Pierre Mariotte

Subordinate plant species enhance community resistance against drought in semi‐natural grasslands

According to the insurance hypothesis, more diverse plant communities are more likely to be resistant to drought. Whilst many experiments have been carried out to determine the effects of plant diversity on plant community insurance, the results are still contradictory. Here, we conducted a drought experiment where we tested whether the presence of subordinate species increases plant community insurance. In Swiss Jura grassland, we combined a removal experiment of subordinate species with a summer drought event using rainout shelters. Plant community composition was determined after the drought and based on biomass measurements; we estimated resistance, recovery and resilience of the plant community for each combination of treatments. Moreover, to assess drought impacts on water-use efficiency (WUE), we analysed carbon isotope ratios (13C values) in plant leaves of two dominants and two subordinates collected at the end of the drought period. We showed that subordinate species are more resistant to drought and increased community resistance by enhancing their above-ground biomass production during the imposed drought. These patterns were associated with decreased competitiveness of dominant species whose biomass decreased during drought. Significant increase in 13C values in plant tissue under drought indicated a better WUE for the measured species. Interestingly, the WUE was significantly higher in plots where subordinates were removed. Recovery and resilience were not affected by the summer drought, but the absence of subordinates reduced overall above-ground biomass in both watered and drought plots. Synthesis. We demonstrated that, independent of plant diversity, the presence of drought-resistant subordinate species increases plant community insurance against drought and, hence, is important for the functioning of grassland ecosystems.

Do subordinate species punch above their weight? Evidence from above‐ and below‐ground

Variation in the relative abundance of species is a ubiquitous feature of ecological communities. Understanding the link between the relative abundance of a species and its contribution to ecosystem function is key to predicting ecosystem stability over time or during perturbations. According to the ‘mass ratio hypothesis’ (Grime, 1998), proportional inputs to primary production act as immediate controls on ecosystem function and sustainability.While dominant species are consideredmore important in ecosystems because of the large amount of biomass they produce, an increasing number of recent studies have shown that subordinate species may have a larger influence on ecosystem functioning than their relative abundance suggests. In particular, subordinate species can have significant impacts on soil microbial communities (Peltzer et al., 2009; Holdaway et al., 2011; Mariotte et al., 2013d). Growing recognition of how belowground processes affect biodiversity, ecosystem functioning and services (De Vries et al., 2013; Grigulis et al., 2013; Van der Putten et al., 2013) likely explains the recent interest in less abundant species. Nevertheless, studies of subordinate species remain scarce, largely because of the influence of the ‘mass ratio hypothesis’, which likely overestimates the role of dominant species, as well as a lack of adequate understanding of the distinguishing features of subordinate species. In addition to differences in relative abundance, other characteristics may differ between dominant and subordinate species including functional groups, traits, resource acquisition strategies, and spatial growth. While these differences remain poorly understood, their implications could be far reaching if subordinate species are capable of buffering ecosystem functions against perturbations, and under climate change. This paper aims to reconcile theories about subordinate species in order to provide a framework for future studies of biodiversity effects on ecosystem functioning. I synthesize the current state of knowledge about subordinate species and give a clear definition for these species, I provide evidence of their functional role in ecosystems, and I show how the importance of this group may increase with global climate change; a topic which has not been addressed in the literature. I address each of these issues by including an aboveand below-ground perspective, which is often missing from studies of functional diversity. While evidence is drawn mainly from experiments in grassland communities, the basic principles can be applied more widely. Reconciling theories of subordinate species

Arbuscular mycorrhizal fungi reduce the differences in competitiveness between dominant and subordinate plant species

In grassland communities, plants can be classified as dominants or subordinates according to their relative abundances, but the factors controlling such distributions remain unclear. Here, we test whether the presence of the arbuscular mycorrhizal (AM) fungus Glomus intraradices affects the competitiveness of two dominant (Taraxacum officinale and Agrostis capillaris) and two subordinate species (Prunella vulgaris and Achillea millefolium). Plants were grown in pots in the presence or absence of the fungus, in monoculture and in mixtures of both species groups with two and four species. In the absence of G. intraradices, dominants were clearly more competitive than subordinates. In inoculated pots, the fungus acted towards the parasitic end of the mutualism–parasitism continuum and had an overall negative effect on the growth of the plant species. However, the negative effects of the AM fungus were more pronounced on dominant species reducing the differences in competitiveness between dominant and subordinate species. The effects of G. intraradices varied with species composition highlighting the importance of plant community to mediate the effects of AM fungi. Dominant species were negatively affected from the AM fungus in mixtures, while subordinates grew identically with and without the fungus. Therefore, our findings predict that the plant dominance hierarchy may flatten out when dominant species are more reduced than subordinate species in an unfavourable AM fungal relationship (parasitism).

