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Mutualistic symbioses between ants and plants are widespread in nature. Ants can deter unwanted pests and provide protection for plants in return for food or housing rewards. Using a long-term demographic dataset in a tropical seasonal rain forest in Southwest China, we found that associations with ants positively influenced seedling survival and adult growth, and also, species with extrafloral nectaries experienced weaker conspecific negative density dependence compared with species without extrafloral nectaries. Furthermore, we found strong evidence suggesting that species in our forest experienced conspecific density dependence, which we interpreted as heavy pest pressure that may drive the development of anti-pest symbioses such as the plant-ant relationship. Our findings suggest that ants and conspecific neighbors play important but inverse roles on plant survival and growth and that ants can buffer tree neighborhood interactions in this tropical forest.
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Nowadays, capacitive deionization (CDI) has emerged as a prominent technology in the desalination field, typically utilizing porous carbons as electrodes. However, the precise significance of electrode properties and operational conditions in shaping desalination performance remains blurry, necessitating numerous time-consuming and resource-intensive CDI experiments. Machine learning (ML) presents an emerging solution, offering the prospect of predicting CDI performance with minimal investment in electrode material synthesis and testing. Herein, four ML models are used for predicting the CDI performance of porous carbons. Among them, the gradient boosting model delivers the best performance on test set with low root mean square error values of 2.13 mg g-1 and 0.073 mg g-1 min-1 for predicting desalination capacity and rate, respectively. Furthermore, SHapley Additive exPlanations is introduced to analyze the significance of electrode properties and operational conditions. It highlights that electrolyte concentration and specific surface area exert a substantially more influential role in determining desalination performance compared to other features. Ultimately, experimental validation employing metal-organic frameworks-derived porous carbons and biomass-derived porous carbons as CDI electrodes is conducted to affirm the prediction accuracy of ML models. This study pioneers ML techniques for predicting CDI performance, offering a compelling strategy for advancing CDI technology.
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Host specialization plays a critical role in the ecology and evolution of plant-microbe symbiosis. Theory predicts that host specialization is associated with microbial genome streamlining and is influenced by the abundance of host species, both of which can vary across latitudes, leading to a latitudinal gradient in host specificity. Here, we quantified the host specificity and composition of plant-bacteria symbioses on leaves across 329 tree species spanning a latitudinal gradient. Our analysis revealed a predominance of host-specialized leaf bacteria. The degree of host specificity was negatively correlated with bacterial genome size and the local abundance of host plants. Additionally, we found an increased host specificity at lower latitudes, aligning with the high prevalence of small bacterial genomes and rare host species in the tropics. These findings underscore the importance of genome streamlining and host abundance in the evolution of host specificity in plant-associated bacteria along the latitudinal gradient.
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Tamaño del Genoma , Especificidad del Huésped , Hojas de la Planta , Simbiosis , Hojas de la Planta/microbiología , Bacterias/genética , Bacterias/clasificación , Genoma Bacteriano , Árboles/microbiologíaRESUMEN
The early growth stage of plants is vital to community diversity and community regeneration. The Janzen-Connell hypothesis predicts that conspecific density dependence lowers the survival of conspecific seedlings by attracting specialist natural enemies, promoting the recruitment and performance of heterospecific neighbors. Recent work has underscored how this conspecific negative density dependence may be mediated by mutualists - such as how mycorrhizal fungi may mediate the accrual of host-specific pathogens beneath the crown of conspecific adult trees. Aboveground mutualist and enemy interactions exist as well, however, and may provide useful insight into density dependence that are as of yet unexplored. Using a long-term seedling demographic dataset in a subtropical forest plot in central China, we confirmed that conspecific neighborhoods had a significant negative effect on seedling survival in this subtropical forest. Furthermore, although we detected more leaf damage in species that were closely related to ants, we found that the presence of ants had significant positive effects on seedling survival. Beside this, we also found a negative effect of ant appearance on seedling growth which may reflect a trade-off between survival and growth. Overall, our findings suggested that ants and conspecific neighborhoods played important but inverse roles on seedling survival and growth. Our results suggest ants may mediate the influence of conspecific negative density dependence on seedling survival at community level.
