Aggregated spatial distributions of species in a subtropical karst forest,southwestern China

Aims Spatial distribution patterns of species reflect not only the ecological processes but also the habitat features that are related to species distribution. In karst topography, species distribution patterns provide more specific information about their environments. The objectives of this study are as follows: (i) to analyse and explain the spatial distribution patterns of conspecific trees in an old-growth subtropical karst forest; (ii) to investigate pattern changes at different spatial scales; (iii) to test the spatial pattern similarity (or dissimilarity) between trees at different abundances, diameter at breast height classes, canopy layers and different functional groups (shade tolerance and seed dispersal mode); (iv) to examine whether habitat heterogeneity has an important effect on the species spatial distribution.Methods The spatial distributions of woody species with ≥20 individuals in a 1-ha subtropical karst forest plot at Maolan in southwestern China were quantified using the relative neighbourhood density Ω based on the average density of conspecific species in a circular neighbourhood around each species.Important findings Aggregated distribution is the dominant pattern in the karst forest, but the ratio of aggregated species in total species number decreases with an increase in spatial scale. Less abundant species are more aggregated than most abundant species. Aggregation is weaker in larger diameter classes, which is consistent with the prediction of self-thinning. Seed dispersal mode influences spatial patterns, with species dispersed by animals being less aggregated than those dispersed by wind and gravity. Other species functional traits (e.g. shade tolerance) also influence the species spatial distributions. Moreover, differences among species habitat associations, e.g. with rocky outcrops, play a significant role in species spatial distributions. These results indicate that habitat heterogeneity, seed dispersal limitation and self-thinning primarily contribute to the species spatial distributions in this subtropical karst forest.     

Density dependence across multiple life stages in a temperate old-growth forest of northeast China

Recent studies on species coexistence suggest that density dependence is an important mechanism regulating plant populations. However, there have been few studies of density dependence conducted for more than one life-history stage or that control for habitat heterogeneity, which may influence spatial patterns of survival and mask density dependence. We explored the prevalence of density dependence across multiple life stages, and the effects of controlling for habitat heterogeneity, in a temperate forest in northeast China. We used generalized linear mixed-effects models to test for density-dependent mortality of seedlings and spatial point pattern analysis to detect density dependence for sapling-to-juvenile transitions. Conspecific neighbors had a negative effect on survival of plants in both life stages. At the seedling stage, we found a negative effect of conspecific seedling neighbors on survival when analyzing all species combined. However, in species-level analyses, only 2 of 11 focal species were negatively impacted by conspecific neighbors, indicating wide variation among species in the strength of density dependence. Controlling for habitat heterogeneity did not alter our findings of density dependence at the seedling stage. For the sapling-to-juvenile transition stage, 11 of 15 focal species showed patterns of local scale (<10 m) conspecific thinning, consistent with negative density dependence. The results varied depending on whether we controlled for habitat heterogeneity, indicating that a failure to account for habitat heterogeneity can obscure patterns of density dependence. We conclude that density dependence may promote tree species coexistence by acting across multiple life-history stages in this temperate forest.     

长白山次生杨桦林种群空间点格局及密度制约效应

以长白山5.2 hm²次生杨桦林样地为研究对象,利用空间点格局分析的双相关函数g（r）以及随机标签零模型和案例-对照设计法,探讨了树木种群空间格局及其密度制约效应。研究结果表明：生境异质性对不同生活型以及不同树种的成熟个体在大尺度范围上具有强烈影响。剔除生境异质性带来的影响后,在14个常见树种中有12个树种在小尺度上呈显著的空间聚集分布格局;随着空间尺度的增加聚集性分布树种数量急剧下降,在18 m尺度上聚集率下降到0。全部14个常见树种均呈现出显著的密度制约效应,表明密度制约是调节温带森林树木种群空间结构的主要作用机制。此外,密度制约与物种多度呈负相关,与亚林层和灌木层树种的同种聚集强度呈正相关。并且随着空间尺度增大受密度制约影响的树种百分比逐渐减小,14个常见树种中有11个在0-1 m处达到最大的密度制约强度。     

