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Based on the framework of attribute diversity (a generalization of Hill numbers of order q), we develop a class of functional diversity measures sensitive not only to species abundances but also to trait-based species-pairwise functional distances. The new method refines and improves on the conventional species-equivalent approach in three areas: (1) the conventional method often gives similar values (close to unity) to assemblages with contrasting levels of functional diversity; (2) when a distance metric is unbounded, the conventional functional diversity depends on the presence/absence of other assemblages in the study; (3) in partitioning functional gamma diversity into alpha and beta components, the conventional gamma is sometimes less than alpha. To resolve these issues, we add to the attribute-diversity framework a novel concept: τ, the threshold of functional distinctiveness between any two species; here, τ can be chosen to be any positive value. Any two species with functional distance ≥ τ are treated as functionally equally distinct. Our functional diversity quantifies the effective number of functionally equally distinct species (or \"virtual functional groups\") with all pairwise distances at least s for different species pairs. We advocate the use of two complementary diversity profiles (τ profile and q profile), which depict functional diversity with varying levels of τ and q, respectively. Both the conventional species-equivalent method (i.e., τ is the maximum of species-pairwise distances) and classic taxonomic diversity (i.e., τ is the minimum of non-zero species-pairwise distances) are incorporated into our proposed τ profile for an assemblage. For any type of species-pairwise distance matrices, our attribute-diversity approach allows proper diversity partitioning, with the desired property gamma ≥ alpha and thus avoids all the restrictions that apply to the conventional diversity decomposition. Our functional alpha and gamma are interpreted as the effective numbers of functionally equally distinct species, respectively, in an assemblage and in the pooled assemblage, while beta is the effective number of equally large assemblages with no shared species and all species in the assemblages being equally distinct. The resulting beta diversity can be transformed to obtain abundance-sensitive Sørensen-and Jaccard-type functional (dis)similarity profiles. Hypothetical and real examples are used to illustrate the framework. Online software and R codes are available to facilitate computations.

The classical environmental control model assumes that species distribution is determined by the spatial variation of underlying habitat conditions. This niche-based model has recently been challenged by the neutral theory of biodiversity which assumes that ecological drift is a key process regulating species coexistence. Understanding the mechanisms that maintain biodiversity in communities critically depends on our ability to decompose the variation of diversity into the contributions of different processes affecting it. Here we investigated the effects of pure habitat, pure spatial, and spatially structured habitat processes on the distributions of species richness and species composition in a recently established 24-ha stem-mapping plot in the subtropical evergreen broad-leaved forest of Gutianshan National Nature Reserve in East China. We used the new spatial analysis method of principal coordinates of neighbor matrices (PCNM) to disentangle the contributions of these processes. The results showed that (1) habitat and space jointly explained ~53% of the variation in richness and ~65% of the variation in species composition, depending on the scale (sampling unit size); (2) tree diversity (richness and composition) in the Gutianshan forest was dominantly controlled by spatially structured habitat (24%) and habitat-independent spatial component (29%); the spatially independent habitat contributed a negligible effect (6%); (3) distributions of richness and species composition were strongly affected by altitude and terrain convexity, while the effects of slope and aspect were weak; (4) the spatial distribution of diversity in the forest was dominated by broad-scaled spatial variation; (5) environmental control on the one hand and unexplained spatial variation on the other (unmeasured environmental variables and neutral processes) corresponded to spatial structures with different scales in the Gutianshan forest plot; and (6) five habitat types were recognized; a few species were statistically significant indicators of three of these habitats, whereas two habitats had no significant indicator species. The results suggest that the diversity of the forest is equally governed by environmental control (30%) and neutral processes (29%). In the fine-scale analysis (10 × 10 m cells), neutral processes dominated (43%) over environmental control (20%).

Native species that forage in farmland may increase their local abundances thereby affecting adjacent ecosystems within their landscape. We used two decades of ecological data from a protected primary rainforest in Malaysia to illutrate how subsidies from neighboring oil palm plantations triggered powerful secondary ‘cascading’ effects on natural habitats located >1.3 km away. We found (i) oil palm fruit drove 100-fold increases in crop-raiding native wild boar ( Sus scrofa ), (ii) wild boar used thousands of understory plants to construct birthing nests in the pristine forest interior, and (iii) nest building caused a 62% decline in forest tree sapling density over the 24-year study period. The long-term, landscape-scale indirect effects from agriculture suggest its full ecological footprint may be larger in extent than is currently recognized. Cross-boundary subsidy cascades may be widespread in both terrestrial and marine ecosystems and present significant conservation challenges. The expansion of agriculture and rangelands can cause ecological spillover effects across cultivated-natural ecosystem boundaries. Here, Luskin et al. show irruptions of oil palm-subsidized wild boar alter the abundance and diversity of understory trees >1 km into a primary forest reserve in Malaysia.

