Elevational turnover in the composition of leaf miners and their interactions with host plants in Australian subtropical rainforest

Leaf miners are specialist herbivorous insects that are potentially vulnerable to environmental change because of their dependency on particular host plants. Little, however, is known about how climate affects the distribution of leaf miner communities and their interactions with host plants. Elevational gradients are useful tools for understanding how ecological communities respond to local clines in climate. Given that plant communities are known to undergo elevational turnover in response to changes in climatic conditions, we expect that leaf miner species will also change with elevation. We repeatedly hand collected leaf miners along three elevational gradients in subtropical rainforest in eastern Australia. Individual leaf miners were counted and identified to species, and their host plants were recorded. We tested if leaf miner species richness and the number of unique interactions among leaf miner and host plant species were affected by elevation. We also tested if the composition of leaf miner species and the composition of interactions between leaf miners and host plants showed a relationship with elevation. The rarefied number of unique leaf miner–host plant interactions significantly decreased with elevation, with a slight peak at approx. 700 m a.s.l., while neither rarefied or observed species richness (species density) of leaf miners nor observed numbers of unique interactions (interaction density) were significantly affected by elevation. The composition of leaf miner species and the composition of leaf miner–host plant interactions (occurrence of pairwise interactions) were significantly related to elevation. Elevational turnover in leaf miner species composition indicated that different species varied in their response to changes in biotic and/or abiotic conditions imposed by increasing elevation. Through our analyses, we identified four leaf miner species that may be locally vulnerable to climate change, as a result of their restricted elevational distribution and level of host specificity.     

内蒙古锡林河流域草原群落植物功能群组成沿水热梯度变化的样带研究

 用样带法调查了草原群落植物功能群组成沿水热梯度的变化。结果表明,随着海拔高度的降低,降水量的减少,热量和干燥度的增加,群落植物生活型功能群组成中,多年生杂类草的相对多度逐渐减少,而多年生丛生禾草的相对多度逐渐增加,多年生丛生禾草对多年生杂类草具有强烈的生态替代作用。在植物生态类群功能群组成中,中旱生植物的作用逐渐减弱,而旱生植物的作用显著增强。草地的利用方式对群落组成具有重要的影响,永久样地和割草场中不耐牧种的相对生物量显著地高于放牧场,而放牧场中耐牧种的相对生物量则显著地高于永久样地和割草场。受草地利用     

Leaf size, specific leaf area and microhabitat distribution of chaparral woody plants: contrasting patterns in species level and community level analyses

We examined variation in leaf size and specific leaf area (SLA) in relation to the distribution of 22 chaparral shrub species on small-scale gradients of aspect and elevation. Potential incident solar radiation (insolation) was estimated from a geographic information system to quantify microclimate affinities of these species across north- and south-facing slopes. At the community level, leaf size and SLA both declined with increasing insolation, based on average trait values for the species found in plots along the gradient. However, leaf size and SLA were not significantly correlated across species, suggesting that these two traits are decoupled and associated with different aspects of performance along this environmental gradient. For individual species, SLA was negatively correlated with species distributions along the insolation gradient, and was significantly lower in evergreen versus deciduous species. Leaf size exhibited a negative but non-significant trend in relation to insolation distribution of individual species. At the community level, variance in leaf size increased with increasing insolation. For individual species, there was a greater range of leaf size on south-facing slopes, while there was an absence of small-leaved species on north-facing slopes. These results demonstrate that analyses of plant functional traits along environmental gradients based on community level averages may obscure important aspects of trait variation and distribution among the constituent species.     

New method for rapid assessment of the functional composition of herbaceous plant communities

Abstract We propose a rapid sampling method to assess the functional composition of herbaceous plant communities without prior knowledge of the floristic composition. To determine the community‐level value of traits (‘aggregated trait values’) for a plant community, a standardized population‐centred method exists, but requires substantial manpower and reliable botanical knowledge. We tested an alternative method, the trait transect, using four subalpine pastures in the Beaufortain region (Northern French Alps) selected along a fertility gradient. We applied both methods to measure five commonly used ‘soft traits’ known to be responsive to soil nutrient availability: plant vegetative and reproductive height, specific leaf area, leaf dry matter and nitrogen contents. We tested whether the variation of these traits along the gradient detected with the population‐centred method was also detected with the trait transect. Both methods detected expected trends in the traits in response to the fertility gradient. The trait transect method was as efficient as the population‐centred method and is recommended as an appropriate tool for monitoring ecosystem changes in response to environmental conditions and management, especially in species‐rich communities.     