Strong plant-soil associations in a heterogeneous subtropical broad-leaved forest

The relative importance of deterministic and neutral processes on community assembly is currently a topic of much debate among ecologists. Analyzing species-environment associations is an effective way to assess the importance of deterministic process such as niche differentiation, but both habitat association and dispersal limitation can produce similar patterns of spatial aggregation in species. Therefore, it is crucial to control for the impact of dispersal limitation on species distributions when analyzing species-environment associations. We sampled soil with high resolutions in a 24 ha stem-mapped subtropical forest and tested plant-soil associations. We controlled for the influence of dispersal limitation by employing the homogeneous Thomas process to simulate the effect of dispersal limitation on the aggregation of tree species. After controlling for the effect of dispersal limitation, we found that the spatial heterogeneity of soil properties was associated with distributions of 88.2% (90 of 102 species) of tree species in this subtropical forest. Furthermore, not only did soil properties influence the distribution of tree species, but also tree species tended to affect properties of the soil around them. The soil factors most strongly influencing species distributions were TC, TN, TP, K, Mg, Si, soil moisture, and bulk density. We found the spatial heterogeneity of soil properties to be strongly associated with tree species distributions. Niche partitioning of soil gradients contributed substantially to species coexistence in this subtropical forest.     

Seedling survival simultaneously determined by conspecific,heterospecific, and phylogenetically related neighbors and habitat heterogeneity in a subtropical forest in Taiwan

Density dependence and habitat heterogeneity have been recognized as important driving mechanisms that shape the patterns of seedling survival and promote species coexistence in species‐rich forests. In this study, we evaluated the relative importance of density dependence by conspecific, heterospecific, and phylogenetically related neighbors and habitat heterogeneity on seedling survival in the Lienhuachih (LHC) Forest, a subtropical, evergreen forest in central Taiwan. Age‐specific effects of different variables were also studied. We monitored the fates of 1,642 newly recruited seedlings of woody plants within a 25‐ha Forest Dynamics Plot for 2 years. The effects of conspecific, heterospecific, and phylogenetically related neighbors and habitat heterogeneity on seedling survival were analyzed by generalized linear mixed models. Our results indicated that conspecific negative density dependence (CNDD) had a strong impact on seedling survival, and the effects of CNDD increased with seedling age. Heterospecific positive density dependence (HPDD) and phylogenetic positive density dependence (PPDD) had a significant influence on the survival of seedlings, and stronger HPDD and PPDD effects were detected for older seedlings. Furthermore, seedling survival differed among habitats significantly. Seedling survival was significantly higher in the plateau, high‐slope, and low‐slope habitats than in the valley. Overall, our results suggested that the effects of CNDD, HPDD, PPDD, and habitat heterogeneity influenced seedling survival simultaneously in the LHC subtropical forest, but their relative importance varied with seedling age. Such findings from our subtropical forest were slightly different from tropical forests, and these contrasting patterns may be attributed to differences in abiotic environments. These findings highlight the importance to incorporate phylogenetic relatedness, seedling age, and habitat heterogeneity when investigating the impacts of density dependence on seedling survival that may contribute to species coexistence in seedling communities.     

Spatial distributions of tree species in a subtropical forest of China

The spatial dispersion of individuals in a species is an important pattern that is controlled by many mechanisms. In this study we analyzed spatial distributions of tree species in a large-scale (20 ha) stem-mapping plot in a species-rich subtropical forest of China. O-ring statistic was used to measure spatial patterns of species with abundance >10. Ω_0–10, the mean conspecific density within 10 m of a tree, was used as a measure of the intensity of aggregation of a species. Our results showed: (1) aggregated distribution was the dominant pattern in the plot. The percentage of aggregated species decreased with increased spatial scale. (2) The percentages of significantly aggregated species decreased from abundant to intermediate and to rare species. Rare species was more strongly aggregated than common species. Aggregation was weaker in larger diameter classes. (3) Seed traits determined the spatial patterns of trees. Seed dispersal mode can influence spatial patterns of species, with species dispersed by both modes being less clumped than species dispersed by animal or wind, respectively. Considering these results, we concluded that seed dispersal limitation, self-thinning and habitat heterogeneity primarily contributed to spatial patterns and species coexistence in the forest.     

