Do assembly history effects attenuate from species to ecosystem properties? A field test with wood‐inhabiting fungi

Assembly history, or the order of species arrival, can have wide-ranging effects on species, communities and ecosystems. However, it remains unclear whether assembly history primarily affects individual species, with effects attenuating at the level of communities and ecosystems or, alternatively, has consistent effect sizes across increasing levels of ecological organisation. We address this question using a field-based manipulation of assembly history of wood-inhabiting fungi. The largest effect sizes were observed for the frequency of some individual species, and mean effect sizes were lower for community metrics of fungi immigrating from the regional species pool. There was little evidence, however, of attenuation in effect sizes at the ecosystem level (carbon, nitrogen, decomposition) in comparison to the species or community level. These results indicate that assembly history can have strong effects on ecosystem properties even under natural levels of environmental variability.     

Environmental parasitology: stressor effects on aquatic parasites

Anthropogenic stressors are causing fundamental changes in aquatic habitats and to the organisms inhabiting these ecosystems. Yet, we are still far from understanding the diverse responses of parasites and their hosts to these environmental stressors and predicting how these stressors will affect host–parasite communities. Here, we provide an overview of the impacts of major stressors affecting aquatic ecosystems in the Anthropocene (habitat alteration, global warming, and pollution) and highlight their consequences for aquatic parasites at multiple levels of organisation, from the individual to the community level. We provide directions and ideas for future research to better understand responses to stressors in aquatic host–parasite systems.     

Environmental and spatial filters of zooplankton metacommunities in shallow pools in high‐elevation peatlands in the tropical Andes

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The consequences of size dependent foraging for food web topology

A general question in biology is how processes at one scale, for example that of individual organisms, influence patterns at larger scales, for example communities of interacting individuals. Here we ask how changing the size‐dependence of the foraging behaviour of individuals can influence the structure of food webs. We assembled communities using a model in which species interactions are determined by allometric foraging rules of (1) handling time and (2) attack rates, and also (3) the distribution of body sizes. We systematically varied these three factors and examined their effects on three community level, food web allometries: the generality ‐ mass correlation, the vulnerability ‐ mass correlation and the trophic height ‐ mass correlation. The results demonstrate how allometries of individual foraging behaviour (handling time and attack rates) are linked across scales of organisation: different community level allometries are influenced by different individual level allometries. For example, generality allometries in the community are most affected by the individual allometric relationships of the attack rate, whereas trophic level allometries in the community are more strongly influenced by variation in individual handling time allometries. Importantly, we also find that the shape of the body size distribution from which species are drawn has a substantial influence on how these links between scales operate. This study suggests that understanding the variation of size structure among ecological networks requires knowledge about the causes of variation in individual foraging behaviour and determinants of the regional body size distribution.     

Simultaneous exposure to a pulsed and a prolonged anthropogenic stressor can alter consumer multifunctionality

Ecosystems face multiple anthropogenic threats globally, and the effects of these environmental stressors range from individual‐level organismal responses to altered system functioning. Understanding the combined effects of stressors on process rates mediated by individuals in ecosystems would greatly improve our ability to predict organismal multifunctionality (e.g. multiple consumer‐mediated functions). We conducted a laboratory experiment to test direct and indirect, as well as immediate and delayed effects of a heat wave (pulsed stress) and micropollutants (MPs) (prolonged stress) on individual consumers (the great pond snail Lymnaea stagnalis) and their multifunctionality (i.e. consumption of basal resources, growth, reproduction, nutrient excretion and organic‐matter cycling). We found that stressful conditions increased the process rates of multiple functions mediated by individual consumers. Specifically, the artificial heat wave increased process rates in the majority of the quantified functions (either directly or indirectly), whereas exposure to MPs increased consumption of basal resources which led to increases in the release of nutrients and fine particulate organic matter. Moreover, snails exposed to a heat wave showed decreased reproduction and nutrient excretion after the heat‐wave, indicating the potential for ecologically relevant delayed effects. Our study indicates that the immediate and delayed effects of stressors on individual organisms may directly and indirectly impact multiple ecosystem functions. In particular, delayed effects of environmental stress on individual consumers may cumulatively impede recovery due to decreased functioning following a perturbation. Reconciling these results with studies incorporating responses at higher levels of biological complexity will enhance our ability to forecast how individual responses upscale to ecosystem multifunctionality.     