天童常绿阔叶林演替系列植物群落的N∶P化学计量特征

 土壤氮磷养分对植物生长的限制性可通过植被的N∶P化学计量特征来反映。该研究以常绿阔叶林演替系列为对象,将N∶P作为诊断指标,揭示 常绿阔叶林次生演替过程中植物群落的N∶P化学计量特征和养分限制作用。结果显示：1)物种水平的N∶P大小不一,但演替系列总体的变化特 征表现出了较高的一致性。2）在群落水平上,次生演替初期的灌草丛N∶P极小(7.38),远远低于14,当演替进入灌丛阶段,N∶P 显著增高到 19.96,在进入演替中期的针叶林(14.29)和针阔混交林(14.21)时,N∶P显著下降到 14～16之间,演替中后期的木荷(Schima superba)群落 (18.77)和栲树(Castanopsis fargesii)群落(20.13)的N∶P发生了显著的升高过程 。根据以往对N∶P临界值的确定,可以认为,常绿阔叶林次 生演替初期的植物群落生产力主要受到氮素的限制作用;演替中期的针叶林和针阔混交林主要受氮磷的共同限制,但以氮素的限制作用更为强 烈;演替中后期植物群落主要受到土壤磷素的限制作用。     

Leaf δ13C reflects ecosystem patterns and responses of alpine plants to the environments on the Tibetan Plateau

Leaf δ¹³C is an indicator of water-use efficiency and provides useful information on the carbon and water balance of plants over longer periods. Variation in leaf δ¹³C between or within species is determined by plant physiological characteristics and environmental factors. We hypothesized that variation in leaf δ¹³C values among dominant species reflected ecosystem patterns controlled by large-scale environmental gradients, and that within-species variation indicates plant adaptability to environmental conditions. To test these hypotheses, we collected leaves of dominant species from six ecosystems across a horizontal vegetation transect on the Tibetan Plateau, as well as leaves of Kobresia pygmaea (herbaceous) throughout its distribution and leaves of two coniferous tree species ( Picea crassifolia, Abies fabri ) along an elevation gradient throughout their distribution in the Qilian Mountains and Gongga Mountains, respectively. Leaf δ¹³C of dominant species in the six ecosystems differed significantly, with values for evergreen coniferous13C values of the dominant species and of K. pygmaea were negatively correlated with annual precipitation along a water gradient, but leaf δ¹³C of A. fabri was not significantly correlated with precipitation in habitats without water-stress. This confirms that variation of δ¹³C between or within species reflects plant responses to environmental conditions. Leaf δ¹³C of the dominant species also reflected water patterns on the Tibetan Plateau, providing evidence that precipitation plays a primary role in controlling ecosystem changes from southeast to northwest on the Tibetan Plateau.     

锡林河流域草原群落植物多样性和初级生产力沿水热梯度变化的样带研究

 用样带法研究了草原群落植物多样性和初级生产力沿海拔和水分梯度的变化,结果表明;在样带梯度上,物种丰富度、多样性和群落初级生产力与海拔高度、年降水量和土壤有机C及全N含量呈正相关,而与年平均气温和干燥度呈负相关。随着海拔高度的降低,降水量的减少,热量和干燥度的增加,以及土壤有机C和全N含量的降低,草原群落的物种丰富度、多样性和初级生产力逐渐降低。典型相关分析的结果揭示出,土壤有机C含量和干燥度是对草原群落物种丰富度和初级生产力具有更大的影响。同时,草地的利用方式和强度对群落植物多样性和生产力也具有较强的影响。     

