The positive effect of plant diversity on soil carbon depends on climate.

Little is currently known about how climate modulates the relationship between plant diversity and soil organic carbon and the mechanisms involved. Yet, this knowledge is of crucial importance in times of climate change and biodiversity loss. Here, we show that plant diversity is positively correlated with soil carbon content and soil carbon-to-nitrogen ratio across 84 grasslands on six continents that span wide climate gradients. The relationships between plant diversity and soil carbon as well as plant diversity and soil organic matter quality (carbon-to-nitrogen ratio) are particularly strong in warm and arid climates. While plant biomass is positively correlated with soil carbon, plant biomass is not significantly correlated with plant diversity. Our results indicate that plant diversity influences soil carbon storage not via the quantity of organic matter (plant biomass) inputs to soil, but through the quality of organic matter. The study implies that ecosystem management that restores plant diversity likely enhances soil carbon sequestration, particularly in warm and arid climates.

Herbivory and nutrients shape grassland soil seed banks.

Anthropogenic nutrient enrichment and shifts in herbivory can lead to dramatic changes in the composition and diversity of aboveground plant communities. In turn, this can alter seed banks in the soil, which are cryptic reservoirs of plant diversity. Here, we use data from seven Nutrient Network grassland sites on four continents, encompassing a range of climatic and environmental conditions, to test the joint effects of fertilization and aboveground mammalian herbivory on seed banks and on the similarity between aboveground plant communities and seed banks. We find that fertilization decreases plant species richness and diversity in seed banks, and homogenizes composition between aboveground and seed bank communities. Fertilization increases seed bank abundance especially in the presence of herbivores, while this effect is smaller in the absence of herbivores. Our findings highlight that nutrient enrichment can weaken a diversity maintaining mechanism in grasslands, and that herbivory needs to be considered when assessing nutrient enrichment effects on seed bank abundance.

Environmental factors regulate soil microbial attributes and their response to drought in rangeland ecosystems.

In ecosystems, soil microbial variables characterization are used to determine soil biological health and the response of soils to environmental stress. Although there are strong associations between plants and soil microorganisms, they may respond asynchronously to environmental factors and severe droughts. We aimed to: I) evaluate the special variation of soil microbiome such as microbial biomass carbon (MBC) and nitrogen (MBN), soil basal respiration (SBR) and microbial indexes in eight rangeland sites located across an aridity gradient (distributed from arid to mesic climates); II) analyze the relative importance of main environmental factors (climate, soils, and plants) and their relationships with microbial variables in the rangelands; and III) assess the effect of drought on microbial and plant variables in field-based manipulative experiments. First, we found significant changes of microbial variables along a precipitation and temperature gradient. The responses of MBC and MBN were strongly dependent on soil pH, soil nitrogen (N), soil organic carbon (SOC), C:N ratio and vegetation cover. In contrast, SBR was influenced by the aridity index (AI), the mean annual precipitation (MAP), the soil pH and vegetation cover. MBC, MBN and SBR were negatively related with soil pH compared to the other factors (C, N, C:N, vegetation cover, MAP and AI) that had a positive relationship. Second, we found a stronger soil microbial variables response to drought in arid sites compared to humid rangelands. Third, the responses of MBC, MBN, and SBR to drought showed positive relationships with vegetation cover and aboveground biomass, but with different regression slopes, this suggest that plant and microbial communities responded differently to drought. The results from this study improve our understanding about the microbial response to drought in different rangelands, and may facilitate the development of predictive models on responses of soil microorganisms in carbon cycle under global change scenarios.

Environmental heterogeneity modulates the effect of plant diversity on the spatial variability of grassland biomass.

Plant productivity varies due to environmental heterogeneity, and theory suggests that plant diversity can reduce this variation. While there is strong evidence of diversity effects on temporal variability of productivity, whether this mechanism extends to variability across space remains elusive. Here we determine the relationship between plant diversity and spatial variability of productivity in 83 grasslands, and quantify the effect of experimentally increased spatial heterogeneity in environmental conditions on this relationship. We found that communities with higher plant species richness (alpha and gamma diversity) have lower spatial variability of productivity as reduced abundance of some species can be compensated for by increased abundance of other species. In contrast, high species dissimilarity among local communities (beta diversity) is positively associated with spatial variability of productivity, suggesting that changes in species composition can scale up to affect productivity. Experimentally increased spatial environmental heterogeneity weakens the effect of plant alpha and gamma diversity, and reveals that beta diversity can simultaneously decrease and increase spatial variability of productivity. Our findings unveil the generality of the diversity-stability theory across space, and suggest that reduced local diversity and biotic homogenization can affect the spatial reliability of key ecosystem functions.