Identifying “vital attributes” for assessing disturbance–recovery potential of seafloor communities

Despite a long history of disturbance–recovery research, we still lack a generalizable understanding of the attributes that drive community recovery potential in seafloor ecosystems. Marine soft‐sediment ecosystems encompass a range of heterogeneity from simple low‐diversity habitats with limited biogenic structure, to species‐rich systems with complex biogenic habitat structure. These differences in biological heterogeneity are a product of natural conditions and disturbance regimes. To search for unifying attributes, we explore whether a set of simple traits can characterize community disturbance–recovery potential using seafloor patch‐disturbance experiments conducted in two different soft‐sediment landscapes. The two landscapes represent two ends of a spectrum of landscape biotic heterogeneity in order to consider multi‐scale disturbance–recovery processes. We consider traits at different levels of biological organization, from the biological traits of individual species, to the traits of species at the landscape scale associated with their occurrence across the landscape and their ability to be dominant. We show that in a biotically heterogeneous landscape (Kawau Bay, New Zealand), seafloor community recovery is stochastic, there is high species turnover, and the landscape‐scale traits are good predictors of recovery. In contrast, in a biotically homogeneous landscape (Baltic Sea), the options for recovery are constrained, the recovery pathway is thus more deterministic and the scale of recovery traits important for determining recovery switches to the individual species biological traits within the disturbed patch. Our results imply that these simple, yet sophisticated, traits can be effectively used to characterize community recovery potential and highlight the role of landscapes in providing resilience to patch‐scale disturbances.     

Recovery dynamics of benthic marine communities from the Lower Triassic Werfen Formation,northern Italy

The Lower Triassic Werfen Formation of northern Italy represents an important archive for Early Triassic ecosystems. Based on quantitative community analysis using species level identifications, we reconstruct the recovery of benthic ecosystems after the end‐Permian mass extinction throughout this unit. The analysis of benthic macrofossil communities shows that incipient recovery has taken place during the Griesbachian and the Griesbachian–Dienerian transition. A probable deterioration in environmental conditions is observed towards the end of the Dienerian. The Smithian part of the Werfen Formation is characterized by high siliciclastic input, which ceases around the Smithian‐Spathian boundary. The Spathian marks the definitive phase of recovery in the Werfen Formation. The comparison of this pattern with other palaoegeographical regions suggests that both the Griesbachian recovery and the Dienerian decline were of inter‐regional if not global extent, whereas the Smithian diversity low in the Werfen Formation is a local signal. In contrast to the recovery dynamics of ammonoids and conodonts, the Smithian–Spathian boundary interval was no caesura for benthic ecosystems. The Spathian recovery pulse is possibly also an inter‐regional event, at least in the palaeotropics. These results are in contrast with the previously proposed scenario of persistent hostile conditions during the Griesbachian time interval and highlight an initial recovery phase restricted to Griesbachian times. Instead, the apparently sluggish recovery of benthic ecosystems was at least partly shaped by set‐backs due to short‐term environmental perturbations during the Dienerian.     

植物性状研究的机遇与挑战:从器官到群落

植物性状(Plant trait)或植物功能性状(Plant functional trait)通常是指植物对外界环境长期适应与进化后所表现出的可量度、且与生产力优化或环境适应等密切相关的属性。近几十年来,植物性状研究在性状-生产力、性状-养分、性状间相互关系、性状-群落结构维持等方面取得了卓越成就。然而,由于大多数性状调查都是以植物群落内优势种或亚优势种为对象,使其在探讨群落尺度的性状-功能关系、性状数据如何用于改进或优化模型、性状数据如何与遥感连接等问题时,存在空间尺度和量纲不匹配的极大挑战。为了破解上述难题,亟需发展新的、基于单位土地面积的群落性状(Community trait)概念体系、数据源和计算方法等,推动植物性状数据与快速发展的宏观生态学新技术(遥感、模型和通量观测等)相结合,既拓展了植物性状研究范畴,又可推动其更好地服务于区域生态环境问题的解决。所定义的群落性状(如叶片氮含量、磷含量、比叶面积、气孔密度、叶绿素含量等),是在充分考虑群落内所有物种的性状实测数据,再结合比叶面积、生物量异速生长方程和群落结构数据等,推导而成的基于单位土地面积的群落性状。受测试方法的影响,传统的直接算术平均法或相对生物量加权平均法所获得的群落水平的植物性状(如叶片氮含量g/kg或%),虽然可以有效地探讨群落结构维持机制,由于无法实现对群落性状在量纲上向单位土地面积转换,使它很难与模型和遥感数据相匹配。基于单位土地面积的群落性状,可在空间尺度匹配(或量纲匹配)的前提下实现个体水平测定的植物性状数据与生态模型和遥感观测相联系,更好地探讨区域尺度下自然生态系统结构和功能的关系及其对全球变化的响应与适应。同时,它也可更好地建立群落水平的性状-功能的定量关系(非物种水平),为更好地探讨自然群落结构维持机制和生产力优化机制提供了新思路。     

