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Species-rich grasslands provide important ecosystem services, and in South Africa, approximately 40% of these grasslands are degraded. Vetiver grass (from India) is often used during rehabilitation efforts to restore soil function without a thorough understanding of the potential negative ecological impacts. Hence, a study was initiated to investigate vetiver’s ecological impacts during grassland rehabilitation. Firstly, a field survey was conducted using a contiguous quadrat method to evaluate the extent of grass secondary succession in these rehabilitated sites. Secondly, the effect of vetiver competition and seed sowing method on the recruitment of two native grasses (Eragrostis curvula and Megathyrsus maximus) was examined using pot trials. The field survey results showed no evidence of grass secondary succession, but rather the abundance of bare ground around vetiver, and a marked increase in grass species richness with increasing distance from planted vetiver. Subsequently, in the pot trial, vetiver facilitated emergence in both native grasses, and soil surface sowing of indigenous grass seeds showed greater emergence than other sowing methods. However, vetiver inhibited native grass seedling establishment, even when root competition was excluded. This study suggests that areas rehabilitated using vetiver are unlikely to become productive grasslands with good grazing, because vetiver inhibits colonisation by native grasses.

The amount of nitrogen (N) derived from symbiotic N2 fixation by legumes in grasslands might be affected by anthropogenic N and phosphorus (P) inputs, but the underlying mechanisms are not known. Methods We evaluated symbiotic N2 fixation in 17 natural and semi-natural grasslands on four continents that are subjected to the same full-factorial N and P addition experiment, using the 15N natural abundance method. Results N as well as combined N and P (NP) addition reduced aboveground legume biomass by 65% and 45%, respectively, compared to the control, whereas P addition had no significant impact. Addition of N and/or P had no significant effect on the symbiotic N2 fixation per unit legume biomass. In consequence, the amount of N fixed annually per grassland area was less than half in the N addition treatments compared to control and P addition, irrespective of whether the dominant legumes were annuals or perennials. Conclusion Our results reveal that N addition mainly impacts symbiotic N2 fixation via reduced biomass of legumes rather than changes in N2 fixation per unit legume biomass. The results show that soil N enrichment by anthropogenic activities significantly reduces N 2 fixation in grasslands, and these effects cannot be reversed by additional P amendment. EEA Santa Cruz Fil: Vázquez, Eduardo. University of Bayreuth. Department of Soil Ecology. Bayreuth Center of Ecology and Environmental Research (BayCEER); Alemania Fil: Vázquez, Eduardo. Swedish University of Agricultural Sciences. Department of Soil and Environment; Suecia Fil: Schleuss, Per‑Marten. University of Bayreuth. Department of Soil Ecology. Bayreuth Center of Ecology and Environmental Research (BayCEER); Alemania Fil: Borer, Elizabeth T. University of Minnesota. Department of Ecology, Evolution, and Behavior; Estados Unidos Fil: Bugalho, Miguel N. University of Lisbon. Centre for Applied Ecology “Prof. Baeta Neves” (CEABN-InBIO). School of Agriculture; Portugal. Fil: Caldeira, Maria. C. University of Lisbon. Forest Research Centre. School of Agriculture; Portugal. Fil: Eisenhauer, Nico. German Centre for Integrative Biodiversity Research; Alemania Fil: Eisenhauer, Nico. Leipzig University. Institute of Biology; Alemania Fil: Eskelinen, Anu. German Centre for Integrative Biodiversity Research; Alemania Fil: Eskelinen, Anu. Physiological Diversity, Helmholtz Centrefor Environmental Research; Alemania Fil: Eskelinen, Anu. University of Oulu. Ecology & Genetics; Finlandia Fil: Fay, Philip A. Grassland Soil and Water Research Laboratory (USDA-ARS); Estados Unidos Fil: Haider, Sylvia. German Centre for Integrative Biodiversity Research; Alemania Fil: Haider, Sylvia. Martin Luther University. Institute of Biology. Geobotany and Botanical Garden; Alemania Fil: Jentsch, Anke. University of Bayreuth. Department of Soil Ecology. Bayreuth Center of Ecology and Environmental Research (BayCEER); Alemania Fil: Kirkman, Kevin P. University of KwaZulu-Natal. School of Life Sciences; Sudáfrica Fil: McCulley, Rebecca L. University of Kentucky. Department of Plant and Soil Sciences; Estados Unidos Fil: Peri, Pablo Luis. Instituto Nacional de Tecnología Agropecuaria (INTA). Estación Experimental Agropecuaria Santa Cruz; Argentina. Fil: Peri, Pablo Luis. Universidad Nacional de la Patagonia Austral; Argentina. Fil: Peri, Pablo Luis. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Fil: Price, Jodi. Charles Sturt University. Institute for Land, Water and Society; Australia. Fil: Richards, Anna E. CSIRO Land and Water. Northern Territory; Australia. Fil: Risch, Anita C. Swiss Federal Institute for Forest, Snow and Landscape Research WSL; Suiza Fil: Roscher, Christiane. German Centre for Integrative Biodiversity Research; Alemania Fil: Roscher, Christiane. Physiological Diversity, Helmholtz Centre for Environmental Research; Alemania Fil: Schütz, Martin. Swiss Federal Institute for Forest, Snow and Landscape Research WSL; Suiza Fil: Seabloom, Eric William. University of Minnesota. Dept. of Ecology, Evolution, and Behavior; Estados Unidos Fil: Standish, Rachel J. Murdoch University. Harry Butler Institute; Australia. Fil: Stevens, Carly J. Lancaster University. Lancaster Environment Centre; Reino Unido Fil: Tedder, Michelle J. University of KwaZulu-Natal. School of Life Sciences; Sudáfrica Fil: Virtanen, Risto. University of Oulu. Ecology & Genetics; Finlandia. Fil: Spohn, Marie. University of Bayreuth. Department of Soil Ecology. Bayreuth Center of Ecology and Environmental Research (BayCEER); Alemania Fil: Spohn, Marie. Swedish University of Agricultural Sciences. Department of Soil and Environment; Suecia

Fire occurs naturally in grasslands and savannas and it is commonly used as a management tool to influence structure and composition. The ability of a grass plant to recover from fire may be influenced by the fire frequency and intensity (influenced by seasonal weather conditions and fuel load), defoliation and competition from neighbouring plants. This study sought to investigate factors influencing the fire and herbivore dynamics of an evergreen C3 grass, Festuca costata, in a high-altitude C4 dominated grassland in the Drakensberg Mountains of South Africa. We experimentally examined the effect of fire frequency, defoliation and competition from neighbours on the growth dynamics of F. costata. Annual burning significantly decreased aboveground biomass production, number of tillers and tuft circumference of F. costata. Defoliation similarly reduced aboveground biomass production, number of tillers, tuft circumference and specific leaf area of F. costata. Competition from neighbouring plants reduced aboveground biomass production, number of tillers and tuft circumference of F. costata. Given the expected variable impacts of increasing CO2 levels and temperature on C3 and C4 grasses, these results provide insights for managing the matrix of C3 and C4 grasses in the Drakensberg grasslands in a changing climate. Keywords: biodiversity; climate change; herbivory

(2021). Montane rangelands in a changing world. African Journal of Range & Forage Science: Vol. 38, Montane Rangelands in a Changing World, pp. iii-vi.

