Identification of root-colonizing AM fungal communities and their responses to short-term climate change and grazing on Tibetan plateau

The molecular diversity of arbuscular mycorrhizal (AM) fungi remains largely unexplored in sedge-dominated ecosystems, much less their responses to global climate change and grazing. To determine the molecular diversity of AM fungi and how they are affected by climate change and grazing, we examined AM fungal communities inside roots in a sedge meadow ecosystem (4875 m a.s.l.) undergoing 4 yr. of fully factorial manipulations of experimental warming, snow addition and grazing on the Tibetan Plateau. A total of 21 AM fungal virtual taxa (VTs) were identified, including one new genus-like clade and one new VT unrecorded in the MaarjAM database. However, we did not observe significant effects of climate change and/or grazing on AM fungal abundance, diversity and community composition. These results indicate that diverse AM fungal taxa inhabit in the sedge-dominated meadows on the Tibetan Plateau, but the AM fungal communities were not sensitive to short-term climate change and grazing.     

Trait structure and redundancy determine sensitivity to disturbance in marine fish communities

Trait diversity is believed to influence ecosystem dynamics through links between organismal traits and ecosystem processes. Theory predicts that key traits and high trait redundancy—large species richness and abundance supporting the same traits—can buffer communities against environmental disturbances. While experiments and data from simple ecological systems lend support, large‐scale evidence from diverse, natural systems under major disturbance is lacking. Here, using long‐term data from both temperate (English Channel) and tropical (Seychelles Islands) fishes, we show that sensitivity to disturbance depends on communities’ initial trait structure and initial trait redundancy. In both ecosystems, we found that increasing dominance by climatically vulnerable traits (e.g., small, fast‐growing pelagics/corallivores) rendered fish communities more sensitive to environmental change, while communities with higher trait redundancy were more resistant. To our knowledge, this is the first study demonstrating the influence of trait structure and redundancy on community sensitivity over large temporal and spatial scales in natural systems. Our results exemplify a consistent link between biological structure and community sensitivity that may be transferable across ecosystems and taxa and could help anticipate future disturbance impacts on biodiversity and ecosystem functioning.     

Metabolic compensation constrains the temperature dependence of gross primary production

Gross primary production (GPP) is the largest flux in the carbon cycle, yet its response to global warming is highly uncertain. The temperature dependence of GPP is directly linked to photosynthetic physiology, but the response of GPP to warming over longer timescales could also be shaped by ecological and evolutionary processes that drive variation in community structure and functional trait distributions. Here, we show that selection on photosynthetic traits within and across taxa dampens the effects of temperature on GPP across a catchment of geothermally heated streams. Autotrophs from cold streams had higher photosynthetic rates and after accounting for differences in biomass among sites, biomass‐specific GPP was independent of temperature in spite of a 20 °C thermal gradient. Our results suggest that temperature compensation of photosynthetic rates constrains the long‐term temperature dependence of GPP, and highlights the importance of considering physiological, ecological and evolutionary mechanisms when predicting how ecosystem‐level processes respond to warming.     

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。     

Evolutionary responses to environmental change: trophic interactions affect adaptation and persistence

According to recent reviews, the question of how trophic interactions may affect evolutionary responses to climate change remains unanswered. In this modelling study, we explore the evolutionary dynamics of thermal and plant–herbivore interaction traits in a warming environment. We find the herbivore usually reduces adaptation speed and persistence time of the plant by reducing biomass. However, if the plant interaction trait and thermal trait are correlated, herbivores can create different coevolutionary attractors. One attractor has a warmer plant thermal optimum, and the other a colder one compared with the environment. A warmer plant thermal strategy is given a head start under warming, the only case where herbivores can increase plant persistence under warming. Persistence time of the plant under warming is maximal at small or large thermal niche width. This study shows that considering trophic interactions is necessary and feasible for understanding how ecosystems respond to climate change.     