• The relative importance of tree mortality risk factors remains unknown, especially in diverse tropical forests where species may vary widely in their responses to particular conditions. • We present a new framework for quantifying the importance of mortality risk factors and apply it to compare 19 risks on 31 203 trees (1977 species) in 14 one-year periods in six tropical forests. We defined a condition as a risk factor for a species if it was associated with at least a doubling of mortality rate in univariate analyses. For each risk, we estimated prevalence (frequency), lethality (difference in mortality between trees with and without the risk) and impact (‘excess mortality’ associated with the risk, relative to stand-level mortality). • The most impactful risk factors were light limitation and crown/trunk loss; the most prevalent were light limitation and small size; the most lethal were leaf damage and wounds. Modes of death (standing, broken and uprooted) had limited links with previous conditions and mortality risk factors. • We provide the first ranking of importance of tree-level mortality risk factors in tropical forests. Future research should focus on the links between these risks, their climatic drivers and the physiological processes to enable mechanistic predictions of future tree mortality.

1. In a unique phenomenon restricted to the ever wet forests of Southeast Asia, hundreds of species from dozens of plant families reproduce synchronously at irregular, multi-year intervals. The proximate environmental cues that synchronize these general flowering events have not been evaluated systematically because there have been no long-term, high temporal resolution, species-level records from the region. 2. We present 13 years of weekly flowering records for five Shorea species as well as daily temperature and rainfall records from the Pasoh Forest Reserve, Peninsular Malaysia. We constructed models to evaluate hypothesized relationships between flowering and cool temperature, drought, and additive and synergistic effects of cool temperature and drought for each species. Model parameters include periods of time for floral cue accumulation and flower development and temperature and/or rainfall thresholds required for floral initiation. Parameters estimated using flowering observations from 2001 to 2011 were used to forecast flowering for 2011-2014. 3. We show that drought and cool temperatures acting synergistically best explain the timing of flowering events for all Shorea species in the section Mutica and forecast the largest general flowering event accurately. Periods estimated for signal accumulation ranged from 54 to 90 days among species. Periods estimated for flowers to develop ranged between 43 and 96 days and closely followed the interspecific sequence of flowering in the Shorea species. Drought and temperature thresholds also varied among species, with Shorea maxwelliana requiring the most severe drought and Shorea leprosula the lowest temperatures. 4. Synthesis. Our results indicate that cool temperatures and low rainfall occurring on seasonal time-scales of about 2-3 months rather than brief cold snaps or brief droughts best explain general flowering in Shorea species at the Pasoh Forest Reserve. Low rainfall is equally likely in winter (December-February) and summer (July–August) and cool temperatures are most likely in winter at Pasoh, which explains why general flowering events are restricted to spring and fall, with more frequent and stronger flowering in spring. In addition, species-specific sensitivity to environmental cues suggests that future climate change will have differential impacts on the frequency of reproduction, with potential consequences for regeneration of these dominant species of lowland tropical forests.

An important goal in plant community ecology is to understand how species traits determine demographic performance. Several functional traits have been shown to correlate with growth and mortality rates in trees, but less is known about how the relationships between functional traits and demographic rates change with tree size. We examined the associations of functional traits with growth and mortality across 43 tree species in the Fushan 25‐ha subtropical rain forest plot in northern Taiwan. We estimated the 95th percentile maximum stem diameter, wood density and six leaf functional traits (leaf area, specific leaf area, thickness, succulence, and mass‐based nitrogen and phosphorus contents) obtained from leaves on juvenile and adult individuals of each species. To quantify size‐dependent changes in growth and mortality, relative growth rate (RGR) and mortality were estimated as a function of stem diameter using hierarchical Bayesian models. These rate estimates were then correlated with functional traits at a range of stem diameter classes. Relationships between functional traits and demographic rates varied with tree size. Maximum size was positively correlated with RGR across a wide range of tree sizes. Wood density was negatively correlated with RGR and mortality for small‐sized trees. Leaf traits such as leaf area and specific leaf area at juvenile and adult stages were associated more strongly with demographic rates for corresponding sizes than from other sizes. Synthesis. The observed size‐dependent changes in the trait–demography relationships are possibly due to the effects of developmental and environmental changes with increasing tree size. The underlying effects of functional traits on demographic performance vary with tree size, and this should influence dynamics in a tree community.

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 oldgrowth 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.

