Stochastic geometry best explains spatial associations among species pairs and plant functional types in species‐rich shrublands

Some conceptual models seeking to explain the coexistence of multiple species in hyperdiverse settings predict that species will not be randomly distributed with respect to each other. In stark contrast the ‘stochastic geometry’ model assumes that a species fine‐scale spatial distribution is independent of that of other species in the community. Empirical tests in temperate and tropical forests have provided support for both perspectives. Using point pattern analyses we assessed the prevalence of heterospecific associations between > 10 500 pairs of species and > 3400 pairs of plant functional types (PFTs) in four biodiverse shrubland communities in southwestern Australia. After controlling for first‐order effects, spatial associations between species and PFTs were rare, but were most prevalent at the least species‐rich of the four sites considered. Individuals tended to have fewer species in their local neighbourhoods than expected under a null model of random relabelling, with this departure most pronounced at the site with fewest species. The consistency of neighbourhood composition experienced by individuals of the same species is, as a result, less than the average under random mixing. Our results demonstrate that the frequency of heterospecific spatial associations is both rare in speciose systems and declines with species richness, and provide further empirical support for the stochastic geometry assumption in species‐rich communities.     

Anthropogenic modification disrupts species co‐occurrence in stream invertebrates

The question of whether species co‐occurrence is random or deterministic has received considerable attention, but little is known about how anthropogenic disturbance mediates the outcomes. By combining experiments, field surveys and analysis against null models, we tested the hypothesis that anthropogenic habitat modification disrupts species co‐occurrence in stream invertebrates across spatial scales. Whereas communities in unmodified conditions were structured deterministically with significant species segregation, catchment‐scale conversion to agriculture and sediment deposition at the patch‐ or micro‐habitat scale apparently randomized species co‐occurrences. This shift from non‐random to random was mostly independent of species richness, abundance and spatial scale. Data on community‐wide life‐history traits (body size, dispersal ability and predatory habits) and beta‐diversity indicated that anthropogenic modification disrupted community assembly by affecting biotic interactions and, to a lesser extent, altering habitat heterogeneity. These data illustrate that the balance between predictable and stochastic patterns in communities can reflect anthropogenic modifications that not only transcend scales but also change the relative forces that determine species coexistence. Research into the effects of habitat modification as a key to understanding global change should extend beyond species richness and composition to include species co‐occurrence, species interactions and any functional consequences.     

Individual Species-Area Relationship of Woody Plant Communities in a Heterogeneous Subtropical Monsoon Rainforest

The spatial structure of species richness is often characterized by the species-area relationship (SAR). However, the SAR approach rarely considers the spatial variability of individual plants that arises from species interactions and species’ habitat associations. Here, we explored how the interactions of individual plants of target species influence SAR patterns at a range of neighborhood distances. We analyzed the data of 113,988 woody plants of 110 species from the Fushan Forest Dynamics Plot (25 ha), northern Taiwan, which is a subtropical rainforest heavily influenced by typhoons. We classified 34 dominant species into 3 species types (i.e., accumulator, repeller, or no effect) by testing how the individual species-area relationship (i.e., statistics describing how neighborhood species richness changes around individuals) of target species departs (i.e., positively, negatively, or with no obvious trend) from a null model that accounts for habitat association. Deviation from the null model suggests that the net effect of species’ interactions increases (accumulate) or decreases (repel) neighborhood species richness. We found that (i) accumulators were dominant at small interaction distances (<10–30 m); (ii) the detection of accumulator species was lower at large interaction distances (>30 m); (iii) repellers were rarely detected; and (iv) large-sized and abundant species tended to be accumulators. The findings suggest that positive species interactions have the potential to accumulate neighborhood species richness, particularly through size- and density-dependent mechanisms. We hypothesized that the frequently disturbed environment of this subtropical rainforest (e.g., typhoon-driven natural disturbances such as landslides, soil erosion, flooding, and windthrow) might create the spatial heterogeneity of species richness and promote positive species interactions.     