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Hormigas , Bosques , Herbivoria , Plantones , China , Animales , Plantones/fisiología , Hormigas/fisiología , Árboles/fisiología , Densidad de Población , SimbiosisRESUMEN
Predicting and managing the structure and function of plant microbiomes requires quantitative understanding of community assembly and predictive models of spatial distributions at broad geographic scales. Here, we quantified the relative contribution of abiotic and biotic factors to the assembly of phyllosphere bacterial communities, and developed spatial distribution models for keystone bacterial taxa along a latitudinal gradient, by analyzing 16S rRNA gene sequences from 1453 leaf samples taken from 329 plant species in China. We demonstrated a latitudinal gradient in phyllosphere bacterial diversity and community composition, which was mostly explained by climate and host plant factors. We found that host-related factors were increasingly important in explaining bacterial assembly at higher latitudes while nonhost factors including abiotic environments, spatial proximity and plant neighbors were more important at lower latitudes. We further showed that local plant-bacteria associations were interconnected by hub bacteria taxa to form metacommunity-level networks, and the spatial distribution of these hub taxa was controlled by hosts and spatial factors with varying importance across latitudes. For the first time, we documented a latitude-dependent importance in the driving factors of phyllosphere bacteria assembly and distribution, serving as a baseline for predicting future changes in plant phyllosphere microbiomes under global change and human activities.
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Bacterias , Microbiota , Humanos , ARN Ribosómico 16S/genética , Bacterias/genética , Plantas/genética , Hojas de la Planta/microbiologíaRESUMEN
Arbuscular mycorrhizal (AM) and ectomycorrhizal (EcM) associations are critical for host-tree performance. However, how mycorrhizal associations correlate with the latitudinal tree beta-diversity remains untested. Using a global dataset of 45 forest plots representing 2,804,270 trees across 3840 species, we test how AM and EcM trees contribute to total beta-diversity and its components (turnover and nestedness) of all trees. We find AM rather than EcM trees predominantly contribute to decreasing total beta-diversity and turnover and increasing nestedness with increasing latitude, probably because wide distributions of EcM trees do not generate strong compositional differences among localities. Environmental variables, especially temperature and precipitation, are strongly correlated with beta-diversity patterns for both AM trees and all trees rather than EcM trees. Results support our hypotheses that latitudinal beta-diversity patterns and environmental effects on these patterns are highly dependent on mycorrhizal types. Our findings highlight the importance of AM-dominated forests for conserving global forest biodiversity.
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Biodiversidad , Bosques , Micorrizas/fisiología , Árboles/fisiología , Interacciones Microbiota-Huesped/fisiología , Dispersión de las Plantas , Microbiología del Suelo , Árboles/microbiologíaRESUMEN
Flueggea acicularis (Phyllanthaceae) is endemic to the karst region of central China. Male specimens of this species were first collected in 1908. In 1989, female plants of F. acicularis were found for the first time, but misidentified as a new species. Throughout this period the male plants of F. acicularis were mismatched with female plants of other species, and male plants had not been collected since 1908. Then, in March, 2009, the authors rediscovered a wild population of F. acicularis consisting of both male and female plants in Wuxi county, Chongqing municipality, China. Based on field investigation and examination of specimens, we matched the correct female and male plants of this species for the first time since its initial publication a century ago. A complete and accurate morphological description, distribution, habitat and phenology of this species are also provided. Furthermore, the conservation status of F. acicularis is assessed as "Near Threatened" (NT) according to the IUCN Red List criteria.
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Foundation species structure forest communities and ecosystems but are difficult to identify without long-term observations or experiments. We used statistical criteria--outliers from size-frequency distributions and scale-dependent negative effects on alpha diversity and positive effects on beta diversity--to identify candidate foundation woody plant species in 12 large forest-dynamics plots spanning 26 degrees of latitude in China. We used these data (1) to identify candidate foundation species in Chinese forests, (2) to test the hypothesis--based on observations of a midlatitude peak in functional trait diversity and high local species richness but few numerically dominant species in tropical forests--that foundation woody plant species are more frequent in temperate than tropical or boreal forests, and (3) to compare these results with data from the Americas to suggest candidate foundation genera in northern hemisphere forests. Using the most stringent criteria, only two species of Acer, the canopy tree Acer ukurunduense and the shrubby treelet Acer barbinerve, were identified in temperate plots as candidate foundation species. Using more relaxed criteria, we identified four times more candidate foundation species in temperate plots (including species of Acer, Pinus, Juglans, Padus, Tilia, Fraxinus, Prunus, Taxus, Ulmus, and Corlyus) than in (sub)tropical plots (the treelets or shrubs Aporosa yunnanensis, Ficus hispida, Brassaiopsis glomerulata, and Orophea laui). Species diversity of co-occurring woody species was negatively associated with basal area of candidate foundation species more frequently at 5- and 10-m spatial grains (scale) than at a 20-m grain. Conversely, Bray-Curtis dissimilarity was positively associated with basal area of candidate foundation species more frequently at 5-m than at 10- or 20-m grains. Both stringent and relaxed criteria supported the hypothesis that foundation species are more common in mid-latitude temperate forests. Comparisons of candidate foundation species in Chinese and North American forests suggest that Acer be investigated further as a foundation tree genus.