Despite their fundamental importance the links between forest productivity, diversity and climate remain contentious. We consider whether variation in productivity across climates reflects adjustment among tree species and individuals, or changes in tree community structure. We analysed data from 60 plots of humid old-growth forests spanning mean annual temperatures (MAT) from 2.0 to 26.6 °C. Comparing forests at equivalent aboveground biomass (160 Mg C ha –1 ), tropical forests ≥24 °C MAT averaged more than double the aboveground woody productivity of forests <12 °C (3.7 ± 0.3 versus 1.6 ± 0.1 Mg C ha –1 yr –1 ). Nonetheless, species with similar standing biomass and maximum stature had similar productivity across plots regardless of temperature. We find that differences in the relative contribution of smaller- and larger-biomass species explained 86% of the observed productivity differences. Species-rich tropical forests are more productive than other forests due to the high relative productivity of many short-stature, small-biomass species. The link between forest productivity, species diversity and climate remains contentious. Here, Kohyama et al. examine stand productivity and tree diversity in old-growth forests from Japan to Indonesia, showing that warmer sites are more productive, largely due to small-biomass species.

Understanding processes underlying spatial distribution of tree species is fundamental to studying species coexistence and diversity. This study modeled point patterns of tree distribution, expressed by Cartesian coordinates of individual trees within a mapped forest stand, for the purpose of identifying processes that may generate spatial patterns of tree communities. We used four primary point pattern processes (homogeneous Poisson process, inhomogeneous Poisson process, homogeneous Thomas process, and inhomogeneous Thomas process) to model tree distribution in two stem-mapped forests in Taiwan, Republic of China. These four models simulate spatial processes of habitat association and seed dispersal, allowing us to evaluate the potential contribution of habitat heterogeneity and dispersal limitation to the formation of spatial patterns of tree species. The results showed that the inhomogeneous Thomas process was the best fit model and described most of the species studied, suggesting that spatial patterns of tree species might be formed by the joint effects of habitat associations and dispersal limitation. The homogeneous Thomas process that models the effect of dispersal limitation was the second best model. We also found that the best fit models could be predicted by species attributes, including species abundance and dispersal mode. The significant traits, however, differed between the two study plots and demonstrated site-specific patterns. This study indicated that the interactive operation of niche-based (habitat heterogeneity) and neutral-based (dispersal limitation) may be important in generating spatial patterns of tree species in forest communities.

High tree species diversity in tropical forests is driven by reduced interspecific competition relative to intraspecific competition, as a result of the asynchronous timing of tree recruitment permitted by long and stable growing seasons. Forest diversity with latitude The number of tree species that coexist on a hectare scale rises by more than two orders of magnitude from boreal to tropical forests. Jacob Usinowicz and colleagues assess the extent to which latitudinal differences in competition between species shape this gradient, using seed production and seedling recruitment data from ten forests spanning the tropics to the boreal zone. They show that the longer and more stable growing seasons in the tropics permit greater differentiation between species in terms of the seasonal timing of reproduction. This asynchrony in the timing of tree recruitment reduces the degree of competition between species in the tropics, relative to forests at higher latitudes, thereby increasing the potential for species coexistence. The findings highlight the contribution that local-scale ecological processes can make to global-scale gradients in biodiversity. The tropical forests of Borneo and Amazonia may each contain more tree species diversity in half a square kilometre than do all the temperate forests of Europe, North America, and Asia combined 1 . Biologists have long been fascinated by this disparity, using it to investigate potential drivers of biodiversity 2 . Latitudinal variation in many of these drivers is expected to create geographic differences in ecological 2 , 3 , 4 and evolutionary processes 4 , 5 , and evidence increasingly shows that tropical ecosystems have higher rates of diversification, clade origination, and clade dispersal 5 , 6 . However, there is currently no evidence to link gradients in ecological processes within communities at a local scale directly to the geographic gradient in biodiversity. Here, we show geographic variation in the storage effect, an ecological mechanism that reduces the potential for competitive exclusion more strongly in the tropics than it does in temperate and boreal zones, decreasing the ratio of interspecific-to-intraspecific competition by 0.25% for each degree of latitude that an ecosystem is located closer to the Equator. Additionally, we find evidence that latitudinal variation in climate underpins these differences; longer growing seasons in the tropics reduce constraints on the seasonal timing of reproduction, permitting lower recruitment synchrony between species and thereby enhancing niche partitioning through the storage effect. Our results demonstrate that the strength of the storage effect, and therefore its impact on diversity within communities, varies latitudinally in association with climate. This finding highlights the importance of biotic interactions in shaping geographic diversity patterns, and emphasizes the need to understand the mechanisms underpinning ecological processes in greater detail than has previously been appreciated.