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 data10-12. 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.

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.

Effectiveness of primate seed dispersers for an "oversized" fruit, Garcinia benthamii.

The largest fruits found in tropical forests may depend on complementary seed dispersal strategies. These fruits are dispersed most effectively by megafauna, but populations can persist where megafauna are absent or erratic visitors. Smaller animals often consume these large fruits, but their capacity to disperse these seeds effectively has rarely been assessed. We evaluated the contributions of gibbons (Hylobates lar) and other frugivores in the seed dispersal of the megafaunal fruit Garcinia benthamii, using the SDE (seed dispersal effectiveness) landscape. Gibbons preferentially consumed G. benthamii fruits and were the main seed disperser that we observed. However, gibbons became satiated when availability was high, with 57% of fruits falling to the ground unhandled. Recruitment of seedlings from gibbon-dispersed seeds was also very low. Elephants consumed G. benthamii fruit, but occurred at low density and were rare visitors to the trees. We suggest that gibbons might complement the seed dispersal role of elephants for G. benthamii, allowing limited recruitment in areas (such as the study site) where elephants occur at low density. Fruit availability varied between years; when availability was low, gibbons reliably consumed most of the crop and dispersed some seeds that established seedlings, albeit at low numbers (2.5 seedlings per crop). When fruit availability was high, the fruit supply overwhelmed the gibbons and other arboreal frugivores, ensuring a large abundance of fruit available to terrestrial seed dispersers. Although gibbons effectively dispersed more seeds at these times (20.7 seedlings per crop), there was the potential for elephants to move many more seeds. Complementary seed dispersal strategies may be important for megafaunal fruit, because they ensure that very large fruits are able to benefit from megafaunal dispersal but also persist where this dispersal becomes erratic. However, our data suggest that smaller seed dispersers might not be capable of replacing large dispersers, leading to potential changes in landscape-scale dispersal patterns where megafauna are absent.

Tropical tree ectomycorrhiza are distributed independently of soil nutrients.

Mycorrhizae, a form of plant-fungal symbioses, mediate vegetation impacts on ecosystem functioning. Climatic effects on decomposition and soil quality are suggested to drive mycorrhizal distributions, with arbuscular mycorrhizal plants prevailing in low-latitude/high-soil-quality areas and ectomycorrhizal (EcM) plants in high-latitude/low-soil-quality areas. However, these generalizations, based on coarse-resolution data, obscure finer-scale variations and result in high uncertainties in the predicted distributions of mycorrhizal types and their drivers. Using data from 31 lowland tropical forests, both at a coarse scale (mean-plot-level data) and fine scale (20 × 20 metres from a subset of 16 sites), we demonstrate that the distribution and abundance of EcM-associated trees are independent of soil quality. Resource exchange differences among mycorrhizal partners, stemming from diverse evolutionary origins of mycorrhizal fungi, may decouple soil fertility from the advantage provided by mycorrhizal associations. Additionally, distinct historical biogeographies and diversification patterns have led to differences in forest composition and nutrient-acquisition strategies across three major tropical regions. Notably, Africa and Asia's lowland tropical forests have abundant EcM trees, whereas they are relatively scarce in lowland neotropical forests. A greater understanding of the functional biology of mycorrhizal symbiosis is required, especially in the lowland tropics, to overcome biases from assuming similarity to temperate and boreal regions.

LiDAR-based reference aboveground biomass maps for tropical forests of South Asia and Central Africa.

Accurate mapping and monitoring of tropical forests aboveground biomass (AGB) is crucial to design effective carbon emission reduction strategies and improving our understanding of Earth's carbon cycle. However, existing large-scale maps of tropical forest AGB generated through combinations of Earth Observation (EO) and forest inventory data show markedly divergent estimates, even after accounting for reported uncertainties. To address this, a network of high-quality reference data is needed to calibrate and validate mapping algorithms. This study aims to generate reference AGB datasets using field inventory plots and airborne LiDAR data for eight sites in Central Africa and five sites in South Asia, two regions largely underrepresented in global reference AGB datasets. The study provides access to these reference AGB maps, including uncertainty maps, at 100 m and 40 m spatial resolutions covering a total LiDAR footprint of 1,11,650 ha [ranging from 150 to 40,000 ha at site level]. These maps serve as calibration/validation datasets to improve the accuracy and reliability of AGB mapping for current and upcoming EO missions (viz., GEDI, BIOMASS, and NISAR).