Vertical stratification influences global patterns of biodiversity

Species distributions in terrestrial ecosystems are three‐dimensional, spanning both the horizontal landscape and the vertical space provided by the physical environment. Classical hypotheses suggest that communities become more vertically stratified with increasing species richness, owing to reduced competition or finer niche subdivision. However, this assertion remains untested in the context of the broader realm of biogeography. Here, integrating traits and distribution data for amphibians globally, we show how vertical strategies interact with the physical and climatic environments to govern global patterns of species richness and community composition. Our results reveal a marked latitudinal shift in strategies of vertical habitat use, from highly arboreal assemblages in the tropics to highly fossorial assemblages in sub‐tropical and temperate regions. Arboreality is strongly associated with precipitation, vegetation structure and climatic stability, whereas fossoriality is more common in dry environments with high diurnal temperature range and low vegetation structure. These analyses shed light on the importance of vertical stratification for species coexistence in species‐rich regions. As certain tropical habitats become drier from climate change, the rich biological diversity that is emblematic of the tropics may transition from vertically stratified to ‘flattened’, with future communities living mostly on or beneath the ground.     

Traits related to species persistence and dispersal explain changes in plant communities subjected to habitat loss

Aim Habitat fragmentation is a major driver of biodiversity loss but it is insufficiently known how much its effects vary among species with different life‐history traits; especially in plant communities, the understanding of the role of traits related to species persistence and dispersal in determining dynamics of species communities in fragmented landscapes is still limited. The primary aim of this study was to test how plant traits related to persistence and dispersal and their interactions modify plant species vulnerability to decreasing habitat area and increasing isolation. Location Five regions distributed over four countries in Central and Northern Europe. Methods Our dataset was composed of primary data from studies on the distribution of plant communities in 300 grassland fragments in five regions. The regional datasets were consolidated by standardizing nomenclature and species life‐history traits and by recalculating standardized landscape measures from the original geographical data. We assessed the responses of plant species richness to habitat area, connectivity, plant life‐history traits and their interactions using linear mixed models. Results We found that the negative effect of habitat loss on plant species richness was pervasive across different regions, whereas the effect of habitat isolation on species richness was not evident. This area effect was, however, not equal for all the species, and life‐history traits related to both species persistence and dispersal modified plant sensitivity to habitat loss, indicating that both landscape and local processes determined large‐scale dynamics of plant communities. High competitive ability for light, annual life cycle and animal dispersal emerged as traits enabling species to cope with habitat loss. Main conclusions In highly fragmented rural landscapes in NW Europe, mitigating the spatial isolation of remaining grasslands should be accompanied by restoration measures aimed at improving habitat quality for low competitors, abiotically dispersed and perennial, clonal species.     

Common spatial patterns of trees in various tropical forests: Small trees are associated with increased diversity at small spatial scales

Tropical forests are notable for their high species diversity, even on small spatial scales, and right‐skewed species and size abundance distributions. The role of individual species as drivers of the spatial organization of diversity in these forests has been explained by several hypotheses and processes, for example, stochastic dilution, negative density dependence, or gap dynamics. These processes leave a signature in spatial distribution of small trees, particularly in the vicinity of large trees, likely having stronger effects on their neighbors. We are exploring species diversity patterns within the framework of various diversity‐generating hypotheses using individual species–area relationships. We used the data from three tropical forest plots (Wanang—Papua New Guinea, Barro Colorado Island—Panama, and Sinharaja—Sri Lanka) and included also the saplings (DBH ≥ 1 cm). Resulting cross‐size patterns of species richness and evenness reflect the dynamics of saplings affected by the distribution of large trees. When all individuals with DBH ≥1 cm are included, ~50% of all tree species from the 25‐ or 50‐ha plot can be found within 35 m radius of an individual tree. For all trees, 72%–78% of species were identified as species richness accumulators, having more species present in their surroundings than expected by null models. This pattern was driven by small trees as the analysis of DBH >10 cm trees showed much lower proportion of accumulators, 14%–65% of species identified as richness repellers and had low richness of surrounding small trees. Only 11%–26% of species had lower species evenness than was expected by null models. High proportions of species richness accumulators were probably due to gap dynamics and support Janzen–Connell hypothesis driven by competition or top‐down control by pathogens and herbivores. Observed species diversity patterns show the importance of including small tree size classes in analyses of the spatial organization of diversity.     