Background and aims:The amount of nitrogen (N) derived from symbiotic N2 fixation by legumes in grasslands might be affected by anthropogenic N and phosphorus (P) inputs, but the underlying mechanisms are not known. Methods:We evaluated symbiotic N2 fixation in 17 grasslands on four continents that are subjected to the same full-factorial N and P addition experiment, using the 15N natural abundance method.Results:N as well as combined N and P (NP) addition reduced legume biomass by 65% and 45%, respectively, compared to the control, whereas P addition had no significant impact. Element addition had no significant effect on the symbiotic N2 fixation per unit legume biomass. In consequence, the amount of N fixed annually per grassland area was less than half in the N addition treatments compared to control and P addition, irrespective of whether the dominant legumes were annuals or perennials. Conclusion:Our results reveal that N addition mainly impacts symbiotic N2 fixation via reduced biomass of legumes rather than changes in N2 fixation per unit legume biomass. The results show that soil N enrichment by anthropogenic activities significantly reduces N2 fixation in the world’s grasslands, and these effects cannot be reversed by additional P amendment.

Spatial rarity is often used to predict extinction risk, but rarity can also occur temporally. Perhaps more relevant in the context of global change is whether a species is core to a community (persistent) or transient (intermittently present), with transient species often susceptible to human activities that reduce niche space. Using 5–12 yr of data on 1,447 plant species from 49 grasslands on five continents, we show that local abundance and species persistence under ambient conditions are both effective predictors of local extinction risk following experimental exclusion of grazers or addition of nutrients; persistence was a more powerful predictor than local abundance. While perturbations increased the risk of exclusion for low persistence and abundance species, transient but abundant species were also highly likely to be excluded from a perturbed plot relative to ambient conditions. Moreover, low persistence and low abundance species that were not excluded from perturbed plots tended to have a modest increase in abundance following perturbance. Last, even core species with high abundances had large decreases in persistence and increased losses in perturbed plots, threatening the long-term stability of these grasslands. Our results demonstrate that expanding the concept of rarity to include temporal dynamics, in addition to local abundance, more effectively predicts extinction risk in response to environmental change than either rarity axis predicts alone. © 2021 by the Ecological Society of America

Vachellianilotica (L.) Willd. Ex Del. is a multipurpose leguminous tree that is common in grassland and savanna ecosystems in southern and eastern Africa. These ecosystem soils are reported to be acidic and nutrient-limited, specifically with regards to nitrogen (N) and phosphorus (P). The presence of this plant in these terrestrial ecosystems improves soil fertility benefiting the surrounding vegetation due to its ability to fix atmospheric N. This study seeks to understand the N-fixing bacteria symbiosis and physiological adaptations of V. nilotica in these acidic and nutrient-deficient KwaZulu-Natal soils. The soils used for this study were collected from the Ukulinga Grassland Nutrient Experiment located at the Ukulinga research farm of the University of KwaZulu-Natal, Pietermaritzburg, South Africa. Due to long-term soil nutrient addition treatments, these soils offered a diverse nutrient variation for better understanding the effects of acidity and nutrient variation on microbial symbiosis, plant nutrition, and biomass accumulation of V. nilotica. V. nilotica was able to maintain growth by relying on both atmospheric and soil-derived N across all treatments decreasing carbon (C) growth costs. There was an increased reliance on atmospheric-derived N of un-nodulated high N-treated plants. The plants grown in high N + P soils were able to nodulate with various species from the Mesorhizobium genus, which resulted in increased biomass compared to other plants. The results of this study show that V. nilotica can alter N sources to reduce C growth costs. In addition, both nodulating and free-living soil N2 fixing bacteria such as Caulobacter rhizosphaerae, Sphingomonas sp. and Burkholderia contaminans identified in the experimental soils may play an important role under P-deficient conditions.

The importance of maintaining biodiversity in natural grasslands under livestock production systems has been recognised. Forbs contribute more to species richness than grasses in South African gras...

Terrestrial ecosystems have experienced rising nitrogen (N) inputs during the last decades with consequences for belowground carbon (C) and N dynamics. This study investigates how long-term N and phosphorus (P) additions affect microbial community composition, and to what extent microbial homeostasis explains changes in different processes involved in soil C and N cycling in response to nutrient addition. We studied a 66-year-old nutrient addition experiment in a mesic grassland in South Africa, consisting of four different levels of N addition (0, 7, 14, and 21 g N m−2 yr−1) with and without P addition (0, and 9 g P m−2 yr−1). Despite strong changes in the microbial community (observed through 16S rRNA gene and ITS amplicon sequencing), the microbial biomass C:N ratio did not change. N addition decreased microbial N acquisition as indicated by reduced leucine-aminopeptidase activity, and increased microbial net N mineralization. In contrast, predicted relative abundances of functional genes involved in degradation of labile C compounds (e.g. cellulose, hemicellulose, and chitin) as well as β-glucosidase and N-acetylglucosaminidase activities increased with elevated N availability. In combination, this pointed to a more intensive investment of microorganisms into C acquisition upon N addition. In contrast, N addition and associated soil acidification decreased microbial biomass and respiration and altered the community composition with prokaryotes being more affected than fungi. Nitrogen addition increased the relative abundance of gram-positive over gram-negative bacteria and favored taxa with low genome-size. Taken together, our findings support the concept that C and N cycling processes can be explained by the property of the soil microbial community to keep the element ratio of its biomass constant and by its reaction to soil acidification. Our findings suggest that predicted elevated N inputs might largely shape soil C and N cycling because the soil microbial community adjusts metabolic processes, which allows it to maintain its biomass stoichiometry constant.

Applications for exploratory shale gas development via hydraulic fracturing (fracking) have raised concern about energy development impacts in South Africa. Initially, focus was on the arid Karoo, but interest now includes KwaZulu-Natal, a populous, agricultural province with high cultural, ecological, and economic diversity. We conducted focus groups and an online survey to determine how some South Africans perceive fracking. Focus group participants were unanimous in their opposition, primarily citing concerns over water quality and rural way-of-life. The survey confirmed broad consistency with focus group responses. When asked which provinces might be affected by fracking, KwaZulu-Natal ranked behind provinces in the Karoo, suggesting an awareness bias towards Karoo projects. Frequently-identified concerns regarding Agriculture and Natural Resources were Reduced quality of water, Negative impacts to ecosystems and natural biodiversity, Reduced quantity of water, and Pollution hazards. Frequent concerns regarding Social, Cultural, and Local Community issues were Impacts to human health, Visual/aesthetic degradation of tourism areas, Degradation of local infrastructure, and Physical degradation of tourism sites. Most survey respondents were pessimistic about potential benefits of fracking to South Africa's domestic energy supply, and did not agree fracking would reduce negative impacts of coal mining or create jobs. Survey respondents were pessimistic about government's preparedness for fracking and agreed fracking created opportunity for corruption. Many respondents agreed they would consider fracking when voting, and identified needs for more research on fracking in South Africa, which focused heavily on environmental impacts, especially water, in addition to the welfare of local citizens and their communities.