Climate change-related growth improvements in a wide niche-breadth tree species across contrasting environments.

BACKGROUND AND AIMS: The vulnerability and responsiveness of forests to drought are immensely variable across biomes. Intraspecific tree responses to drought in species with wide niche breadths that grow across contrasting climatically environments might provide key information regarding forest resistance and changes in species distribution under climate change. Using a species with an exceptionally wide niche breath, we tested the hypothesis that tree populations thriving in dry environments are more resistant to drought than those growing in moist locations. METHODS: We determined temporal trends in tree radial growth of 12 tree populations of Nothofagus antarctica (Nothofagaceae) located across a sharp precipitation gradient (annual precipitation of 500-2000 mm) in Chile and Argentina. Using dendrochronological methods, we fitted generalized additive mixed-effect models to predict the annual basal area increment as a function of year and dryness (De Martonne aridity index). We also measured carbon and oxygen isotope signals (and estimated intrinsic water-use efficiency) to provide potential physiological causes for tree growth responses to drought. KEY RESULTS: We found unexpected improvements in growth during 1980-1998 in moist sites, while growth responses in dry sites were mixed. All populations, independent of site moisture, showed an increase in their intrinsic water-use efficiency in recent decades, a tendency that seemed to be explained by an increase in the photosynthetic rate instead of drought-induced stomatal closure, given that δ18O did not change with time. CONCLUSIONS: The absence of drought-induced negative effects on tree growth in a tree species with a wide niche breadth is promising because it might relate to the causal mechanisms tree species possess to face ongoing drought events. We suggest that the drought resistance of N. antarctica might be attributable to its low stature and relatively low growth rate.

Leaf wettability, anatomy and ultra-structure of Nothofagus antarctica and N. betuloides grown under a CO2 enriched atmosphere.

Increasing CO2 air concentration may affect wettability, anatomy and ultra-structure of leaves of Patagonian forest species, evergreen and deciduous plants potentially responding differently to such CO2 increases. In this study, we analysed the wettability, anatomy and ultra-structure of leaves of Nothofagus antarctica (deciduous) and N. betuloides (evergreen) grown under high CO2 concentrations. Leaf wettability was affected by increasing CO2, in different directions depending on species and leaf side. In both species, soluble cuticular lipid concentrations per unit leaf area raised with higher CO2 levels. Stomatal parameters (density, size of guard cells and pores) showed different responses to CO2 increasing depending on the species examined. In both species, leaf tissues showed a general trend to diminish with higher CO2 concentration. Cuticle thickness was modified with higher CO2 concentration in N. betuloides, but not in N. antarctica leaves. In both species, chloroplasts were often damaged with the increase in CO2 concentration. Our results show that several surface and internal leaf parameters can be modified in association with an increase in atmospheric CO2 concentration which may very among plant species.

Linking changes in species composition and biomass in a globally distributed grassland experiment.

Global change drivers, such as anthropogenic nutrient inputs, are increasing globally. Nutrient deposition simultaneously alters plant biodiversity, species composition and ecosystem processes like aboveground biomass production. These changes are underpinned by species extinction, colonisation and shifting relative abundance. Here, we use the Price equation to quantify and link the contributions of species that are lost, gained or that persist to change in aboveground biomass in 59 experimental grassland sites. Under ambient (control) conditions, compositional and biomass turnover was high, and losses (i.e. local extinctions) were balanced by gains (i.e. colonisation). Under fertilisation, the decline in species richness resulted from increased species loss and decreases in species gained. Biomass increase under fertilisation resulted mostly from species that persist and to a lesser extent from species gained. Drivers of ecological change can interact relatively independently with diversity, composition and ecosystem processes and functions such as aboveground biomass due to the individual contributions of species lost, gained or persisting.

Co-limitation towards lower latitudes shapes global forest diversity gradients.