Effects of local biotic neighbors and habitat heterogeneity on tree and shrub seedling survival in an old-growth temperate forest

Seedling dynamics play a crucial role in determining species distributions and coexistence. Exploring causes of variation in seedling dynamics can therefore provide key insights into the factors affecting these phenomena. We examined the relative importance of biotic neighborhood processes and habitat heterogeneity using survival data for 5,827 seedlings in 39 tree and shrub species over 2?years from an old-growth temperate forest in northeastern China. We found significant negative density-dependence effects on survival of tree seedlings, and limited effects of habitat heterogeneity (edaphic and topographic variables) on survival of shrub seedlings. The importance of negative density dependence on young tree seedling survival was replaced by habitat in tree seedlings ??4?years old. As expected, negative density dependence was more apparent in gravity-dispersed species compared to wind-dispersed and animal-dispersed species. Moreover, we found that a community compensatory trend existed for trees. Therefore, although negative density dependence was not as pervasive as in other forest communities, it is an important mechanism for the maintenance of community diversity in this temperate forest. We conclude that both negative density dependence and habitat heterogeneity drive seedling survival, but their relative importance varies with seedling age classes and species traits.     

Density dependence is not very prevalent in a heterogeneous subtropical forest

There is a growing body of evidence demonstrating that tree survival is influenced by negative density‐dependence, but it is still controversial how the effect may vary with life‐stage, and to what extent it plays a role in regulating tree survival in heterogeneous subtropical forests. In this study, we investigated density‐dependent effects on tree survival of six tree species in a 5‐ha subtropical forest in eastern China. The roughly 45 000 individuals in the forest were fully censused in 2003 and 2008. For each of these species, we used an inhomogeneous pair‐correlation function to quantify the change in spatial distribution for different size classes, and a case‐control design to study seedling–adult associations in 2003. Autologistic regression was used to determine the influence of neighborhood factors on individual survival from 2003 to 2008. We found that seedlings of five species were repulsed by distance to nearest conspecific adults in terms of their survival, consistent with predictions of the Janzen–Connell mechanism. By contrast, only the least shade‐tolerant Schima superba had a negative relationship with individual survival and conspecific distance‐weighted basal area. This suggests that the Janzen–Connell effect is only prevalent at the early seedling stage or seed‐to‐seedling phase. The strength of clustering significantly declined at sapling–pole and pole–adult transitions for Sycopsis sinensis and at seedling–sapling transition for Cleyera pachyphylla. Correlations between individual survival and conspecific abundance for these species were consistent with trends in the strength of clustering. These results suggest that density dependence plays a limited role in individual survival and species spatial structure beyond the early seedling stage (i.e. after true leaves growing) in this forest. In addition, this study indicates that including individuals from early life‐stages and factoring out potential confounding factors such as habitat preference are important in studies that seek evidence for density dependence in forest trees.     