Vachellianilotica (L.) Willd. Ex Del. is a multipurpose leguminous tree that is common in grassland and savanna ecosystems in southern and eastern Africa. These ecosystem soils are reported to be acidic and nutrient-limited, specifically with regards to nitrogen (N) and phosphorus (P). The presence of this plant in these terrestrial ecosystems improves soil fertility benefiting the surrounding vegetation due to its ability to fix atmospheric N. This study seeks to understand the N-fixing bacteria symbiosis and physiological adaptations of V. nilotica in these acidic and nutrient-deficient KwaZulu-Natal soils. The soils used for this study were collected from the Ukulinga Grassland Nutrient Experiment located at the Ukulinga research farm of the University of KwaZulu-Natal, Pietermaritzburg, South Africa. Due to long-term soil nutrient addition treatments, these soils offered a diverse nutrient variation for better understanding the effects of acidity and nutrient variation on microbial symbiosis, plant nutrition, and biomass accumulation of V. nilotica. V. nilotica was able to maintain growth by relying on both atmospheric and soil-derived N across all treatments decreasing carbon (C) growth costs. There was an increased reliance on atmospheric-derived N of un-nodulated high N-treated plants. The plants grown in high N + P soils were able to nodulate with various species from the Mesorhizobium genus, which resulted in increased biomass compared to other plants. The results of this study show that V. nilotica can alter N sources to reduce C growth costs. In addition, both nodulating and free-living soil N2 fixing bacteria such as Caulobacter rhizosphaerae, Sphingomonas sp. and Burkholderia contaminans identified in the experimental soils may play an important role under P-deficient conditions.

Biodiversity—both above‐ and belowground—influences multiple functions in terrestrial ecosystems. Yet, it is unclear whether differences in above‐ and belowground species composition (β‐diversity) are associated with differences in multiple ecosystem functions (e.g., spatial turnover in ecosystem function). Here, we partitioned the contributions of above‐ and belowground β‐diversity and abiotic factors (geographic distance, differences in environments) on the spatial turnover of multiple grassland ecosystem functions. We compiled a dataset of plant and soil microbial communities and six indicators of grassland ecosystem functions (i.e., plant aboveground live biomass, plant nitrogen [N], plant phosphorus [P], root biomass, soil total N, and soil extractable P) from 18 grassland sites on four continents contributing to the Nutrient Network experiment. We used Mantel tests and structural equation models to disentangle the relationship between above‐ and belowground β‐diversity and spatial turnover in grassland ecosystem functions. We found that the effects of abiotic factors on the spatial turnover of ecosystem functions were largely indirect through their influences on above‐ and belowground β‐diversity, and that spatial turnover of ecosystem function was more strongly associated with plant β‐diversity than with soil microbial β‐diversity. These results indicate that changes in above‐ and belowground species composition are one mechanism that interacts with environmental change to determine variability in multiple ecosystem functions across spatial scales. As grasslands face global threats from shrub encroachment, conversion to agriculture, or are lost to development, the functions and services they provide will more strongly converge with increased aboveground community homogenization than with soil microbial community homogenization.

Fire and herbivory are important natural disturbances in grassy biomes. Both drivers are likely to influence be- lowground microbial communities but no studies have unravelled the long-term impact of both fire and herbiv- ory on bacterial and fungal communities. We hypothesized that soil bacterial communities change through disturbance-induced shifts in soil properties (e.g. pH, nutrients) while soil fungal communities change through vegetation modification (biomass and species composition). To test these ideas, we characterised soil physico- chemical properties (pH, acidity, C, N, P and exchangeable cations content, texture, bulk density, moisture), plant species richness and biomass, microbial biomass and bacterial and fungal community composition and di- versity (using 16S and ITS rRNA amplicon sequencing, respectively) in six long-term (18 to 70 years) ecological research sites in South African savanna and grassland ecosystems. We found that fire and herbivory regimes pro- foundly modified soil physico-chemical properties, plant species richness and standing biomass. In all sites, an increase in woody biomass (ranging from 12 to 50%) was observed when natural disturbances were excluded. The intensity and direction of changes in soil properties were highly dependent on the topo-pedo-climatic con- text. Overall, fire and herbivory shaped bacterial and fungal communities through distinct driving forces: edaphic properties (including Mg, pH, Ca) for bacteria, and vegetation (herbaceous biomass and woody cover) for fungi. Fire and herbivory explained on average 7.5 and 9.8% of the fungal community variability, respectively, compared to 6.0 and 5.6% for bacteria. The relatively small changes in microbial communities due to natural disturbance is in stark contrast to dramatic vegetation and edaphic changes and suggests that soil microbial communities, having evolved with disturbance, are resistant to change. This represents both a buffer to short-term anthropogenic- induced changes and a restoration challenge in the face of long-term changes. Ecological drivers Grassland Next generation sequencing Savanna Soil microbial diversity South Africa

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Plant growth is often co-limited by nitrogen (N) and phosphorus (P). Plants might use one element to acquire another (i.e., trading N for P and P for N), which potentially explains synergistic growth responses to NP addition. We studied a 66-yr-old grassland experiment in South Africa that consists of four levels of N addition with and without P addition. We investigated the response of aboveground net primary production (ANPP) to N and P addition over the last 66 yr. Further, we tested whether phosphatase activity and plant P uptake depend on N availability, and vice versa, whether non-symbiotic N2 fixation and plant N uptake depend on P availability. We expected that the interaction of both elements promote processes of nutrient acquisition and contribute to synergistic plant growth effects in response to NP addition. We found synergistic N and P co-limitation of ANPP for the period from 1951 to 2017 but the response to N and P addition diminished over time. In 2017, aboveground P stocks, relative rRNA operon abundance of arbuscular mycorrhizal fungi, and soil organic P storage increased with N fertilization rate when N was added with P compared to the treatment in which only N was added. Further, N addition increased phosphatase activity, which indicates that plants used N to acquire P from organic sources. In contrast, aboveground N stocks and non-symbiotic N2 fixation did not change significantly due to P addition. Taken together, our results indicate that trading N for P likely contributes to synergistic plant-growth response. Plants used added N to mobilize and take up P from organic sources, inducing stronger recycling of P and making the plant community less sensitive to external nutrient inputs. The latter could explain why indications of synergistic co-limitation diminished over time, which is usually overlooked in short-term nutrient addition experiments.

