Grégoire T. Freschet

A global meta‐analysis of the relative extent of intraspecific trait variation in plant communities

Recent studies have shown that accounting for intraspecific trait variation (ITV) may better address major questions in community ecology. However, a general picture of the relative extent of ITV compared to interspecific trait variation in plant communities is still missing. Here, we conducted a meta-analysis of the relative extent of ITV within and among plant communities worldwide, using a data set encompassing 629 communities (plots) and 36 functional traits. Overall, ITV accounted for 25% of the total trait variation within communities and 32% of the total trait variation among communities on average. The relative extent of ITV tended to be greater for whole-plant (e.g. plant height) vs. organ-level traits and for leaf chemical (e.g. leaf N and P concentration) vs. leaf morphological (e.g. leaf area and thickness) traits. The relative amount of ITV decreased with increasing species richness and spatial extent, but did not vary with plant growth form or climate. These results highlight global patterns in the relative importance of ITV in plant communities, providing practical guidelines for when researchers should include ITV in trait-based community and ecosystem studies.

Linking litter decomposition of above‐ and below‐ground organs to plant–soil feedbacks worldwide

Conceptual frameworks relating plant traits to ecosystem processes such as organic matter dynamics are progressively moving from a leaf-centred to a whole-plant perspective. Through the use of meta-analysis and global literature data, we quantified the relative roles of litters from above- and below-ground plant organs in ecosystem labile organic matter dynamics. We found that decomposition rates of leaves, fine roots and fine stems were coordinated across species worldwide although less strongly within ecosystems. We also show that fine roots and stems had lower decomposition rates relative to leaves, with large differences between woody and herbaceous species. Further, we estimated that on average below-ground litter represents approximately 33 and 48% of annual litter inputs in grasslands and forests, respectively. These results suggest a major role for below-ground litter as a driver of ecosystem organic matter dynamics. We also suggest that, given that fine stem and fine root litters decompose approximately 1.5 and 2.8 times slower, respectively, than leaf litter derived from the same species, cycling of labile organic matter is likely to be much slower than predicted by data from leaf litter decomposition only. Synthesis. Our results provide evidence that within ecosystems, the relative inputs of above- versus below-ground litter strongly control the overall quality of the litter entering the decomposition system. This in turn determines soil labile organic matter dynamics and associated nutrient release in the ecosystem, which potentially feeds back to the mineral nutrition of plants and therefore plant trait values and plant community composition.

Contrasting effects of plant inter‐ and intraspecific variation on community‐level trait measures along an environmental gradient

Summary 1. Despite widespread focus on interspecific variation in trait-based ecology, there is growing evidence that intraspecific trait variability can play a fundamental role in plant community responses to environmental change and community assembly. 2. Here, we quantify the strength and direction of inter- and intraspecific plant community trait responses along a 900 m elevation gradient spanning alpine and subalpine plant communities in southern New Zealand. We measured five commonly used leaf traits (i.e. dry matter content, N and P concentrations, leaf area and specific leaf area) on all 31 dominant and subordinate species recorded along the gradient, and examined their species-specific and community-level responses to elevation using both abundance-weighted and nonweighted averages of trait values. 3. By decomposing the variance of community-level measures of these traits across the gradient, we showed that the contribution of interspecific variation to the response of plant assemblages to elevation was stronger than that of intraspecific variation, for all traits except specific leaf area. Further, the relative contributions of interspecific effects were greater when abundance-weighted rather than nonweighted measures were used. We also observed contrasting intraspecific trait responses to the gradient among species (particularly for leaf N and P concentrations), and found both positive and negative covariation between inter- and intraspecific effects on community-level trait values. 4. The weak community-average trait responses to elevation, as found for specific leaf area (SLA) and leaf N and P concentrations, resulted from strong but opposing responses among vs. within species, which are not typically accounted for in species-based measures of plant community responses. For instance, increasing elevation (and associated factors such as a decrease in soil nutrient availability) favoured the dominance of species with relatively high leaf nutrient concentrations while simultaneously triggering an intraspecific decrease in the leaf nutrient concentrations of these species. 5. The context dependency of positive and negative covariation between inter- and intraspecific trait variability, and the species-specific nature of intraspecific shifts in functional trait values, reveal highly complex plastic responses of plants to environmental changes, and highlights the need for greater consideration of the role that intraspecific variation plays in community-level processes.

Coordinated variation in leaf and root traits across multiple spatial scales in Chinese semi-arid and arid ecosystems.

• Variation in plant functional traits is the product of evolutionary and environmental drivers operating at different scales. Little is known about whether, or how, this variation is coordinated between aboveground and belowground organs across and within spatial scales. • We address these questions using a hierarchically designed dataset of pairwise leaf and root traits related to carbon and nutrient economy of 64 species belonging to 14 plant communities in northern Chinese semi-arid and arid regions. • While both root and leaf traits showed most of their variance among (individuals and) species within communities, leaf trait variance tended to be relatively higher at coarser spatial scales than root trait variance. While leaf nitrogen (N) per area to root N per length ratio increased and specific leaf area to specific root length and leaf [N] to root [N] ratios decreased from semi-arid to arid environments owing to climatic/edaphic shifts, the matching pairs showed a strong pattern of positive correlation that was upheld across spatial scales and geographic areas. • Thus, trade-offs in plant resource investment across organs within individual vascular plants are constrained within a rather narrow range of variation. A new challenge will be to test whether and how such trait coordination is also seen within and across other biomes of the world.

