Determinants of woody encroachment and cover in African savannas

Savanna ecosystems are an integral part of the African landscape and sustain the livelihoods of millions of people. Woody encroachment in savannas is a widespread phenomenon but its causes are widely debated. We review the extensive literature on woody encroachment to help improve understanding of the possible causes and to highlight where and how future scientific efforts to fully understand these causes should be focused. Rainfall is the most important determinant of maximum woody cover across Africa, but fire and herbivory interact to reduce woody cover below the maximum at many locations. We postulate that woody encroachment is most likely driven by CO₂ enrichment and propose a two-system conceptual framework, whereby mechanisms of woody encroachment differ depending on whether the savanna is a wet or dry system. In dry savannas, the increased water-use efficiency in plants relaxes precipitation-driven constraints and increases woody growth. In wet savannas, the increase of carbon allocation to tree roots results in faster recovery rates after disturbance and a greater likelihood of reaching sexual maturity. Our proposed framework can be tested using a mixture of experimental and earth observational techniques. At a local level, changes in precipitation, burning regimes or herbivory could be driving woody encroachment, but are unlikely to be the explanation of this continent-wide phenomenon.     

Savanna woody encroachment is widespread across three continents

Tropical savannas are a globally extensive biome prone to rapid vegetation change in response to changing environmental conditions. Via a meta‐analysis, we quantified savanna woody vegetation change spanning the last century. We found a global trend of woody encroachment that was established prior the 1980s. However, there is critical regional variation in the magnitude of encroachment. Woody cover is increasing most rapidly in the remaining uncleared savannas of South America, most likely due to fire suppression and land fragmentation. In contrast, Australia has experienced low rates of encroachment. When accounting for land use, African savannas have a mean rate annual woody cover increase two and a half times that of Australian savannas. In Africa, encroachment occurs across multiple land uses and is accelerating over time. In Africa and Australia, rising atmospheric CO₂, changing land management and rainfall are likely causes. We argue that the functional traits of each woody flora, specifically the N‐fixing ability and architecture of woody plants, are critical to predicting encroachment over the next century and that African savannas are at high risk of widespread vegetation change.     

A patch-dynamics approach to savanna dynamics and woody plant encroachment – Insights from an arid savanna

The coexistence of woody and grassy plants in savannas has often been attributed to a rooting-niche separation (two-layer hypothesis). Water was assumed to be the limiting resource for both growth forms and grasses were assumed to extract water from the upper soil layer and trees and bushes from the lower layers. Woody plant encroachment (i.e. an increase in density of woody plants often unpalatable to domestic livestock) is a serious problem in many savannas and is believed to be the result of overgrazing in ‘two-layer systems’. Recent research has questioned the universality of both the two-layer hypothesis and the hypothesis that overgrazing is the cause of woody plant encroachment.

We present an alternative hypothesis explaining both tree–grass coexistence and woody plant encroachment in arid savannas. We propose that woody plant encroachment is part of a cyclical succession between open savanna and woody dominance and is driven by two factors: rainfall that is highly variable in space and time, and inter-tree competition. In this case, savanna landscapes are composed of many patches (a few hectares in size) in different states of transition between grassy and woody dominance, i.e. we hypothesize that arid savannas are patch-dynamic systems. We summarize patterns of tree distribution observed in an arid savanna in Namibia and show that these patterns are in agreement with the patch-dynamic savanna hypothesis. We discuss the applicability of this hypothesis to fire-dominated savannas, in which rainfall variability is low and fire drives spatial heterogeneity.

We conclude that field studies are more likely to contribute to a general understanding of tree–grass coexistence and woody plant encroachment if they consider both primary (rain and nutrients) and secondary (fire and grazing) determinants of patch properties across different savannas.     