江西九连山亚热带常绿阔叶林优势种空间分布格局

于江西九连山国家自然保护区设置4 hm2亚热带常绿阔叶林固定监测样地,用样方法对亚热带常绿阔叶林进行调查分析。应用点格局方法分析生境异质性是否影响树木分布并分析优势种红钩栲(Castanopsis lamontii)、米槠(Castanopsis carlesii)、罗浮柿(Diospyros morrisiana)和细枝柃(Eurya loquaiana)分布格局,对比存在生境异质性和排除生境异质性两种情况下优势树种不同生长阶段(幼树、小树、成年树)的空间分布格局以及不同生长阶段之间的空间关联性,探讨生境异质性之外的其它种群分布影响因子。结果表明:1)生境异质性效应显著影响九连山树木分布,4个优势树种在大尺度上存在明显的生境偏好;2)用完全随机零模型不排除生境异质性时,4个优势种总体及其不同生长阶段在0—30 m所有尺度上主要呈现聚集分布;3)用异质性随机零模型排除生境异质性后,4个优势种及其不同生长阶段的聚集程度显著下降,只在小尺度上(0—5 m)呈现聚集分布;4)4个优势种的幼树与小树均表现显著的正相关。红钩栲的成年树与幼树及成年树与小树总体表现不相关。米槠的成年树与幼树总体以无相关为主;成年树与小树之间总体呈正相关。小乔木罗浮柿的成年树与幼树之间在小尺度上(0—5 m)表现负相关或无相关,在较大尺度范围上总体表现正相关关系;小树与成年树在研究尺度上表现出正相关。灌木细枝柃的成年树与幼树,成年树与小树及小树与幼树之间在研究尺度范围内均呈现正相关关系;5)研究发现九连山优势乔木物种通过密度制约和Janzen-Connell效应释放空间,为其他物种共存提供条件,而优势灌木物种细枝柃没有表现出这两个效应。     

Distribution patterns of tree species in an evergreen broadleaved forest in eastern China

Ecological assembly rules in evergreen broad-leaved forest are far from clear understanding. Spatial dispersion of individuals in a species is central in ecological theory. We analyzed the spatial patterns as well as associations between adult and juvenile of each tree species in a 5-ha subtropical evergreen broad-leaved forest plot in eastern China. Out of the 74 species occurring with more than 10 individuals, 88.4% of these species are aggregated. Most of them are aggregated from small to large scales. Spatial distributions of some species corre-spond with topography. Many bad dispersed species in the Baishanzu exhibit a highly aggregated distribution at small scales. These suggest that environmental heterogeneity and/or dispersal limitation may be the most important mechanisms that control the distribution patterns of these species. Our observations of the aggregations of abundant species basically support the hypothesis that dispersal limitation decreases as the number of reproductive trees increases. The rest species are randomly distributed, with less than 10 individuals. For most common species, spatial aggregation is weaker in larger diameter classes, and the distance between adults is larger than that between juveniles and adults, suggesting that density-dependence works on loosing aggregation and excluding conspecific juveniles away from adult trees. However, the density-dependent effect is not strong enough to eliminate all seedlings near adult trees and to result in a regular distribution of trees; thus the density dependence is usually masked by the refuge effect.     

Distribution patterns of tree species in an evergreen broadleaved forest in eastern China

Ecological assembly rules in evergreen broad-leaved forest are far from clear understanding. Spatial dispersion of individuals in a species is central in ecological theory. We analyzed the spatial patterns as well as associations between adult and juvenile of each tree species in a 5-ha subtropical evergreen broad-leaved forest plot in eastern China. Out of the 74 species occurring with more than 10 individuals, 88.4% of these species are aggregated. Most of them are aggregated from small to large scales. Spatial distributions of some species correspond with topography. Many bad dispersed species in the Baishanzu exhibit a highly aggregated distribution at small scales. These suggest that environmental heterogeneity and/or dispersal limitation may be the most important mechanisms that control the distribution patterns of these species. Our observations of the aggregations of abundant species basically support the hypothesis that dispersal limitation decreases as the number of reproductive trees increases. The rest species are randomly distributed, with less than 10 individuals. For most common species, spatial aggregation is weaker in larger diameter classes, and the distance between adults is larger than that between juveniles and adults, suggesting that density-dependence works on loosing aggregation and excluding conspecific juveniles away from adult trees. However, the density-dependent effect is not strong enough to eliminate all seedlings near adult trees and to result in a regular distribution of trees; thus the density dependence is usually masked by the refuge effect.     

