Land use modulates resistance of grasslands against future climate and inter-annual climate variability in a large field experiment.

Climate and land-use change are key drivers of global change. Full-factorial field experiments in which both drivers are manipulated are essential to understand and predict their potentially interactive effects on the structure and functioning of grassland ecosystems. Here, we present 8 years of data on grassland dynamics from the Global Change Experimental Facility in Central Germany. On large experimental plots, temperature and seasonal patterns of precipitation are manipulated by superimposing regional climate model projections onto background climate variability. Climate manipulation is factorially crossed with agricultural land-use scenarios, including intensively used meadows and extensively used (i.e., low-intensity) meadows and pastures. Inter-annual variation of background climate during our study years was high, including three of the driest years on record for our region. The effects of this temporal variability far exceeded the effects of the experimentally imposed climate change on plant species diversity and productivity, especially in the intensively used grasslands sown with only a few grass cultivars. These changes in productivity and diversity in response to alterations in climate were due to immigrant species replacing the target forage cultivars. This shift from forage cultivars to immigrant species may impose additional economic costs in terms of a decreasing forage value and the need for more frequent management measures. In contrast, the extensively used grasslands showed weaker responses to both experimentally manipulated future climate and inter-annual climate variability, suggesting that these diverse grasslands are more resistant to climate change than intensively used, species-poor grasslands. We therefore conclude that a lower management intensity of agricultural grasslands, associated with a higher plant diversity, can stabilize primary productivity under climate change.

Functional traits influence patterns in vegetative and reproductive plant phenology - a multi-botanical garden study.

Phenology has emerged as key indicator of the biological impacts of climate change, yet the role of functional traits constraining variation in herbaceous species' phenology has received little attention. Botanical gardens are ideal places in which to investigate large numbers of species growing under common climate conditions. We ask whether interspecific variation in plant phenology is influenced by differences in functional traits. We recorded onset, end, duration and intensity of initial growth, leafing out, leaf senescence, flowering and fruiting for 212 species across five botanical gardens in Germany. We measured functional traits, including plant height, absolute and specific leaf area, leaf dry matter content, leaf carbon and nitrogen content and seed mass and accounted for species' relatedness. Closely related species showed greater similarities in timing of phenological events than expected by chance, but species' traits had a high degree of explanatory power, pointing to paramount importance of species' life-history strategies. Taller plants showed later timing of initial growth, and flowered, fruited and underwent leaf senescence later. Large-leaved species had shorter flowering and fruiting durations. Taller, large-leaved species differ in their phenology and are more competitive than smaller, small-leaved species. We assume climate warming will change plant communities' competitive hierarchies with consequences for biodiversity.

Conservation with elevated elephant densities sequesters carbon in soils despite losses of woody biomass.

Nature conservation and restoration in terrestrial ecosystems is often focused on increasing the numbers of megafauna, expecting them to have positive impacts on ecological self-regulation processes and biodiversity. In sub-Saharan Africa, conservation efforts also aspire to protect and enhance biodiversity with particular focus on elephants. However, elephant browsing carries the risk of woody biomass losses. In this context, little is known about how increasing elephant numbers affects carbon stocks in soils, including the subsoils. We hypothesized that (1) increasing numbers of elephants reduce tree biomass, and thus the amount of C stored therein, resulting (2) in a loss of soil organic carbon (SOC). If true, a negative carbon footprint could limit the sustainability of elephant conservation from a global carbon perspective. To test these hypotheses, we selected plots of low, medium, and high elephant densities in two national parks and adjacent conservancies in the Namibian component of the Kavango Zambezi Transfrontier Area (KAZA), and quantified carbon storage in both woody vegetation and soils (1 m). Analyses were supplemented by the assessment of soil carbon isotopic composition. We found that increasing elephant densities resulted in a loss of tree carbon storage by 6.4 t ha-1 . However, and in contrast to our second hypothesis, SOC stocks increased by 4.7 t ha-1 with increasing elephant densities. These higher SOC stocks were mainly found in the topsoil (0-30 cm) and were largely due to the formation of SOC from woody biomass. A second carbon input source into the soils was megaherbivore dung, which contributed with 0.02-0.323 t C ha-1  year-1 to ecosystem carbon storage in the low and high elephant density plots, respectively. Consequently, increasing elephant density does not necessarily lead to a negative C footprint, as soil carbon sequestration and transient C storage in dung almost compensate for losses in tree biomass.

Linking genes with ecological strategies in Arabidopsis thaliana.

