Examining differences in phylogenetic composition enhances understanding of the phylogenetic structure of the shrub community in the northeastern Qinghai‐Tibetan Plateau

Periodic climatic oscillations and species dispersal during the postglacial period are two important causes of plant assemblage and distribution on the Qinghai‐Tibet Plateau (QTP). To improve our understanding of the bio‐geological histories of shrub communities on the QTP, we tested two hypotheses. First, the intensity of climatic oscillations played a filtering role during community structuring. Second, species dispersal during the postglacial period contributed to the recovery of species and phylogenetic diversity and the emergence of phylogenetic overdispersion. To test these hypotheses, we investigated and compared the shrub communities in the alpine and desert habitats of the northeastern QTP. Notably, we observed higher levels of species and phylogenetic diversity in the alpine habitat than in the desert habitat, leading to phylogenetic overdispersion in the alpine shrub communities versus phylogenetic clustering in the desert shrub communities. This phylogenetic overdispersion increased with greater climate anomalies. These results suggest that (a) although climate anomalies strongly affect shrub communities, these phenomena do not act as a filter for shrub community structuring, and (b) species dispersal increases phylogenetic diversity and overdispersion in a community. Moreover, our investigation of the phylogenetic community composition revealed a larger number of plant clades in the alpine shrub communities than in the desert shrub communities, which provided insights into plant clade‐level differences in the phylogenetic structures of alpine and desert shrub communities in the northeastern QTP.     

藏东木本植物群落功能性状分布与环境的关系

研究植物群落功能性状间的相关关系及其对环境变化的响应,能够有效揭示植物功能性状的权衡模式及其对环境的适应策略。藏东昌都地区位于横断山脉西北部,复杂气候地貌孕育了丰富的植物资源,是青藏高原森林灌丛生态系统主要组分和国际生物多样性保护的热点地区。以藏东森林灌丛群落优势木本植物为研究对象,在大量野外调查基础上,采用相关分析、主成分分析、线性回归和方差分析等方法,研究了该区域植物功能性状间的相关关系、功能性状对环境变量的响应规律以及功能性状的变异来源。结果表明：(1)藏东木本植物表现出适应高寒环境的性状权衡模式,即：比叶面积、叶体积较小而叶干物质含量较大,叶磷含量和叶钾含量协同变化;(2)海拔和气候变量共同驱动着藏东木本植物功能性状的变化,并且藏东木本植物倾向于采取“高投入—慢回报”提高御寒能力的保守型适应策略;(3)海拔是影响藏东植物功能性状变异最显著的环境变量,种间变异在藏东植物群落功能性状随环境变化中起主要作用。研究结果揭示了藏东木本植物功能性状的权衡模式及其对高寒环境的适应策略,有助于加深对藏东自然植物资源分布规律和生态功能的认识,为区域生态系统功能和生物多样性保护提供科学依据。     

The Importance of Biotic vs. Abiotic Drivers of Local Plant Community Composition Along Regional Bioclimatic Gradients

We assessed if the relative importance of biotic and abiotic factors for plant community composition differs along environmental gradients and between functional groups, and asked which implications this may have in a warmer and wetter future. The study location is a unique grid of sites spanning regional-scale temperature and precipitation gradients in boreal and alpine grasslands in southern Norway. Within each site we sampled vegetation and associated biotic and abiotic factors, and combined broad- and fine-scale ordination analyses to assess the relative explanatory power of these factors for species composition. Although the community responses to biotic and abiotic factors did not consistently change as predicted along the bioclimatic gradients, abiotic variables tended to explain a larger proportion of the variation in species composition towards colder sites, whereas biotic variables explained more towards warmer sites, supporting the stress gradient hypothesis. Significant interactions with precipitation suggest that biotic variables explained more towards wetter climates in the sub alpine and boreal sites, but more towards drier climates in the colder alpine. Thus, we predict that biotic interactions may become more important in alpine and boreal grasslands in a warmer future, although more winter precipitation may counteract this trend in oceanic alpine climates. Our results show that both local and regional scales analyses are needed to disentangle the local vegetation-environment relationships and their regional-scale drivers, and biotic interactions and precipitation must be included when predicting future species assemblages.     