放牧对内蒙古锡林河流域草地群落植物茎叶生物量资源分配的影响

以内蒙古锡林河流域沿水分梯度分布的灰脉苔草(Carex appendiculata)、贝加尔针茅(Stipa baicalensis)、羊草(Leymus chinensis)、大针茅(Stipa grandis)、小叶锦鸡儿(Caragana microphylla)和冷蒿(Artemisia frigida) 6个草地群落为对象, 研究了围封禁牧与放牧样地中144个共有植物种的高度、丛幅面积、茎、叶和株(丛)生物量、茎叶比等性状。结果表明: 1)在个体水平上, 放牧样地中植物的生殖枝高度、营养枝高度、丛幅面积、单株(丛)生物量、茎、叶生物量和茎叶比均显著低于围封禁牧样地, 植物在放牧干扰下表现出明显的小型化现象; 2)在群落水平上, 放牧亦显著降低了群落总生物量和茎、叶生物量; 3)过度放牧显著改变了物种的资源分配策略, 使生物量向叶的分配比例增加, 向茎的分配比例减少。资源优先向同化器官分配可能是植物对长期放牧干扰的一种重要适应对策; 4)轻度放牧对物种的资源分配没有显著影响, 单株(丛)生物量和群落茎、叶及总生物量均表现出增加趋势, 这与过度放牧的影响正好相反。过度放牧引起的植物个体小型化改变了生态系统中物种的资源分配策略, 进而对生态系统功能产生重要的影响。     

Leaf absorptances of Mohave and Sonoran desert plants

Summary Leaf absorptances to solar radiation in the 400–700 nm (photosynthetically useful wavelengths) are presented for a diversity of species in the Mohave and Sonoran Deserts of North America. As a life form shrubs are shown to have the widest range of absorptance, with perennial herbs and cacti exhibiting a smaller range, and very little variation in leaf absorptances among trees and annuals. The measurement of leaf absorptance at 625 nm is shown to be the same as the leaf absorptance to solar radiation over the 400–700 nm. Over a wide range of species and absorptances, the total solar leaf absorptance (400–3,000 nm) is shown to be closely related to the 400–700 nm leaf absorptance.     

Trait variation and functional diversity maintenance of understory herbaceous species coexisting along an elevational gradient in Yulong Mountain,Southwest China

Characterizing trait variation across different ecological scales in plant communities has been viewed as a way to gain insights into the mechanisms driving species coexistence. However, little is known about how changes in intraspecific and interspecific traits across sites influence species richness and community assembly, especially in understory herbaceous communities. Here we partitioned the variance of four functional traits (maximum height, leaf thickness, leaf area and specific leaf area) across four nested biological scales: individual, species, plot, and elevation to quantify the scale-dependent distributions of understory herbaceous trait variance. We also integrated the comparison of the trait variance ratios to null models to investigate the effects of different ecological processes on community assembly and functional diversity along a 1200-m elevational gradient in Yulong Mountain. We found interspecific trait variation was the main trait variation component for leaf traits, although intraspecific trait variation ranged from 10% to 28% of total variation. In particular, maximum height exhibited high plasticity, and intraspecific variation accounted for 44% of the total variation. Despite the fact that species composition varied across elevation and species richness decreased dramatically along the elevational gradient, there was little variance at our largest (elevation) scale in leaf traits and functional diversity remained constant along the elevational gradient, indicating that traits responded to smaller scale influences. External filtering was only observed at high elevations. However, strong internal filtering was detected along the entire elevational gradient in understory herbaceous communities, possibly due to competition. Our results provide evidence that species coexistence in understory herbaceous communities might be structured by differential niche-assembled processes. This approach--integrating different biological scales of trait variation--may provide a better understanding of the mechanisms involved in the structure of communities.     

Different trends in phylogenetic and functional structure of plant communities along an elevation gradient

The study of diversity gradients due to elevation dates back to the foundation of biogeography and ecology. Although elevation-driven patterns of plant diversity have been reported for centuries, uncertainty still exists about the assembly rules that drive these patterns. In this study, we revealed the causal factor of community assemblies for the diversity of tree and herb species along an elevation. To this end, we applied an integrated method using both functional traits and phylogeny, called the mean pairwise functional-phylogenetic distance, to understand the assembly rules for woody and herbaceous species communities along an elevation gradient. At higher elevation sites, woody and herbaceous communities were comprised of species having similar traits. The phylogenetic trends for woody species were consistent with the functional trends; closely related species co-occurred more frequently than expected at higher elevations. Phylogenetic trends for herb species were opposite to the functional trends; species with similar traits but having a random phylogenetic distribution co-occurred at higher elevations. We suggest that the community assembly rules for woody and herb species vary with elevation; and functional constraints due to environmental filtering at higher elevation act as assembly rules along gradients in both woody and herbaceous communities, even though their phylogenetic backgrounds differ.     