Towards understanding the organisation of metacommunities in highly dynamic ecological systems

Community ecology recognises today that local biological communities are not only affected by local biotic interactions and abiotic environmental conditions, but also by regional processes (e.g. dispersal). While much is known about how metacommunities are organised in space in terrestrial, marine and freshwater ecological systems, their temporal variations remain poorly studied. Here, we address the question of the dynamics of metacommunities in highly variable systems, using intermittent rivers (IRs), those rivers which temporarily stop flowing or dry up, as a model system. We first review how habitat heterogeneity in space and time influences metacommunity organisation. Second, we compare the metacommunities in IRs to those in perennial rivers (PRs) and develop the idea that IRs could undergo highly dynamic shifts due to the temporal variability in local and regional community processes. Third, we develop the idea that in IRs, metacommunities of the wet and dry phases of IRs are closely intertwined, thereby increasing even more their respective temporal dynamics. Last, we provide a roadmap to stimulate further conceptual and empirical developments of metacommunity research and identify possible applications for improving the management of IRs and other highly dynamic ecological systems. Synthesis Extensive research has examined the importance of local biotic interactions, environmental filtering, and regional processes on community assembly. Movement of organisms between sites, i.e. dispersal, is a major set of processes within this framework. However, subsequent metacommunity organisation also varies over time in ecosystems because habitat characteristics such as configuration and composition continuously shift. Intermittent rivers are an ideal set of systems to examine these ideas because these freshwater systems temporarily cease flowing thereby limiting dispersal events. We proposed the hypothesis that metacommunities in dynamic ecosystems will undergo frequent shifts in structure and composition in response to the temporal variability in environmental filtering and dispersal. In addition to providing a roadmap for developing a more dynamic perspective for community ecology, these framework provides direct insights for the management of intermittent rivers.     

青藏高原草地植物群落冠层叶片氮磷化学计量学分析

叶片氮(N)和磷(P)的化学计量学研究涉及到植物生态学的众多领域与多个尺度, 然而各个尺度上的化学计量学研究并未同步展开。通过对青藏高原47个草地样地连续3年的调查, 分析了当地群落水平上的植物叶片N、P含量及其化学计量学特征, 并结合温度和降水气候数据研究了N、P含量及N:P比值与这两个气候因子的相关关系。研究结果显示: 青藏高原草地群落水平的叶片N含量变化范围为14.8-36.7 mg·g^-1, 平均为23.2 mg·g^-1; P含量变化范围为0.8-2.8 mg·g^-1, 平均为1.7 mg·g^-1; N:P比值变化范围为6.8-25.6, 平均为13.5。群落叶片N含量与P含量呈显著正相关关系, 叶片的N:P比值与P含量呈显著负相关关系, N:P比值的变化主要由P含量变化决定。另外发现: 群落水平叶片N、P含量及N:P比值存在着显著的年际变化, 叶片的N、P含量及N:P比值与年平均气温之间存在着极显著的相关关系。通过该研究结果推测: P含量较高的变异系数及其与环境因子表现出的显著相关性, 在一定程度上体现了植物群落对当地气候条件的一种适应。     