How life‐history traits affect ecosystem properties: effects of dispersal in meta‐ecosystems

The concept of life‐history traits and the study of these traits are the hallmark of population biology. Acknowledging their variability and evolution has allowed us to understand how species adapt in response to their environment. The same traits are also involved in how species alter ecosystems and shape their dynamics and functioning. Some theories, such as the metabolic theory of ecology, ecological stoichiometry or pace‐of‐life theory, already recognize this junction, but only do so in an implicitly non‐spatial context. Meanwhile, for a decade now, it has been argued that ecosystem properties have to be understood at a larger scale using meta‐ecosystem theory because source–sink dynamics, community assembly and ecosystem stability are all modified by spatial structure. Here, we argue that some ecosystem properties can be linked to a single life‐history trait, dispersal, i.e. the tendency of organisms to live, compete and reproduce away from their birth place. By articulating recent theoretical and empirical studies linking ecosystem functioning and dynamics to species dispersal, we aim to highlight both the known connections between life‐history traits and ecosystem properties and the unknown areas, which deserve further empirical and theoretical developments.     

Individual species–area relationships and spatial patterns of species diversity in a Great Basin,semi‐arid shrubland

Traditional biodiversity metrics operate at the level of a plant community but do not capture spatial variation in diversity from a ‘plant's‐eye view’ of a community. Recently‐developed statistics consider the spatial patterns of plants as well as the number and distribution of species in local plant neighborhoods to quantitatively assess multispecies spatial patterns from a ‘plant's‐eye view’. We used one such statistic, the individual species area relationship (ISAR), to assess spatial patterns of species diversity in a Great Basin (USA) semi‐arid shrubland through an analysis of a spatial dataset on shrub species and locations. In conjunction with appropriate null models, the ISAR blends species area relationships with second‐order spatial statistics to measure the expected species richness in local neighborhoods of variable size around the individuals of a focal species within a community. We found that, contrary to a previous analysis using more traditional methods, the community was well‐mixed with a typical shrub surrounded on average by 4.9 shrub neighbors of 2.1 species at a neighborhood scale of 1.0 m. We also found statistically significant fine‐scale variation in diversity patterns, such that neighborhoods of two species were more diverse than expected by a heterogeneous Poisson null model that accounted for larger‐scale habitat heterogeneity. However, this effect was caused by intraspecific aggregation of these species and was not due to positive interspecific association. Contrary to previous findings in other semi‐arid shrublands, our analysis suggests that the spatial pattern of the shrub community was not significantly structured by interspecific facilitation. This result supports growing evidence for balanced species patterns of adult plants in multispecies communities. Our approach may be used in other communities to describe complex multispecies spatial patterns, quantify species‐specific associations with diversity patterns, and to generate hypotheses regarding relationships between patterns and community‐structuring processes.     

Putting insects on the map: near‐global variation in sphingid moth richness along spatial and environmental gradients

Despite their vast diversity and vital ecological role, insects are notoriously underrepresented in biogeography and conservation, and key broad‐scale ecological hypotheses about them remain untested – largely due to generally incomplete and very coarse spatial distribution knowledge. Integrating records from publications, field work and natural history collections, we used a mixture of species distribution models and expert estimates to provide geographic distributions and emergent richness patterns for all ca 1000 sphingid moth species found outside the Americas in high spatial detail. Total sphingid moth richness, the first for a higher insect group to be documented at this scale, shows distinct maxima in the wet tropics of Africa and the Oriental with notable decay toward Australasia. Using multivariate models controlling for spatial autocorrelation, we found that primary productivity is the dominant environmental variable associated with moth richness, while temperature, contrary to our predictions, is an unexpectedly weak predictor. This is in stark contrast to the importance we identify for temperature as a niche variable of individual species. Despite divergent life histories, both main sub‐groups of moths exhibit these relationships. Tribal‐level deconstruction of richness and climatic niche patterns indicate idiosyncratic effects of biogeographic history for some of the less species‐rich tribes, which in some cases exhibit distinct richness peaks away from the tropics. The study confirms, for a diverse insect group, overall richness associations of remarkable similarity to those documented for vertebrates and highlights the significant within‐taxon structure that underpins emergent macroecological patterns. Results do not, however, meet predictions from vertebrate‐derived hypotheses on how thermoregulation affects the strength of temperature–richness effects. Our study thus broadens the taxonomic focus in this data‐deficient discourse. Our procedures of processing incomplete, scattered distribution data are a template for application to other taxa and regions.     