Intraspecific and phylogenetic density-dependent seedling recruitment in a subtropical evergreen forest

Recent evidence suggests that plant performance can be influenced by the phylogenetic diversity of neighboring plants. However, no study to date has examined the effect of such phylogenetic density dependence on the transition from seed to seedling. Using 6 years of data on seedling recruitment and seed rain of 13 species from 130 stations (one 0.5 m² seed trap and three adjacent 1 m² seedling plots) in a subtropical evergreen forest, we asked: (1) Does negative density dependence act across seed to seedling stages? (2) Is there evidence for phylogenetic density dependence during the seed to seedling transition? (3) Does the strength of density dependence vary among years? Generalized linear mixed-effects models were used to model seed to seedling transition as a function of conspecific seed and seedling densities, heterospecific seed and seedling densities, and mean phylogenetic distance of heterospecific seeds and seedling. Conspecific seed density had a significant negative effect on seedling transition rates for 12 of 13 focal species. In contrast, conspecific seedling density had a positive effect for 7 species, suggesting species-specific habitat preferences. Few species were significantly affected by the density or phylogenetic relatedness of heterospecific seeds and seedlings. Only conspecific seed density effects varied among years for most focal species. Overall, our results reveal that conspecific seed and seedling densities play a more important role than the density or relatedness of heterospecific seeds and seedlings during the seed to seedling stage, suggesting that species-specific seed predators, along with habitat preferences, may contribute to diversity maintenance in this forest.     

Conspecific density dependence and community structure: Insights from 11 years of monitoring in an old‐growth temperate forest in Northeast China

Forest community structure may be influenced by seedling density dependence, however, the effect is loosely coupled with population dynamics and diversity in the short term. In the long term the strength of conspecific density dependence may fluctuate over time because of seedling abundance, yet few long‐term studies exist. Based on 11 years of seedling census data and tree census data from a 25‐ha temperate forest plot in Northeast China, we used generalized linear mixed models to test the relative effects of local neighborhood density and abiotic factors on seedling density and seedling survival. Spatial point pattern analysis was used to determine if spatial patterns of saplings and juveniles, in relation to conspecific adults, were in accordance with patterns uncovered by conspecific negative density dependence at the seedling stage. Our long‐term results showed that seedling density was mainly positively affected by conspecific density, suggesting dispersal limitation of seedling development. The probability of seedling survival significantly decreased over 1 year with increasing conspecific density, indicating conspecific negative density dependence in seedling establishment. Although there was variation in conspecific negative density dependence at the seedling stage among species and across years, a dispersed pattern of conspecific saplings relative to conspecific adults at the local scale (<10 m) was observed in four of the 11 species examined. Overall, sapling spatial patterns were consistent with the impacts of conspecific density on seedling dynamics, which suggests that conspecific negative density dependence is persistent over the long term. From the long‐term perspective, conspecific density dependence is an important driver of species coexistence in temperate forests.     

Spatial Patterns and Associations between Species Belonging to Four Genera of the Lauraceae Family

Spatial distribution pattern of biological related species present unique opportunities and challenges to explain species coexistence. In this study, we explored the spatial distributions and associations among congeneric species at both the species and genus levels to explain their coexistence through examining the similarities and differences at these two levels. We first used DNA and cluster analysis to confirmed the relative relationship of eight species within a 20 ha subtropical forest in southern China. We compared Diameter at breast height (DBH) classes, aggregation intensities and spatial patterns, associations, and distributions of four closely related species pairs to reveal similarities and differences at the species and genus levels. These comparisons provided insight into the mechanisms of coexistence of these congeners. O-ring statistics were used to measure spatial patterns of species. Ω _0–10, the mean conspecific density within 10 m of a tree, was used as a measure of the intensity of aggregation of a species, and g-function was used to analyze spatial associations. Our results suggested that spatial aggregations were common, but the differences between spatial patterns were reduced at the genus level. Aggregation intensity clearly reduced at the genus level. Negative association frequencies decreased at the genus level, such that independent association was commonplace among all four genera. Relationships between more closely related species appeared to be more competitive at both the species and genus levels. The importance of competition on interactions is most likely influenced by similarity in lifestyle, and the habitat diversity within the species’ distribution areas. Relatives with different lifestyles likely produce different distribution patterns through different interaction process. In order to fully understand the mechanisms generating spatial distributions of coexisting siblings, further research is required to determine the spatial patterns and associations at other classification levels.     