Arabidopsis thaliana is the most prominent model system in plant molecular biology and genetics. Although its ecology was initially neglected, collections of various genotypes revealed a complex population structure, with high levels of genetic diversity and substantial levels of phenotypic variation. This helped identify the genes and gene pathways mediating phenotypic change. Population genetics studies further demonstrated that this variation generally contributes to local adaptation. Here, we review evidence showing that traits affecting plant life history, growth rate, and stress reactions are not only locally adapted, they also often co-vary. Co-variation between these traits indicates that they evolve as trait syndromes, and reveals the ecological diversification that took place within A. thaliana. We argue that examining traits and the gene that control them within the context of global summary schemes that describe major ecological strategies will contribute to resolve important questions in both molecular biology and ecology.

Temperature and aridity regulate spatial variability of soil multifunctionality in drylands across the globe.

The relationship between the spatial variability of soil multifunctionality (i.e., the capacity of soils to conduct multiple functions; SVM) and major climatic drivers, such as temperature and aridity, has never been assessed globally in terrestrial ecosystems. We surveyed 236 dryland ecosystems from six continents to evaluate the relative importance of aridity and mean annual temperature, and of other abiotic (e.g., texture) and biotic (e.g., plant cover) variables as drivers of SVM, calculated as the averaged coefficient of variation for multiple soil variables linked to nutrient stocks and cycling. We found that increases in temperature and aridity were globally correlated to increases in SVM. Some of these climatic effects on SVM were direct, but others were indirectly driven through reductions in the number of vegetation patches and increases in soil sand content. The predictive capacity of our structural equation modelling was clearly higher for the spatial variability of N- than for C- and P-related soil variables. In the case of N cycling, the effects of temperature and aridity were both direct and indirect via changes in soil properties. For C and P, the effect of climate was mainly indirect via changes in plant attributes. These results suggest that future changes in climate may decouple the spatial availability of these elements for plants and microbes in dryland soils. Our findings significantly advance our understanding of the patterns and mechanisms driving SVM in drylands across the globe, which is critical for predicting changes in ecosystem functioning in response to climate change.

Quantifying drylands' drought resistance and recovery: the importance of drought intensity, dominant life history and grazing regime.

Projected global change will increase the level of land-use and environmental stressors such as drought and grazing, particularly in drylands. Still, combined effects of drought and grazing on plant production are poorly understood, thus hampering adequate projections and development of mitigation strategies. We used a large, cross-continental database consisting of 174 long-term datasets from >30 dryland regions to quantify ecosystem responses to drought and grazing with the ultimate goal to increase functional understanding in these responses. Two key aspects of ecosystem stability, resistance to and recovery after a drought, were evaluated based on standardized and normalized aboveground net primary production (ANPP) data. Drought intensity was quantified using the standardized precipitation index. We tested effects of drought intensity, grazing regime (grazed, ungrazed), biome (grassland, shrubland, savanna) or dominant life history (annual, perennial) of the herbaceous layer to assess the relative importance of these factors for ecosystem stability, and to identify predictable relationships between drought intensity and ecosystem resistance and recovery. We found that both components of ecosystem stability were better explained by dominant herbaceous life history than by biome. Increasing drought intensity (quasi-) linearly reduced ecosystem resistance. Even though annual and perennial systems showed the same response rate to increasing drought intensity, they differed in their general magnitude of resistance, with annual systems being ca. 27% less resistant. In contrast, systems with an herbaceous layer dominated by annuals had substantially higher postdrought recovery, particularly when grazed. Combined effects of drought and grazing were not merely additive but modulated by dominant life history of the herbaceous layer. To the best of our knowledge, our study established the first predictive, cross-continental model between drought intensity and drought-related relative losses in ANPP, and suggests that systems with an herbaceous layer dominated by annuals are more prone to ecosystem degradation under future global change regimes.

Local density effects on individual production are dynamic: insights from natural stands of a perennial savanna grass.

Perennial grasses are a dominant component of grasslands, and provide important ecosystem services. However, most knowledge of grasslands' functioning and production comes from plot-level studies, and drivers of individual-level production remain poorly explored. Extrapolation from existing experiments is hampered by the fact that these are mostly concentrated on even-aged cohorts, and/or on the early stages of a plant's life cycle. Here we explored how local density regulates individual production in mono-specific natural grassland, focusing on adult individuals of a perennial savanna grass (Stipagrostis uniplumis). We found individual production to increase with individuals' size, but to decrease with neighbour abundance. A metric of neighbour abundance that considered size was superior to a metric based solely on the number of individuals. This finding is particularly important for studying competitive effects in natural populations, where plants are normally not even-sized. The inferred competition kernel, i.e. the function describing how competitive strength varies with spatial distance from a target plant, was hump-shaped, indicating strongest intraspecific competition at intermediate distances (10-30 cm). The spatial signature of competitive effects changed with time since fire; peak effects moved successively away from the target plant. Our results suggest that inferred competition kernels of long-lived plant populations may have shapes that differ from exponential or sigmoidal decreases. More generally, results underline that competition among neighbouring plants is dynamic. Studies that address density-dependent and density-independent (fire-related) population dynamics of perennial grasses in their fire-prone environment may thus shed new light on the functioning and production of grasslands.