高原鼢鼠干扰强度对祁连山东段高寒草甸大型土壤动物功能群特征及空间分布的影响

为明晰高原鼢鼠干扰对大型土壤动物类群分布和功能群特征影响,选取祁连山东段高原鼢鼠典型分布高寒草甸,依据鼠丘密度将研究区划分4个干扰梯度,调查各干扰区植物、群落结构、土壤理化性质和大型土壤动物功能群组成。采用冗余分析方法探讨植物土壤因子对土壤动物功能群组成和分布的影响。结果表明：研究区植食性土壤动物为优势功能群,极重度干扰区植食性功能群丰富度和Shannon多样性指数显著高于中度和重度干扰区(P<0.05);高原鼢鼠干扰对高寒草甸土壤动物群落稳定和相似性影响较小;冗余分析表明土壤温度、紧实度、全氮和全钾以及莎草科生物量和植物物种均匀度均显著影响高寒草甸土壤动物功能群的空间分布(P<0.05),其中土壤温度对土壤动物功能群分布影响最为显著。     

Using functional trait diversity to evaluate the contribution of multiple ecological processes to community assembly during succession

Successional chronosequences provide a unique opportunity to study the effects of multiple ecological processes on plant community assembly. Using a series of 0.5 × 0.5 m² plots (n = 30) from five successional sub‐alpine meadow plant communities (ages 3, 5, 9, 12, and undisturbed) in the Qinghai‐Tibetan Plateau, we investigated whether community assembly is stochastic or deterministic for species and functional traits. We tested directional change in species composition, functional trait composition, and then functional trait diversity measured by Rao's quadratic entropy for four traits – plant height, leaf dry matter content, specific leaf area, and seed mass – along two comparable successional chronosequences. We then evaluated the importance of species interactions, habitat filtering and stochasticity by comparing with random communities and partitioning the environmental and spatial components of Rao's quadratic entropy. We found no directional change in species composition, but clear directionality in functional trait composition. None of the abiotic environmental variables (except P) showed linear change with successional age, but soil moisture and nitrogen were positively related to functional diversity within meadows. Functional trait diversity increased significantly with the increase in successional age. Comparison with random communities showed a significant shift from trait divergence in early stages of succession (3‐ and 5‐yr) to convergence in the later stages of succession 9‐, 12‐yr and undisturbed). The relative importance of abiotic variables and spatial structure for functional trait diversity changed in a predictable manner with successional age. Stochasticity at the species level may indicate dispersal limitation, but deterministic effects on functional trait distributions show the role of both habitat effects and biotic interactions.     

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.     

Frequency and intensity of facilitation reveal opposing patterns along a stress gradient

Disentangling the different processes structuring ecological communities is a long‐standing challenge. In species‐rich ecosystems, most emphasis has so far been given to environmental filtering and competition processes, while facilitative interactions between species remain insufficiently studied. Here, we propose an analysis framework that not only allows for identifying pairs of facilitating and facilitated species, but also estimates the strength of facilitation and its variation along environmental gradients. Our framework combines the analysis of both co‐occurrence and co‐abundance patterns using a moving window approach along environmental gradients to control for potentially confounding effects of environmental filtering in the co‐abundance analysis. We first validate our new approach against community assembly simulations, and exemplify its potential on a large 1,134 plant community plots dataset. Our results generally show that facilitation intensity was strongest under cold stress, whereas the proportion of facilitating and facilitated species was higher under drought stress. Moreover, the functional distance between individual facilitated species and their facilitating species significantly changed along the temperature–moisture gradient, and seemed to influence facilitation intensity, although no general positive or general negative trend was discernible among species. The main advantages of our robust framework are as follows: It enables detecting facilitating and facilitated species in species‐rich systems, and it allows identifying the directionality and intensity of facilitation in species pairs as well as its variation across long environmental gradients. It thus opens numerous opportunities for incorporating functional (and phylogenetic) information in the analysis of facilitation patterns. Our case study indicated high complexity in facilitative interactions across the stress gradient and revealed new evidence that facilitation, similarly to competition, can operate between functionally similar and dissimilar species. Extending the analyses to other taxa and ecosystems will foster our understanding how complex interspecific interactions promote biodiversity.     