Elevation,space and host plant species structure Ericaceae root-associated fungal communities in Papua New Guinea

Our study aimed to identify significant predictors (spatial distance, elevation, host plant taxonomy) which shape the structure of endophytic fungal (ENDF) and putative ericoid mycorrhizal (ErMF) communities associated with roots of Ericaceae in Papua New Guinea. Roots of five Ericaceae together with one non-Ericaceae species were sampled at an experimental site and one common Ericaceae species was chosen for sampling along an elevation gradient. ENDF and putative ErMF communities were determined using the 454-sequencing approach. ENDF as well as putative ErMF communities were affected by interacting host plant. While the putative ErMF community was structured by host plants at the genus level, the ENDF community was affected by host plant subfamily level. Composition of ENDF as well as putative ErMF communities were affected by elevation. Non-Ericaceae plant species (Hypericum sp.) harbored similar communities of ENDF as well as putative ErMF as Ericaceae plants. Our study provides a first insight into ErMF and ENDF community ecology of Ericaceae in Papua New Guinea.     

Leaf trait relationships of native and invasive plants: community- and global-scale comparisons

Leaf carbon capture strategies of native and exotic invasive plants were compared by examining leaf traits and their scaling relationships at community and global scales. Community-level leaf trait data were obtained for 55 vascular plant species from nutrient-enriched and undisturbed bushland in Sydney, Australia. Global-scale leaf trait data were compiled from the literature for 75 native and 90 exotic invasive coexisting species. At the community level, specific leaf area (SLA), foliar nitrogen and phosphorus (N(mass) and P(mass)) and N:P ratio were significantly higher for exotics at disturbed sites compared with natives at undisturbed sites, with natives at disturbed sites being intermediate. SLA, N(mass) and P(mass) were positively correlated, with significant shifts in group means along a common standardized major axis (SMA) slope. At the global scale, invasives had significantly higher N(mass), P(mass), assimilation rate (A(mass) and A(area)) and leaf area ratio (LAR) than natives. All traits showed positive correlations, with significant shifts in group means along a common slope. For a given SLA, invasives had higher A(mass) (7.7%) and N(mass) (28%). Thus, exotic invasives do not have fundamentally different carbon capture strategies from natives but are positioned further along the leaf economics spectrum towards faster growth strategies. Species with leaf traits enabling rapid growth will be successful invaders when introduced to novel environments where resources are not limited.     

大青沟自然保护区主要森林群落优势种的叶性状

植物功能性状是近年来生态学研究的热点。其中叶功能性状与植株生物量和植物对资源的获得、利用及利用效率的关系最为密切。大青沟森林植物群落分布于科尔沁沙地,生境条件非常特殊,在沙沟里存在着一片茂密的森林,与周围浩瀚无垠的沙坨景观形成极为鲜明的对照。从沟底到沟顶,虽然海拔高度仅相差六、七十米,但由于距离沟底水源不同,土壤条件差异大,形成了不同的森林植物群落类型。大果榆群落、蒙古栎群落和水曲柳群落分别分布在大青沟自然保护区的沟顶、沟中和沟底。为了对大青沟自然保护区,不同环境梯度下的森林植物群落叶片功能性状进行研究,以大青沟自然保护区大果榆、蒙古栎、水曲柳3种主要森林群落为研究对象,分别测定不同群落优势种的叶厚度、比叶面积、叶干物质含量、叶大小和叶干重等5项叶功能性状,研究叶功能性状之间的关系,并对不同生长型、不同群落叶功能性状进行比较。相关分析结果表明,叶厚度与比叶面积呈极显著负相关,与叶大小和叶干重呈极显著正相关;比叶面积与叶干物质含量、叶干重呈极显著负相关,与叶大小呈显著正相关;叶干物质含量与叶大小呈极显著负相关,与叶干重呈极显著正相关;叶大小与叶干重呈极显著正相关。不同生长型植物叶片性状的分析表明,草本植物的叶干物质含量比乔木和灌木低,而其比叶面积高于乔木和灌木;对不同群落叶功能性状进行比较发现,大果榆群落和蒙古栎群落乔木、灌木叶厚度与干物质含量显著高于水曲柳群落的叶厚度与干物质含量,二者比叶面积显著低于水曲柳群落。大果榆群落和蒙古栎群落从叶功能性状的角度,它们具有较高的叶干物质含量和较低的比叶面积,体现出适应干旱生境叶片的特征。水曲柳群落呈现出低叶干物质含量、高比叶面积的特征,体现出适应湿润、土壤水分较好生境的特征。不同群落通过调节自身的物种组成,形成不同的功能性状组合来适应环境。     