Land management trumps the effects of climate change and elevated CO2 on grassland functioning

1. Grasslands cover similar to 30% of the Earths terrestrial surface and provide many ecosystem services. Many grasslands are heavily managed to maximize these services for human benefit, but the outcome of management is anticipated to be increasingly influenced by various aspects of climate change and elevated atmospheric CO2. The relative importance of global change vs. land management on grasslands is largely unknown. 2. A meta-analysis is used here to examine drivers at both scales primarily targeting services provided by grasslands relating to plant productivity (above- and below-ground biomass) and soil processes (nutrients and soil respiration) in 38 manipulative experiments published in the last decade. We specifically target effects of (i) single and combined land management practices (LMs), (ii) single and combined factors relating to broad-scale climate change and elevated CO2, and (iii) combined management practices and changes to climate and CO2. Collectively, this examines the general efficacy of global change models in predicting changes to grassland functioning. 3. We found that combinations of management practices had approximately double the explanatory power for variation in grassland services compared with individual or interactive effects of factors associated with climate change and CO2. These interacting management practices such as nutrient additions and defoliation predominantly influenced functions associated with productivity or biomass both below and above ground. The effects of interacting factors of climate and CO2 influenced a wider range of ecosystem functions, but the magnitude of these effects was relatively smaller. 4. Interactions between management practices or between climate change/CO2 factors always had higher explanatory power than any factor in isolation indicating that multivariate synergistic models of environmental change can better describe impacts on ecosystem function in plant communities (e.g. relative to univariate climate-based models). Given that the magnitude and direction (positive or negative) of the interactions varied widely, this also implies that the outcomes of these multivariate interactions can vary spatially, temporally or by immediate context (e.g. management prescriptions). 5. Synthesis. Although our work confirms how climate change and CO2 can affect many ecosystem-based functional attributes, it suggests that combinations of LMs remain the dominant set of factors in determining the performance of grassland plant communities. Land management may thus be critical for influencing projected responses to future climate change and elevated CO2 in models of grassland function at least for factors relating to primary production.

Stoichiometric N:P flexibility and mycorrhizal symbiosis favour plant resistance against drought

1. Drought induces changes in the nitrogen (N) and phosphorus (P) cycle but most plant species have limited flexibility to take up nutrients under such variable or unbalanced N and P availability. Both the degree of flexibility in plant N:P ratio and of root symbiosis with arbuscular mycorrhizal fungi might control plant resistance to drought-induced changes in nutrient availability, but this has not been directly tested. 2. Here, we examined the role of plant N:P stoichiometric status and mycorrhizal symbiosis in the drought-resistance of dominant and subordinate species in a semi-natural grassland. 3. We reduced water availability using rainout shelters (control vs. drought) and measured how plant biomass responded for the dominant and subordinate species. We then selected a dominant (Paspalum dilatatum) and a subordinate species (Cynodon dactylon), for which we investigated the N:P stoichiometric status, mycorrhizal root colonization and water-use efficiency. 4. The biomass of all dominant plant species, but not subordinate species, decreased under drought. Drought increased soil available nitrogen, and thus increased soil N:P ratio, due to decreasing plant N uptake. The dominant P. dilatatum showed a high degree of plant N:P homeostasis and a considerable reduction in biomass under drought. At the opposite, the more flexible subordinate species C. dactylon increased its N uptake and water-use efficiency, apparently due to stronger symbiosis with mycorrhizae, and maintained its biomass. 5. Synthesis. We conclude that the maintenance of N:P homeostasis in dominant species, possibly because of a large root nutrient foraging capacity, becomes inefficient when water stress limits N mobility in the soil. By contrast, we demonstrate that higher stoichiometric N:P flexibility coupled with stronger mutualistic association with mycorrhizae allow subordinate species to better withstand drought perturbations. Using a stoichiometric approach in a field experiment, our study provides for the first time clear and novel understandings of the mechanisms involved in drought-resistance within the plant-mycorrhizae-soil system.

Cattle as Dispersal Vectors of Invasive and Introduced Plants in a California Annual Grassland.

ABSTRACT Plant invasions are a threat to rangelands in California. Understanding how seeds of invasive plants are dispersed is critical to developing sound management plans. Domestic livestock can transport seeds long distances by ingesting and passing seeds in dung (endozoochory) or by the attachment of seeds to skin and fur (epizoochory). Our objective was to characterize the role of cattle as seed dispersers of both invasive and noninvasive species via endozoochory and epizoochory in a Sierra foothills rangeland. To quantify endozoochory, we sampled dung from two dry-season grazing periods and evaluated seed content by growing dung for 3 months in a greenhouse. To quantify epizoochory, we collected seeds directly from the fur of 40 cattle. We categorized the invasion status and functional groups of all species found and quantified landscape-scale vegetation composition in order to determine whether dispersal mode was associated with functional group, invasion status, or vegetation composition. Finally, we evaluated the potential for the noxious weed medusahead (Taeniatherum caput-medusae [L.] Nevski) to travel long distances on cattle fur using a detachment experiment with a model cow. We found that forbs were more likely to be dispersed by endozoochory, and invasive species were more likely to be dispersed by epizoochory. Medusahead was dispersed exclusively by epizoochory and was able to travel up to 160 m on a model cow. Our results suggest that cattle may be an important dispersal vector for both invasive and noninvasive plants.