Recovery of three arctic stream reaches from experimental nutrient enrichment

1. Nutrient enrichment and resulting eutrophication is a widespread anthropogenic influence on freshwater ecosystems, but recovery from nutrient enrichment is poorly understood, especially in stream environments. We examined multi‐year patterns in community recovery from experimental low‐concentration nutrient enrichment (N + P or P only) in three reaches of two Arctic tundra streams (Kuparuk River and Oksrukuyik Creek) on the North Slope of Alaska (U.S.A.). 2. Rates of recovery varied among community components and depended on duration of enrichment (2–13 consecutive growing seasons). Biomass of epilithic algae returned to reference levels rapidly (within 2 years), regardless of nutrients added or enrichment duration. Aquatic bryophyte cover, which increased greatly in the Kuparuk River only after long‐term enrichment (8 years), took 8 years of recovery to approach reference levels, after storms had scoured most remnant moss in the recovering reach. 3. Multi‐year persistence of bryophytes in the Kuparuk River appeared to prevent recovery of insect populations that had either been positively (e.g. the mayfly Ephemerella, most chironomid midge taxa) or negatively (e.g. the tube‐building chironomid Orthocladius rivulorum) affected by this shift in dominant primary producer. These lags in recovery (of >3 years) were probably driven by the persistent effect of bryophytes on physical benthic habitat. 4. Summer growth rates of Arctic grayling (both adults and young‐of‐year) in Oksrukuyik Creek (fertilised for 6 years with no bryophyte colonisation), which were consistently increased by nutrient addition, returned to reference rates within 1–2 years. 5. Rates of recovery of these virtually pristine Arctic stream ecosystems from low‐level nutrient enrichment appeared to be controlled largely by duration of enrichment, mediated through physical habitat shifts caused by eventual bryophyte colonisation, and subsequent physical disturbance that removed bryophytes. Nutrient enrichment of oligotrophic Arctic stream ecosystems caused by climate change or local anthropogenic activity may have dramatic and persistent consequences if it results in the colonisation of long‐lived primary producers that alter physical habitat.     

Environmental and spatial controls of taxonomic versus trait composition of stream biota

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Primates as Predictors of Mammal Community Diversity in the Forest Ecosystems of Madagascar

The geographic distribution of species is the typical metric for identifying priority areas for conservation. Since most biodiversity remains poorly studied, a subset of charismatic species, such as primates, often stand as surrogates for total biodiversity. A central question is therefore, how effectively do primates predict the pooled species richness of other mammalian taxa? We used lemurs as indicator species to predict total non-primate mammal community richness in the forest ecosystems of Madagascar. We combine environmental and species occurrence data to ascertain the extent to which primate diversity can predict (1) non-primate mammal α-diversity (species richness), (2) non-primate complementarity, and (3) non-primate β-diversity (species turnover). Our results indicate that primates are effective predictors of non-primate mammal community diversity in the forest ecosystems of Madagascar after controlling for habitat. When individual orders of mammals are considered, lemurs effectively predict the species richness of carnivorans and rodents (but not afrosoricids), complementarity of rodents (but not carnivorans or afrosoricids), and all individual components of β-diversity. We conclude that lemurs effectively predict total non-primate community richness. However, surrogate species alone cannot achieve complete representation of biodiversity.     

Trophic-dynamic considerations in relating species diversity to ecosystem resilience

Complexity in the networks of interactions among and between the living and abiotic components forming ecosystems confounds the ability of ecologists to predict the economic consequences of perturbations such as species deletions in nature. Such uncertainty hampers prudent decision making about where and when to invest most intensively in species conservation programmes. Demystifying ecosystem responses to biodiversity alterations may be best achieved through the study of the interactions allowing biotic communities to compensate internally for population changes in terms of contributing to ecosystem function, or their intrinsic functional redundancy. Because individual organisms are the biologically discrete working components of ecosystems and because environmental changes are perceived at the scale of the individual, a mechanistic understanding of functional redundancy will hinge upon understanding how individuals' behaviours influence population dynamics in the complex community setting. Here, I use analytical and graphical modelling to construct a conceptual framework for predicting the conditions under which varying degrees of interspecific functional redundancy can be found in dynamic ecosystems. The framework is founded on principles related to food web successional theory, which provides some evolutionary insights for mechanistically linking functional roles of discrete, interacting organisms with the dynamics of ecosystems because energy is the currency both for ecological fitness and for food web commerce. Net productivity is considered the most contextually relevant ecosystem process variable because of its socioeconomic significance and because it ultimately subsumes all biological processes and interactions. Redundancy relative to productivity is suggested to manifest most directly as compensatory niche shifts among adaptive foragers in exploitation ecosystems, facilitating coexistence and enhancing ecosystem recovery after disturbances which alter species' relative abundances, such as extinctions. The framework further explicates how resource scarcity and environmental stochasticity may constitute 'ecosystem legacies' influencing the emergence of redundancy by shaping the background conditions for foraging behaviour evolution and, consequently, the prevalence of compensatory interactions. Because it generates experimentally testable predictions for a priori hypothesis testing about when and where varying degrees of functional redundancy are likely to be found in food webs, the framework may be useful for advancing toward the reliable knowledge of biodiversity and ecosystem function relations necessary for prudent prioritization of conservation programmes. The theory presented here introduces explanation of how increasing diversity can have a negative influence on ecosystem sustainability by altering the environment for biotic interactions and thereby changing functional compensability among biota--under particular conditions.     