Leaf-trait responses to environmental gradients in moorland communities: contribution of intraspecific variation,species replacement and functional group replacement

The values of many important traits of plants in a community change along environmental gradients. Such changes may involve intraspecific variation and replacement by species that have different trait values. We hypothesized that they also involve the variation within and among functional groups (FGs) to the environmental dependence of trait values at the community level. We studied environmental dependence of trait values in 27 moorlands at various scales and analyzed to what extent intraspecific variation, species replacement within FGs and FG replacement contribute to the gradient of community trait values. The community structure in moorlands was influenced mainly by two environmental factors: temperature and water condition. Plants inhabiting sites with low temperature and low-pH generally tended to have lower maximum leaf height, greater leaf mass per area, and smaller leaf size. At the community level, site-mean of maximum leaf height and leaf size generally increased with increasing temperature and water pH. Our analysis demonstrated that the relative contributions of intraspecific variation, species replacement within FGs and FG replacement differed depending on combinations of the traits and environments. The contribution of FG replacement varied considerably among cases (0.6–34.5 %). Species replacement within FGs, which has received little attention in previous studies, was most responsible for the community-level changes (31.6–65.3 %) and intraspecific variation also made a large contribution (22.9–57.9 %). Understanding such various mechanisms involving intraspecific variation and species replacement should help us better predict how the structure and functioning of moorland plant communities will respond to climate change.     

Characterizing the contribution of plasticity and genetic differentiation to community‐level trait responses to environmental change

The match between functional trait variation in communities and environmental gradients is maintained by three processes: phenotypic plasticity and genetic differentiation (intraspecific processes), and species turnover (interspecific). Recently, evidence has emerged suggesting that intraspecific variation might have a potentially large role in driving functional community composition and response to environmental change. However, empirical evidence quantifying the respective importance of phenotypic plasticity and genetic differentiation relative to species turnover is still lacking. We performed a reciprocal transplant experiment using a common herbaceous plant species (Oxalis montana) among low‐, mid‐, and high‐elevation sites to first quantify the contributions of plasticity and genetic differentiation in driving intraspecific variation in three traits: height, specific leaf area, and leaf area. We next compared the contributions of these intraspecific drivers of community trait–environment matching to that of species turnover, which had been previously assessed along the same elevational gradient. Plasticity was the dominant driver of intraspecific trait variation across elevation in all traits, with only a small contribution of genetic differentiation among populations. Local adaptation was not detected to a major extent along the gradient. Fitness components were greatest in O. montana plants with trait values closest to the local community‐weighted means, thus supporting the common assumption that community‐weighted mean trait values represent selective optima. Our results suggest that community‐level trait responses to ongoing climate change should be mostly mediated by species turnover, even at the small spatial scale of our study, with an especially small contribution of evolutionary adaptation within species.     

Grazing effects on intraspecific trait variability vary with changing precipitation patterns in Mongolian rangelands

Functional traits are proxies for plant physiology and performance, which do not only differ between species but also within species. In this work, we hypothesized that (a) with increasing precipitation, the percentage of focal species which significantly respond to changes in grazing intensity increases, while under dry conditions, climate‐induced stress is so high that plant species hardly respond to any changes in grazing intensity and that (b) the magnitude with which species change their trait values in response to grazing, reflected by coefficients of variation (CVs), increases with increasing precipitation. Chosen plant traits were canopy height, plant width, specific leaf area (SLA), chlorophyll fluorescence, performance index, stomatal pore area index (SPI), and individual aboveground biomass of 15 species along a precipitation gradient with different grazing intensities in Mongolian rangelands. We used linear models for each trait to assess whether the percentage of species that respond to grazing changes along the precipitation gradient. To test the second hypothesis, we assessed the magnitude of intraspecific trait variability (ITV) response to grazing, per species, trait, and precipitation level by calculating CVs across the different grazing intensities. ITV was most prominent for SLA and SPI under highest precipitation, confirming our first hypothesis. Accordingly, CVs of canopy height, SPI, and SLA increased with increasing precipitation, partly confirming our second hypothesis. CVs of the species over all traits increased with increasing precipitation only for three species. This study shows that it remains challenging to predict how plant performance will shift under changing environmental conditions based on their traits alone. In this context, the implications for the use of community‐weighted mean trait values are discussed, as not only species abundances change in response to changing environmental conditions, but also values of traits considerably change. Including this aspect in further studies will improve our understanding of processes acting within and among communities.     

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.     