Background Recently there has been considerable focus on the ecosystem services concept which has resulted in important advancements in biodiversity conservation across land management scales. Many have, however, cautioned against the ecosystem services approach because of its focus only on certain aspects of the ecosystem which may be unsustainable in the long term. This has encouraged calls for deeper study into ecosystem functioning using an holistic ecosystem multifunctionality framework. Here greater biodiversity is thought to facilitate greater functioning leading to more sustainable ecosystems. Although ecosystem multifunctionality is a relatively recent development, the general premise is based on the hypothesis that diversity begets stability. However, several key review syntheses have consistently called for ecosystem stability driver-outcome relationship studies to extend beyond traditional measurements. Understanding these relationships requires holistic approaches which are often challenging to investigate experimentally due to resource constraints. Systematically mapping out the relationships between various stability drivers and outcomes could provide a more empirical basis on which both the ecosystem multifunctionality and services land management frameworks could be based. This work outlines the protocol for the first systematic map which will identify and catalogue diversity–stability related studies within the grassland biome. The outcomes of this study will produce a searchable database of the body of literature relevant to the debate and suggest future research directions in both empirical and applied ecology fields. Methods Relevant studies will be sourced from online databases. Inclusion criteria will be applied to the returned articles to identify studies relevant to the primary question; what evidence is available on the drivers of grassland ecosystem stability across a range of outcome measurements. These inclusion criteria will be based on (1) subject population—the grassland biome; (2) possible ecosystem stability drivers and comparators (i.e. measures of diversity, functioning, food web connectedness, and disturbances); and (3) stability outcomes considering all measures of ecosystem stability (i.e. coefficients of variation, changes in ecosystem functionality, resistance to disturbances and invasions, return rates following disturbance). Studies will be screened for relevance and included articles will be critically appraised for meta-analysis and systematic review potential. A narrative synthesis, together with a searchable and expandable database, will be compiled to catalogue the relevant studies. Descriptive summary statistics and bibliometric network analyses will also be presented.

Although most fire research in plant ecology focuses on vegetation responses to burning, shifts in plant community composition wrought by climate change can change wildland fuelbeds and affect fire behaviour such that the nature of fire in these systems is altered. Changes that introduce substantially different fuel types can alter the spatial extent of fire, with potential impacts on community succession and biodiversity. Montane grasslands of sub-Saharan Africa are threatened by climate change because species distributions can shift with climatically determined ranges. We studied the impact of patches of the temperate C3 grass Festuca costata in C4-dominated grassland at the transition between their subalpine ranges in South Africa’s Drakensberg. We used empirical data on fuel moisture and fuel load across F. costata-dominated patches in a C4-dominated matrix in fire spread models to predict the effect of larger, higher-moisture F. costata patches on the spatial extent of fire. Results indicate F. costata reduces fire spread and burn probability in F. costata patches, and the effect increases as live fuel moisture increases and patches get larger. However, as a native species, F. costata does not appear to have the extreme, fire-suppressing effect of non-native C3 grasses in other C4 grasslands. Instead, F. costata patches likely increase variability in the spatial extent of fire in this C4-dominated grassland, which likely translates to spatial variability on vegetation succession.

(2021). Primer of Ecological Restoration - Karen D Holl, published in cooperation with the Society for Ecological Restoration. African Journal of Range & Forage Science: Vol. 38, No. 3, pp. 244-245.

Leucaena leucocephala (Fabaceae) is native to Central America and has invaded many climatic regions of the tropics. In South Africa, the species is categorized as an emerging or incipient weed used as fodder, timber, firewood and in erosion control on degraded habitats. The species is common along the eastern subtropical regions of KwaZulu-Natal (KZN) Province, where it invades grasslands, savannas and edges of forests. Soils of these ecosystems are characterized as nutrient deficient and acidic. Using a pot trial, we determined the effects of the nutrient addition treatments on microbial symbiosis, N nutrition and biomass accumulation of L. leucocephala under greenhouse conditions. After 180 days of growth, plants were harvested, and their utilization of N derived from the atmosphere and from the soil was quantified through determination of δ15N values. L. leucocephala maintained growth and N nutrition by relying on both atmospheric- and soil-derived N across all soil treatments. The NDFA was significantly higher in high P (N1 + P, N2 + P and N3 + P) soils. L. leucocephala was able to nodulate with intermediate and fast-growing strains from the Mesorhizobium and Rhizobium genus in N2 + P grown plants. This shows that L. leucocephala possesses traits that are successful in acquiring nutrients, especially in nutrient limited conditions, by establishing plant symbiosis with multiple bacteria and relying on extracting N from the soil and from the atmosphere through the symbiosis.

A suite of new technologies for developing unconventional petroleum resources, collectively known as fracking, has stirred debate over energy production in South Africa. As an arid country with a growing human population, concern about fracking has understandably focused on water consumption and pollution. But recent research on public perceptions of fracking in South Africa indicates surface impacts to agricultural and nature conservation resources are not given sufficient consideration. We recommend agricultural and conservation organisations understand these potential impacts and draw on lessons learned from rapid fracking development abroad to prepare for production-scale fracking in South Africa (even though it is likely far off). We apply results from research on landowner and operator perspectives on fracking in the rural, agricultural Bakken region of North Dakota, USA, which describe substantial impacts on farm and ranch management and rural way-of-life. We present these experiences as lessons learned that South Africans can use to prepare for fracking from the individual farm to national legislation and regulations.

Soil respiration is often used as an index of fertility because the majority of nutrients are cycled through the microbial biomass. We assessed the role of soil respiration as a measure of resource productivity in three long-term grassland experiments near Pietermaritzburg, South Africa. All of these experiments have shown significant changes in grass species composition and productivity. An ongoing Veld (=field) Fertilizer Experiment (VFE) that manipulated the level of nitrogen fertilizer, phosphorus and lime has been running since 1951. A Burning and Mowing Experiment (BME) has been running since 1950. The third experiment is part of the Nutrient Network (NutNet), a global fertilizer experiment that has been manipulating nitrogen, phosphorus, potassium and micronutrients and has been running since 2009. We found that longer-term experiments were more likely to show significant effects on soil respiration. We found several significant effects in the VFE but no significant differences in soil respiration among fertilization treatments in the shortest-term experiment (NutNet). In the VFE, we found significant differences in soil respiration due to levels of nitrogen fertilizer, form of nitrogen fertilizer (limestone ammonium nitrate and ammonium sulphate), phosphorus and lime. We found no significant relationship between above-ground net primary productivity and soil respiration despite the frequent detection of such a pattern due to the link between soil respiration, soil fertility and productivity. We found that, while there was a consistent increase in total soil nitrogen with increasing levels of nitrogen fertilizer applied, there was a consistent decrease in soil microbial respiration. There was a significant positive correlation between soil respiration and pH. Possible mechanisms behind this are unclear but may involve changes in dominant enzymes and possibly switches between dominance of bacteria and fungi. We also found significant effects of the timing of burning in the BME, but not due to the frequency of burning or the occurrence of mowing. Our results suggest that studies may need to be long-term, for example here at least 10 years, before key functional relationships with soil fertility can be reliably understood.