Dataset on woody aboveground biomass, disturbance losses, and wood density from an African savanna ecosystem.

This dataset comprises tree inventories and damage assessments performed in Namibia's semi-arid Zambezi Region. Data were sampled in savannas and savanna woodlands along steep gradients of elephant population densities to capture the effects of those (and other) disturbances on individual-level and stand-level aboveground woody biomass (AGB). The dataset contains raw data on dendrometric measures and processed data on specific wood density (SWD), woody aboveground biomass, and biomass losses through disturbance impacts. Allometric proxies (height, canopy diameters, and in adult trees also stem circumferences) were recorded for n = 6,179 tree and shrub individuals. Wood samples were taken for each encountered species to measure specific wood density. These measurements have been used to estimate woody aboveground biomass via established allometric models, advanced through our improved methodologies and workflows that accounted for tree and shrub architecture shaped by disturbance impacts. To this end, we performed a detailed damage assessment on each woody individual in the field. In addition to estimations of standing biomass, our new method also delivered data on biomass losses to different disturbance agents (elephants, fire, and others) on the level of plant individuals and stands. The data presented here have been used within a study published with Ecological Indicators (Kindermann et al., 2022) to evaluate the benefits of our improved methodology in comparison to a standard reference method of aboveground biomass estimations. Additionally, it has been employed in a study on carbon storage and sequestration in vegetation and soils (Sandhage-Hofmann et al., 2021). The raw data of dendrometric measurements can be subjected to other available allometric models for biomass estimation. The processed data can be used to analyze disturbance impacts on woody aboveground biomass, or for regional carbon storage estimates. The data on species-specific wood density can be used for application to other dendrometric datasets to (re-) estimate biomass through allometric models requiring wood density. It can further be used for plant functional trait analyses.

Site-specific distribution of oak rhizosphere-associated oomycetes revealed by cytochrome c oxidase subunit II metabarcoding.

The phylum Oomycota comprises important tree pathogens like Phytophthora quercina, involved in central European oak decline, and Phytophthora cinnamomi shown to affect holm oaks among many other hosts. Despite the importance to study the distribution, dispersal and niche partitioning of this phylum, metabarcoding surveys, and studies considering environmental factors that could explain oomycete community patterns are still rare. We investigated oomycetes in the rhizosphere of evergreen oaks in a Spanish oak woodland using metabarcoding based on Illumina sequencing of the taxonomic marker cytochrome c oxidase subunit II (cox2). We developed an approach amplifying a 333 bp long fragment using the forward primer Hud-F (Mycologia, 2000) and a reverse primer found using DegePrime (Applied and Environmental Microbiology, 2014). Factors reflecting topo-edaphic conditions and tree health were linked to oomycete community patterns. The majority of detected OTUs belonged to the Peronosporales. Most taxa were relatives of the Pythiaceae, but relatives of the Peronosporaceae and members of the Saprolegniales were also found. The most abundant OTUs were related to Globisporangium irregulare and P. cinnamomi, both displaying strong site-specific patterns. Oomycete communities were strongly correlated with the environmental factors: altitude, crown foliation, slope and soil skeleton and soil nitrogen. Our findings illustrate the significance of small scale variation in habitat conditions for the distribution of oomycetes and highlight the importance to study oomycete communities in relation to such ecological patterns.

The impact of rain events on CO2 emissions from contrasting land use systems in semi-arid West African savannas.