Structural and functional responses of invertebrate communities to climate change and flow regulation in alpine catchments

Understanding and predicting how biological communities respond to climate change is critical for assessing biodiversity vulnerability and guiding conservation efforts. Glacier‐ and snow‐fed rivers are one of the most sensitive ecosystems to climate change, and can provide early warning of wider‐scale changes. These rivers are frequently used for hydropower production but there is minimal understanding of how biological communities are influenced by climate change in a context of flow regulation. This study sheds light on this issue by disentangling structural (water temperature preference, taxonomic composition, alpha, beta and gamma diversities) and functional (functional traits, diversity, richness, evenness, dispersion and redundancy) effects of climate change in interaction with flow regulation in the Alps. For this, we compared environmental and aquatic invertebrate data collected in the 1970s and 2010s in regulated and unregulated alpine catchments. We hypothesized a replacement of cold‐adapted species by warming‐tolerant ones, high temporal and spatial turnover in taxa and trait composition, along with reduced taxonomic and functional diversities in consequence of climate change. We expected communities in regulated rivers to respond more drastically due to additive or synergistic effects between flow regulation and climate change. We found divergent structural but convergent functional responses between free‐flowing and regulated catchments. Although cold‐adapted taxa decreased in both of them, greater colonization and spread of thermophilic species was found in the free‐flowing one, resulting in higher spatial and temporal turnover. Since the 1970s, taxonomic diversity increased in the free flowing but decreased in the regulated catchment due to biotic homogenization. Colonization by taxa with new functional strategies (i.e. multivoltine taxa with small body size, resistance forms, aerial dispersion and reproduction by clutches) increased functional diversity but decreased functional redundancy through time. These functional changes could jeopardize the ability of aquatic communities facing intensification of ongoing climate change or new anthropogenic disturbances.     

Impact of climate change on alpine vegetation of mountain summits in Norway

Climate change is affecting the composition and functioning of ecosystems across the globe. Mountain ecosystems are particularly sensitive to climate warming since their biota is generally limited by low temperatures. Cryptogams such as lichens and bryophytes are important for the biodiversity and functioning of these ecosystems, but have not often been incorporated in vegetation resurvey studies. Hence, we lack a good understanding of how vascular plants, lichens and bryophytes respond interactively to climate warming in alpine communities. Here we quantified long-term changes in species richness, cover, composition and thermophilization (i.e. the increasing dominance of warm-adapted species) of vascular plants, lichens and bryophytes on four summits at Dovrefjell, Norway. These summits are situated along an elevational gradient from the low alpine to high alpine zone and were surveyed for all species in 2001, 2008 and 2015. During the 15-year period, a decline in lichen richness and increase in bryophyte richness was detected, whereas no change in vascular plant richness was found. Dwarf-shrub abundance progressively increased at the expense of lichens, and thermophilization was most pronounced for vascular plants, but occurred only on the lowest summits and northern aspects. Lichens showed less thermophilization and, for the bryophytes, no significant thermophilization was found. Although recent climate change may have primarily caused the observed changes in vegetation, combined effects with non-climatic factors (e.g. grazing and trampling) are likely important as well. At a larger scale, alpine vegetation shifts could have a profound impact on biosphere functioning with feedbacks to the global climate.     

Legacy effects and memory loss: how contingencies moderate the response of rocky intertidal biofilms to present and past extreme events

Understanding how historical processes modulate the response of ecosystems to perturbations is becoming increasingly important. In contrast to the growing interest in projecting biodiversity and ecosystem functioning under future climate scenarios, how legacy effects originating from historical conditions drive change in ecosystems remains largely unexplored. Using experiments in combination with stochastic antecedent modelling, we evaluated how extreme warming, sediment deposition and grazing events modulated the ecological memory of rocky intertidal epilithic microphytobenthos (EMPB). We found memory effects in the non‐clustered scenario of disturbance (60 days apart), where EMPB biomass fluctuated in time, but not under clustered disturbances (15 days apart), where EMPB biomass was consistently low. A massive grazing event impacted on EMPB biomass in a second run of the experiment, also muting ecological memory. Our results provide empirical support to the theoretical expectation that stochastic fluctuations promote ecological memory, but also show that contingencies may lead to memory loss.     

Nurse species and indirect facilitation through grazing drive plant community functional traits in tropical alpine peatlands