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Biodiversidad , Ecosistema , China , Bosques , ÁrbolesRESUMEN
Investigating the drivers of phenotypic and genetic divergence can reveal the underlying processes and strategies that species adopt in rapidly changing environments. However, knowledge of adaptive strategies and the underlying mechanisms is lacking for the majority of taxa, especially those living in habitats sensitive to climate change. Here, we investigated 20 populations of a Tertiary-relict tree species, Euptelea pleiospermum (Eupteleaceae), scattered in a mountain riparian habitat in China. We integrated genetic, growth, and reproductive traits, and evaluated the relative contributions of climatic and soil factors on genetic and functional trait divergence. The E. pleiospermum populations were divided into south and north genetic clusters, and there were significant differences in leaf density and seed mass of adult trees between the two. The spatial pattern of genetic divergence resulted from effects of both isolation by distance (IBD) and isolation by environment (IBE), whereas the divergence of growth and reproductive traits resulted solely from IBE effects. Spatial distance and selection by temperature and soils played dominant roles in genetic divergence. Precipitation drove the spatial divergence of sprouting. Both divergence of leaf density and seed mass were prominently induced by genetic divergence, and the influences might be enhanced by temperature and soil nutrients. We infer that E. pleiospermum populations adopt a resource-conservative strategy with low growth rates and higher sprouting under flooding disturbance, with larger seeds for improved seedling recruitment at lower latitudes. In contrast, high growth rate and sexual reproduction with small seeds are strategies adopted by populations at higher latitudes. We conclude that sprouting reflects a plastic response to precipitation, and leaf density and seed mass reflect local adaption under selection by temperature and soil factors. The underlying mechanisms of species adaptation strategies were trait-specific. Temperature and soil conditions are likely the main ecological factors shaping plant divergence in montane riparian regions.
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Ecosistema , Árboles , China , Fenotipo , PlantonesRESUMEN
Ex-situ conservation is widely used to protect wild plant species from extinction. However, it remains unclear how genetic variation of ex-situ plant collections reflects wild source population diversity. Here, we conduct a global meta-analysis of the genetic representativeness of ex-situ populations by comparing genetic diversity (i.e. AR, allelic richness; HE , expected heterozygosity; PPB, percentage polymorphic bands; and SWI, Shannon-Winner index), inbreeding coefficient (FIS ), and genetic differentiation between ex-situ plant collections and their wild source populations. Genetic diversity (i.e., HE , PPB, and SWI) was significantly lower in ex-situ populations than their wild source populations, while genetic differentiation between ex-situ and wild populations (ex-situ-wild FST ), but not that among ex-situ populations, was significantly higher than among wild populations. Outcrossing species, but not those with mixed mating system, had significantly lower genetic diversity in ex-situ populations, and significantly higher ex-situ-wild FST . When the collection size for ex-situ conservation was ≥30 or 50, PPB, HE , and ex-situ-wild FST were not significantly different between ex-situ and wild populations, indicating a relatively high genetic representativeness. Collecting from the entire natural distribution range and mixing collections from different sources could significantly increase the genetic representativeness of ex-situ populations. Type of ex-situ conservation (i.e., planting or seed bank) had no effect on genetic representativeness. The effect size of HE decreased and the effect size of ex-situ-wild FST increased with the duration of ex-situ conservation. Our results suggest that current ex-situ plant collections do not effectively capture the genetic variation of wild populations. Low genetic representativeness of ex-situ populations was caused by both initial incomplete sampling from wild populations and genetic erosion during ex-situ conservation. We emphasize that it is necessary to employ more thorough sampling strategies in future collecting efforts and to add new individuals where needed. Article impact statement: Low genetic representativeness of living plant collections is a worldwide problem in ex situ conservation. This article is protected by copyright. All rights reserved.