Shrub encroachment in the Alps gives rise to the convergence of sub‐alpine communities on a regional scale

Question: How does Alnus alnobetula colonization affect plant communities in sub‐alpine grasslands undergoing land abandonment on a regional scale? Location: Savoie, French Alps. Methods: Variations within and between communities were monitored in 243 plots within three types of stands representing an A. alnobetula colonization chronosequence from grassland to dense vegetation. They were tested for species richness, species strategies and species distribution through multivariate analysis. Regional variations were assessed at three sites along a rain shadow gradient. Results: We detected a convergence of the understorey communities along an A. alnobetula colonization gradient at all sites, associated with a reduction in species richness and the replacement of stress related strategists by competitor strategists. This convergence was unaffected by the rain shadow gradient although it was a key factor of plant distribution. Conclusions: The dense cover and the soil properties provided by A. alnobetula stands buffered the heterogeneity of sub‐alpine communities induced by a rain shadow gradient throughout the study region. It facilitated the development a homogeneous understorey community dominated by competitor related strategists while excluding grasslands stress adapted species through competition processes. The resulting community, although species‐poor, has an interesting conservation value. Accordingly, as a ‘native invader’, A. alnobetula does not display typical traits of biological invaders. Future experimental research on the interactions between A. alnobetula and components of the understorey would be particularly interesting in a severe environment that promotes facilitation among plants.     

高寒草甸不同草地类型功能群多样性及组成对植物群落生产力的影响

对不同类型草地功能群多样性和组成与植物群落生产力之间的关系进行了探讨。结果表明：(1)在矮嵩草(Kobresia humlis)草甸和金露梅(Potentilla froticosa)灌丛中,豆科植物的作用比较明显,而其他功能群植物的作用较弱。(2)在藏嵩草(Kobresia tibetica)沼泽化草甸和小嵩草(K．pygmaca)草甸中,虽然杂类草、C3植物和莎草科植物功能群的生产力占群落初级生产力的比例较大,但二者在统计上没有显著性差异,这表明群落生产力除受物种多样性的影响外,也受物种本身特征和环境资源的影响,更主要的是受到功能群内物种密度和均匀度的影响,即功能群组成比功能群多样性更能说明对生态系统过程的影响。(3)不同类型草地群落植物功能群盖度与群落初级生产力呈显著的线性相关。(4)不同类型草地群落生产力与功能群内物种数的变化均表现为单峰曲线关系,即功能群内物种数处于中间水平时,群落生产力最高。     

吉林灌木群落物种多样性与气候和局域环境因子的关系

为了研究气候和局域环境因子对物种多样性的相对作用大小,以及验证两种均匀度地理格局的假说在半湿润地区次生灌丛的适用性,对吉林东、南部地区的灌木群落进行了研究。共调查森林破坏后形成的次生灌丛样方45个,结合气候数据和局域环境因子数据,研究了气候、局域环境因子对群落、灌木层、草本层的物种丰富度、均匀度的影响,以及对不同水分生态型(旱生、旱中生、湿中生)灌木影响的差异。结果表明:1)吉林次生灌丛的群落、草本层物种丰富度,以及草本层均匀度,随纬度增加而显著上升。2)对物种多样性和气候、局域环境因子的分析表明,群落、草本层物种数主要受局域环境因子而不是气候的影响;其物种丰富度与纬度的反常关系,是由于灌木层盖度随降水增加而上升,从而导致物种数下降。灌木层物种数与纬度、气候因子的相关性不显著,则是由于不同水分生态型对气候梯度的响应不一致,反映出功能群对多样性格局的影响。3)群落、灌木层均匀度主要受气候因子的影响;而草本层均匀度主要受局域环境因子的影响,降水同样通过对灌木层盖度的影响间接作用于草本均匀度。但群落、灌木和草本层的结果,都支持均匀度随着环境条件改善而增加的假说,而不支持随着生产力增加、竞争加剧,从而导致均匀度下降的假说。结果表明,物种丰富度和均匀度的影响机制存在很大差异,但二者都受到局域环境因子的强烈影响。气候通过局域生物因素(如盖度、生活型)间接作用于多样性格局,是气候对多样性影响的一个重要方面,但尚未得到应有的重视。由于局域生物因素也随气候而变化,仅研究多样性和气候的表面关系,将无法准确预测气候变化对多样性的影响。     