Ecological filtering of exotic plants in an Australian sub‐alpine environment

We investigated some of the factors influencing exotic invasion of native sub‐alpine plant communities at a site in southeast Australia. Structure, floristic composition and invasibility of the plant communities and attributes of the invasive species were studied. To determine the plant characteristics correlated with invasiveness, we distinguished between roadside invaders, native community invaders and non‐invasive exotic species, and compared these groups across a range of traits including functional group, taxonomic affinity, life history, mating system and morphology. Poa grasslands and Eucalyptus‐Poa woodlands contained the largest number of exotic species, although all communities studied appeared resilient to invasion by most species. Most community invaders were broad‐leaved herbs while roadside invaders contained both herbs and a range of grass species. Over the entire study area the richness and cover of native and exotic herbaceous species were positively related, but exotic herbs were more negatively related to cover of specific functional groups (e.g. trees) than native herbs. Compared with the overall pool of exotic species, those capable of invading native plant communities were disproportionately polycarpic, Asteracean and cross‐pollinating. Our data support the hypothesis that strong ecological filtering of exotic species generates an exotic assemblage containing few dominant species and which functionally converges on the native assemblage. These findings contrast with those observed in the majority of invaded natural systems. We conclude that the invasion of closed sub‐alpine communities must be viewed in terms of the unique attributes of the invading species, the structure and composition of the invaded communities and the strong extrinsic physical and climatic factors typical of the sub‐alpine environment.     

The coincidence of rarity and richness and the potential signature of history in centres of endemism

We investigate the relative importance of stochastic and environmental/topographic effects on the occurrence of avian centres of endemism, evaluating their potential historical importance for broad‐scale patterns in species richness across Sub‐Saharan Africa. Because species‐rich areas are more likely to be centres of endemism by chance alone, we test two null models: Model 1 calculates expected patterns of endemism using a random draw from the occurrence records of the continental assemblage, whereas Model 2 additionally implements the potential role of geometric constraints. Since Model 1 yields better quantitative predictions we use it to identify centres of endemism controlled for richness. Altitudinal range and low seasonality emerge as core environmental predictors for these areas, which contain unusually high species richness compared to other parts of sub‐Saharan Africa, even when controlled for environmental differences. This result supports the idea that centres of endemism may represent areas of special evolutionary history, probably as centres of diversification.     

Landscape‐scale determinants of non‐native fish communities

Aim Predicting and preventing invasions depends on knowledge of the factors that make ecosystems susceptible to invasion. Current studies generally rely on non‐native species richness (NNSR) as the sole measure of ecosystem invasibility; however, species identity is a critical consideration, given that different ecosystems may have environmental characteristics suitable to different species. Our aim was to examine whether non‐native freshwater fish community composition was related to ecosystem characteristics at the landscape scale. Location United States. Methods We described spatial patterns in non‐native freshwater fish communities among watersheds in the Mid‐Atlantic region of the United States based on records of establishment in the U.S. Geological Survey’s Nonindigenous Aquatic Species Database. We described general relationships between non‐native species and ecosystem characteristics using canonical correspondence analysis. We clustered watersheds by non‐native fish community and described differences among clusters using indicator species analysis. We then assessed whether non‐native communities could be predicted from ecosystem characteristics using random forest analysis and predicted non‐native communities for uninvaded watersheds. We estimated which ecosystem characteristics were most important for predicting non‐native communities using conditional inference trees. Results We identified four non‐native fish communities, each with distinct indicator species. Non‐native communities were predicted based on ecosystem characteristics with an accuracy of 80.6%, with temperature as the most important variable. Relatively uninvaded watersheds were predicted to be invasible by the most diverse non‐native community. Main conclusions Non‐native species identity is an important consideration when assessing ecosystem invasibility. NNSR alone is an insufficient measure of invasibility because ecosystems with equal NNSR may not be equally invasible by the same species. Our findings can help improve predictions of future invasions and focus management and policy decisions on particular species in highly invasible ecosystems.     