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Despite several decades of study in community ecology, the relative importance of the ecological processes that determine species co-occurrence across spatial scales remains uncertain. Some of this uncertainty may be reduced by studying the scale dependency of community assembly in the light of environmental variation. Phylogenetic information and functional trait information are often used to provide potentially valuable insights into the drivers of community assembly. Here, we combined phylogenetic and trait-based tests to gain insights into community processes at four spatial scales in a large stem-mapped subtropical forest dynamics plot in central China. We found that all of the six leaf economic traits measured in this study had weak, but significant, phylogenetic signal. Nonrandom phylogenetic and trait-based patterns associated with topographic variables indicate that deterministic processes tend to dominate community assembly in this plot. Specifically, we found that, on average, co-occurring species were more phylogenetically and functionally similar than expected throughout the plot at most spatial scales and assemblages of less similar than expected species could only be found on finer spatial scales. In sum, our results suggest that the trait-based effects on community assembly change with spatial scale in a predictable manner and the association of these patterns with topographic variables, indicates the importance of deterministic processes in community assembly relatively to random processes.
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Vegetation maps are important sources of information for biodiversity conservation, ecological studies, vegetation management and restoration, and national strategic decision making. The current Vegetation Map of China (1:1000000) was generated by a team of more than 250 scientists in an effort that lasted over 20 years starting in the 1980s. However, the vegetation distribution of China has experienced drastic changes during the rapid development of China in the last three decades, and it urgently needs to be updated to better represent the distribution of current vegetation types. Here, we describe the process of updating the Vegetation Map of China (1:1000000) generated in the 1980s using a "crowdsourcing-change detection-classification-expert knowledge" vegetation mapping strategy. A total of 203,024 field samples were collected, and 50 taxonomists were involved in the updating process. The resulting updated map has 12 vegetation type groups, 55 vegetation types/subtypes, and 866 vegetation formation/sub-formation types. The overall accuracy and kappa coefficient of the updated map are 64.8% and 0.52 at the vegetation type group level, 61% and 0.55 at the vegetation type/subtype level and 40% and 0.38 at the vegetation formation/sub-formation level. When compared to the original map, the updated map showed that 3.3 million km2 of vegetated areas of China have changed their vegetation type group during the past three decades due to anthropogenic activities and climatic change. We expect this updated map to benefit the understanding and management of China's terrestrial ecosystems.
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Tropical and subtropical forest ecosystems play an important role in the global carbon regulation. Despite increasing evidence for effects of biodiversity (species diversity, functional diversity and functional dominance), stand structural attributes, stand age and environmental conditions (climate and topography) on tree carbon storage, the relative importance of these drivers at large scale is poorly understood. It is also still unclear whether biodiversity effects on tree carbon storage work through niche complementarity (i.e. increased tree carbon storage due to interspecific resource partitioning) or through the mass-ratio effect (tree carbon storage regulated by dominant traits within communities). Here we analyze tree carbon storage and its drivers using data of 480 plots sampled across subtropical forests in China. We use multiple regression models to test the relative effects of biodiversity, stand structural attributes, stand age and environmental conditions on tree carbon storage, and use a partial least squares path model to test how these variables directly and/or indirectly affect tree carbon storage. Our results show that tree carbon storage is most strongly affected by stand age, followed by climate, biodiversity and stand structural attributes. Stand age and climate had both direct and indirect (through species diversity, functional dominance and stand structural attributes) effects. We find that tree carbon storage correlates with both species diversity and functional dominance after stand age and environmental drivers are accounted for. Our results suggest that niche complementarity and the mass-ratio effect, not necessarily mutually exclusive, both play a role in maintaining ecosystem functioning. Our results further indicate that biodiversity conservation might be an effective way for enhancing tree carbon storage in natural, species-rich forest ecosystems.