The paradigmatic hypothesis for the effect of fertilisation on plant diversity represents a one-dimensional trade-off for plants competing for below-ground nutrients (generically) and above-ground light: fertilisation reduces competition for nutrients while increasing biomass and thereby shifts competition for depleted available light. The essential problem of this simple paradigm is that it misses both the multivariate and mechanistic nature of the factors that determine biodiversity as well as their causal relationships. We agree that light limitation, as DeMalach and Kadmon argue, can indeed be an important factor associated with diversity loss, and we presented it as an integral part of our tests of the niche dimension hypothesis. We disagree with DeMalach and Kadmon that light is the ‘main’ factor explaining diversity, because this misrepresents the causal structure represented in the design of our experiment in which multiple nutrient addition was the ultimate causal driver of a suite of correlated responses that included diversity and light, and especially live and dead biomass, which are the factors that control light depletion. Our findings highlight that multiple nutrient limitations can structure plant diversity and composition independently of changes in light and biomass. For example, approximately one-third of our sites showed no significant increase in biomass with greater number of added nutrients yet still lost diversity when nutrients were added. The important message is that while light limitation can be an important contributor to diversity loss, it is not a necessary mechanism. Fil: Harpole, W. Stanley. Helmholtz Center for Environmental Research; Alemania Fil: Sullivan, Lauren L.. University of Minnesota; Estados Unidos Fil: Lind, Eric M.. University of Minnesota; Estados Unidos Fil: Firn, Jennifer. Queensland University of Technology; Australia Fil: Adler, Peter B.. State University of Utah; Estados Unidos Fil: Borer, Elizabeth T.. University of Minnesota; Estados Unidos Fil: Chase, Jonathan. Martin Luther University Halle Wittenberg; Alemania Fil: Fay, Philip A.. USDA-ARS Grassland Soil and Water Research Lab; Estados Unidos Fil: Hautier, Yann. Utrecht University; Países Bajos Fil: Hillebrand, Helmut. University of Oldenburg; Alemania Fil: MacDougall, Andrew S.. University of Guelph; Canadá Fil: Seabloom, Eric W.. University of Minnesota; Estados Unidos Fil: Bakker, Jonathan D.. University of Washington; Estados Unidos Fil: Cadotte, Marc W.. University of Toronto; Canadá Fil: Chaneton, Enrique Jose. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Instituto de Investigaciones Fisiológicas y Ecológicas Vinculadas a la Agricultura. Universidad de Buenos Aires. Facultad de Agronomía; Argentina Fil: Chu, Chengjin. Sun Yat-sen University; China Fil: Hagenah, Nicole. University of KwaZulu-Natal; Sudáfrica Fil: Kirkman, Kevin. University of KwaZulu-Natal; Sudáfrica Fil: La Pierre, Kimberly J.. Smithsonian Environmental Research Center; Estados Unidos Fil: Moore, Joslin L.. Monash University; Australia Fil: Morgan, John W.. La Trobe University; Australia Fil: Prober, Suzanne M.. CSIRO Land and Water; Australia Fil: Risch, Anita C.. Swiss Federal Institute for Forest, Snow and Landscape Research; Suiza Fil: Schuetz, Martin. Swiss Federal Institute for Forest, Snow and Landscape Research; Suiza Fil: Stevens, Carly J.. Lancaster University; Reino Unido

Livestock production is an appropriate land use for mainstreaming biodiversity conservation, but little is known about the impact of grazing strategies on forbs that contribute most species, in grasslands. This study compared the effects of high-density, short-duration stocking (HDG) with no grazing (control) on vegetation structure, composition and demography of selected perennial forbs in a South African mesic grassland in a three-year experiment in KwaZulu-Natal, South Africa. HDG resulted in four-fold more and deeper litter accumulation, which reduced irradiance. Cover of three forb species, including the alien Bidens pilosa, was promoted and cover of another three forb species was reduced under HDG. There were no short-term effects of HDG on diversity, evenness and species richness. Ninety percent of forb species were damaged by HDG; type of damage varied widely from leaves and stems grazed to tearing and shredding by the hooves of cattle. Study of demography revealed that HDG threatened future popul...

The balance of papers in this Special Issue show that how Holistic Planned Grazing™ is managed and where it is used impacts the efficacy of the approach. While we will do well to develop more mechanistic models that can identify these thresholds and test them in real-life situations, it is certain that broad generalisations will not do. We can neither dismiss Holistic Planned Grazing out of hand nor claim that it will work anywhere. Both land-users and scientists should consider the evidence at hand along with their management goals (production, conservation or restoration) before deciding what livestock management approach is appropriate.Keywords: management (natural resources), modelling, Savory, thresholds

High-density, short-duration stocking (HDG) is gaining popularity amongst farmers in the South African mesic grasslands, but little is known about its potential impact on natural resources. Using a fence-line contrast approach, this study compared the long-term effects of HDG with those of other rotational grazing systems at lower densities (LDG) on soil properties and vegetation composition at two sites. Soils at Kokstad and Cedarville were 31% and 19% more compacted under HDG than LDG, respectively, but did not differ in total nitrogen, total carbon, available phosphorus or pH. At Kokstad, HDG had promoted unpalatable grass species at the expense of palatable grass species. Forb species composition differed substantially between HDG and LDG, with HDG having 13 and LDG having 16 unique species. A radical prostrate forb growth habit was favoured over a cauline erect habit under HDG. At Cedarville, few grass or forb species showed differences between HDG and LDG, possibly because of an overriding influence...

Rangeland management strategies must be based on robust ecological and economic concepts if they are to be effective and profitable. Thus, the aim of this paper was to examine concepts related to grazing and resting of grassland and associated effects on grassland productivity and energy flow to livestock. Our review and models demonstrate that grassland and livestock productivity is determined by lagged effects of the previous year’s grazing and by grassland maturation, which influence grass growth rates, as well as forage quality. Livestock productivity is also influenced by the scale at which livestock are able to make foraging decisions in time and space, which is restricted by increasing intensity of management (increasing rotation and declining paddock size). Finally, overstocking will override key management initiatives, such as effective recovery periods, leading to rangeland degradation. Thus, in variable climates, stocking rate should be set conservatively to allow easier adaptation of animal numbers to rainfall variability from year to year. We suggest several key concepts that can be used to design and assess grazing management strategies and make suggestions on how to implement these concepts.Keywords: forage quality, grassland productivity, Holistic Planned Grazing™, livestock performance, rotational grazing

Soil nitrogen mineralisation (Nmin), the conversion of organic into inorganic N, is important for productivity and nutrient cycling. The balance between mineralisation and immobilisation (net Nmin) varies with soil properties and climate. However, because most global-scale assessments of net Nmin are laboratory-based, its regulation under field-conditions and implications for real-world soil functioning remain uncertain. Here, we explore the drivers of realised (field) and potential (laboratory) soil net Nmin across 30 grasslands worldwide. We find that realised Nmin is largely explained by temperature of the wettest quarter, microbial biomass, clay content and bulk density. Potential Nmin only weakly correlates with realised Nmin, but contributes to explain realised net Nmin when combined with soil and climatic variables. We provide novel insights of global realised soil net Nmin and show that potential soil net Nmin data available in the literature could be parameterised with soil and climate data to better predict realised Nmin., Nitrogen mineralisation (Nmin), an important index of soil fertility, is often determined in the laboratory, with an uncertain relationship to Nmin under field conditions. Here the authors show that combining laboratory measurements with environmental data greatly improves predictions of field Nmin at a global scale.