Direct and indirect interactions among plants explain counterintuitive positive drought effects on an eastern Mediterranean shrub species

Coexistence of woody and herbaceous plants may be governed by a complex set of direct and indirect interactions, whose relative importance have been rarely assessed. We experimentally studied woody species establishment in a mixed plant community by disentangling the potential role of such biotic interactions and the effect of environmental variations on them. Seedling establishment of the common eastern Mediterranean shrub species Sarcopoterium spinosum was investigated under different rainfall and light conditions, combined with the effect of the presence of adult shrubs and annual neighbors. We predicted that seedlings will be directly affected by competition with annuals with increasing water availability, while direct effects of adult shrubs will be positive via amelioration of water stress. Indirect effects were expected beneath shrub canopies due to reduced water stressed and light availability for both annuals and shrub seedlings, which may intensify competition between annuals and shrub seedlings. To test these predictions we performed field and garden experiments in which we combined manipulation of shrub and annual presence with manipulations of water availability and light conditions to simulate the effect of shrub canopy. In contrast to our prediction, shrub seedling establishment was not facilitated but inhibited by adult shrubs because of light limitation. As expected, annuals had direct negative effects on shrub seedlings under wet conditions, which shifted to neutral or positive effects under dry conditions. Thus, interactions among shrubs and annuals, and in particular the release from competition during drought years, leads to a counterintuitive positive effect of drought on shrub seedling establishment. Our findings point to the importance of experimentally studying multidimensional interactions for coexistence of different life forms and to the underestimated role of light for success in water‐limited ecosystems.     

灌木年轮学研究进展

灌木往往分布在树线以上或以北的高海拔和高纬度地区以及干旱、半干旱区,是把传统上以乔木为主的树轮研究扩展至森林分布界限以外的唯一选择.尽管灌木具有以上研究潜力,迄今用于树木年代学研究的灌木种类仅有30种左右.介绍了灌木年轮研究方法,综述了过去几十年来环北极高纬度地区,干旱、半干旱区以及高海拔地区的灌木年轮研究的主要进展.主要研究进展如下:(1)发掘一些灌木的树木年代学潜力;(2)揭示限制灌木生长的主要环境因子,并尝试利用灌木年轮宽度等指标重建过去区域气候变化历史;(3)探讨全球变暖的背景下,灌木的生长或分布范围的变化;(4)通过人为控制增温来揭示变暖对灌木生理特征和生长的影响.这些研究展示了灌木在扩展传统乔木树轮研究网络方面的潜力,也是树木年代学研究中最有前景的研究方向之一.目前的灌木年轮学研究多集中于环北极苔原带.作为地球的第三极,青藏高原具有广泛的高山灌木分布,具有把青藏高原边缘区以乔木为主的树木年轮网络扩展至更高海拔和高原内部的潜力.青藏高原高山灌木的年轮学研究并没有引起足够的重视.青藏高原高山灌木的生长是如何适应极端环境条件的,全球变暖的背景下,青藏高原高山灌木的分布和生长正在发生哪些变化等,都有待深入研究.     

Experimental comparison of competition and facilitation in alpine communities varying in productivity