Water-use characteristics of Syzygium antisepticum and Adinandra integerrima in a secondary forest of Khao Yai National Park in Thailand with implications for environmental management.

Background: Southeast Asia has experienced widespread deforestation and change in land use. Consequently, many reforestation projects have been initiated in this region. However, it is imperative to carefully choose the tree species for planting, especially in light of the increasing climate variability and the potential alteration of plantation on the watershed water balance. Thus, the information regarding water-use characteristics of various tree species and sizes is critical in the tree species selection for reforestation. Methods: We estimated tree water use (T) of dominant species including Syzygium antisepticum and Adinandra integerrima, hereafter Sa and Ai, respectively, in a secondary tropical forest in Khao Yai National Park, Thailand, using sap flow data, and compared T between species and size classes. Additionally, we evaluated the responses of T of both species in each size class to environmental factors including soil moisture and vapor pressure deficit (VPD). Results: Results showed consistently higher T in Sa compared to Ai across ranges of VPD and soil moisture. Under low soil moisture, T of Sa responded to VPD, following a saturating exponential pattern while Ai maintained T across different VPD levels, irrespective of tree size. No responses of T to VPD were observed in either species when soil water was moderate. When soil moisture was high, T of both species significantly increased and saturated at high VPD, albeit the responses were less sensitive in large trees. Our results imply that Ai may be suitable for reforestation in water-limited areas where droughts frequently occur to minimize reforestation impact on water availability to downstream ecosystems. In contrast, Sa should be planted in regions with abundant and reliable water resources. However, a mixed species plantation should be generally considered to increase forest resilience to increasing climate variation.

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.

Variations in leaf water status and drought tolerance of dominant tree species growing in multi-aged tropical forests in Thailand.

Large-scale abandoned agricultural areas in Southeast Asia resulted in patches of forests of multiple successions and characteristics, challenging the study of their responses to environmental changes, especially under climatic water stress. Here, we investigated seasonal variation in leaf water status and drought tolerance of dominant tree species in three multi-aged tropical forests, ranging from 5 to > 200 years old, with contrasting soil moisture in Thailand. Seasonal variation in leaf water status differed among the forests with trees in young and intermediate sites demonstrating larger differences between seasons than the old-growth forest. Although vulnerability to embolism curves revealed that trees in old-growth forest were potentially more sensitive to declining leaf water status than others, they were predicted to lose < 5% of their hydraulic capacity as opposed to 13% for the trees in the younger sites. Our results suggest that the responses to water stress of tree species in different forest ages greatly vary with a tendency of trees in younger sites to be more resilience than those in older sites. Such information would benefit the selection of tree species that could adapt well to specific environments, thus improving the strategies for managing forests of different ages under a warmer future.

Seed Size Variation of Trees and Lianas in a Tropical Forest of Southeast Asia: Allometry, Phylogeny, and Seed Trait - Plant Functional Trait Relationships.

Seed size is a key trait for understanding and predicting ecological processes in a plant community. In a tropical forest, trees and lianas are major components driving ecosystem function and biogeochemical processes. However, seed ecological research on both components remains limited, particularly phylogenetic patterns and relationships with other traits. Here, we compiled a unique dataset of seed size (seed mass and geometrical size metrics) based on collections of more than 5,200 seeds of 196 woody plant species, covering >98 and 70% of tree and liana stems, respectively, located on a 30-ha plot in a tropical evergreen forest in central Thailand. We aimed to (1) develop allometric equations among seed size metrics to predict seed mass; (2) examine phylogenetic influence on seed size variation; and (3) examine relationships among seed traits and several other functional plant traits. Our allometric equations relating seed mass, seed volume, and width were well-fitted with data (R 2 = 0.94, 0.87 respectively). A phylogenetic signal test found that seed size was randomly distributed across the phylogeny. To study the functional trait relationships, we separately tested seed size data of the tree and liana communities (146 and 50 species, respectively), against mean body size of frugivores, successional niches, leaf, and structural traits. For the tree community, seed size was significantly related to mean body size of frugivores, which we believe is a basic driver of seed size because it is related to the gape width affecting dispersal effectiveness. Nearly all leaf traits were significantly positively correlated with seed size (p < 0.03). The significant positive correlation of leaf area and greenness suggested the high-energy demand of large-seeded species. We found a strong positive correlation between seed size and leaf toughness, suggesting a coordination between seed size and leaf defense. However, all these patterns disappeared in the same analysis applied to the liana community. Liana seed size variation was lower than that of trees, perhaps because lianas grow in relatively more uniform conditions in the forest canopy. Frugivore size was the strongest driver of seed size variation. Our study shows a surprising contrast between trees and lianas that is worth further investigation.

