Climate and forest loss interactively restructure trait composition across a human-modified landscape.

Traits determine species response to climate conditions and the match between phenotypes and climate mediates spatial variation in species composition. These trait-climate linkages can be disrupted in human-modified landscapes. Human land use creates forest fragments where dispersal limitation or edge effects exclude species that may otherwise suit a given macroclimate. Furthermore, stressful macroclimate can limit viable trait combinations such that only a subset of values of any given trait occurs with respect to another trait, resulting in stronger trait covariance. Because forest loss can compound climatic stress, trait covariance from benign to harsher climates is expected to be stronger in fragments compared to contiguous forests. In a wet tropical forest landscape in the Western Ghats Biodiversity Hotspot of peninsular India, I compared fragments with adjacent contiguous forests for signatures of trait-mediated assembly of tree communities. Using four key plant traits-seed size, specific leaf area (SLA), wood density, and maximum height-I evaluated trait-abundance associations and trait covariance across climate, soil, and elevation gradients. In the contiguous forest, smaller-seeded, shorter, thinner-leaved species became more abundant from low to high elevations. In fragments, species with higher SLA were more abundant at sites with more seasonal climates and lower precipitation, and larger seeded species were less abundant at warmer sites. However, traits only weakly predicted abundances in both habitats. Moreover, only contiguous forests exhibited significant compositional change via traits, driven by trait syndromes varying along a composite gradient defined by elevation, water deficit, and soil C:N ratio. Site-level trait covariance revealed that warmer, wetter conditions in fragments favored taller species for given seed size, as compared to similar conditions in contiguous forests. Overall, trait syndromes and trait covariance, rather than single traits, determined the phenotypes best suited to macroclimate conditions and should inform management or restoration goals in fragments.

Environment shapes the spatial organization of tree diversity in fragmented forests across a human-modified landscape.

Biodiversity patterns are shaped by the combination of dispersal, environment, and stochasticity, but how the influence of these drivers changes in fragmented habitats remains poorly understood. We examined patterns and relationships among total (γ) and site-level (α) diversity, and site-to-site variation in composition (ß-diversity) of tree communities in structurally contiguous and fragmented tropical rainforests within a human-modified landscape in India's Western Ghats. First, for the entire landscape, we assessed the extent to which habitat type (fragment or contiguous forest), space and environment explained variation in α-diversity and composition. Next, within fragments and contiguous forest, we assessed the relative contribution of spatial proximity, environmental similarity, and their joint effects in explaining ß-diversity. We repeated these assessments with ß-diversity values corrected for the confounding effects of α- and γ-diversity using null models (ß-deviation). Lower γ-diversity of fragments resulted from both lower α- and ß-diversity compared to contiguous forests. However, ß-deviation did not differ between contiguous forests and fragments. Fragmented and contiguous forest clearly diverged in floristic composition, which was attributable to ß-diversity being driven by differences in elevation and MAP. Within fragmented forest, neither space nor environment explained ß-diversity, but ß-deviation increased with greater elevational differences. In contiguous forests by comparison, environment alone (mainly elevation) explained the most variation in ß-diversity and ß-deviation of both species' occurrences and abundances. Spatial gradients in environmental conditions played a larger role than dispersal limitation in shaping diversity and composition of tree communities across forest fragments. Thus, location of remnant patches at different elevations was a key factor underlying site-to-site variability in species abundances across fragments. Understanding the environmental characteristics of remnant forests in human-modified landscapes, combined with knowledge of species-environment relationships across different functional groups, would therefore be important considerations for management and restoration planning in human-modified landscapes.

Edge effects alter the role of fungi and insects in mediating functional composition and diversity of seedling recruits in a fragmented tropical forest.

BACKGROUND AND AIMS: In fragmented forests, proximity to forest edges can favour the establishment of resource-acquisitive species over more resource-conservative species. During seedling recruitment, resource-acquisitive species may benefit from either higher light availability or weaker top-down effects of natural enemies. The relative importance of light and enemies for recruitment has seldom been examined with respect to edge effects. METHODS: In a human-modified wet tropical forest in India, we first examined how functional traits indicative of resource-acquisitive vs. resource-conservative strategies, i.e. specific leaf area (SLA), leaf dry matter content, wood density and seed size, explained interspecific differences in densities of seedling recruits with distance to the forest edge. Then, we checked whether fungicide and insecticide treatments and canopy openness (proxy for light availability) explained edge effects on trait-mediated changes in seedling density. Finally, we examined whether light availability and natural enemy activity explained edge effects on functional diversity of seedling recruits. KEY RESULTS: Up to 60 m from edges, recruit densities increased with decreasing seed size, but not at 90-100 m, where recruit densities increased with higher SLA. Trait-mediated variation in recruit densities changed with pesticides only at 90-100 m: compared with control plots, fungicide increased recruit densities for low SLA species and insecticide increased smaller seeded species. For SLA, wood density and seed size, functional diversity of recruits was higher at 90-100 m than at 0-5 m. At 90-100 m, fungicide decreased functional diversity for SLA and insecticide reduced seed size diversity compared with control plots. Canopy openness explained neither variation in recruit density in relation to traits nor functional diversity. CONCLUSIONS: Altered biotic interactions can mediate local changes to trait composition and functional diversity during seedling recruitment in forest fragments, hinting at downstream effects on the structure and function of human-modified forests.