Environmental heterogeneity among lakes promotes hyper β‐diversity across phytoplankton communities

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Testing a ‘genes‐to‐ecosystems’ approach to understanding aquatic–terrestrial linkages

A ‘genes‐to‐ecosystems’ approach has been proposed as a novel avenue for integrating the consequences of intraspecific genetic variation with the underlying genetic architecture of a species to shed light on the relationships among hierarchies of ecological organization (genes → individuals → communities → ecosystems). However, attempts to identify genes with major effect on the structure of communities and/or ecosystem processes have been limited and a comprehensive test of this approach has yet to emerge. Here, we present an interdisciplinary field study that integrated a common garden containing different genotypes of a dominant, riparian tree, Populus trichocarpa, and aquatic mesocosms to determine how intraspecific variation in leaf litter alters both terrestrial and aquatic communities and ecosystem functioning. Moreover, we incorporate data from extensive trait screening and genome‐wide association studies estimating the heritability and genes associated with litter characteristics. We found that tree genotypes varied considerably in the quality and production of leaf litter, which contributed to variation in phytoplankton abundances, as well as nutrient dynamics and light availability in aquatic mesocosms. These ‘after‐life’ effects of litter from different genotypes were comparable to the responses of terrestrial communities associated with the living foliage. We found that multiple litter traits corresponding with aquatic community and ecosystem responses differed in their heritability. Moreover, the underlying genetic architecture of these traits was complex, and many genes contributed only a small proportion to phenotypic variation. Our results provide further evidence that genetic variation is a key component of aquatic–terrestrial linkages, but challenge the ability to predict community or ecosystem responses based on the actions of one or a few genes.     

Shifts in trait means and variances in North American tree assemblages: species richness patterns are loosely related to the functional space

One of the key hypothesized drivers of gradients in species richness is environmental filtering, where environmental stress limits which species from a larger species pool gain membership in a local community owing to their traits. Whereas most studies focus on small‐scale variation in functional traits along environmental gradient, the effect of large‐scale environmental filtering is less well understood. Furthermore, it has been rarely tested whether the factors that constrain the niche space limit the total number of coexisting species. We assessed the role of environmental filtering in shaping tree assemblages across North America north of Mexico by testing the hypothesis that colder, drier, or seasonal environments (stressful conditions for most plants) constrain tree trait diversity and thereby limit species richness. We assessed geographic patterns in trait filtering and their relationships to species richness pattern using a comprehensive set of tree range maps. We focused on four key plant functional traits reflecting major life history axes (maximum height, specific leaf area, seed mass, and wood density) and four climatic variables (annual mean and seasonality of temperature and precipitation). We tested for significant spatial shifts in trait means and variances using a null model approach. While we found significant shifts in mean species’ trait values at most grid cells, trait variances at most grid cells did not deviate from the null expectation. Measures of environmental harshness (cold, dry, seasonal climates) and lower species richness were weakly associated with a reduction in variance of seed mass and specific leaf area. The pattern in variance of height and wood density was, however, opposite. These findings do not support the hypothesis that more stressful conditions universally limit species and trait diversity in North America. Environmental filtering does, however, structure assemblage composition, by selecting for certain optimum trait values under a given set of conditions.     

It is about time: genetic variation in the timing of leaf‐litter inputs influences aquatic ecosystems

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Community assembly along a successional gradient in sub‐alpine meadows of the Qinghai‐Tibetan Plateau,China

Community assembly is a dynamic progression that reflects the interaction of several processes functioning at multiple scales. Understanding how these processes work in communities at different successional stages is important for identifying when regional or local processes are more important for community assembly, and for developing effective preservation and restoration strategies. We examined community assembly using a chronosequence of sub‐alpine meadows in Qinghai‐Tibetan Plateau that range from ‘natural’ (never farmed), to those that have been protected from agricultural exploitation for 1 to 10 years. We tested for shifts in species and traits among meadows and also for changes in environmental and spatial correlates of species distributions within meadows. We found that species richness increased and species composition returned to natural conditions within ten years of protection. These changes coincided with shifts in species traits; abundant species had high seed mass and specific leaf area in late‐successional meadows, whereas the opposite occurred in early‐successional meadows. Despite these shifts among meadows of different ages, spatial distributions of species within meadows did not change – when associated with abiotic variables, these spatial patterns reflected changes in soil pH and nitrogen. There was also no consistent change in the relative importance of environmental and spatial correlates of species distributions within meadows. These trends indicate that local processes of community assembly are similar within meadows even when species in those meadows differ. We conclude that successional change is a large‐scale process that alters the species pool and resulting suite of traits that are present within meadows. As a result, regional planning that incorporates successional age should be the focus for the conservation of diversity in this area. In contrast, local processes work within the constraints of the species pool set by successional age, producing consistent patterns within meadows of different ages.     

