropical forests. We tested patterns of size-dependent tree survival across the tropics using data from 1,781 species and over 2 million individuals to assess whether tropical forests can be characterized by size-dependent life-history survival strategies. We found that species were classifiable into four 'survival modes' that explain life-history variation that shapes carbon cycling and the relative abundance within forests. Frequently collected functional traits, such as wood density, leaf mass per area and seed mass, were not generally predictive of the survival modes of species. Mean annual temperature and cumulative water deficit predicted the proportion of biomass of survival modes, indicating important links between evolutionary strategies, climate and carbon cycling. The application of survival modes in demographic simulations predicted biomass change across forest sites. Our results reveal globally identifiable size-dependent survival strategies that differ across diverse systems in a consistent way. The abundance of survival modes and interaction with climate ultimately determine forest structure, carbon storage in biomass and future forest trajectories.

Response to Comment on "Plant diversity increases with the strength of negative density dependence at the global scale".

Hülsmann and Hartig suggest that ecological mechanisms other than specialized natural enemies or intraspecific competition contribute to our estimates of conspecific negative density dependence (CNDD). To address their concern, we show that our results are not the result of a methodological artifact and present a null-model analysis that demonstrates that our original findings-(i) stronger CNDD at tropical relative to temperate latitudes and (ii) a latitudinal shift in the relationship between CNDD and species abundance-persist even after controlling for other processes that might influence spatial relationships between adults and recruits.

Response to Comment on "Plant diversity increases with the strength of negative density dependence at the global scale".

Chisholm and Fung claim that our method of estimating conspecific negative density dependence (CNDD) in recruitment is systematically biased, and present an alternative method that shows no latitudinal pattern in CNDD. We demonstrate that their approach produces strongly biased estimates of CNDD, explaining why they do not detect a latitudinal pattern. We also address their methodological concerns using an alternative distance-weighted approach, which supports our original findings of a latitudinal gradient in CNDD and a latitudinal shift in the relationship between CNDD and species abundance.

Plant diversity increases with the strength of negative density dependence at the global scale.

Theory predicts that higher biodiversity in the tropics is maintained by specialized interactions among plants and their natural enemies that result in conspecific negative density dependence (CNDD). By using more than 3000 species and nearly 2.4 million trees across 24 forest plots worldwide, we show that global patterns in tree species diversity reflect not only stronger CNDD at tropical versus temperate latitudes but also a latitudinal shift in the relationship between CNDD and species abundance. CNDD was stronger for rare species at tropical versus temperate latitudes, potentially causing the persistence of greater numbers of rare species in the tropics. Our study reveals fundamental differences in the nature of local-scale biotic interactions that contribute to the maintenance of species diversity across temperate and tropical communities.

Linking leaf veins to growth and mortality rates: an example from a subtropical tree community.

A fundamental goal in ecology is to link variation in species function to performance, but functional trait-performance investigations have had mixed success. This indicates that less commonly measured functional traits may more clearly elucidate trait-performance relationships. Despite the potential importance of leaf vein traits, which are expected to be related to resource delivery rates and photosynthetic capacity, there are few studies, which examine associations between these traits and demographic performance in communities. Here, we examined the associations between species traits including leaf venation traits and demographic rates (Relative Growth Rate, RGR and mortality) as well as the spatial distributions of traits along soil environment for 54 co-occurring species in a subtropical forest. Size-related changes in demographic rates were estimated using a hierarchical Bayesian approach. Next, Kendall's rank correlations were quantified between traits and estimated demographic rates at a given size and between traits and species-average soil environment. Species with denser venation, smaller areoles, less succulent, or thinner leaves showed higher RGR for a wide range of size classes. Species with leaves of denser veins, larger area, cheaper construction costs or thinner, or low-density wood were associated with high mortality rates only in small size classes. Lastly, contrary to our expectations, acquisitive traits were not related to resource-rich edaphic conditions. This study shows that leaf vein traits are weakly, but significantly related to tree demographic performance together with other species traits. Because leaf traits associated with an acquisitive strategy such as denser venation, less succulence, and thinner leaves showed higher growth rate, but similar leaf traits were not associated with mortality, different pathways may shape species growth and survival. This study suggests that we are still not measuring some of key traits related to resource-use strategies, which dictate the demography and distributions of species.

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.

Ecological facilitation between two epiphytes through drought mitigation in a subtropical rainforest.

Positive species interactions (facilitation) play an important role in shaping the structures and species diversity of ecological communities, particularly under stressful environmental conditions. Epiphytes in rainforests often grow in multiple-species clumps, suggesting interspecies facilitation. However, little is known about the patterns and mechanisms of epiphyte co-occurrence. We assessed the interactions of two widespread epiphyte species, Asplenium antiquum and Haplopteris zosterifolia, by examining their co-occurrence and size-class association in the field. To elucidate factors controlling their interactions, we conducted reciprocal-removal and greenhouse-drought experiments, and nutrient and isotope analyses. Forty-five percent of H. zosterifolia co-occurred with A. antiquum, whereas only 17% of A. antiquum co-occurred with H. zosterifolia. Removing the fronds plus substrate of A. antiquum reduced the relative frond length and specific leaf area of H. zosterifolia, but removing fronds only had little effect. Removing H. zosterifolia had no significant effects on the growth of A. antiquum. H. zosterifolia co-occurring and not co-occurring with A. antiquum had similar foliar nutrient concentrations and δ(15)N values, suggesting that A. antiquum does not affect the nutrient status of H. zosterifolia. Reduced growth of H. zosterifolia with the removal of A. antiquum substrate, together with higher foliar δ(13)C for H. zosterifolia growing alone than those co-occurring with A. antiquum, suggest that A. antiquum enhances water availability to H. zosterifolia. This enhancement probably resulted from water storage in the substrate of A. antiquum, which could hold water up to 6.2 times its dry weight, and from reduced evapotranspiration due to shading of A. antiquum fronds. Greater water loss occurred in the frond-clipped group than the unclipped group between days 3-13 of the drought treatment. Our results imply that drought mitigation by substrate-forming epiphytes is important for maintaining epiphyte diversity in tropic and subtropic regions with episodic water limitations, especially in the context of anthropogenic climate change.

Effects of climate change and shifts in forest composition on forest net primary production.

Forests are dynamic in both structure and species composition, and these dynamics are strongly influenced by climate. However, the net effects of future tree species composition on net primary production (NPP) are not well understood. The objective of this work was to model the potential range shifts of tree species (DISTRIB Model) and predict their impacts on NPP (PnET-II Model) that will be associated with alterations in species composition. We selected four 200 x 200 km areas in Wisconsin, Maine, Arkansas, and the Ohio-West Virginia area, representing focal areas of potential species range shifts. PnET-II model simulations were carried out assuming that all forests achieved steady state, of which the species compositions were predicted by DISTRIB model with no migration limitation. The total NPP under the current climate ranged from 552 to 908 g C/m(2) per year. The effects of potential species redistributions on NPP were moderate (-12% to +8%) compared with the influence of future climatic changes (-60% to +25%). The direction and magnitude of climate change effects on NPP were largely dependent on the degree of warming and water balance. Thus, the magnitude of future climate change can affect the feedback system between the atmosphere and biosphere.