Edge effects reduce α-diversity but not ß-diversity during community assembly in a human-modified tropical forest.

Edge effects can alter the spatial organization of diversity in fragmented habitats. For tropical forests, however, there has been large variation in the strength and direction of such effects reported by different studies. For long-lived organisms like trees, one reason for inconsistent patterns might be due to most studies having examined patterns of diversity and compositional variation in older life stages that bear the legacy of a forest past. Younger life stages can reveal ongoing processes of assembly, but multi-stage examinations are rare. For seedling, sapling, and adult life stages of trees in a human-modified wet tropical forest in the Western Ghats Biodiversity Hotspot (India), we examined how proximity to forest edges (edge influence) modified the spatial organization of diversity. Specifically, for each life stage we tested whether edge influence led to loss of α- and γ-diversity and decreased ß-diversity in this landscape. We found lower α- and γ-diversity closer to forest edges, but only for seedlings. Seedling composition at 90-100 m from forest edges diverged from composition of sites within 60 m, suggesting that edge influence restricted the recruitment of some species to interior sites. In contrast, ß-diversity was greater near edges than interior forest for all life stages and most prominently for seedlings. Furthermore, ß-diversity at edges was primarily driven by species turnover, suggesting either marked species-environment associations or dispersal limitation. Low turnover at 90-100 m implies that ß-diversity arose from stochastic fluctuations in occurrences and abundances of the same species set. Overall, we find that high ß-diversity structured spatial patterns of diversity near edges, but recruitment bottlenecks are likely to reduce alpha diversity of forest fragments. Our results also corroborate the need to maintain sufficiently large areas of tropical forest free from edge effects to avoid the loss of interior forest species. To improve landscape-scale diversity of fragmented landscapes, restoration efforts should focus on recovery of species that are unable to regenerate near forest edges.

Weaker plant-enemy interactions decrease tree seedling diversity with edge-effects in a fragmented tropical forest.

In fragmented forests, tree diversity declines near edges but the ecological processes underlying this loss of diversity remain poorly understood. Theory predicts that top-down regulation of seedling recruitment by insect herbivores and fungal pathogens contributes to maintaining tree diversity in forests, but it is unknown whether proximity to forest edges compromises these diversity-enhancing biotic interactions. Here we experimentally demonstrate that weakened activity of fungal pathogens and insect herbivores reduced seedling diversity, despite similar diversity of seed rain, during recruitment near forest edges in a human-modified tropical landscape. Only at sites farthest from forest edges (90-100 m) did the application of pesticides lower seedling diversity relative to control plots. Notably, lower seedling diversity corresponded with weaker density-dependent mortality attributable to insects and fungi during the seed-to-seedling transition. We provide mechanistic evidence that edge-effects can manifest as cryptic losses of crucial biotic interactions that maintain diversity.

Environment and past land use together predict functional diversity in a temperate forest.

Environment and human land use both shape forest composition. Abiotic conditions sift tree species from a regional pool via functional traits that influence species' suitability to the local environment. In addition, human land use can modify species distributions and change functional diversity of forests. However, it is unclear how environment and land use simultaneously shape functional diversity of tree communities. Land-use legacies are especially prominent in temperate forest landscapes that have been extensively modified by humans in the last few centuries. Across a 900-ha temperate deciduous forest in the northeastern United States, comprising a mosaic of different-aged stands due to past human land use, we used four key functional traits-maximum height, rooting depth, wood density, and seed mass-to examine how multiple environmental and land-use variables influenced species distributions and functional diversity. We sampled ~40,000 trees >8 cm DBH within 485 plots totaling 137 ha. Species within plots were more functionally similar than expected by chance when we estimated functional diversity using all traits together (multi-trait), and to a lesser degree, with each trait separately. Multi-trait functional diversity was most strongly correlated with distance from the perennial stream, elevation, slope, and forest age. Environmental and land-use predictors varied in their correlation with functional diversities of the four individual traits. Landscape-wide change in abundances of individual species also correlated with both environment and land-use variables, but magnitudes of trait-environment interactions were generally stronger than trait interactions with land use. These findings can be applied for restoration and assisted regeneration of human-modified temperate forests by using traits to predict which tree species would establish well in relation to land-use history, topography, and soil conditions.