The latitudinal diversity gradient (LDG) is one of the most recognized global patterns of species richness exhibited across a wide range of taxa. Numerous hypotheses have been proposed in the past two centuries to explain LDG, but rigorous tests of the drivers of LDGs have been limited by a lack of high-quality global species richness data. Here we produce a high-resolution (0.025° × 0.025°) map of local tree species richness using a global forest inventory database with individual tree information and local biophysical characteristics from ~1.3 million sample plots. We then quantify drivers of local tree species richness patterns across latitudes. Generally, annual mean temperature was a dominant predictor of tree species richness, which is most consistent with the metabolic theory of biodiversity (MTB). However, MTB underestimated LDG in the tropics, where high species richness was also moderated by topographic, soil and anthropogenic factors operating at local scales. Given that local landscape variables operate synergistically with bioclimatic factors in shaping the global LDG pattern, we suggest that MTB be extended to account for co-limitation by subordinate drivers.

Multi-taxon biodiversity assessment of Southern Patagonia: Supporting conservation strategies at different landscapes.

In the last years, different spatial analyses were developed to support multi-taxon biodiversity conservation strategies. In fact, the use of species distribution models as input allowed to create spatial decision-support maps. Of special interest are maps of potential biodiversity (MPB), which define distribution and ecological requirements of relevant species and maps of priority conservation areas (MPCA), which define priority areas considering endemism and richness. The objective of this paper was to assess multi-taxon biodiversity based on two different spatial analyses and to test their efficiency to support conservation decision at Patagonia. We computed 119 potential habitat suitability maps (one deer, birds, lizards, darkling-beetles, plants) with ENFA (Environmental Niche Factor Analysis) and 15 environmental variables, using Biomapper software. ENFA calculate two ecological indexes (marginality and specialization) which describe the narrowness of species niches and how extreme are the optimum environmental conditions related to the whole study area. These maps were combined obtaining a MPB and MPCA using Zonation software. Multivariate analyses were performed to compare methodologies, analysing environmental variables, ecological areas, forest types and protected areas. Multivariate and ecological indexes showed that deer, lizards and darkling-beetles presented a narrow range, while birds and plants presented a large range of marginality and specialization mainly related to vegetation and climate. At provincial level, highest potential biodiversity and conservation priority values were related to shrublands and humid steppes. However, MPCA showed higher values related to forests and alpine vegetation due to endemism, while MPB showed differences among forest types. These analyses showed that the most valuable areas were not represented in the protected areas, however, many higher conservation priority values were found inside the protected compared with unprotected areas. Different spatial decision-support maps presented similar outputs at provincial scale, but differed in the forest landscape matrix. Both methodologies can be used to plan conservation strategies depending on the specific objectives (e.g. highlighting richness or endemism).

Opposing community assembly patterns for dominant and nondominant plant species in herbaceous ecosystems globally.

Biotic and abiotic factors interact with dominant plants-the locally most frequent or with the largest coverage-and nondominant plants differently, partially because dominant plants modify the environment where nondominant plants grow. For instance, if dominant plants compete strongly, they will deplete most resources, forcing nondominant plants into a narrower niche space. Conversely, if dominant plants are constrained by the environment, they might not exhaust available resources but instead may ameliorate environmental stressors that usually limit nondominants. Hence, the nature of interactions among nondominant species could be modified by dominant species. Furthermore, these differences could translate into a disparity in the phylogenetic relatedness among dominants compared to the relatedness among nondominants. By estimating phylogenetic dispersion in 78 grasslands across five continents, we found that dominant species were clustered (e.g., co-dominant grasses), suggesting dominant species are likely organized by environmental filtering, and that nondominant species were either randomly assembled or overdispersed. Traits showed similar trends for those sites (<50%) with sufficient trait data. Furthermore, several lineages scattered in the phylogeny had more nondominant species than expected at random, suggesting that traits common in nondominants are phylogenetically conserved and have evolved multiple times. We also explored environmental drivers of the dominant/nondominant disparity. We found different assembly patterns for dominants and nondominants, consistent with asymmetries in assembly mechanisms. Among the different postulated mechanisms, our results suggest two complementary hypotheses seldom explored: (1) Nondominant species include lineages adapted to thrive in the environment generated by dominant species. (2) Even when dominant species reduce resources to nondominant ones, dominant species could have a stronger positive effect on some nondominants by ameliorating environmental stressors affecting them, than by depleting resources and increasing the environmental stress to those nondominants. These results show that the dominant/nondominant asymmetry has ecological and evolutionary consequences fundamental to understand plant communities.

General destabilizing effects of eutrophication on grassland productivity at multiple spatial scales.