Subordinate plants mitigate drought effects on soil ecosystem processes by stimulating fungi

1. The subordinate insurance hypothesis suggests that highly diverse communities contain greater numbers of subordinate species than less diverse communities. It has previously been reported that subordinate species can improve grassland productivity during drought, but the underlying mechanisms remain undetermined. 2. Using a combination of subordinate species removal and summer drought, we show that soil processes play a critical role in community resistance to drought. Interestingly, subordinate species drive soil microbial community structure and largely mitigate the effect of drought on grassland soil functioning. Our results highlight subordinate species in shifting the balance within the phospholipid fatty acid (PLFA) microbial community towards more fungal dominance. 3. Fungal communities promoted by subordinate species were more resistant to drought and maintained higher rates of litter decomposition and soil respiration. These results emphasize the important role of subordinate species in mitigating drought effects on soil ecosystem functions. Reciprocal effects between fungi and subordinate species explain also how subordinate species better resisted to drought conditions. 4. Our results point to a delayed plant-soil feedback following environmental perturbation. Additionally, they extend the diversity insurance hypothesis by showing that more diverse communities not only contain species well adapted to perturbations, but also species with higher impacts on soil microbial communities and related ecosystem functions.

Plant–Soil Feedback: Bridging Natural and Agricultural Sciences

In agricultural and natural systems researchers have demonstrated large effects of plant-soil feedback (PSF) on plant growth. However, the concepts and approaches used in these two types of systems have developed, for the most part, independently. Here, we present a conceptual framework that integrates knowledge and approaches from these two contrasting systems. We use this integrated framework to demonstrate (i) how knowledge from complex natural systems can be used to increase agricultural resource-use efficiency and productivity and (ii) how research in agricultural systems can be used to test hypotheses and approaches developed in natural systems. Using this framework, we discuss avenues for new research toward an ecologically sustainable and climate-smart future.

Separating sources of density‐dependent and density‐independent establishment limitation in invading species

Summary Successful colonization by invasive species depends on both the ability to disperse seeds to a site and an ability to establish once seeds have arrived. While seed and establishment limitation are known to jointly influence colonization, decomposing establishment limitation into density-dependent and density-independent components has remained challenging. Here, we couple theoretical models of recruitment with a multispecies invasion experiment conducted within a natural gradient of soil moisture and productivity to assess how variation in establishment limitation shapes outcomes for invasion. Recruitment was affected by both density-dependent and density-independent sources of establishment limitation in three of four species. Soil moisture stress and productivity both increased density-independent mortality in one species, whereas density-dependent mortality increased in locations with favourable soil moisture. Synthesis. Successful establishment of invading species can be limited by both density-dependent and density-independent mechanisms. In particular, the strength of density-independent limitation may depend on natural gradients in abiotic factors. The varying strengths of establishment limitation suggest that patterns of invasion are likely to be uneven both in space and in time. Understanding how intraspecific competitive constraints and density-independent limitation vary with abiotic gradients can assist with predicting when invasions are likely to occur, information that can be harnessed in the development of better methods for control.

Long-term regeneration of a tropical plant community after sand mining

Abstract Sandy coastal plant communities in tropical regions have been historically under strong anthropic pressure. In Brazil, these systems shelter communities with highly plastic plant species. However, the potential of these systems to regenerate without human assistance after disturbances has hardly been examined. We determined the natural regeneration of a coastal sandy plain vegetation (restinga) in Brazil, 16 years after the end of sand removal. We inventoried 38 plots: 20 within a sand‐mined site and 18 in an adjacent undisturbed site. We expected lower diversity values in the sand‐mined site compared to the undisturbed site, but similar species composition between the two sites due to the spatial proximity of the two sites and the high plasticity of restinga species. Species were ranked using abundance and importance value index in both sites, and comparisons were performed using Rényi entropy profiles, rarefaction curves, principal component analysis, and redundancy analysis. Species composition and dominant species differed markedly between the two sites. Bromeliads and Clusia hilariana, well‐known nurse plants, dominated the undisturbed site but were almost absent in the regenerating site. Species richness did not differ between both sites, but diversity was higher in the undisturbed site. Within‐site composition differences in the mined area were associated with field characteristics. Interestingly, species classified as subordinate or rare in the undisturbed site became dominants in the regenerating site. These newer dominants in the sand‐mined site are not those known as nurse plants in other restingas, thus yielding strong implications for restoration.