Commercial seed mixes and various mechanical planting methods have been implemented to overcome barriers to successful restoration of degraded grasslands but their effectiveness in aiding the restoration process has been understudied. Harvested seed must be cleaned of chaff to allow mechanical planting, which can exclude small seeds. We propose that fluid seed drilling techniques (suspending uncleaned harvested seed in a cellulose-based gel matrix, which may then be injected into the soil with a mechanical planter) may overcome these challenges. As a preliminary investigation of the proposed method we designed a greenhouse pot trial experiment aimed at quantifying restoration success through the measurement of seedling recruitment, biomass production and composition responses of commercial and harvested seed mixes under simulated fluid seed drilling and smoke water priming (collectively termed application method). These data were analysed using generalised linear modelling and multivariate analyses. Commercial mixed seed resulted in greater recruitment, whereas harvested seed produced the greatest biomass production. There was large multivariate separation between seed types and seed type × application method centroids for recruitment responses and only seed type for biomass responses. Application method had minimal effects on restoration measurements whilst seed type greatly affected these measurements. Careful selection of the restoration seed mix is advised.Keywords: biomass, cellulose gel planting method, germination, multivariate diversity, restoration seed mixes, smoke water seed priming

Human activities have increased nitrogen (N) and phosphorus (P) inputs in terrestrial ecosystems and altered carbon (C) availability, shifting the stoichiometry of microbial substrates in soils, such as the C:N:P ratios of the dissolved organic matter pool. These stoichiometric deviations between microbial biomass and its substrate may control microbial processes of N cycling. We studied the effects of this stoichiometric mismatch using a full factorial N and P addition experiment replicated in six grassland ecosystems in South Africa, the USA, and the UK. We found that N and P addition changed the dissolved organic matter C:N ratio, but not the C:N ratio of the soil microbial biomass. Compared to P addition, N addition decreased microbial N acquisition via non-symbiotic N2 fixation by −55% and increased microbial N release via net N mineralization by +134%. A possible explanation is that the dissolved elements, e.g., dissolved organic C (DOC) and dissolved total N (DN), serve as the main microbial substrate and its C:N ratio defines whether N is scarce or abundant with respect to microbial demands. If N is available in excess relative to microbial demands, net N mineralization increases. In contrast, when N is scarce, immobilization outweighs release decreasing net N mineralization. However, the activity of leucine aminopeptidases, which decompose peptides, was not affected by nutrient additions. Further, C rather than P availability may control the rates of non-symbiotic N2 fixation in the six studied grassland sites. In conclusion, globally increasing nutrient inputs change processes of microbial N acquisition and release in grassland ecosystems and these changes are largely driven by shifts in substrate stoichiometry.

Summary 1.Productivity and herbivory often interact to shape plant community composition and species richness with levels of production mediating the impact of herbivory. Yet, differences in herbivore traits such as size, feeding guild, and dietary requirements may result in different impacts of diverse herbivore guilds across productivity gradients. 2.We used size-selective herbivore exclosures to separate the effects of herbivory by larger herbivores, such as elephant, Burchell's zebra, and blue wildebeest from those of medium/smaller herbivores, such as impala and warthog, on herbaceous plant communities. These exclosures were established along a 10-fold productivity gradient, ranging from 90-950 g m−2 of standing plant biomass in the Kruger National Park, South Africa. 3.Exclusion of all herbivores generally increased plant species richness at low productivity but decreased richness at high productivity. Exclusion of medium/smaller herbivores (e.g. impala, warthog) showed stronger effects on plant richness, particularly loss of forbs, at higher productivity rather than at lower productivity. In contrast, exclusion of larger herbivores had stronger effects on plant richness, typically with increasing forb richness, at low rather than high productivity. 4.The change in species richness appeared linked to changes in light availability following herbivore exclusion. Strong increases in shading led to declines in species richness while more moderate increases in shading led in increases in species richness, possibly due to amelioration of heat and water stress by modest increases in shading. 5.Increasing plant dominance, which likely alters multiple mechanisms of plant interactions, was correlated with declines in plant richness following herbivore exclusion. The impact of increasing dominance on plant richness operated independent of productivity, with the exclusion of impala appearing particularly important in driving this relationship. 6.Synthesis. We show that the impact of herbivore losses on plant diversity will be strongly situation dependent and will vary with the herbivores lost (e.g. larger vs. smaller, grazers vs. browsers), plant functional type (e.g. grasses vs. forbs), and environmental context (e.g. productivity). Although larger herbivores are often emphasized for their strong impacts on community dynamics and ecosystem processes, we show that smaller, abundant herbivores can exert strong top-down control on plant communities. This article is protected by copyright. All rights reserved.

In wetlands the effects of fire on vegetation dynamics are somewhat uncertain. A change detection analysis in the herbaceous wetlands of KwaMbonambi, South Africa, which were subject to frequent fires, revealed that in 1937 the study area comprised grassland (69%), herbaceous wetland (25%), indigenous swamp forest (4%) and tree plantations (1%). However, by 1970, tree plantations occupied 78% of the landscape and grasslands and herbaceous wetlands had declined to 9% and 6%, respectively, whereas indigenous swamp forest had increased to 6%. By 2009 tree plantations had been removed from the wetland areas. Despite this opportunity for herbaceous wetlands to recover their historical extent, they decreased to only 2%, mostly changing to indigenous swamp forest or to an herbaceous/fern (Stenochlaena tenuifolia)/woodland mosaic. Fire records showed suppression of fire to be an important contributing factor, particularly in wetlands that had been disturbed by tree plantations, although subsequently removed. A pi...

Fire and grazing, key determinants of structure and function of savanna grasslands worldwide, have been extensively altered by humans. We used existing long-term manipulations of fire and grazing in North American and South African mesic savanna grasslands, as well as new experiments, to determine whether the impacts of fire and grazing by large herbivores differed between these systems. We found that despite a body of literature suggesting that these savanna grasslands respond uniquely to fire and grazing, their ecosystem responses (aboveground productivity) were generally similar. In contrast, plant-community responses to fire and herbivores diverged strongly between systems. The differences in plant-community responses, as well as convergence in ecosystem function, were underpinned by a common mechanism: the response of grass dominance to changing fire and grazing regimes. As a result, divergent responses of plant communities to altered fire and grazing regimes did not preclude convergence in ecosystem function.

In areas with diverse herbivore communities such as African savannas, the frequency of disturbance by fire may alter the top–down role of different herbivore species on plant community dynamics. In a seven year experiment in the Kruger National Park, South Africa, we examined the habitat use of nine common herbivore species across annually burned, triennially burned and unburned areas. We also used two types of exclosures (plus open access controls) to examine the impacts of different herbivores on plant community dynamics across fire disturbance regimes. Full exclosures excluded all herbivores > 0.5 kg (e.g. elephant, zebra, impala) while partial exclosures allowed access only to animals with shoulder heights = 0.85 m (e.g. impala, steenbok). Annual burns attracted a diverse suite of herbivores, and exclusion of larger herbivores (e.g. elephant, zebra, wildebeest) increased plant abundance. When smaller species, mainly impala, were also excluded there were declines in plant diversity, likely mediated by a decline in open space available for colonization of uncommon plant species. Unburned areas attracted the least diverse suite of herbivores, dominated by impala. Here, herbivore exclusion, especially of impala, led to strong declines in plant richness and diversity. With no fire disturbance, herbivore exclusion led to competitive exclusion via increases in plant dominance and light limitation. In contrast, on triennial burns, herbivore exclusion had no effect on plant richness or diversity, potentially due to relatively little open space for colonization across exclosure treatments but also little competitive exclusion due to the intermediate fire disturbance. Further, the diverse suite of grazers and browsers on triennial burns may have had a compensating effect of on the diversity of grasses and forbs. Ultimately, our work shows that differential disturbance regimes can result in differential consumer pressure across a landscape and result in heterogeneous patterns in top–down control of community dynamics.