Density dependence and habitat preference shape seedling survival in a subtropical forest in central China

Aims Seedlings are vulnerable to many kinds of fatal abiotic and biotic agents, and examining the causes of seedling dynamics can help understand mechanisms of species coexistence. To disentangle the relative importance of neighborhood densities, habitat factors and phylogenetic relatedness on focal seedling survival, we monitored the survival of 5306 seedlings of 104 species>15 months. We address the following questions: (i) How do neighborhood densities, habitat variables and phylogenetic relatedness affect seedling survival? What is the relative importance of conspecific densities, habitat variables and phylogenetic relatedness to seedling survival? (ii) Does the importance of the neighborhood densities, habitat variables and phylogenetic relatedness vary among growth forms, leaf habits or dispersal modes? Specially, does the conspecific negative density dependence inhibit tree and deciduous seedlings more compared with shrub and evergreen species? Does density dependence affect the wind and animal-dispersed species equally?Methods We established 135 census stations to monitor seedling dynamics in a 25-ha subtropical forest plot in central China. Conspecific and heterospecific seedling density in the 1-m 2 seedling plot and adult basal area within a 20-m radius provided neighborhood density variables. Mean elevation, convexity and aspect of every 5- × 5-m grid with seedling plots were used to quantify habitat characteristics. We calculated the relative average phylodiversity between focal seedling and heterospecific neighbors to quantify the species relatedness in the neighborhood. Eight candidate generalized linear mixed models with binominal error distribution were used to compare the relative importance of these variables to seedling survival. Akaike's information criteria were used to identify the most parsimonious models.Important findings At the community level, both the neighborhood densities and phylogenetic relatedness were important to seedling survival. We found negative effects of increasing conspecific seedlings, which suggested the existence of species-specific density-dependent mortality. Phylodiversity of heterospecific neighbors was negatively related to survival of focal seedlings, indicating similar habitat preference shared among phylogenetically closely related species may drive seedling survival. The relative importance of neighborhood densities, habitat variables and phylogenetic relatedness varied among ecological guilds. Conspecific densities had significant negative effect for deciduous and wind-dispersed species, and marginally significant for tree seedlings>10cm tall and animal-dispersed species. Habitat variables had limited effects on seedling survival, and only elevation was related to the survival of evergreen species in the best-fit model. We conclude that both negative density-dependent mortality and habitat preference reflected by the phylogenetic relatedness shape the species coexistence at seedling stage in this forest.     

Species–habitat associations and demographic rates of forest trees

Niche‐driven effects on demographic processes generated in response to habitat heterogeneity partly shape local distributions of species. Thus, tree distributions are commonly studied in relation to habitat conditions to understand how niche differentiation contributes to species coexistence in forest communities. Many such studies implicitly assume that local abundance reflects habitat suitability, and that abundance is relatively stable over time. We compared models based on abundance with those based on demographic performance for making inferences about habitat association for 287 tree species from three large dynamic plots located in tropical, subtropical and temperate forests. The correlation between the predictions of the abundance‐based models and the demography‐based models varied widely, with correlation coefficients ranging nearly from ?1 to 1.This suggests that the two types of models capture different information about species–habitat associations. Demography‐based models evaluate habitat quality by focusing on population processes and thus should be preferred for understanding responses of tree species to habitat conditions, especially when habitat conditions are changing and species–habitat interactions cannot be considered to be at equilibrium.     

Species co-existence in an Australian subtropical rain forest: evidence for compensatory mortality

1 The relationship between the distribution of juveniles and conspecific trees was examined in a subtropical rain forest in south-east Queensland (Australia) to determine the role of compensatory mortality in maintaining the species richness of the forest.
2 Two species, which were among the most common canopy trees at the site, showed some evidence that the abundance of juveniles was reduced in the vicinity of conspecific trees. This spacing effect was particularly marked in Sloanea woollsii , but also present in Doryphora sassafras . The probability of finding juveniles of these species decreased as a function of (i) decreasing distance from the quadrats to the nearest conspecific tree and (ii) increasing density of conspecific trees ( Sloanea woollsii only). In addition, these species accounted for a smaller proportion of the total number of juveniles close to conspecifics than they did at greater distances.
3 In contrast, the juveniles of a number of other species were more commonly found in the vicinity of conspecific trees. Two species had juveniles that were comparatively poorly dispersed: Acmena ingens and Polyosma cunninghamii juveniles were seldom found more than 10 m and 15 m away from conspecific trees.
4 Compensatory mortality might be necessary to prevent other species from being excluded by the two most common species, but it does not appear to be sufficient to explain how the majority of species co-exist with each other.     