Fire‐sensitive species dominate seed rain after fire suppression: Implications for plant community diversity and woody encroachment in the Cerrado

Woody encroachment is becoming common in tropical savannas. We studied natural seed rain and performed seed addition experiments in a Brazilian savanna that had not been burned for several decades. We found greater abundance of fire‐sensitive species in the seed rain, likely contributing to woody encroachment. Flexible fire management policies that allow for natural and prescribed fires may be required to maintain savanna diversity.     

Woody cover in African savannas: the role of resources, fire and herbivory

Aim To determine the functional relationships between, and the relative importance of, different driver variables (mean annual precipitation, soil properties, fire and herbivory) in regulating woody plant cover across broad environmental gradients in African savannas. Location Savanna grasslands of East, West and Southern Africa. Methods The dependence of woody cover on mean annual precipitation (MAP), soil properties (texture, nitrogen mineralization potential and total phosphorus), fire regimes, and herbivory (grazer, browser + mixed feeder, and elephant biomass) was determined for 161 savanna sites across Africa using stochastic gradient boosting, a refinement of the regression tree analysis technique. Results All variables were significant predictors of woody cover, collectively explaining 71% of the variance in our data set. However, their relative importance as regulators of woody cover varied. MAP was the most important predictor, followed by fire return periods, soil characteristics and herbivory regimes. Woody cover showed a strong positive dependence on MAP between 200 and 700 mm, but no dependence on MAP above this threshold when the effects of other predictors were accounted for. Fires served to reduce woody cover below rainfall‐determined levels. Woody cover showed a complex, non‐linear relationship with total soil phosphorus, and was negatively correlated with clay content. There was a strong negative dependence of woody cover on soil nitrogen (N) availability, suggesting that increased N‐deposition may cause shifts in savannas towards more grassy states. Elephants, mixed feeders and browsers had negative effects on woody cover. Grazers, on the other hand, depressed woody cover at low biomass, but favoured woody vegetation when their biomass exceeded a certain threshold. Main conclusions Our results indicate complex and contrasting relationships between woody cover, rainfall, soil properties and disturbance regimes in savannas, and suggest that future environmental changes such as altered precipitation regimes, N‐enrichment and elevated levels of CO₂ are likely to have opposing, and potentially interacting, influences on the tree–grass balance in savannas.     

Short‐term effects of herbicides and a prescribed fire on restoration of a shrub‐encroached pine savanna

Shrub encroachment occurring worldwide in savannas and grasslands has commonly been hypothesized to result from anthropogenically altered environments. Two disturbance‐based approaches to restoration have involved: (1) application of selective herbicides to reduce density/cover of shrubs; (2) reinstatement of natural fire regimes to generate environmental conditions favoring herbaceous species. We studied short‐term responses of native shrubs, vines, and grasses in a Louisiana pine savanna to herbicides followed by a prescribed fire and fire alone. Plots established in the summer, 2013, were hand‐sprayed in the fall with Imazapyr and Triclopyr, Triclopyr alone, or no herbicide, then prescribed burned the following spring. Numbers of species of shrubs and vines at scales of 1 and 100 m², numbers of stems and regrowth of stems produced by six common species of shrubs, and the number of flowering culms of perennial C₄ grasses were assessed postfire in 2014. Compared with fire alone, herbicides followed by fire resulted in (1) small reductions in species richness of shrubs and no effects on vines, (2) fewer stems comprising shrub genets, but similar postfire regrowth of resprouting shrub stems, and (3) fewer flowering culms of C₄ grasses. Underground storage organs of savanna shrubs and vines survived both aboveground disturbances. Thus, single applications of herbicides followed by fires reduced, but did not reverse shrub encroachment and negatively affected grasses. Because effects of herbicides overrode those of prescribed fires, these disturbances did not act synergistically, suggesting that reinstating natural fire regimes should be a priority in restoration of savannas and grasslands.     