Integrated plant phenotypic responses to contrasting above- and below-ground resources: key roles of specific leaf area and root mass fraction

Plants adapt phenotypically to different conditions of light and nutrient supply, supposedly in order to achieve colimitation of these resources. Their key variable of adjustment is the ratio of leaf area to root length, which relies on plant biomass allocation and organ morphology. We recorded phenotypic differences in leaf and root mass fractions (LMF, RMF), specific leaf area (SLA) and specific root length (SRL) of 12 herbaceous species grown in factorial combinations of high/low irradiance and fertilization treatments. Leaf area and root length ratios, and their components, were influenced by nonadditive effects between light and nutrient supply, and differences in the strength of plant responses were partly explained by Ellenbergs species values representing ecological optima. Changes in allocation were critical in plant responses to nutrient availability, as the RMF contribution to changes in root length was 2.5× that of the SRL. Contrastingly, morphological adjustments (SLA rather than LMF) made up the bulk of plant response to light availability. Our results suggest largely predictable differences in responses of species and groups of species to environmental change. Nevertheless, they stress the critical need to account for adjustments in below-ground mass allocation to understand the assembly and responses of communities in changing environments.

Controls on Coarse Wood Decay in Temperate Tree Species: Birth of the LOGLIFE Experiment

Dead wood provides a huge terrestrial carbon stock and a habitat to wide-ranging organisms during its decay. Our brief review highlights that, in order to understand environmental change impacts on these functions, we need to quantify the contributions of different interacting biotic and abiotic drivers to wood decomposition. LOGLIFE is a new long-term ‘common-garden’ experiment to disentangle the effects of species’ wood traits and site-related environmental drivers on wood decomposition dynamics and its associated diversity of microbial and invertebrate communities. This experiment is firmly rooted in pioneering experiments under the directorship of Terry Callaghan at Abisko Research Station, Sweden. LOGLIFE features two contrasting forest sites in the Netherlands, each hosting a similar set of coarse logs and branches of 10 tree species. LOGLIFE welcomes other researchers to test further questions concerning coarse wood decay that will also help to optimise forest management in view of carbon sequestration and biodiversity conservation.

Plasticity in above‐ and belowground resource acquisition traits in response to single and multiple environmental factors in three tree species

Functional trait plasticity is a major component of plant adjustment to environmental stresses. Here, we explore how multiple local environmental gradients in resources required by plants (light, water, and nutrients) and soil disturbance together influence the direction and amplitude of intraspecific changes in leaf and fine root traits that facilitate capture of these resources. We measured population-level analogous above- and belowground traits related to resource acquisition, i.e. “specific leaf area”–“specific root length” (SLA–SRL), and leaf and root N, P, and dry matter content (DMC), on three dominant understory tree species with contrasting carbon and nutrient economics across 15 plots in a temperate forest influenced by burrowing seabirds. We observed similar responses of the three species to the same single environmental influences, but partially species-specific responses to combinations of influences. The strength of intraspecific above- and belowground trait responses appeared unrelated to species resource acquisition strategy. Finally, most analogous leaf and root traits (SLA vs. SRL, and leaf versus root P and DMC) were controlled by contrasting environmental influences. The decoupled responses of above- and belowground traits to these multiple environmental factors together with partially species-specific adjustments suggest complex responses of plant communities to environmental changes, and potentially contrasting feedbacks of plant traits with ecosystem properties. We demonstrate that despite the growing evidence for broadly consistent resource-acquisition strategies at the whole plant level among species, plants also show partially decoupled, finely tuned strategies between above- and belowground parts at the intraspecific level in response to their environment. This decoupling within species suggests a need for many species-centred ecological theories on how plants respond to their environments (e.g. competitive/stress-tolerant/ruderal and response-effect trait frameworks) to be adapted to account for distinct plant-environment interactions among distinct individuals of the same species and parts of the same individual.

