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Title: Latitudinal patterns in stabilizing density dependence of forest communities

Numerous studies have shown reduced performance in plants that are surrounded by neighbours of the same species1,2, a phenomenon known as conspecific negative density dependence (CNDD)3. A long-held ecological hypothesis posits that CNDD is more pronounced in tropical than in temperate forests4,5, which increases community stabilization, species coexistence and the diversity of local tree species6,7. Previous analyses supporting such a latitudinal gradient in CNDD8,9have suffered from methodological limitations related to the use of static data10–12. Here we present a comprehensive assessment of latitudinal CNDD patterns using dynamic mortality data to estimate species-site-specific CNDD across 23 sites. Averaged across species, we found that stabilizing CNDD was present at all except one site, but that average stabilizing CNDD was not stronger toward the tropics. However, in tropical tree communities, rare and intermediate abundant species experienced stronger stabilizing CNDD than did common species. This pattern was absent in temperate forests, which suggests that CNDD influences species abundances more strongly in tropical forests than it does in temperate ones13. We also found that interspecific variation in CNDD, which might attenuate its stabilizing effect on species diversity14,15, was high but not significantly different across latitudes. Although the consequences of these patterns for latitudinal diversity gradients are difficult to evaluate, we speculate that a more effective regulation of population abundances could translate into greater stabilization of tropical tree communities and thus contribute to the high local diversity of tropical forests.

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564 to 571
Medium: X
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National Science Foundation
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  1. Abstract

    The relationship between biodiversity and stability, or its inverse, temporal variability, is multidimensional and complex. Temporal variability in aggregate properties, like total biomass or abundance, is typically lower in communities with higher species diversity (i.e., the diversity–stability relationship [DSR]). At broader spatial extents, regional‐scale aggregate variability is also lower with higher regional diversity (in plant systems) and with lower spatial synchrony. However, focusing exclusively on aggregate properties of communities may overlook potentially destabilizing compositional shifts. It is not yet clear how diversity is related to different components of variability across spatial scales, nor whether regional DSRs emerge across a broad range of organisms and ecosystem types. To test these questions, we compiled a large collection of long‐term metacommunity data spanning a wide range of taxonomic groups (e.g., birds, fish, plants, invertebrates) and ecosystem types (e.g., deserts, forests, oceans). We applied a newly developed quantitative framework for jointly analyzing aggregate and compositional variability across scales. We quantified DSRs for composition and aggregate variability in local communities and metacommunities. At the local scale, more diverse communities were less variable, but this effect was stronger for aggregate than compositional properties. We found no stabilizing effect of γ‐diversity on metacommunity variability, but β‐diversity played a strong role in reducing compositional spatial synchrony, which reduced regional variability. Spatial synchrony differed among taxa, suggesting differences in stabilization by spatial processes. However, metacommunity variability was more strongly driven by local variability than by spatial synchrony. Across a broader range of taxa, our results suggest that high γ‐diversity does not consistently stabilize aggregate properties at regional scales without sufficient spatial β‐diversity to reduce spatial synchrony.

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  2. Yavitt, Joseph B. (Ed.)
    Conspecific negative density dependence (CNDD) promotes tree species diversity by reducing recruitment near conspecific adults due to biotic feedbacks from herbivores, pathogens, or competitors. While this process is well-described in tropical forests, tests of temperate tree species range from strong positive to strong negative density dependence. To explain this, several studies have suggested that tree species traits may help predict the strength and direction of density dependence: for example, ectomycorrhizal-associated tree species typically exhibit either positive or weaker negative conspecific density dependence. More generally, the strength of density dependence may be predictably related to other species-specific ecological attributes such as shade tolerance, or the relative local abundance of a species. To test the strength of density dependence and whether it affects seedling community diversity in a temperate forest, we tracked the survival of seedlings of three ectomycorrhizal-associated species experimentally planted beneath conspecific and heterospecific adults on the Prospect Hill tract of the Harvard Forest, in Massachusetts, USA. Experimental seedling survival was always lower under conspecific adults, which increased seedling community diversity in one of six treatments. We compared these results to evidence of CNDD from observed sapling survival patterns of 28 species over approximately 8 years in an adjacent 35-ha forest plot. We tested whether species-specific estimates of CNDD were associated with mycorrhizal association, shade tolerance, and local abundance. We found evidence of significant, negative conspecific density dependence (CNDD) in 23 of 28 species, and positive conspecific density dependence in two species. Contrary to our expectations, ectomycorrhizal-associated species generally exhibited stronger (e.g., more negative) CNDD than arbuscular mycorrhizal-associated species. CNDD was also stronger in more shade-tolerant species but was not associated with local abundance. Conspecific adult trees often have a negative influence on seedling survival in temperate forests, particularly for tree species with certain traits. Here we found strong experimental and observational evidence that ectomycorrhizal-associating species consistently exhibit CNDD. Moreover, similarities in the relative strength of density dependence from experiments and observations of sapling mortality suggest a mechanistic link between negative effects of conspecific adults on seedling and sapling survival and local tree species distributions. 
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  3. Abstract Aim

    We examined tree beta diversity in four biogeographical regions with contrasting environmental conditions, latitude, and diversity. We tested: (a) the influence of the species pool on beta diversity; (b) the relative contribution of niche‐based and dispersal‐based assembly to beta diversity; and (c) differences in the importance of these two assembly mechanisms in regions with differing productivity and species richness.