Impacts of Intensive Logging on the Trophic Organisation of Ant Communities in a Biodiversity Hotspot

Trophic organisation defines the flow of energy through ecosystems and is a key component of community structure. Widespread and intensifying anthropogenic disturbance threatens to disrupt trophic organisation by altering species composition and relative abundances and by driving shifts in the trophic ecology of species that persist in disturbed ecosystems. We examined how intensive disturbance caused by selective logging affects trophic organisation in the biodiversity hotspot of Sabah, Borneo. Using stable nitrogen isotopes, we quantified the positions in the food web of 159 leaf-litter ant species in unlogged and logged rainforest and tested four predictions: (i) there is a negative relationship between the trophic position of a species in unlogged forest and its change in abundance following logging, (ii) the trophic positions of species are altered by logging, (iii) disturbance alters the frequency distribution of trophic positions within the ant assemblage, and (iv) disturbance reduces food chain length. We found that ant abundance was 30% lower in logged forest than in unlogged forest but changes in abundance of individual species were not related to trophic position, providing no support for prediction (i). However, trophic positions of individual species were significantly higher in logged forest, supporting prediction (ii). Consequently, the frequency distribution of trophic positions differed significantly between unlogged and logged forest, supporting prediction (iii), and food chains were 0.2 trophic levels longer in logged forest, the opposite of prediction (iv). Our results demonstrate that disturbance can alter trophic organisation even without trophically-biased changes in community composition. Nonetheless, the absence of any reduction in food chain length in logged forest suggests that species-rich arthropod food webs do not experience trophic downgrading or a related collapse in trophic organisation despite the disturbance caused by logging. These food webs appear able to bend without breaking in the face of some forms of anthropogenic disturbance.     

Control of plant species diversity and community invasibility by species immigration: seed richness versus seed density

Immigration rates of species into communities are widely understood to influence community diversity, which in turn is widely expected to influence the susceptibility of ecosystems to species invasion. For a given community, however, immigration processes may impact diversity by means of two separable components: the number of species represented in seed inputs and the density of seed per species. The independent effects of these components on plant species diversity and consequent rates of invasion are poorly understood. We constructed experimental plant communities through repeated seed additions to independently measure the effects of seed richness and seed density on the trajectory of species diversity during the development of annual plant communities. Because we sowed species not found in the immediate study area, we were able to assess the invasibility of the resulting communities by recording the rate of establishment of species from adjacent vegetation. Early in community development when species only weakly interacted, seed richness had a strong effect on community diversity whereas seed density had little effect. After the plants became established, the effect of seed richness on measured diversity strongly depended on seed density, and disappeared at the highest level of seed density. The ability of surrounding vegetation to invade the experimental communities was decreased by seed density but not by seed richness, primarily because the individual effects of a few sown species could explain the observed invasion rates. These results suggest that seed density is just as important as seed richness in the control of species diversity, and perhaps a more important determinant of community invasibility than seed richness in dynamic plant assemblages.     

Clockwise and counterclockwise hysteresis characterize state changes in the same aquatic ecosystem

Incremental increases in a driver variable, such as nutrients or detritus, can trigger abrupt shifts in aquatic ecosystems that may exhibit hysteretic dynamics and a slow return to the initial state. A model system for understanding these dynamics is the microbial assemblage that inhabits the cup‐shaped leaves of the pitcher plant Sarracenia purpurea. With enrichment of organic matter, this system flips within three days from an oxygen‐rich state to an oxygen‐poor state. In a replicated greenhouse experiment, we enriched pitcher‐plant leaves at different rates with bovine serum albumin (BSA), a molecular substitute for detritus. Changes in dissolved oxygen (DO) and undigested BSA concentration were monitored during enrichment and recovery phases. With increasing enrichment rates, the dynamics ranged from clockwise hysteresis (low), to environmental tracking (medium), to novel counter‐clockwise hysteresis (high). These experiments demonstrate that detrital enrichment rate can modulate a diversity of hysteretic responses within a single aquatic ecosystem, and suggest different management strategies may be needed to mitigate the effects of high vs. low rates of detrital enrichment.     