Eutrophication is a widespread environmental change that usually reduces the stabilizing effect of plant diversity on productivity in local communities. Whether this effect is scale dependent remains to be elucidated. Here, we determine the relationship between plant diversity and temporal stability of productivity for 243 plant communities from 42 grasslands across the globe and quantify the effect of chronic fertilization on these relationships. Unfertilized local communities with more plant species exhibit greater asynchronous dynamics among species in response to natural environmental fluctuations, resulting in greater local stability (alpha stability). Moreover, neighborhood communities that have greater spatial variation in plant species composition within sites (higher beta diversity) have greater spatial asynchrony of productivity among communities, resulting in greater stability at the larger scale (gamma stability). Importantly, fertilization consistently weakens the contribution of plant diversity to both of these stabilizing mechanisms, thus diminishing the positive effect of biodiversity on stability at differing spatial scales. Our findings suggest that preserving grassland functional stability requires conservation of plant diversity within and among ecological communities.

Microbial processing of plant remains is co-limited by multiple nutrients in global grasslands.

Microbial processing of aggregate-unprotected organic matter inputs is key for soil fertility, long-term ecosystem carbon and nutrient sequestration and sustainable agriculture. We investigated the effects of adding multiple nutrients (nitrogen, phosphorus and potassium plus nine essential macro- and micro-nutrients) on decomposition and biochemical transformation of standard plant materials buried in 21 grasslands from four continents. Addition of multiple nutrients weakly but consistently increased decomposition and biochemical transformation of plant remains during the peak-season, concurrent with changes in microbial exoenzymatic activity. Higher mean annual precipitation and lower mean annual temperature were the main climatic drivers of higher decomposition rates, while biochemical transformation of plant remains was negatively related to temperature of the wettest quarter. Nutrients enhanced decomposition most at cool, high rainfall sites, indicating that in a warmer and drier future fertilized grassland soils will have an even more limited potential for microbial processing of plant remains.

Author Correction: Leaf nutrients, not specific leaf area, are consistent indicators of elevated nutrient inputs.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Nutrient availability controls the impact of mammalian herbivores on soil carbon and nitrogen pools in grasslands.

Grasslands are subject to considerable alteration due to human activities globally, including widespread changes in populations and composition of large mammalian herbivores and elevated supply of nutrients. Grassland soils remain important reservoirs of carbon (C) and nitrogen (N). Herbivores may affect both C and N pools and these changes likely interact with increases in soil nutrient availability. Given the scale of grassland soil fluxes, such changes can have striking consequences for atmospheric C concentrations and the climate. Here, we use the Nutrient Network experiment to examine the responses of soil C and N pools to mammalian herbivore exclusion across 22 grasslands, under ambient and elevated nutrient availabilities (fertilized with NPK + micronutrients). We show that the impact of herbivore exclusion on soil C and N pools depends on fertilization. Under ambient nutrient conditions, we observed no effect of herbivore exclusion, but under elevated nutrient supply, pools are smaller upon herbivore exclusion. The highest mean soil C and N pools were found in grazed and fertilized plots. The decrease in soil C and N upon herbivore exclusion in combination with fertilization correlated with a decrease in aboveground plant biomass and microbial activity, indicating a reduced storage of organic matter and microbial residues as soil C and N. The response of soil C and N pools to herbivore exclusion was contingent on temperature - herbivores likely cause losses of C and N in colder sites and increases in warmer sites. Additionally, grasslands that contain mammalian herbivores have the potential to sequester more N under increased temperature variability and nutrient enrichment than ungrazed grasslands. Our study highlights the importance of conserving mammalian herbivore populations in grasslands worldwide. We need to incorporate local-scale herbivory, and its interaction with nutrient enrichment and climate, within global-scale models to better predict land-atmosphere interactions under future climate change.

Essential Nutrient and Trace Element Foliar Resorption of Two Co-Existing Nothofagus Species Grown Under Different Environmental Conditions in Southern Patagonia.