Different responses of soil respiration to environmental factors across forest stages in a Southeast Asian forest.

Soil respiration (SR) in forests contributes significant carbon dioxide emissions from terrestrial ecosystems and is highly sensitive to environmental changes, including soil temperature, soil moisture, microbial community, surface litter, and vegetation type. Indeed, a small change in SR may have large impacts on the global carbon balance, further influencing feedbacks to climate change. Thus, detailed characterization of SR responses to changes in environmental conditions is needed to accurately estimate carbon dioxide emissions from forest ecosystems. However, data for such analyses are still limited, especially in tropical forests of Southeast Asia where various stages of forest succession exist due to previous land-use changes. In this study, we measured SR and some environmental factors including soil temperature (ST), soil moisture (SM), and organic matter content (OM) in three successional tropical forests in both wet and dry periods. We also analyzed the relationships between SR and these environmental variables. Results showed that SR was higher in the wet period and in older forests. Although no response of SR to ST was found in younger forest stages, SR of the old-growth forest significantly responded to ST, plausibly due to the nonuniform forest structure, including gaps, that resulted in a wide range of ST. Across forest stages, SM was the limiting factor for SR in the wet period, whereas SR significantly varied with OM in the dry period. Overall, our results indicated that the responses of SR to environmental factors varied temporally and across forest succession. Nevertheless, these findings are still preliminary and call for detailed investigations on SR and its variations with environmental factors in Southeast Asian tropical forests where patches of successional stages dominate.

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.

Different megafauna vary in their seed dispersal effectiveness of the megafaunal fruit Platymitra macrocarpa (Annonaceae).

The world's largest terrestrial animals (megafauna) can play profound roles in seed dispersal. Yet, the term 'megafauna' is often used to encompass a diverse range of body sizes and physiologies of, primarily, herbivorous animals. To determine the extent to which these animals varied in their seed dispersal effectiveness (SDE), we compared the contribution of different megafauna for the large-fruited Platymitra macrocarpa (Annonaceae), in a tropical evergreen forest in Thailand. We quantified 'seed dispersal effectiveness' by measuring the quantity and quality contributions of all consumers of P. macrocarpa fruit. Seed dispersal quantity was the proportion of the crop consumed by each species. Quality was defined as the proportion of seeds handled by each animal taxon that survived to produce a 2-month seedling. Megafauna (elephants, sambar deer, bears) dispersed 78% of seeds that produced seedlings, with 21% dispersed by gibbons (a medium-sized frugivore). The main megafaunal consumers displayed different dispersal strategies. Elephants were the most effective dispersers (37% of seedlings) and they achieved this by being high-quality and low-quantity dispersers. Bears displayed a similar strategy but were especially rare visitors to the trees (24% of the total seedlings produced). Sambar were high-quantity dispersers, but most seeds they handled did not survive and they were responsible for only 17% of seedlings. Gibbons displayed a high SDE relative to their body size, but they probably cannot match the role of elephants despite being more regular consumers of the fruit. The low density and poor regeneration of P. macrocarpa in the study site suggest that current dispersal rates by megafauna are insufficient, possibly reflecting reduced or missing megafauna populations. We show that different megafaunal species disperse seeds in different ways and may make unique contributions to the reproductive success of the plant species.

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.