Question. Competitive and facilitative interactions among plant species in different abiotic environments potentially link productivity, vegetation structure, species composition and functional diversity. We investigated these interactions among four alpine communities along an environmental productivity gradient in a generally harsh climate. We hypothesised that the importance of competition would be higher in more productive sites. Location. Mt. M. Khatipara (43°27′N, 41°41′E, altitude 2750 m), NW Caucasus, Russia. Communities ranged from low‐productivity alpine lichen heath (ALH) and snowbed communities (SBC), to intermediate productivity Festuca grassland (FVG), and high‐productivity Geranium‐Hedysarum meadow (GHM). Methods. We quantified the relative influence of competition and facilitation on community structure by expressing biomass of target species within each natural community proportionally to biomass of the species in a “null community” with experimental release from interspecific competition by removing all other species (for 6 years). An overall index of change in community composition due to interspecific interactions was calculated as the sum of absolute or proportional differences of the component species. Results. Species responses to neighbour removal ranged from positive to neutral. There was no evidence of facilitation among the selected dominant species. As expected, competition was generally most important in the most productive alpine community (GHM). The intermediate position for low‐productivity communities of stressful environments (ALH, SBC) and the last position of intermediately productive FVG were unexpected. Conclusions. Our results appear to support the Fretwell‐Oksanen hypothesis in that competition in communities of intermediate productivity was less intense than in low‐ or high‐productive communities. However, the zero net effect of competition and facilitation in FVG might be the result of abiotic stress due to strong sun exposure and high soil temperatures after neighbour removal. Thus, non‐linear relationships between soil fertility, productivity and different abiotic stresses may also determine the balance between competition and facilitation.     

Shrub range expansion alters diversity and distribution of soil fungal communities across an alpine elevation gradient

Global climate and land use change are altering plant and soil microbial communities worldwide, particularly in arctic and alpine biomes where warming is accelerated. The widespread expansion of woody shrubs into historically herbaceous alpine plant zones is likely to interact with climate to affect soil microbial community structure and function; however, our understanding of alpine soil ecology remains limited. This study aimed to (i) determine whether the diversity and community composition of soil fungi vary across elevation gradients and to (ii) assess the impact of woody shrub expansion on these patterns. In the White Mountains of California, sagebrush (Artemisia rothrockii) shrubs have been expanding upwards into alpine areas since 1960. In this study, we combined observational field data with a manipulative shrub removal experiment along an elevation transect of alpine shrub expansion. We utilized next‐generation sequencing of the ITS1 region for fungi and joint distribution modelling to tease apart effects of the environment and intracommunity interactions on soil fungi. We found that soil fungal diversity declines and community composition changes with increasing elevation. Both abiotic factors (primarily soil moisture and soil organic C) and woody sagebrush range expansion had significant effects on these patterns. However, fungal diversity and relative abundance had high spatial variation, overwhelming the predictive power of vegetation type, elevation and abiotic soil conditions at the landscape scale. Finally, we observed positive and negative associations among fungal taxa which may be important in structuring community responses to global change.     

喀斯特高原山区灌丛草地昆虫群落多样性研究

为探讨喀斯特高原山区灌丛草地昆虫群落多样性.本研究于2018年9月-2019年8月使用"Z"字样方网捕法、陷阱法、黄盘诱集法三种相结合的方法对贵州省大方县星宿乡崔苏坝灌丛草地进行昆虫群落多样性的调查,采用单因素分析、线性回归分析、相关性分析探讨了灌丛草地昆虫群落多样性.本研究共获昆虫标本12481号,隶属8目77科,其...     

新疆平茶渍属地衣生态分布与地理区系成分分析

根据多年的实地调查资料和前人研究资料,对新疆平茶渍属（Aspicilia）地衣的种类以及它们的分布区、区系特征进行了初步研究。结果表明,分布在新疆的平茶渍属地衣共有33种,主要分布在天山、阿勒泰山、昆仑山和准噶尔盆地。根据它们对环境的适应特征和选择性,将新疆平茶渍属地衣的地理区系划分为广布种、环极北极高山种、环极低北极及北方种、北美东部种、中亚种、中国特有种、分布范围不清楚的种等7种类型。同时把新疆平茶渍属地衣的地理分布区域划分为欧亚森林植物亚区、亚洲荒漠植物亚区、青藏高原亚区等三种分布区域。研究还发现分布在阿勒泰山、天山、昆仑山和准噶尔盆地的平茶渍属主要生长在岩石上面,生长基物比较单一。     

Extreme drought stress shifts net facilitation to neutral interactions between shrubs and sub‐canopy plants in an arid desert