    Lowland and montane tropical forests in the Madidi region (Bolivia), lowland temperate forests in the Ozarks (USA), and montane temperate forests in the Cantabrian Mountains (Spain).


    We surveyed woody plants with a diameter ≥2.5 cm following a standardized protocol in 236 0.1‐ha forest plots in four different biogeographical regions. We estimated the species pool at each region and used it to recreate null communities determined entirely by the species pool. Observed patterns of beta diversity smaller or greater than the null‐expected patterns of beta diversity implies the presence of local assembly mechanisms beyond the influence of the species pool. We used variation‐partitioning analyses to compare the contribution of niche‐based and dispersal‐based assembly to patterns of observed beta diversity and their deviations from null models among the four regions.


    (a) Differences in species pools alone did not explain observed differences in beta diversity among biogeographic regions. (b) In 3/4 regions, the environment explained more of the variation in beta diversity than spatial variables. (c) Spatial variables explained more of the beta diversity in more diverse and more productive regions with more rare species (tropical and lower‐elevation regions) compared to less diverse and less productive regions (temperate and higher‐elevation regions). (d) Greater alpha or gamma diversity did not result in higher beta diversity or stronger correlations with the environment.


    Overall, the observed differences in beta diversity are better explained by differences in community assembly mechanism than by biogeographical processes that shaped the species pool.

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  4. Abstract Aim

    Soil microorganisms are essential for the functioning of terrestrial ecosystems. Although soil microbial communities and functions are linked to tree species composition and diversity, there has been no comprehensive study of the generality or context dependence of these relationships. Here, we examine tree diversity–soil microbial biomass and respiration relationships across environmental gradients using a global network of tree diversity experiments.


    Boreal, temperate, subtropical and tropical forests.

    Time period


    Major taxa studied

    Soil microorganisms.


    Soil samples collected from 11 tree diversity experiments were used to measure microbial respiration, biomass and respiratory quotient using the substrate‐induced respiration method. All samples were measured using the same analytical device, method and procedure to reduce measurement bias. We used linear mixed‐effects models and principal components analysis (PCA) to examine the effects of tree diversity (taxonomic and phylogenetic), environmental conditions and interactions on soil microbial properties.


    Abiotic drivers, mainly soil water content, but also soil carbon and soil pH, significantly increased soil microbial biomass and respiration. High soil water content reduced the importance of other abiotic drivers. Tree diversity had no effect on the soil microbial properties, but interactions with phylogenetic diversity indicated that the effects of diversity were context dependent and stronger in drier soils. Similar results were found for soil carbon and soil pH.

    Main conclusions

    Our results indicate the importance of abiotic variables, especially soil water content, for maintaining high levels of soil microbial functions and modulating the effects of other environmental drivers. Planting tree species with diverse water‐use strategies and structurally complex canopies and high leaf area might be crucial for maintaining high soil microbial biomass and respiration. Given that greater phylogenetic distance alleviated unfavourable soil water conditions, reforestation efforts that account for traits improving soil water content or select more phylogenetically distant species might assist in increasing soil microbial functions.

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  5. Abstract Aim

    Trees associating with ectomycorrhizal (ECM) fungi typically occur in infertile soils and use nutrients more conservatively than arbuscular mycorrhizal (AM) trees. We hypothesized that ECM trees would have greater nutrient resorption (i.e., proportion of nutrients resorbed during leaf senescence) than AM trees.




    We synthesized nitrogen (N) and phosphorus (P) resorption data from 378 species from sub/tropical, temperate and boreal forests, including 43 studies where ECM and AM trees co‐occurred, and conducted a meta‐analysis. Additionally, we quantified N resorption in 45 plots varying in ECM‐AM tree abundances in the temperate deciduous forests of southern Indiana, USA.


    Overall, resorption patterns were driven primarily by mycorrhizal type, climate zone, and to a lesser degree, leaf habit. In the boreal forest, P resorption was 76% greater for ECM than AM trees (p < .05). In the sub/tropics, AM trees resorbed 30% more N than ECM trees. At the sites where AM and ECM trees co‐occurred, ECM trees resorbed more N in temperate forests (15% greater;p < .001) whereas AM trees tended to resorb more N in sub/tropical forests (by 29%;p = .08). Besides, deciduous ECM trees resorbed more N (10%) and P (15%) than deciduous AM trees, while evergreen ECM and AM trees did not differ. In the deciduous forests of Indiana, where ECM and AM trees co‐occurred, the relative abundance of ECM trees in a plot was positively correlated to plot‐scale N resorption (R2 = .25,p = .001), indicating greater nutrient conservatism with increasing ECM‐dominance.

    Main conclusions

    Our results indicate that mycorrhizal association – in addition to other factors – is correlated with the degree to which trees recycle nutrients, with the strongest effects occurring for N resorption by temperate deciduous trees.

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