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

Temporal population variability in local forest communities has mixed effects on tree species richness across a latitudinal gradient

Among the local processes that determine species diversity in ecological communities, fluctuation‐dependent mechanisms that are mediated by temporal variability in the abundances of species populations have received significant attention. Higher temporal variability in the abundances of species populations can increase the strength of temporal niche partitioning but can also increase the risk of species extinctions, such that the net effect on species coexistence is not clear. We quantified this temporal population variability for tree species in 21 large forest plots and found much greater variability for higher latitude plots with fewer tree species. A fitted mechanistic model showed that among the forest plots, the net effect of temporal population variability on tree species coexistence was usually negative, but sometimes positive or negligible. Therefore, our results suggest that temporal variability in the abundances of species populations has no clear negative or positive contribution to the latitudinal gradient in tree species richness

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 CNDD8,9 have suffered from methodological limitations related to the use of static data10,11,12. 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

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

Suwanvecho et al.Gibbon foods_2006-2011

This file contains data on fruit species eaten by the gibbon study group A during April-May follows for the six years 2006-2011. It contains for each visit to a fruit source: the tree species and tree number visited, date, time and fruit species code; and a summary of sources, no. of visits and no. of days for each species. The full species names are given in a separate sheet, copied from the SI on line file associated with Suwanvecho et al. 2017 in Biotropica

Size‐dependent intraspecific variation in wood traits has little impact on aboveground carbon estimates in a tropical forest landscape

International audienceThere is increasing evidence that intraspecific trait variation plays a role in governing rates of ecosystem functioning. While wood traits such as wood specific gravity (WSG) and wood carbon concentration (WCC) are key drivers of forest aboveground carbon (AGC) stocks, the sources of intraspecific variation in these wood traits and the consequences of this variation on AGC are poorly known, especially in the tropics. Here, we investigated intraspecific variation in wood specific gravity (WSG) and wood carbon concentration (WCC) from 556 individual trees belonging to 15 species that well characterize different successional stages of seasonal evergreen forests in Southeast Asia. Specifically, we tested the contribution of individual or species characteristics (tree size, growth rate and regeneration guilds) and local environmental conditions (topographic wetness index and successional stages) to intraspecific variation in WSG and WCC, and assessed the consequences of intraspecific variation in these wood traits on AGC estimates in 14 permanent forest plots established along a successional gradient in Khao Yai National park, Thailand. We found that tree size was the main driver of intraspecific variation in WSG and WCC as tree sizes increased from 10–100 cm in diameter, WSG increased by 7.3%, while WCC increased by 2.4% in heartwood, 1.6% and 2.7% in sapwood without and with volatile carbon included. There was no effect of the topographic wetness and other local environment condition in wood traits led to a slight overestimation of AGC in young secondary forests (+0.09% to +1.29%) and a small underestimation in older forests (−0.86% to −2.87%), but overall AGC estimates (13 of 14 forest plots) remained within error margins (the 95% interval). Our study provides evidence that tree size variation translates into intraspecific variability in wood traits, whereas local environmental conditions related to topography successional stages had no effect on wood trait variability. While size-dependent variation in WSG and WCC have largely been undocumented and thus ignored in forest carbon assessment approaches, we highlight that it has a limited impact on AGC estimates, indicating that it does not invalidate current forest carbon stock estimation approaches. Read the free Plain Language Summary for this article on the Journal blog

Dispersal success of a specialized tropical tree depends on complex interactions among diverse mammalian frugivores

The study of seed dispersal in tropical forest communities is complicated by the high diversity of frugivores and the complex interactions among species and their environments. Determining which species are effective dispersers and which are opportunists with neutral or even negative effects on fruiting plants is a major problem which requires detailed studies focused on particular plant species. In this study we focused on seed dispersal of the wild rambutan (Nephelium melliferum) which supplies energy-rich fruits to primates and other mammals in a seasonal evergreen forest in central Thailand. We hypothesized that gibbons (Hylobates lar) were the most important dispersers and were capable of carrying and defecating seeds away from the tree crown where seeds or seedlings could escape increased distance or density-dependent mortality. We determined the seed dispersal effectiveness of all major arboreal consumers of Nephelium fruit (gibbons, pig-tail monkeys, squirrels) by using data on fruit production and consumption from fruit/seed traps under the canopies of eight sample trees, observations on animals feeding in fruiting trees, and seed deposition data at varying distances from tree crowns. We assessed the survival of seeds and seedlings in relation to distance from the crown by experiments and transect counts. Camera traps were used to detect consumption of seeds on the ground by terrestrial consumers. About half of the ripe fruit crop was harvested by squirrels which gnawed off the outer husk, ate the pulp and dropped intact seeds to the ground. Monkeys chewed and sucked off the pulp and dropped most seeds to the ground not far from the tree. Seeds and intact fruits dropped by squirrels and monkeys supported a large variety of terrestrial mammals most of which were seed predators. Gibbons, which swallow the seed with pulp attached, consumed only 16 % of the fruit crop but were the most effective and reliable seed dispersers overall, and appear to be the most specialized seed dispersal mutualist. Distance-dependent mortality was found in first-year seedlings, as well as evidence that further mortality must space out older saplings prior to recruitment into the tree population, which validates the importance of dispersal by gibbons. However, the great majority of fruits of Nephelium melliferum were consumed by mammalian opportunists and seed predators which interacted with one another in ways which depend on composition of the local community