Influence of soil pathogens on early regeneration success of tropical trees varies between forest edge and interior.

Soil fungi are key mediators of negative density-dependent mortality in seeds and seedlings, and the ability to withstand pathogens in the shaded understory of closed-canopy forests could reinforce light gradient partitioning by tree species. For four species of tropical rainforest trees-two shade-tolerant and two shade-intolerant-we conducted a field experiment to examine the interactive effects of fungal pathogens, light, and seed density on germination and early seedling establishment. In a fully factorial design, seeds were sown into 1 m2 plots containing soil collected from underneath conspecific adult trees, with plots assigned to forest edge (high light) or shaded understory, high or low density, and fungicide or no fungicide application. We monitored total seed germination and final seedling survival over 15 weeks. Shade-intolerant species were strongly constrained by light; their seedlings survived only at the edge. Fungicide application significantly improved seedling emergence and/or survival for three of the four focal species. There were no significant interactions between fungicide and seed density, suggesting that pathogen spread with increased aggregation of seeds and seedlings did not contribute to pathogen-mediated mortality. Two species experienced significant edge-fungicide interactions, but fungicide effects in edge vs. interior forest varied with species and recruitment stage. Our results suggest that changes to plant-pathogen interactions could affect plant recruitment in human-impacted forests subject to fragmentation and edge-effects.

Testing predictions of the Janzen-Connell hypothesis: a meta-analysis of experimental evidence for distance- and density-dependent seed and seedling survival.

The Janzen-Connell hypothesis proposes that specialist natural enemies, such as herbivores and pathogens, maintain diversity in plant communities by reducing survival rates of conspecific seeds and seedlings located close to reproductive adults or in areas of high conspecific density. Variation in the strength of distance- and density-dependent effects is hypothesized to explain variation in plant species richness along climatic gradients, with effects predicted to be stronger in the tropics than the temperate zone and in wetter habitats compared to drier habitats.We conducted a comprehensive literature search to identify peer-reviewed experimental studies published in the 40+ years since the hypothesis was first proposed. Using data from these studies, we conducted a meta-analysis to assess the current weight of evidence for the distance and density predictions of the Janzen-Connell hypothesis.Overall, we found significant support for both the distance- and density-dependent predictions. For all studies combined, survival rates were significantly reduced near conspecifics compared to far from conspecifics, and in areas with high densities of conspecifics compared to areas with low conspecific densities. There was no indication that these results were due to publication bias.The strength of distance and density effects varied widely among studies. Contrary to expectations, this variation was unrelated to latitude, and there was no significant effect of study region. However, we did find a trend for stronger distance and density dependence in wetter sites compared to sites with lower annual precipitation. In addition, effects were significantly stronger at the seedling stage compared to the seed stage.Synthesis. Our study provides support for the idea that distance- and density-dependent mortality occurs in plant communities world-wide. Available evidence suggests that natural enemies are frequently the cause of such patterns, consistent with the Janzen-Connell hypothesis, but additional studies are needed to rule out other mechanisms (e.g. intraspecific competition). With the widespread existence of density and distance dependence clearly established, future research should focus on assessing the degree to which these effects permit species coexistence and contribute to the maintenance of diversity in plant communities.

The response of the frugivorous lion-tailed macaque (Macaca silenus) to a period of fruit scarcity.

Tropical rainforests show seasonal fluctuations in the abundance of fruits resulting in periods of resource scarcity for frugivores. We examined the response of an obligate frugivore, the lion-tailed macaque (LTM) (Macaca silenus), to a period of fruit scarcity in a rainforest in the Western Ghats, India. We estimated the abundance and distribution of fruit resources from food tree densities obtained from 348 point centered quadrats, and fruit availability from phenological monitoring of 195 trees of 15 reported major food species. Macronutrient content was estimated for fruits of 15 major food species. We estimated time spent feeding on different food items from 1,853 individual scans spanning 120 hr of observation of one habituated study group. There was a distinct period of fruit scarcity during the drier months of February to mid-March (Period 1) compared with late March and April (Period 2), separated by summer showers. Fruits available in Period 1 had lower soluble carbohydrate and lipid content and overall caloric value compared with Period 2. During the lean period, the LTM fed more on fruits of Drypetes wightii, which had the highest carbohydrate content, than on nectar of Palaquium ellipticum or Ficus spp., which had low carbohydrate content. During this period, the resource availability in a location significantly influenced the occurrence of feeding there. In Period 2, the group fed most on the seeds of Cullenia exarillata, the most abundant tree in the home-range and with the highest content of soluble carbohydrates. During this period, the abundance of food trees in fruit in a location did not seem to influence the occurrence of feeding. Low abundance, stochastic fruiting and, low quality might make Ficus spp. a poor fallback option for the LTM.