Climate, soil and plant functional types as drivers of global fine‐root trait variation

Summary Ecosystem functioning relies heavily on below-ground processes, which are largely regulated by plant fine-roots and their functional traits. However, our knowledge of fine-root trait distribution relies to date on local- and regional-scale studies with limited numbers of species, growth forms and environmental variation. We compiled a world-wide fine-root trait dataset, featuring 1115 species from contrasting climatic areas, phylogeny and growth forms to test a series of hypotheses pertaining to the influence of plant functional types, soil and climate variables, and the degree of manipulation of plant growing conditions on species fine-root trait variation. Most particularly, we tested the competing hypotheses that fine-root traits typical of faster return on investment would be most strongly associated with conditions of limiting versus favourable soil resource availability. We accounted for both data source and species phylogenetic relatedness. We demonstrate that: (i) Climate conditions promoting soil fertility relate negatively to fine-root traits favouring fast soil resource acquisition, with a particularly strong positive effect of temperature on fine-root diameter and negative effect on specific root length (SRL), and a negative effect of rainfall on root nitrogen concentration; (ii) Soil bulk density strongly influences species fine-root morphology, by favouring thicker, denser fine-roots; (iii) Fine-roots from herbaceous species are on average finer and have higher SRL than those of woody species, and N2-fixing capacity positively relates to root nitrogen; and (iv) Plants growing in pots have higher SRL than those grown in the field. Synthesis. This study reveals both the large variation in fine-root traits encountered globally and the relevance of several key plant functional types and soil and climate variables for explaining a substantial part of this variation. Climate, particularly temperature, and plant functional types were the two strongest predictors of fine-root trait variation. High trait variation occurred at local scales, suggesting that wide-ranging below-ground resource economics strategies are viable within most climatic areas and soil conditions.

Increasing soil carbon storage: mechanisms, effects of agricultural practices and proxies. A review

The international 4 per 1000 initiative aims at supporting states and non-governmental stakeholders in their efforts towards a better management of soil carbon (C) stocks. These stocks depend on soil C inputs and outputs. They are the result of fine spatial scale interconnected mechanisms, which stabilise/destabilise organic matter-borne C. Since 2016, the CarboSMS consortium federates French researchers working on these mechanisms and their effects on C stocks in a local and global change setting (land use, agricultural practices, climatic and soil conditions, etc.). This article is a synthesis of this consortium’s first seminar. In the first part, we present recent advances in the understanding of soil C stabilisation mechanisms comprising biotic and abiotic processes, which occur concomitantly and interact. Soil organic C stocks are altered by biotic activities of plants (the main source of C through litter and root systems), microorganisms (fungi and bacteria) and ‘ecosystem engineers’ (earthworms, termites, ants). In the meantime, abiotic processes related to the soil-physical structure, porosity and mineral fraction also modify these stocks. In the second part, we show how agricultural practices affect soil C stocks. By acting on both biotic and abiotic mechanisms, land use and management practices (choice of plant species and density, plant residue exports, amendments, fertilisation, tillage, etc.) drive soil spatiotemporal organic inputs and organic matter sensitivity to mineralisation. Interaction between the different mechanisms and their effects on C stocks are revealed by meta-analyses and long-term field studies. The third part addresses upscaling issues. This is a cause for major concern since soil organic C stabilisation mechanisms are most often studied at fine spatial scales (mm–μm) under controlled conditions, while agricultural practices are implemented at the plot scale. We discuss some proxies and models describing specific mechanisms and their action in different soil and climatic contexts and show how they should be taken into account in large scale models, to improve change predictions in soil C stocks. Finally, this literature review highlights some future research prospects geared towards preserving or even increasing C stocks, our focus being put on the mechanisms, the effects of agricultural practices on them and C stock prediction models.

Aboveground and belowground legacies of native Sami land use on boreal forest in northern Sweden 100 years after abandonment

Human activities that involve land-use change often cause major transformations to community and ecosystem properties both aboveground and belowground, and when land use is abandoned, these modifications can persist for extended periods. However, the mechanisms responsible for rapid recovery vs. long-term maintenance of ecosystem changes following abandonment remain poorly understood. Here, we examined the long-term ecological effects of two remote former settlements, regularly visited for -300 years by reindeer-herding Sami and abandoned -100 years ago, within an old-growth boreal forest that is considered one of the most pristine regions in northern Scandinavia. These human legacies were assessed through measurements of abiotic and biotic soil properties and vegetation characteristics at the settlement sites and at varying distances from them. Low-intensity land use by Sami is characterized by the transfer of organic matter towards the settlements by humans and reindeer herds, compaction of soil through trampling, disappearance of understory vegetation, and selective cutting of pine trees for fuel and construction. As a consequence, we found a shift towards early successional plant species and a threefold increase in soil microbial activity and nutrient availability close to the settlements relative to away from them. These changes in soil fertility and vegetation contributed to 83% greater total vegetation productivity, 35% greater plant biomass, and 23% and 16% greater concentrations of foliar N and P nearer the settlements, leading to a greater quantity and quality of litter inputs. Because decomposer activity was also 40% greater towards the settlements, soil organic matter cycling and nutrient availability were further increased, leading to likely positive feedbacks between the aboveground and belowground components resulting from historic land use. Although not all of the activities typical of Sami have left visible residual traces on the ecosystem after 100 years, their low-intensity but long-term land use at settlement sites has triggered a rejuvenation of the ecosystem that is still present. Our data demonstrates that aboveground-belowground interactions strongly control ecosystem responses to historical human land use and that medium- to long-term consequences of even low-intensity human activities must be better accounted for if we are to predict and manage ecosystems succession following land-use abandonment.