Defaunation of large-bodied frugivores reduces carbon storage in a tropical forest of Southeast Asia

Recent studies have suggested that defaunation of large-bodied frugivores reduces above-ground carbon storage in tropical forests of South America and Africa, but not, or less so, in Southeast Asian tropical forests. Here we analyze the issue using the seed dispersal network (data of interaction between trees and animal seed dispersers) and forest composition of a 30-ha forest dynamics plot in central Thailand, where an intact fauna of primates, ungulates, bears and birds of all sizes still exists. We simulate the effect of two defaunation scenarios on forest biomass: 1) only primates extirpated (a realistic possibility in near future), and 2) extirpation of all large-bodied frugivores (LBF) including gibbons, macaques, hornbills and terrestrial mammals, the main targets of poachers in this region. For each scenario, we varied the population size reduction of the LBF dispersed tree species from 20% to 100%. We find that tree species dependent on seed dispersal by large-bodied frugivores (LBF) account for nearly one-third of the total carbon biomass on the plot, and that the community turnover following a complete defaunation would result in a carbon reduction of 2.4% to 3.0%, depending on the defaunation scenario and the model assumptions. The reduction was always greater than 1% when the defaunation intensity was at least 40%. These effect sizes are comparable to values reported for Neotropical forests, suggesting that the impact of defaunation on carbon deficit is not necessarily lower in Southeast Asian forests. The problem of defaunation in Asia, and the mutual benefits between biodiversity conservation and climate change mitigation, should therefore not be neglected by global policies to reduce carbon emissions

Data from: High interannual variation in the diet of a tropical forest frugivore (Hylobates lar)

Frugivores must deal with seasonal changes in fruit availability and changes from year to year, as most species of tropical forest fruiting trees have considerable interannual variation in phenology and many are mast fruiters. We quantified seasonal and interannual changes in the fruit diet in a frugivore and important seed disperser, the white-handed gibbon, Hylobates lar, in Thailand. We used 40-day following data during April and May replicated in six consecutive years to study interannual variability in the diet, and compared it with seasonal changes measured in monthly samples of the same size collected in three successive years. The 40-day periods of following also allowed us to measure the decline in dietary similarity with time over a finer scale. We measured fruit diet similarity between replicated 5-day periods using the percentage overlap (Renkonen’s) index, and Jaccard’s similarity index. Seasonally, average dietary overlap between adjacent months was low, and similarity approached zero after four months. Average rate of decline in similarity exceeded 20 percent per five day period. Variation in fruit species in the diet between years was high, and was correlated with interannual variation in fruiting phenology. The strongest correlation occurred in the case of Nephelium melliferum, a highly preferred species that dominated the diet in good fruiting years. It is difficult to separate changes in food-species preference from changes in availability from year to year. We devised a relative measure of preference that depends on the degree to which the gibbons rely on prior knowledge to find sources

Whole-Plant Seedling Functional Traits Suggest Lianas Also Support “Fast-Slow” Plant Economics Spectrum

Lianas are predicted to perform better than trees during seasonal drought among tropical forests, which has substantial implications for tree and forest dynamics. Here, we use whole-plant trait comparison to test whether lianas allocated on the resource acquisitive end of the continuum of woody plant strategies. We measured morphological and biomass allocation traits for seedlings of 153 species of trees and lianas occurring in a tropical forest in Thailand during the dry season. We first compared trait differences between lianas and trees directly, and then classified all species based on their trait similarities. We found that liana seedlings had significantly higher specific leaf areas and specific stem lengths than co-occurring tree seedlings. Trait similarity classification resulted in a liana-dominated cluster and a tree-dominated cluster. Compared to the tree-dominated cluster, species in the liana-dominated cluster were characterized by a consistent pattern of lower dry matter content and cheaper and more efficient per dry mass unit investment in both above-and below-ground organs. The consistency of all organs operating in tandem for dry matter content, together with optimized investment in them per mass unit, implied that the lianas and trees can be highly overlapped on the strategy gradient of the resource acquisition continuum