Exploring the dynamics of ecological indicators using food web models fitted to time series of abundance and catch data

Previously, standardized snap-shot models of the Southern Benguela (1980–1989), Southern Humboldt (1992) and Southern Catalan Sea (1994) ecosystems were examined and found to facilitate assessment of ecosystem characteristics related to the gradient in exploitation status of the ecosystems; highest level of exploitation in the South Catalan Sea (North-western Mediterranean), high in the Southern Humboldt and lower in the Southern Benguela. Subsequently, these models were calibrated and fitted using available catch, fishing effort/mortality and abundance data series and incorporated environmental and internal drivers. This study furthers the previous comparative analyses by comparing changes in ecosystem structure using a selection of ecosystem indicators from the calibrated models and assessing how these indicators change over time in these three contrasting ecosystems. Indicators examined include community turnover rates (production/biomass), trophic level of landings and the community, biodiversity indicators, ratios of predatory/forage fish and pelagic/demersal fish biomass, catch ratios, and network analysis indicators. Using the set of model-derived indicators, the three ecosystems were ranked in terms of exploitation level. This ranking was performed using the values of these indicators in recent years (ecosystem state) as well as their trends over time (ecosystem trend). The non-parametric Kruskal–Wallis and Median tests were used to test for significance of the difference between indicators from the three ecosystems in the last 5 years of the simulation to compare present ecosystem states. We compared the slope of the lineal trend and its significance between ecosystems using the generalized least-squares regression taking auto-correlation into consideration to analyse ecosystem trends. The indicators that capture better the high impacts of fishing prevalent in the Mediterranean and Humboldt ecosystems, and the more conservative exploitation of the Southern Benguela, are the fish/invertebrates biomass and catch ratio, the demersal/pelagic fish biomass and catch ratio (depending on the ecosystem and the fishery being developed), flows to detritus, and the mean trophic level of the community (when large, poorly quantified groups such as zooplankton and detritus are excluded). This study suggests that the best option for classifying ecosystems according to the impact of fishing is to consider a broad range of indicators to understand how and why an ecosystem is responding to particular environmental or fishing drivers (or more likely a combination of these). Our results highlight the importance of including indicators capturing trends over time as well as recent ecosystem states. We also identified 23 pairs of indicators that correlated similarly in the three ecosystems (they showed a significant correlation with same sign). Further comparisons may contribute towards generalization of this list, progressing towards a better understanding of the behaviour of ecological indicators.     

Parallels and contrasts between intermittently freezing and drying streams: From individual adaptations to biodiversity variation

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Inter- and intraspecific variability of plant individual growth and its role on species ranking in grasslands

植物个体的生长是其适合度的重要组成部分。然而,在个体水平上非破坏性地测量草本植物的生长速度非常困难。本研究基于准确的非破坏性地面生物量估算方法,旨在评估某些物种的个体相对生长速率,识别其环境驱动因子,并检验其对群落模式的影响。我们提出以下三个具体的科学问题：(1)环境条件可以在多大程度上解释个体植物生长在不同样地间的差异;(2)在相同样地内和不同样地间,植物个体生长的种内差异有多大,以及(3)相同样地内,平均每个物种的个体生长相较于整个植被的优势/劣势,与群落水平的物种等级是否有关？为了回答上述问题,我们在法国中央高地沿古气候梯度的18 个永久草地内监测了4种常见的多年生植物的个体生长。我们测量了土壤性质、资源水平和气象参数,以表征样地的环境条件。这使我们能够评估环境条件对个体生长的影响,以及在相同和不同样地之间个体间生长变异的相对程度。我们通过植物学调查确定了每个样地内四个物种的等级,以评估物种平均个体生长相对于整个群落的优势劣势与物种在群落中的等级之间的关系。研究结果表明,环境条件解释了个体生长变异性很重要的一部分,而且这部分变异在物种之间也有很大差异。光照有效性是影响植物生长的主要因素,其次是降雨量和潜在的蒸散量,而土壤性质对植物生长的影响较小。我们的结果进一步强调了相同样地内个体间生长变异的程度是中到高,而且物种等级与物种平均个体生长之间没有相关性。