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Secuestro de Carbono , Carbono/metabolismo , Bosques , Árboles/metabolismo , Biodiversidad , Biomasa , China , Análisis MultivarianteRESUMEN
Climate is widely recognised as an important determinant of the latitudinal diversity gradient. However, most existing studies make no distinction between direct and indirect effects of climate, which substantially hinders our understanding of how climate constrains biodiversity globally. Using data from 35 large forest plots, we test hypothesised relationships amongst climate, topography, forest structural attributes (stem abundance, tree size variation and stand basal area) and tree species richness to better understand drivers of latitudinal tree diversity patterns. Climate influences tree richness both directly, with more species in warm, moist, aseasonal climates and indirectly, with more species at higher stem abundance. These results imply direct limitation of species diversity by climatic stress and more rapid (co-)evolution and narrower niche partitioning in warm climates. They also support the idea that increased numbers of individuals associated with high primary productivity are partitioned to support a greater number of species.
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Biodiversidad , Árboles , ClimaRESUMEN
The relationship between ß-diversity and latitude still remains to be a core question in ecology because of the lack of consensus between studies. One hypothesis for the lack of consensus between studies is that spatial scale changes the relationship between latitude and ß-diversity. Here, we test this hypothesis using tree data from 15 large-scale forest plots (greater than or equal to 15 ha, diameter at breast height ≥ 1 cm) across a latitudinal gradient (3-30o) in the Asia-Pacific region. We found that the observed ß-diversity decreased with increasing latitude when sampling local tree communities at small spatial scale (grain size ≤0.1 ha), but the observed ß-diversity did not change with latitude when sampling at large spatial scales (greater than or equal to 0.25 ha). Differences in latitudinal ß-diversity gradients across spatial scales were caused by pooled species richness (γ-diversity), which influenced observed ß-diversity values at small spatial scales, but not at large spatial scales. Therefore, spatial scale changes the relationship between ß-diversity, γ-diversity and latitude, and improving sample representativeness avoids the γ-dependence of ß-diversity.
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Niche and neutral theories emphasize different processes contributing to the maintenance of species diversity. In this study, we calculated the local contribution to beta diversity (LCBD) of every cell, using variation partitioning in combination with spatial distance and environmental variables of the 25-ha Badagongshan plot (BDGS), to determine the contribution of environmentally-related variation versus pure spatial variation. We used topography and soil characteristics as environmental variables, distance-based Moran's eigenvectors maps (dbMEM) to describe spatial relationships among cells and redundancy analysis (RDA) to apportion the variation in beta diversity into three components: pure environmental, spatially-structured environmental, and pure spatial. Results showed LCBD values were negatively related to number of common species and positively related to number of rare species. Environment and space jointly explained ~60% of the variation in species composition; soil variables alone explained 21.6%, slightly more than the topographic variables that explained 15.7%; topography and soil together explained 27%, slightly inferior to spatial variables that explained 34%. The BDGS forest was controlled both by the spatial and environmental variables, and the results were consistent across different life forms and life stages.
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Biodiversidad , Conservación de los Recursos Naturales , Árboles/fisiología , China , Bosques , Suelo/química , Especificidad de la Especie , Árboles/clasificación , Clima TropicalRESUMEN
Global change is impacting forests worldwide, threatening biodiversity and ecosystem services including climate regulation. Understanding how forests respond is critical to forest conservation and climate protection. This review describes an international network of 59 long-term forest dynamics research sites (CTFS-ForestGEO) useful for characterizing forest responses to global change. Within very large plots (median size 25 ha), all stems ≥ 1 cm diameter are identified to species, mapped, and regularly recensused according to standardized protocols. CTFS-ForestGEO spans 25 °S-61 °N latitude, is generally representative of the range of bioclimatic, edaphic, and topographic conditions experienced by forests worldwide, and is the only forest monitoring network that applies a standardized protocol to each of the world's major forest biomes. Supplementary standardized measurements at subsets of the sites provide additional information on plants, animals, and ecosystem and environmental variables. CTFS-ForestGEO sites are experiencing multifaceted anthropogenic global change pressures including warming (average 0.61 °C), changes in precipitation (up to ± 30% change), atmospheric deposition of nitrogen and sulfur compounds (up to 3.8 g N m(-2) yr(-1) and 3.1 g S m(-2) yr(-1)), and forest fragmentation in the surrounding landscape (up to 88% reduced tree cover within 5 km). The broad suite of measurements made at CTFS-ForestGEO sites makes it possible to investigate the complex ways in which global change is impacting forest dynamics. Ongoing research across the CTFS-ForestGEO network is yielding insights into how and why the forests are changing, and continued monitoring will provide vital contributions to understanding worldwide forest diversity and dynamics in an era of global change.