Using functional diversity patterns to explore metacommunity dynamics: a framework for understanding local and regional influences on community structure

The metacommunity concept, describing how local and regional scale processes interact to structure communities, has been successfully applied to patterns of taxonomic diversity. Functional diversity has proved useful for understanding local scale processes, but has less often been applied to understanding regional scale processes. Here, we explore functional diversity patterns within a metacommunity context to help elucidate how local and regional scale processes influence community assembly. We detail how each of the four metacommunity perspectives (species sorting, mass effects, patch dynamics, neutral) predict different patterns of functional beta‐ and alpha‐diversity and spatial structure along two key gradients: dispersal limitation and environmental conditions. We then apply this conceptual model to a case study from alpine tundra plant communities. We sampled species composition in 17 ‘sky islands’ of alpine tundra in the Colorado Rocky Mountains, USA that differed in geographic isolation and area (key factors related to dispersal limitation) and temperature and elevation (key environmental factors). We quantified functional diversity in each site based on specific leaf area, leaf area, stomatal conductance, plant height and chlorophyll content. We found that colder high elevation sites were functionally more similar to each other (decreased functional beta‐diversity) and had lower functional alpha‐diversity. Geographic isolation and area did not influence functional beta‐ or alpha‐diversity. These results suggest a strong role for environmental conditions structuring alpine plant communities, patterns consistent with the species sorting metacommunity perspective. Incorporating functional diversity into metacommunity theory can help elucidate how local and regional factors structure communities and provide a framework for observationally examining the role of metacommunity dynamics in systems where experimental approaches are less tractable.     

Light and nitrogen competition limit Lolium perenne in experimental grasslands of increasing plant diversity

Positive species richness effects on aboveground community productivity in experimental grasslands have been reported to correlate with variable responses of individual species. So far, it is largely unknown whether more complete use of resources at the community level correlates with resource limitation of particular species and may explain their decreasing performance with increasing plant diversity. Using the subordinate grass species Lolium perenne L. as a model, we monitored populations in 82 experimental grasslands of different plant diversity (Jena Experiment) from year 2 to 6 after establishment, and measured ecophysiological leaf traits related to light and nutrient acquisition and use. Population and plant individual sizes of L. perenne decreased with increasing species richness. A decrease in transmitted light with increasing species richness and legume proportion correlated with increasing specific leaf area (SLA). Despite this morphological adaptation to lower light availability, decreasing foliar δ(13) C signatures with increasing species richness and low variation in leaf gas exchange and chlorophyll concentrations suggested a low capacity of L. perenne for adjustment to canopy shade. Leaf nitrogen concentrations and foliar δ(15) N signatures indicated a better N supply in communities with legumes and a shift in the uptake of different N forms with increasing species richness. Leaf blade nitrate and carbohydrate concentrations as indicators of plants nutritional status supported that light limitation with increasing species richness and legume proportions, combined with a N limitation in communities with increasing proportions of non-legumes, correlated with the decreasing performance of L. perenne in communities of increasing plant diversity.     

Mycorrhizal features and leaf traits covary at the community level during primary succession

The mycorrhizal association between fungi and plants is thought to be an important component of the functional diversity of plant communities, although evidence is weak. We explored community-level changes to mycorrhizal features and leaf traits along a primary successional gradient on a sedimentation coast. We found that community weighted mean values of mycorrhizal features and leaf traits varied systematically along the successional series, representing a transition from communities dominated by more stress-tolerant plant species (less mycorrhizal colonization and lower specific leaf area) to less stress-tolerant plant species (higher mycorrhizal colonization and specific leaf area). Despite the co-variation in leaf traits and mycorrhizal features at the community level, only mycorrhizal features showed an increasing functional diversity with successional age. These results suggest that mycorrhizal fungi should not be forgotten in trait-based plant community ecology.     