Positive and negative effects of grass,cattle, and wild herbivores on <Emphasis Type="Italic">Acacia</Emphasis> saplings in an East African savanna

Plant–plant interactions can be a complex mixture of positive and negative interactions, with the net outcome depending on abiotic and community contexts. In savanna systems, the effects of large herbivores on tree–grass interactions have rarely been studied experimentally, though these herbivores are major players in these systems. In African savannas, trees often become more abundant under heavy cattle grazing but less abundant in wildlife preserves. Woody encroachment where cattle have replaced wild herbivores may be caused by a shift in the competitive balance between trees and grasses. Here we report the results of an experiment designed to quantify the positive, negative, and net effects of grasses, wild herbivores, and cattle on Acacia saplings in a Kenyan savanna. Acacia drepanolobium saplings under four long-term herbivore regimes (wild herbivores, cattle, cattle + wild herbivores, and no large herbivores) were cleared of surrounding grass or left with the surrounding grass intact. After two years, grass-removal saplings exhibited 86% more browse damage than control saplings, suggesting that grass benefited saplings by protecting them from herbivory. However, the negative effect of grass on saplings was far greater; grass-removal trees accrued more than twice the total stem length of control trees. Where wild herbivores were present, saplings were browsed more and produced more new stem growth. Thus, the net effect of wild herbivores was positive, possibly due to the indirect effects of lower competitor tree density in areas accessible to elephants. Additionally, colonization of saplings by symbiotic ants tracked growth patterns, and colonized saplings experienced lower rates of browse damage. These results suggest that savanna tree growth and woody encroachment cannot be predicted by grass cover or herbivore type alone. Rather, tree growth appears to depend on a variety of factors that may be acting together or antagonistically at different stages of the tree’s life cycle.     

Trade‐offs between savanna woody plant diversity and carbon storage in the Brazilian Cerrado

Incentivizing carbon storage can be a win‐win pathway to conserving biodiversity and mitigating climate change. In savannas, however, the situation is more complex. Promoting carbon storage through woody encroachment may reduce plant diversity of savanna endemics, even as the diversity of encroaching forest species increases. This trade‐off has important implications for the management of biodiversity and carbon in savanna habitats, but has rarely been evaluated empirically. We quantified the nature of carbon‐diversity relationships in the Brazilian Cerrado by analyzing how woody plant species richness changed with carbon storage in 206 sites across the 2.2 million km² region at two spatial scales. We show that total woody plant species diversity increases with carbon storage, as expected, but that the richness of endemic savanna woody plant species declines with carbon storage both at the local scale, as woody biomass accumulates within plots, and at the landscape scale, as forest replaces savanna. The sharpest trade‐offs between carbon storage and savanna diversity occurred at the early stages of carbon accumulation at the local scale but the final stages of forest encroachment at the landscape scale. Furthermore, the loss of savanna species quickens in the final stages of forest encroachment, and beyond a point, savanna species losses outpace forest species gains with increasing carbon accumulation. Our results suggest that although woody encroachment in savanna ecosystems may provide substantial carbon benefits, it comes at the rapidly accruing cost of woody plant species adapted to the open savanna environment. Moreover, the dependence of carbon‐diversity trade‐offs on the amount of savanna area remaining requires land managers to carefully consider local conditions. Widespread woody encroachment in both Australian and African savannas and grasslands may present similar threats to biodiversity.     

Ecosystem responses to woody plant encroachment in a semiarid savanna rangeland

Woody plant encroachment alters the structure and function of rangeland ecosystems. The objective of this study was to explore the association between woody plant encroachment and various ecosystem properties (i.e. vascular plant species diversity, richness, evenness, soil organic matter, herbaceous biomass, leaf litter and bare ground cover) in a semiarid savanna rangeland, and also to test whether the relationships were influenced by woody species composition, elevation and site. We carried out a vegetation survey in four rangeland sites in the lower Omo region of southwestern Ethiopia, and regressed each one of the ecosystem properties, separately, against woody plant density, elevation and site using multiple linear regressions. We found that vascular plant species diversity, richness and evenness increased with woody plant density, most likely due to increased spatial heterogeneity and soil microclimate improvement. Bare ground cover increased significantly, whereas herbaceous biomass and soil organic matter did not respond to woody encroachment. In a subsequent investigation, we used a redundancy analysis to assess whether ecosystem properties were influenced by the identity of encroaching woody plant species. Species diversity and richness responded positively to Lannea triphylla, whereas leaf litter responded positively to Grewia tenax and G. villosa. Our findings suggest that woody plant encroachment in a semiarid rangeland does alter ecosystem properties. However, its impact is highly variable, influenced by a set of factors including the level of encroachment and identity of encroaching woody species.     