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Cambio Climático , Conservación de los Recursos Naturales , Monitoreo del Ambiente , BosquesRESUMEN
The stochastic dilution hypothesis has been proposed to explain species coexistence in species-rich communities. The relative importance of the stochastic dilution effects with respect to other effects such as competition and habitat filtering required to be tested. In this study, using data from a 25-ha species-rich subtropical forest plot with a strong topographic structure at Badagongshan in central China, we analyzed overall species associations and fine-scale species interactions between 2,550 species pairs. The result showed that: (1) the proportion of segregation in overall species association analysis at 2 m neighborhood in this plot followed the prediction of the stochastic dilution hypothesis that segregations should decrease with species richness but that at 10 m neighborhood was higher than the prediction. (2) The proportion of no association type was lower than the expectation of stochastic dilution hypothesis. (3) Fine-scale species interaction analyses using Heterogeneous Poisson processes as null models revealed a high proportion (47%) of significant species effects. However, the assumption of separation of scale of this method was not fully met in this plot with a strong fine-scale topographic structure. We also found that for species within the same families, fine-scale positive species interactions occurred more frequently and negative ones occurred less frequently than expected by chance. These results suggested effects of environmental filtering other than species interaction in this forest. (4) We also found that arbor species showed a much higher proportion of significant fine-scale species interactions (66%) than shrub species (18%). We concluded that the stochastic dilution hypothesis only be partly supported and environmental filtering left discernible spatial signals in the spatial associations between species in this species-rich subtropical forest with a strong topographic structure.
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Distribución Animal , Biodiversidad , Biota , Bosques , Modelos Biológicos , China , Especificidad de la Especie , Procesos EstocásticosRESUMEN
We used landscape genetics and statistical models to test how landscape features influence connectivity or create barriers to dispersal for a mountain riparian tree species, Euptelea pleiospermum. Young leaves from 1078 individuals belonging to 36 populations at elevations of 900-2000 m along upper reaches of four rivers were genotyped using eight nuclear microsatellite markers. We found no evidence for the unidirectional dispersal hypothesis in E. pleiospermum within each river. The linear dispersal pattern along each river valley is mostly consistent with the "classical metapopulaton" model. Mountain ridges separating rivers were genetic barriers for this wind-pollinated tree species with anemochorous seeds, whereas river valleys provided important corridors for dispersal. Gene flow among populations along elevational gradients within each river prevails over gene flow among populations at similar elevations but from different rivers. This pattern of gene flow is likely to promote elevational range shifts of plant populations and to hinder local adaptation along elevational gradients. This study provides a paradigm to determine which of the two strategies (migration or adaptation) will be adopted by mountain riparian plants under climate warming.
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Altitud , Ríos , Tracheophyta/genética , Árboles/genética , Desequilibrio de Ligamiento , Repeticiones de Microsatélite/genética , Modelos Genéticos , Polimorfismo GenéticoRESUMEN
⢠This study aimed to reveal species-genetic diversity correlations (SGDCs) and their underlying mechanisms in natural and disturbed forests. ⢠A community survey and molecular analyses were carried out to compare species diversity (SD), the genetic diversity of the dominant tree species Euptelea pleiospermum (GD), the altitudinal patterns of SD and GD, SGDC, genetic differentiation (F(ST) ), community divergence (F(ST) -C), effective population size (N(e) ), and recent migration rate between mountain riparian forests along the Yandu (natural) and Nan (disturbed) rivers. ⢠In natural forests, both SD and GD showed a unimodal altitudinal pattern and GD was positively correlated with SD, whereas a unimodal pattern and positive SGDC were not found in the disturbed forests. SD and F(ST) at the natural sites were higher than those at the disturbed sites. However, there were no significant differences in GD, F(ST) -C, N(e) or recent migration rate between the natural and disturbed sites. ⢠A correlation between the patterns of SD and GD along a geographical gradient (e.g. altitude) is an important driver of positive SGDC. The absence of positive SGDC in the disturbed forests may result from reduced SD but unaffected GD, indicating nonparallel changes in SD and GD. This study furthermore cautions against generalizations about changes in SD and GD following disturbance.