Determinant factors influencing the spatial distributions of subtropical lianas are correlated with components of functional trait spectra

Lianas are important vegetation components that control structure and function, especially in tropical and subtropical forests. To explore the spatial assembly mechanisms of a subtropical liana community, we tested the following hypotheses: spatial distributions of subtropical lianas are determined by forest structures and topographic features, which are surrogates for host/light availability and edaphic/water conditions, respectively, and these effects are mediated through species functional traits. We examined the spatial distribution of lianas in two plots (areas 9 and 16 ha) representing landscapes in an intact forest and a secondary forest, and analyzed spatial distribution pattern at the species level using a simple, spatially explicit model. We also examined the correlations between determinant factors for species distribution and species functional traits, including climbing habits, leaf traits and wood density. The spatial distribution of lianas was controlled mainly by topographic gradient. Most species had preferences for concave topographies, i.e., valley habitats. Any covariates related to the host (or to light) had little influence on the distribution of most liana species. Distributional responses to topography were different among species, and associated significantly with leaf nitrogen content and climbing habit, but not with wood density. The correlation between variation in habitat preferences and leaf economic spectrum suggests that an environmental filter for physiological response to topography is the important mechanism shaping the spatial patterns of this subtropical liana community.     

A spatial theory for characterizing predator–multiprey interactions in heterogeneous landscapes

Trophic interactions in multiprey systems can be largely determined by prey distributions. Yet, classic predator–prey models assume spatially homogeneous interactions between predators and prey. We developed a spatially informed theory that predicts how habitat heterogeneity alters the landscape-scale distribution of mortality risk of prey from predation, and hence the nature of predator interactions in multiprey systems. The theoretical model is a spatially explicit, multiprey functional response in which species-specific advection–diffusion models account for the response of individual prey to habitat edges. The model demonstrates that distinct responses of alternative prey species can alter the consequences of conspecific aggregation, from increasing safety to increasing predation risk. Observations of threatened boreal caribou, moose and grey wolf interacting over 378 181 km² of human-managed boreal forest support this principle. This empirically supported theory demonstrates how distinct responses of apparent competitors to landscape heterogeneity, including to human disturbances, can reverse density dependence in fitness correlates.     

Nature and Age of Neighbours Matter: Interspecific Associations among Tree Species Exist and Vary across Life Stages in Tropical Forests

Detailed information about interspecific spatial associations among tropical tree species is scarce, and hence the ecological importance of those associations may have been underestimated. However, they can play a role in community assembly and species diversity maintenance. This study investigated the spatial dependence between pairs of species. First, the spatial associations (spatial attraction and spatial repulsion) that arose between species were examined. Second, different sizes of trees were considered in order to evaluate whether the spatial relationships between species are constant or vary during the lifetime of individuals. Third, the consistency of those spatial associations with the species-habitat associations found in previous studies was assessed. Two different tropical ecosystems were investigated: a montane cloud forest and a lowland moist forest. The results showed that spatial associations among species exist, and these vary among life stages and species. The rarity of negative spatial interactions suggested that exclusive competition was not common in the studied forests. On the other hand, positive interactions were common, and the results of this study strongly suggested that habitat associations were not the only cause of spatial attraction among species. If this is true, habitat associations and density dependence are not the only mechanisms that explain species distribution and diversity; other ecological interactions, such as facilitation among species, may also play a role. These spatial associations could be important in the assembly of tropical tree communities and forest succession, and should be taken into account in future studies.