Facilitation among plants mediated by grazers occurs when an unpalatable plant extends its protection against grazing to another plant. This type of indirect facilitation impacts species coexistence and ecosystem functioning in a large array of ecosystems worldwide. It has nonetheless generally been understudied so far in comparison with the role played by direct facilitation among plants. We aimed at providing original data on indirect facilitation at the community scale to determine the extent to which indirect facilitation mediated by grazers can shape plant communities. Such experimental data are expected to contribute to refining the conceptual framework on plant–plant–herbivore interactions in stressful environments. We set up a 2‐year grazing exclusion experiment in tropical alpine peatlands in Bolivia. Those ecosystems depend entirely on a few, structuring cushion‐forming plants (hereafter referred to as “nurse” species), in which associated plant communities develop. Fences have been set over two nurse species with different strategies to cope with grazing (direct vs. indirect defenses), which are expected to lead to different intensities of indirect facilitation for the associated communities. We collected functional traits which are known to vary according to grazing pressure (LDMC, leaf thickness, and maximum height), on both the nurse and their associated plant communities in grazed (and therefore indirect facilitation as well) and ungrazed conditions. We found that the effect of indirectly facilitated on the associated plant communities depended on the functional trait considered. Indirect facilitation decreased the effects of grazing on species relative abundance, mean LDMC, and the convergence of the maximum height distribution of the associated communities, but did not affect mean height or cover. The identity of the nurse species and grazing jointly affected the structure of the associated plant community through indirect facilitation. Our results together with the existing literature suggest that the “grazer–nurse–beneficiary” interaction module can be more complex than expected when evaluated in the field.     

A trait-based framework for predicting when and where microbial adaptation to climate change will affect ecosystem functioning

As the earth system changes in response to human activities, a critical objective is to predict how biogeochemical process rates (e.g. nitrification, decomposition) and ecosystem function (e.g. net ecosystem productivity) will change under future conditions. A particular challenge is that the microbial communities that drive many of these processes are capable of adapting to environmental change in ways that alter ecosystem functioning. Despite evidence that microbes can adapt to temperature, precipitation regimes, and redox fluctuations, microbial communities are typically not optimally adapted to their local environment. For example, temperature optima for growth and enzyme activity are often greater than in situ temperatures in their environment. Here we discuss fundamental constraints on microbial adaptation and suggest specific environments where microbial adaptation to climate change (or lack thereof) is most likely to alter ecosystem functioning. Our framework is based on two principal assumptions. First, there are fundamental ecological trade-offs in microbial community traits that occur across environmental gradients (in time and space). These trade-offs result in shifting of microbial function (e.g. ability to take up resources at low temperature) in response to adaptation of another trait (e.g. limiting maintenance respiration at high temperature). Second, the mechanism and level of microbial community adaptation to changing environmental parameters is a function of the potential rate of change in community composition relative to the rate of environmental change. Together, this framework provides a basis for developing testable predictions about how the rate and degree of microbial adaptation to climate change will alter biogeochemical processes in aquatic and terrestrial ecosystems across the planet.     

Does rapid evolution matter? Measuring the rate of contemporary evolution and its impacts on ecological dynamics

Rapid contemporary evolution due to natural selection is common in the wild, but it remains uncertain whether its effects are an essential component of community and ecosystem structure and function. Previously we showed how to partition change in a population, community or ecosystem property into contributions from environmental and trait change, when trait change is entirely caused by evolution (Hairston et al. 2005). However, when substantial non-heritable trait change occurs (e.g. due to phenotypic plasticity or change in population structure) that approach can mis-estimate both contributions. Here, we demonstrate how to disentangle ecological impacts of evolution vs. non-heritable trait change by combining our previous approach with the Price Equation. This yields a three-way partitioning into effects of evolution, non-heritable phenotypic change and environment. We extend the approach to cases where ecological consequences of trait change are mediated through interspecific interactions. We analyse empirical examples involving fish, birds and zooplankton, finding that the proportional contribution of rapid evolution varies widely (even among different ecological properties affected by the same trait), and that rapid evolution can be important when it acts to oppose and mitigate phenotypic effects of environmental change. Paradoxically, rapid evolution may be most important when it is least evident.     

The influence of fire on phylogenetic and functional structure of woody savannas: Moving from species to individuals

Fire is a key determinant of tropical savanna structure and functioning. High fire frequencies are expected to assemble closely related species with a restricted range of functional trait values. Here we determined the effect of fire on phylogenetic and functional diversity of woody species and individuals in savanna communities under different fire frequencies. We found phylogenetic signals for one third of the functional traits studied. High numbers of fires simultaneously led to phylogenetic overdispersion and functional clustering when communities were represented by mean trait values with all traits that putatively should be affected or respond to fire. This finding is important, because it shows that the relationship between ecological processes and the phylogenetic structure of communities is not straightforward. Thus, we cannot always assume that close relatives are more similar in their ecological features. However, when considering a different set of traits representing different plant strategies (fire resistance/avoidance, physiological traits and regeneration traits), the results were not always congruent. When asking how communities are assembled in terms of individuals (not species) the outcome was different from the species-based approach, suggesting that the realised trait values – rather than mean species trait values – have an important role in driving community assembly. Thus, intraspecific trait variability should be taken into account if we want fully to improve our mechanistic understanding of assembly rules in plant communities.     