In the future the Sudanian savanna - one of West Africa's high-potential "bread baskets" - will likely face shorter rainy seasons with more extreme rains and droughts. That could have serious impacts on the vegetation and its carbon dioxide (CO2) exchange with potentially increasing CO2 emissions accelerating climate warming. Understanding how the CO2 fluxes in this area respond to environmental variables, in particular rain events, is essential, but available data are scarce. In this study, we monitored net ecosystem exchange (NEE) of CO2, rainfall and other environmental parameters during four years at three savannas. Savannas were characterized by different vegetation due to different land use: i) woody and nearly pristine, ii) mixture of cropland and grassland and iii) intensive grazing. The impact of rain events on CO2 exchange for these contrasting ecosystems were analyzed for single rain events (short-term) and on a yearly time scale (long-term) using three eddy covariance towers. We found that the woody pristine savanna site was a prominent sink of CO2 (-864 to -1299 g CO2 m-2 y-1) while the degraded sites were net CO2 sources (118 to 605 g CO2 m-2 y-1) with a complicated relation with annual rainfall amounts. The NEE responses to single rain events revealed that daytime rain systematically decreased the sink strengths at all sites, which might be associated with decreased light availability. At the degraded sites, additional factors increasing CO2 losses were rain duration and dry spell length. The observed patterns of immediate CO2 flux responses to rainfall at differently used savannas indicate strong internal feedbacks between vegetation and land use changes and raise the question whether the CO2 sink strengths might be overestimated with possible implications for global CO2 budgets. Sustainable adaptation strategies need to be developed for West Africa.

Learning from local knowledge: modeling the pastoral-nomadic range management of the Himba, Namibia.

It is widely accepted that successful grazing management strategies in semiarid ecosystems need to be adapted to the highly temporal and spatially heterogeneous forage production. Nevertheless, a full understanding of the key factors and processes for sustainable adaptive management has yet to be reached. The investigation of existing, successful range management systems by simulation models may help to derive general understanding and basic principles. The semi-nomadic Himba in northern Namibia applied a sophisticated management system until the mid-1990s which combined season-dependent pasture use (resulting in rainy-season pastures and dry-season pastures), preservation of reserves for drought and sanctions for rule breaking. A stochastic ecological simulation model has been developed here which represents the main aspects of this management system. With this model we analyze (1) which components of the traditional Himba strategy are essential for sustainability and (2) what happens to the state of the rangeland system under socioeconomic changes. This study shows that temporally and spatially heterogeneous pasture use yields higher productivity and quality of a pasture area than the pressure of homogeneous permanent grazing. Two aspects are of importance: (1) intra-annual heterogeneous use (resting of the dry-season pastures during the rainy season) and (2) interannual heterogeneous use (spatial extension of grazing in years of drought). This management system leads to an effective build-up and use of a buffer in the system: the reserve biomass (the non-photosynthetic reserve organs of the plants), an indicator for grazing and management history. Analyzing purchase as one form of socioeconomic change, we demonstrate that easier market access to purchase livestock may lead to a decline in vegetation quality. However, cattle production increases as long as rest periods on parts of the pasture during the rainy season are granted. Methodologically, we emphasize that simulation models offer an excellent framework for analyzing and depicting basic principles in sustainable range management derived from local knowledge. They provide the opportunity of testing whether these basic principles are also valid under different ecological and socioeconomic settings.

Are there consistent grazing indicators in Drylands? Testing plant functional types of various complexity in South Africa's Grassland and Savanna Biomes.

Despite our growing knowledge on plants' functional responses to grazing, there is no consensus if an optimum level of functional aggregation exists for detecting grazing effects in drylands. With a comparative approach we searched for plant functional types (PFTs) with a consistent response to grazing across two areas differing in climatic aridity, situated in South Africa's grassland and savanna biomes. We aggregated herbaceous species into PFTs, using hierarchical combinations of traits (from single- to three-trait PFTs). Traits relate to life history, growth form and leaf width. We first confirmed that soil and grazing gradients were largely independent from each other, and then searched in each biome for PFTs with a sensitive response to grazing, avoiding confounding with soil conditions. We found no response consistency, but biome-specific optimum aggregation levels. Three-trait PFTs (e.g. broad-leaved perennial grasses) and two-trait PFTs (e.g. perennial grasses) performed best as indicators of grazing effects in the semi-arid grassland and in the arid savanna biome, respectively. Some PFTs increased with grazing pressure in the grassland, but decreased in the savanna. We applied biome-specific grazing indicators to evaluate if differences in grazing management related to land tenure (communal versus freehold) had effects on vegetation. Tenure effects were small, which we mainly attributed to large variability in grazing pressure across farms. We conclude that the striking lack of generalizable PFT responses to grazing is due to a convergence of aridity and grazing effects, and unlikely to be overcome by more refined classification approaches. Hence, PFTs with an opposite response to grazing in the two biomes rather have a unimodal response along a gradient of additive forces of aridity and grazing. The study advocates for hierarchical trait combinations to identify localized indicator sets for grazing effects. Its methodological approach may also be useful for identifying ecological indicators in other ecosystems.