Soil microbial carbon use efficiency (CUE), defined as the ratio between carbon (C) allocated to growth and C taken up by microorganisms, is pivotal for the understanding of C cycling in terrestrial ecosystems. Soil microbial CUE is thought to increase under nitrogen (N) addition, thereby mediating the effects of atmospheric N deposition on C cycling in soils. We studied the effects of N, phosphorus (P), and combined N and P addition on soil microbial CUE from a total of six grassland soils from South Africa, USA, and UK. Microbial CUE varied between 25 and 57% with a mean value of 40% across all sites, depth increments, and treatments. Most of the site variability in microbial CUE was explained by sand content, mean annual precipitation and temperature, and the dissolved organic C:dissolved N ratio. Soil microbial CUE as well as microbial biomass turnover time were robust to changes in N, P, and NP supply. However, N addition significantly reduced microbial respiration and C uptake in the topsoil. Taken together, N, P, and NP addition did not influence microbial CUE and biomass turnover time in grassland soils on different continents, indicating that microbial CUE varies little despite large changes in element inputs. Consequently, increased N inputs to soil may have a smaller impact on microbial CUE and biomass turnover time, and therefore C cycling in grassland soils, than expected and models assuming increased CUE with increasing N inputs could overestimate future C storage.

Although indigenous to southern Africa, Pechuel-loeschea leubnitziae is considered a problematic weed as it forms dense monotypic stands in the grasslands and woodlands of the Okavango Delta, Botswana, and is associated with veld degradation and a dominance of shade-tolerant, poor-quality grass species. Generalised linear modelling was used to determine the effect of the following environmental conditions on the likelihood of occurrence of P. leubnitziae: type of land use, vegetation type, burning history, total woody species density, soil pH and soil texture. There was no difference in P. leubnitziae presence under wildlife, large livestock and small livestock land use (p = 0.227). Only vegetation type (p

Savanna vegetation is characterized by tree–grass co-existence that can experience intense water limitation, yet the water relations of these savanna plants are poorly understood. We examined the water sources for trees and grasses in different seasons and investigated the importance of hydraulic redistribution in three tree species inhabiting a semi-arid savanna in South Africa. We used natural variation in H and O stable isotope composition of source waters to identify the principal water sources for these plants. We conducted an experiment by labelling deep-soil (2.5-m depth) with a deuterium tracer. Seasonal differences in the stable isotope composition of water in trees and grasses indicated that there was water-source use partitioning as well as overlap. Trees and grasses used water from the topsoil after rainfall indicating overlap of water-source use. All tree species shifted to groundwater or subsoil water use when there was no water in the topsoil indicating partitioning of water use. Grasses always used water from the topsoil. The seasonal changes in water-source use by trees and grasses indicated possible shifts in tree–grass interactions during different periods of the year. The tracer experiment confirmed hydraulic redistribution in all the three tree species and water transfer to grasses via the topsoil. However, this occurred only in the dry season. Our observations and experimental results indicate the potential for facilitation effects by trees to their understory grasses and show that dry season hydraulic redistribution from trees to grasses could be an important facilitative mechanism maintaining tree–grass co-existence in savannas.

Theory predicts that nutrient addition to soil reduces the benefit of mycorrhizal associations to plants. This study sought to determine how long-term management of soil fertility influences the abundance of fungal mycelium. Ingrowth mesh bags were used to measure mycelial biomass in grassland plots fertilised (nitrogen [N] and phosphorus [P]) and limed for 60 years. Mycelial biomass more than tripled with N addition, whereas P addition showed modest increases in biomass. Applied in combination, N and P fertilisers had no effect on mycelial biomass. Liming alone had no effect but increased biomass when it was applied in combination with N. The results of this study suggest that the fertiliser affected the biomass of the fungal community, and may also change the composition of the fungal community.

Biodiversity is declining in many local communities while also becoming increasingly homogenized across space. Experimentsshow that local plant species loss reduces ecosystem functioning and services, but the role of spatial homogenization of communitycomposition and the potential interaction between diversity at different scales in maintaining ecosystem functioningremains unclear, especially when many functions are considered (ecosystem multifunctionality). We present an analysis ofeight ecosystem functions measured in 65 grasslands worldwide. We find that more diverse grasslands?those with both species-rich local communities (α-diversity) and large compositional differences among localities (β-diversity)?had higher levelsof multifunctionality. Moreover, α- and β-diversity synergistically affected multifunctionality, with higher levels of diversity at one scale amplifying the contribution to ecological functions at the other scale. The identity of species influencing ecosystem functioning differed among functions and across local communities, explaining why more diverse grasslands maintained greater functionality when more functions and localities were considered. These results were robust to variation in environmental drivers. Our findings reveal that plant diversity, at both local and landscape scales, contributes to the maintenance of multiple ecosystem services provided by grasslands. Preserving ecosystem functioning therefore requires conservation of biodiversity both within and among ecological communities. Fil: Hautier, Yann. University of Utrecht; Países Bajos Fil: Isbell, Forest. University of Minnesota; Estados Unidos Fil: Borer, Elizabeth T.. University of Minnesota; Estados Unidos Fil: Seabloom, Eric W.. University of Minnesota; Estados Unidos Fil: Harpole, Stanley. Martin Luther University Halle-Wittenberg; Alemania Fil: Lind, Eric M.. University of Minnesota; Estados Unidos Fil: MacDougall, Andrew. University of Guelph; Canadá Fil: Stevens, Carly. Lancaster University, ; Reino Unido Fil: Adler, Peter. Utah State University; Estados Unidos Fil: Alberti, Juan. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Bakker, Jonathan D. University of Washington, ; Estados Unidos Fil: Brudvig, Lars A. Michigan State University; Estados Unidos Fil: Buckley, Yvonne M. Trinity College Dublin; Irlanda Fil: Cadotte, Marc. University of Toronto Scarborough,; Canadá Fil: Caldeira, María C. University of Lisbon; Portugal Fil: Chaneton, Enrique Jose. Universidad de Buenos Aires; Argentina Fil: Chu, Chengjin. Sun Yat-sen University; China Fil: Daleo, Pedro. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Dickman, Christopher R. The University of Sydney; Australia Fil: Dwyer, John M. The University of Queensland; Australia Fil: Eskelinen, Anu. University of Oulu; Finlandia Fil: Fay, Philip. USDA-ARS, Grassland Soil and Water Lab; Estados Unidos Fil: Firn, Jennifer. Queensland University of Technology; Australia Fil: Hagenah, Nicole. University of KwaZulu-Natal; Sudáfrica Fil: Hillebrand, Helmut. University Oldenburg; Alemania Fil: Iribarne, Oscar Osvaldo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Kirkman, Kevin P.. University of KwaZulu-Natal; Sudáfrica Fil: Knops, Johannes M. H. University of Nebraska, Lincoln; Estados Unidos Fil: La Pierre, Kimberly J. Smithsonian Environmental Research Center; Estados Unidos Fil: McCulley, Rebecca L. University of Kentucky; Estados Unidos Fil: Morgan, John W.. La Trobe University; Australia Fil: Pärtel, Meelis. University of Tartu; Estonia Fil: Pascual, Jesus Maria. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Price, Jodi N. Charles Sturt University; Australia Fil: Prober, Suzanne M. CSIRO Land and Water; Australia Fil: Risch, Anita C. Swiss Federal Institute of Forest, Snow and Landscape Research; Suiza Fil: Sankaran, Mahesh. National Centre for Biological Sciences; India Fil: Schuetz, Martin. Swiss Federal Institute of Forest, Snow and Landscape Research; Suiza Fil: Standish, Rachel J. Murdoch University; Australia Fil: Virtanen, Risto. University of Oulu; Finlandia Fil: Wardle, Glenda M. The University of Sydney; Australia Fil: Yahdjian, María Laura. UBA; Argentina Fil: Hector, Andy. University of Oxford; Reino Unido