Community functional trait composition at the continental scale: the effects of non‐ecological processes

Ecological communities and their response to environmental gradients are increasingly being described by measures of trait composition at the community level – the trait‐based approach. Whether ecological or non‐ecological processes influence trait composition between communities has been debated. Understanding the processes that influence trait composition is important for reconstructing paleoenvironmental conditions from fossil deposits and for understanding changes in community functionality through time. Here, we assess the influence of ecological and non‐ecological processes on the distribution of traits within North American mammals. We found that non‐ecological processes including historical contingency, spatial autocorrelation, and evolutionary history do not influence trait composition; however, the variance in trait composition is highly explained by climate gradients. Our results suggest that habitat breadth, terrestriality, diet breadth, and reproductive traits are strong candidates as proxies for measuring functional aspects of environments in the past and present.     

Consistent physiological,ecological and evolutionary effects of fire regime on conservative leaf economics strategies in plant communities

The functional response of plant communities to disturbance is hypothesised to be controlled by changes in environmental conditions and evolutionary history of species within the community. However, separating these influences using direct manipulations of repeated disturbances within ecosystems is rare. We evaluated how 41 years of manipulated fire affected plant leaf economics by sampling 89 plant species across a savanna-forest ecotone. Greater fire frequencies created a high-light and low-nitrogen environment, with more diverse communities that contained denser leaves and lower foliar nitrogen content. Strong trait–fire coupling resulted from the combination of significant intraspecific trait–fire correlations being in the same direction as interspecific trait differences arising through the turnover in functional composition along the fire-frequency gradient. Turnover among specific clades helped explain trait–fire trends, but traits were relatively labile. Overall, repeated burning led to reinforcing selective pressures that produced diverse plant communities dominated by conservative resource-use strategies and slow soil nitrogen cycling.     

Fish communities diverge in species but converge in traits over three decades of warming

Describing the spatial and temporal dynamics of communities is essential for understanding the impacts of global environmental change on biodiversity and ecosystem functioning. Trait‐based approaches can provide better insight than species‐based (i.e. taxonomic) approaches into community assembly and ecosystem functioning, but comparing species and trait dynamics may reveal important patterns for understanding community responses to environmental change. Here, we used a 33‐year database of fish monitoring to compare the spatio‐temporal dynamics of taxonomic and trait structure in North Sea fish communities. We found that the majority of variation in both taxonomic and trait structure was explained by a pronounced spatial gradient, with distinct communities in the southern and northern North Sea related to depth, sea surface temperature, salinity and bed shear stress. Both taxonomic and trait structure changed significantly over time; however taxonomically, communities in the south and north diverged towards different species, becoming more dissimilar over time, yet they converged towards the same traits regardless of species differences. In particular, communities shifted towards smaller, faster growing species with higher thermal preferences and pelagic water column position. Although taxonomic structure changed over time, its spatial distribution remained relatively stable, whereas in trait structure, the southern zone of the North Sea shifted northward and expanded, leading to homogenization. Our findings suggest that global environmental change, notably climate warming, will lead to convergence towards traits more adapted for novel environments regardless of species composition.     

Evolutionary community ecology of plant-associated arthropods in terrestrial ecosystems

In the 21st century, researchers have attempted a synthesis between community ecology and evolutionary biology. This emerging research area, which aims to synthesize community ecology and evolutionary biology, is evolutionary community ecology. Evolutionary community ecology addresses how intraspecific trait variation in community members is essential for predicting community properties and, how community properties are a key component of the selective forces that determine genetic and phenotypic variation in a community member. In this paper, I review recent findings in evolutionary community ecology in plant-associated arthropods in terrestrial ecosystems. I discuss roles of both genetic variation and phenotypic plasticity as a source of trait variation in plants in shaping plant-associated arthropod communities. Also, I discuss effects of genetic variation in herbivores on plant-associated arthropod communities. Furthermore, I highlight community context evolution in which multiple species interactions and community composition affect trait evolution of a community member. Finally, I argue that future studies should investigate a feedback loop between community and evolutionary dynamics beyond unidirectional studies on effects of evolution on a community or vice versa. This approach will provide major insights into mechanistic principles for making predictions of community ecology.