Nutrient resorption is crucial for mineral element conservation and efficiency of forest species, but knowledge on its significance and the mechanisms involved is still limited for most species and habitats. Focusing on the harsh conditions for plant growth and survival of southern Patagonia, a field study for comparing the rate of foliar resorption of macro-, micro-nutrients, and trace elements in coexisting Nothofagus pumilio and Nothofagus antarctica forests was performed. Forests located in three contrasting productivity sites (with different soil and climatic conditions) were selected, and mature, functional versus senescent leaves of both species were collected at two different dates of the growing season. Macro- (N, P, Ca, K, S, and Mg), micronutrients (B, Cu, Fe, Mn, Zn, and Ni), and trace elements (Al, Li, Pb, Rb, Sr, Ti, and Tl) were determined in foliar tissues. The mineral element concentrations of mature and senescent leaves were used for calculating the nutrient resorption efficiency (NuR). In general, and making an average of all sites and species, macro-nutrient resorption showed a decreasing trend for N > S = K > P > Mg, being Ca the only macro-nutrient with negative values (i.e., no resorption). Resorption of the majority of the elements did not vary between species in any of the evaluated sites. Variation across sites in nutrient resorption efficiency for most macronutrients, some micronutrients, and trace elements was observed for N. antarctica, whereas N. pumilio had a similar NuR for all experimental sites. On the other hand, regardless of the site or the species, some elements were not resorbed (e.g., B, Cu, Fe, Mn, Al, and Ti). It is concluded that both Nothofagus species performed similarly concerning their nutrient conservation strategy, when coexisting in the same mixed forest. However, no evidence was gained for an increased rate of foliar NuR in association with the sites subjected to more limiting soil and climatic conditions for plant growth.

Land sharing in South Patagonia: Conservation of above-ground beetle diversity in forests and non-forest ecosystems.

Land-sharing strategies, as variable retention silvicultural proposals, are useful to mitigate harmful effects of economic activities on forest biodiversity; benefits have been reported worldwide for several organisms. However, we suggest that this approach could be useful to improve beetle conservation not only in forests but also in other ecosystem types, based on the results from Southern Patagonia (Argentina). We studied above-ground beetle communities using pitfall traps in Nothofagus pumilio forests, Mulguraea tridens shrublands, and magellanic steppes. The forests were located in Tierra del Fuego Province, while the shrublands and the steppes were in Santa Cruz Province. In forests and shrublands, we compared retention approaches (aggregated/dispersed retention harvesting in forests, and managed cut and retention strips in shrublands) vs. control situations (without harvesting/cuttings). In dry and humid steppes, both impacted by livestock, we evaluated grazed and exclusion paddocks, comparable to structural retentions (reference areas without grazing do not exist). Richness, abundance, frequency, Shannon-Wiener diversity and Pielou evenness indices, and similarity among assemblages were evaluated using univariate and multivariate statistical tests. In forests and shrublands, retention approaches (aggregated/dispersed and strips) allowed the partial or total maintenance of beetle community richness, preserving them similar to natural and non-impacted ecosystems. In dry and humid steppes, exclusion areas presented significantly different richness, abundance and diversity of arthropod assemblages, but with inverse trends: lower values in grazed areas than in exclusions in dry steppe, and higher values in grazed areas than in exclusions in humid steppe. We concluded that land-sharing could be implemented in forests and non-forest ecosystems to preserve beetle communities, being the variable retention approaches and the grazing exclusion areas good alternatives for private or public lands. Likewise, we consider that legislation to promote conservation (like National Law 26331) should not be only applicable for and implemented in forests, but also in non-forest ecosystems.

Potential biodiversity map of understory plants for Nothofagus forests in Southern Patagonia: Analyses of landscape, ecological niche and conservation values.

The role of understory plants in native forests is critical for ecosystem function, wildlife protection and ecosystem productivity. The interest to estimate biodiversity increased during the last decades at landscape level. The objective was to elaborate a map of potential biodiversity (MPB) of understory species of Nothofagus forest using potential habitat suitability maps (PHS) of 15 plants in Santa Cruz province, Argentina. Additionally, we asked the following questions: (i) Were plant species differentially distributed according to the forest types?, (ii) do forest types represent different plant species assemblage with specific ecological niche requirements?, and (iii) is it possible to detect hotspots in the MBP according to the forest types? We used 721 plots database of vascular plants, from where 15 indicator species were identified. The assemblage species for different forests (Nothofagus antarctica, N. pumilio and evergreen mixed) were analysed using a detrended correspondence analysis. Also, we explored 41 potential explanatory variables to develop PHS, and combined these maps to obtain one MPB (1-100%). Finally, we analysed the outputs into a GIS through different landscapes alternatives to detect hotspot areas. Marginality and specialization values allowed identifying species assemblage that presented similar variability in the habitat requirements. MPB varied across the landscape, with higher values in the south and lower values near glaciers. MPB had the highest values in N. antarctica forest with >50% cover at landscape level. N. antarctica present more hotspots than N. pumilio forests, mainly in the south, compared to mixed evergreen forests which present few hotspots near glaciers. These results can be used as a tool to design new management and conservation strategies at landscape level.