Herbivory and fire shape plant community structure in grass-dominated ecosystems, but these disturbance regimes are being altered around the world. To assess the consequences of such alterations, we excluded large herbivores for seven years from mesic savanna grasslands sites burned at different frequencies in North America (Konza Prairie Biological Station, Kansas, USA) and South Africa (Kruger National Park). We hypothesized that the removal of a single grass-feeding herbivore from Konza would decrease plant community richness and shift community composition due to increased dominance by grasses. Similarly, we expected grass dominance to increase at Kruger when removing large herbivores, but because large herbivores are more diverse, targeting both grasses and forbs, at this study site, the changes due to herbivore removal would be muted. After seven years of large-herbivore exclusion, richness strongly decreased and community composition changed at Konza, whereas little change was evident at Kruger. We found that this divergence in response was largely due to differences in the traits and numbers of dominant grasses between the study sites rather than the predicted differences in herbivore assemblages. Thus, the diversity of large herbivores lost may be less important in determining plant community dynamics than the functional traits of the grasses that dominate mesic, disturbance-maintained savanna grasslands.

The interaction, both above and belowground, between tree saplings and the surrounding grass sward is dependent on solar radiation, temperature, rainfall, soil depth, tree species and sward composition. These factors, as well as level of sward defoliation, influence whether the system will remain as savanna or move towards a woodland or grassland state. The effects of above- and belowground competition between grasses and two Acacia species and the effects of soil depth on these interactions were examined by planting A. karroo and A. nilotica seedlings into a natural sward on three different soil depths. Three aboveground treatments: full shading, reduced shading by tying back the neighboring grasses and reduced shading by clipping, and two belowground treatments: full and no belowground competition, were used. Plant size increased with increasing soil depth, while belowground competitive intensity was unaffected. Removing belowground competition increased sapling biomass by half (P < 0.05) on all soil depths. By contrast, reduced shading had little effect at all soil depths, whereas sward clipping increased sapling biomass (47%) on shallow soils only (P = 0.027), indicating that encroachment on shallow soils may result from factors that decrease root vigor of the surrounding grasses rather than light competition. Irrespective of soil depth, root competition appeared to be the major factor influencing sapling growth rates, thus grazing management practices that reduce grass root productivity are expected to result in woody encroachment.

Savanna plant communities change considerably across time and space. The processes driving savanna plant species diversity, coexistence and turnover along environmental gradients are still unclear. Understanding how species respond differently to varying environmental conditions during the seedling stage, a critical stage for plant population dynamics, is needed to explain the current composition of plant communities and to enable us to predict their responses to future environmental changes. Here we investigate whether seedling response to changes in resource availability, and to competition with grass, varied between two functional groups of African savanna trees: species with small leaves, spines and N-fixing associations (fine-leaved species), and species with broad leaves, no spines, and lacking N-fixing associations (broad-leaved species). We show that while tree species were strongly suppressed by grass, the effect of resource availability on seedling performance varied considerably between the two functional groups. Nutrient inputs increased stem length only of broad-leaved species and only under an even watering treatment. Low light conditions benefited mostly broad-leaved species’ growth. Savannas are susceptible to ongoing global environment changes. Our results suggest that an increase in woody cover is only likely to occur in savannas if grass cover is strongly suppressed (e.g. by fire or overgrazing). However, if woody cover does increase, broad-leaved species will benefit most from the resulting shaded environments, potentially leading to an expansion of the distribution of these species. Eutrophication and changes in rainfall patterns may also affect the balance between fine- and broad-leaved species.

Aims With the increase of game farming in woody areas, as well as the pressing need to assess the spatial distribution of carbon, the quantification of available browse biomass has become an important subject to refine. We aimed to (1) calculate tree canopy volume (CVol) for varying tree canopy shapes and for strata within those shapes to cater for different herbivore browse heights, (2) develop a relation to estimate available browse of deciduous (micro- and mesophyllous) and evergreen (macrophyllous) semi-arid savanna tree species using our calculated CVols, and (3) demonstrate the application of the use of partial volumes and the resultant regression models in the determination of available browse biomass for browsing game species. Location Four key savanna tree species were sampled in the open wooded grassland areas of three game reserves in northeastern KwaZulu-Natal, South Africa: Bonamanzi Game Park, Mun-Ya-Wana Private Game Reserve and Pongola Game Reserve. Methods Tree CVol determination involved calculation of partial volumes, using the integration of a series of circles, allowing a wider range of tree shapes to be included. Non-linear regression was used to investigate the relation between available browse biomass of deciduous microphyllous Acacia nilotica, evergreen macrophyllous Euclea divinorum and Gymnosporia senegalensis and deciduous mesophyllous Spirostachys africana and their respective calculated partial CVols. Results Exponential non-linear regression of the natural logarithm of CVol data provided an accurate and precise description of the tree CVol–available browse biomass relation. Significant relations were obtained over a 2-yr period (2003–2004) for species–site models and general species models. The incorporation of tree foliage densities in available browse biomass estimation further improved model accuracy and precision. Conclusion The general models provided an objective means of determining the browsing capacity of a tree population. In addition, estimation of carbon sequestration could be related to the model output, particularly when changes over time are recorded. Regular measurement of available browse comprises a useful monitoring tool to evaluate impact of animals on trees and shrubs.

Themeda triandra is a perennial tussock grass endemic to Africa, Australia and Asia. Within these regions it is found across a broad range of climates, geological substrates and ecosystems. Because it is widespread across these areas it has great economic and ecological value, as it is a relatively palatable species across most of its range. It is of critical importance in supporting local populations of both native and introduced herbivores, and is thus central to wildlife and livestock production, and consequently rural livelihoods. It is an important climax or subclimax species that is well adapted to fire, a common element of many areas where it is found. Inappropriate grazing management, however, can result in a decline of Themeda, as it is not well adapted to an uninterrupted, selective grazing regime. A decline in abundance of Themeda in a grassland is usually coupled to a decline in grazing value, species richness, cover and ecosystem function. In spite of its significant ecological and economic i...