Climate‐based empirical models show biased predictions of butterfly communities along environmental gradients

A better understanding of the factors that mould ecological community structure is required to accurately predict community composition and to anticipate threats to ecosystems due to global changes. We tested how well stacked climate‐based species distribution models (S‐SDMs) could predict butterfly communities in a mountain region. It has been suggested that climate is the main force driving butterfly distribution and community structure in mountain environments, and that, as a consequence, climate‐based S‐SDMs should yield unbiased predictions. In contrast to this expectation, at lower altitudes, climate‐based S‐SDMs overpredicted butterfly species richness at sites with low plant species richness and underpredicted species richness at sites with high plant species richness. According to two indices of composition accuracy, the Sorensen index and a matching coefficient considering both absences and presences, S‐SDMs were more accurate in plant‐rich grasslands. Butterflies display strong and often specialised trophic interactions with plants. At lower altitudes, where land use is more intense, considering climate alone without accounting for land use influences on grassland plant richness leads to erroneous predictions of butterfly presences and absences. In contrast, at higher altitudes, where climate is the main force filtering communities, there were fewer differences between observed and predicted butterfly richness. At high altitudes, even if stochastic processes decrease the accuracy of predictions of presence, climate‐based S‐SDMs are able to better filter out butterfly species that are unable to cope with severe climatic conditions, providing more accurate predictions of absences. Our results suggest that predictions should account for plants in disturbed habitats at lower altitudes but that stochastic processes and heterogeneity at high altitudes may limit prediction success of climate‐based S‐SDMs.     

Host functional and phylogenetic composition rather than host diversity structure plant–herbivore networks

Declining plant diversity alters ecological networks, such as plant–herbivore interactions. However, our knowledge of the potential mechanisms underlying effects of plant species loss on plant–herbivore network structure is still limited. We used DNA barcoding to identify herbivore–host plant associations along declining levels of tree diversity in a large‐scale, subtropical biodiversity experiment. We tested for effects of tree species richness, host functional and phylogenetic diversity, and host functional (leaf trait) and phylogenetic composition on species, phylogenetic and network composition of herbivore communities. We found that phylogenetic host composition and related palatability/defence traits but not tree species richness significantly affected herbivore communities and interaction network complexity at both the species and community levels. Our study indicates that evolutionary dependencies and functional traits of host plants determine the composition of higher trophic levels and corresponding interaction networks in species‐rich ecosystems. Our findings highlight that characteristics of the species lost have effects on ecosystem structure and functioning across trophic levels that cannot be predicted from mere reductions in species richness.     

长白山暗针叶林群落物种多样性维持机制

植物群落物种多样性维持机制一直是生态学研究的热点话题,其中生态位理论和中性理论是被普遍接受的两种理论观点,但是目前关于生态位理论和中性理论在群落物种多样性维持中的相对重要性还没有统一定论。基于长白山暗针叶林群落数据,采用单物种-面积关系模型探究特定树种对邻域物种丰富度的影响,并借助同质性和异质性泊松零模型检验其显著性。(1)群落水平上,在3—15 m空间尺度上,促进种占据优势地位,在>15 m空间尺度上,中性种逐渐取代促进种起主导作用,抑制种比例较低,并且随着空间尺度变化幅度不大。(2)物种水平上,采用同质性泊松零模型检验树种对邻域物种丰富度的影响,臭冷杉、花楷槭、青楷槭在0—20 m空间尺度上对邻域物种丰富度增加起促进作用,黄花落叶松、鱼鳞云杉在0—20 m空间尺度上抑制了邻域物种丰富度增加。花楸树、黑桦和硕桦在全部研究尺度上表现为中性种,髭脉槭、大青杨、红松等在不同研究尺度上表现为不同的作用效果。剔除了生境过滤作用的异质性泊松零模型检验结果与同质性泊松零模型结果差异不显著,表明研究样地内生境过滤作用对多样性格局形成影响不大,各树种间的相互作用对群落物种组成影响较大,进一步证明...