The stress gradient hypothesis (SGH) predicts that the importance or intensity of competition and facilitation will change inversely along abiotic stress gradients. It was originally postulated that increasing environmental stress can induce a monotonic increase in facilitation. However, more recent models predicted that the relationship between severity and interaction exhibits a hump‐shaped pattern, in which positive interactions prevail under moderate stress but decline at the extreme ends of stress gradients. In the present study, we conducted a field experiment along a temporal rainfall gradient for five consecutive years, in order to investigate interactions in a shrub‐herbaceous plant community at the southern edge of the Badain Jaran Desert, and, more specifically, investigated the effects of Calligonum mongolicum, a dominant shrub species, on both abiotic environmental variables and the performance of sub‐canopy plant species. We found that shrubs can improve sub‐canopy water regimes, soil properties, plant biomass, density, cover, and richness and, more importantly, that the positive effect of shrubs on sub‐canopy soil moisture during the summer diminishes as rainfall decreases, a pattern that partly explains the collapse of the positive interaction between shrubs and their understory plants. These results provide empirical evidence that the positive effect of shrubs on understory plant communities in extreme arid environments may decline and become neutral with increasing drought stress.     

青藏高原高寒灌丛植被对长期放牧强度试验的响应特征

在青藏高原中国科学院海北高寒草甸生态系统定位研究站对金露梅高寒灌丛草场植被开展了长期不同放牧强度试验,分别在短期(4年)、中期(11年)和长期(18年)放牧阶段研究不同放牧干扰强度对草地植物物种多样性、群落结构、地上生物量和草场质量的影响.研究表明,在不同放牧阶段,随着放牧强度增加植物群落的高度和盖度都降低.在中期放牧干扰阶段,物种多样性数和均匀度指数随着放牧强度增加呈现典型的单峰曲线模式;在长期放牧干扰阶段,随着放牧强度增加,占优势地位的灌木和禾草被典型杂类草替代,其中的重度放牧干扰简化了高寒灌丛植被群落结构,减少了地上现存生物量,特别是可食优良牧草生物量.植被对放牧的响应除了与放牧强度和放牧时间阶段密切相关外,还与该地区水热条件的变化有一定的相关性.针对长期放牧干扰的反应特性可将金露梅灌丛草场中植物划分为增加型、敏感型、忍耐型和无反应型4种类型.除了丰富度指数、多样性指数和均匀度指数外,其它一些特征参数并不支持著名的中度干扰假说.本研究发现,长期重度放牧促进了青藏高原高寒草地退化,适度放牧有利于高寒灌丛草场的生物多样性保护和牧草利用;"取半留半"的放牧原则在青藏高原草场放牧管理实践中值得推荐,它将有利于防止草场退化,提高牧草利用率和维持较高的生物多样性.     

锡林河流域羊草草原植物种群和功能群的长期变异性及其对群落稳定性的影响

 分析了内蒙古锡林河流域羊草(Leymus chinensis)草原群落植物种群、功能群和群落生物量的长期变异性,以及植物功能群间的补偿作用对群落稳定性的影响,结果表明：从种群水平到功能群和群落水平,地上生物量的年度间变异性逐渐降低,而稳定性则逐渐增加。植物功能群组成对群落稳定性具有更强烈的影响,在植物生活型功能群组成中多年生根茎禾草与多年生丛生禾草和多年生杂类草功能群,在生物量的年度变化上具有补偿作用,在生态类群组成中,旱生植物与中旱生植物也具有补偿作用。植物种群多度的长期变异性可能依赖于物种对环境波动的敏感性和种间相互作用对环境波动的放大效应,生物多样性使物种对环境波动响应的多样性增加,并通过不同响应类型(功能群)间的补偿作用使群落稳定性增加。     

The hidden complexity of multi‐species interactions

In this issue of the Journal of Vegetation Science, Michalet et al. used species removal experiments to detect direct and indirect species interactions in a sub‐alpine grassland. They found evidence for competition, facilitation and a range of indirect interactions at the species level, but no measurable effects when aggregated at the community level. Their results raise fundamental questions regarding the role and importance of indirect interactions in structuring ecological communities.