Altitudinal Change in LAI and Stand Leaf Biomass in Tropical Montane Forests: a Transect Study in Ecuador and a Pan-Tropical Meta-Analysis

Abstract Leaf area index (LAI) is a key parameter controlling plant productivity and biogeochemical fluxes between vegetation and the atmosphere. Tropical forests are thought to have comparably high LAIs; however, precise data are scarce and environmental controls of leaf area in tropical forests are not understood. We studied LAI and stand leaf biomass by optical and leaf mass-related approaches in five tropical montane forests along an elevational transect (1,050–3,060 m a.s.l.) in South Ecuador, and conducted a meta-analysis of LAI and leaf biomass data from tropical montane forests around the globe. Study aims were (1) to assess the applicability of indirect and direct approaches of LAI determination in tropical montane forests, (2) to analyze elevation effects on leaf area, leaf mass, SLA, and leaf lifespan, and (3) to assess the possible consequences of leaf area change with elevation for montane forest productivity. Indirect optical methods of LAI determination appeared to be less reliable in the complex canopies than direct leaf mass-related approaches based on litter trapping and a thorough analysis of leaf lifespan. LAI decreased by 40–60% between 1,000 and 3,000 m in the Ecuador transect and also in the pan-tropical data set. This decrease indicates that canopy carbon gain, that is, carbon source strength, decreases with elevation in tropical montane forests. Average SLA decreased from 88 to 61 cm² g⁻¹ whereas leaf lifespan increased from 16 to 25 mo between 1,050 and 3,060 m in the Ecuador transect. In contrast, stand leaf biomass was much less influenced by elevation. We conclude that elevation has a large influence not only on the leaf traits of trees but also on the LAI of tropical montane forests with soil N (nitrogen) supply presumably being the main controlling factor.     

Elevational Variation of Leaf Traits in Montane Rain Forest Tree Species at La Chinantla, Southern México1

Variation in leaf traits of dominant tree species in six montane rain forest communities was analyzed along an elevational gradient ranging from 1220 to 2560 m within a single basin at La Chinantla, Oaxaca, México. Three groups of characters were used: morphological (leaf shape, margin, blade configuration, and phyllotaxy), morphometric (leaf area, leaf mass per area, stomatal density, and blade length/width ratio), and anatomical (thicknesses of blade, palisade [PP], and spongy [SP] parenchymae, PP/SP ratio, and epidermis and cuticle thicknesses). The variation of morphological characteristics was only evident at the highest elevations; in contrast, thickness of leaf blade, PP, SP, as well as leaf mass per area clearly increased along the gradient, whereas leaf area was the only variable that significantly decreased with elevation. Thicknesses of epidermis and of the two cuticles were not significantly correlated with elevation. A classification analysis based on a leaf trait matrix led to the distinction between low and high elevation communities, with an approximate limit between them at ca 2300 to 2400 m. The results are discussed in light of environmental changes occurring along elevational gradients. Leaf characteristics of montane rain forest plants offer important insights about the complex roles of abiotic factors operating in these environments and supplement the traditional physiognomic classification schemes for these communities.     

Community Functional Responses to Soil and Climate at Multiple Spatial Scales: When Does Intraspecific Variation Matter?

Despite increasing evidence of the importance of intraspecific trait variation in plant communities, its role in community trait responses to environmental variation, particularly along broad-scale climatic gradients, is poorly understood. We analyzed functional trait variation among early-successional herbaceous plant communities (old fields) across a 1200-km latitudinal extent in eastern North America, focusing on four traits: vegetative height, leaf area, specific leaf area (SLA), and leaf dry matter content (LDMC). We determined the contributions of species turnover and intraspecific variation to between-site functional dissimilarity at multiple spatial scales and community trait responses to edaphic and climatic factors. Among-site variation in community mean trait values and community trait responses to the environment were generated by a combination of species turnover and intraspecific variation, with species turnover making a greater contribution for all traits. The relative importance of intraspecific variation decreased with increasing geographic and environmental distance between sites for SLA and leaf area. Intraspecific variation was most important for responses of vegetative height and responses to edaphic compared to climatic factors. Individual species displayed strong trait responses to environmental factors in many cases, but these responses were highly variable among species and did not usually scale up to the community level. These findings provide new insights into the role of intraspecific trait variation in plant communities and the factors controlling its relative importance. The contribution of intraspecific variation to community trait responses was greatest at fine spatial scales and along edaphic gradients, while species turnover dominated at broad spatial scales and along climatic gradients.