A continental-scale analysis of tree cover in African savannas

Aim We present a continental‐scale analysis that explores the processes controlling woody community structure in tropical savannas. We analyse how biotic and abiotic factors interact to promote and modify tree cover, examine alternative ecological hypotheses and quantify disturbance effects using satellite estimates of tree cover. Location African savannas. Methods Tree cover is represented as a resource‐driven potential cover related to rainfall and soil characteristics perturbed by natural and human factors such as fire, cattle grazing, human population and cultivation. Within this framework our approach combines semi‐empirical modelling and information theory to identify the best models. Results Woody community structure across African savannas is best represented by a sigmoidal response of tree cover to mean annual precipitation (MAP), with a dependency on soil texture, which is modified by the separate effects of fire, domestic livestock, human population density and cultivation intensity. This model explains c. 66% of the variance in tree cover and appears consistent across the savanna regions of Africa. Main conclusions The analysis provides a new understanding of the importance and interaction of environmental and disturbance factors that create the broad spatial patterns of tree cover observed in African savannas. Woody cover increases with rainfall, but is modified by disturbances. These ‘perturbation’ effects depend on MAP regimes: in arid savannas (MAP < 400 mm) they are generally small (< 1% decrease in cover), while in semi‐arid and mesic savannas (400–1600 mm), perturbations result in an average 2% (400 mm) to 23% (1600 mm) decrease in cover; fire frequency and human population have more influence than cattle, and cultivation appears, on average, to lead to small increases in woody cover. Wet savannas (1600–2200 mm) are controlled by perturbations that inhibit canopy closure and reduce tree cover by, on average, 24–34%. Full understanding of the processes determining savanna structure requires consideration of resource limitation and disturbance dynamics.     

Complex facilitation and competition in a temperate grassland: loss of plant diversity and elevated CO2 have divergent and opposite effects on oak establishment

Encroachment of woody vegetation into grasslands is a widespread phenomenon that alters plant community composition and ecosystem function. Woody encroachment is often the result of fire suppression, but it may also be related to changes in resource availability associated with global environmental change. We tested the relative strength of three important global change factors (CO₂ enrichment, nitrogen deposition, and loss of herbaceous plant diversity) on the first 3 years of bur oak (Quercus macrocarpa) seedling performance in a field experiment in central Minnesota, USA. We found that loss of plant diversity decreased initial oak survival but increased overall oak growth. Conversely, elevated CO₂ increased initial oak seedling survival and reduced overall growth, especially at low levels of diversity. Nitrogen deposition surprisingly had no net effect on survival or growth. The magnitude of these effects indicates that long-term woody encroachment trends may be most strongly associated with those few individuals that survive, but grow much larger in lower diversity patches. Further, while the CO₂ results and the species richness results appear to describe opposing trends, this is due only to the fact that the natural drivers are moving in opposite directions (decreasing species richness and increasing CO₂). Interestingly, the mechanisms that underlie both patterns are very similar, increased CO₂ and increased species richness both increase herbaceous biomass which (1) increases belowground competition for resources and (2) increases facilitation of early plant survival under a more diverse plant canopy; in other words, both competition and facilitation help determine community composition in these grasslands.     

Multi‐scale patterns and bush encroachment in an arid savanna with a shallow soil layer

Abstract. Question: Bush encroachment (i.e. an increase in density of woody plants often unpalatable to domestic livestock) is a serious problem in many savannas and threatens the livelihood of many pastoralists. Can we derive a better understanding of the factors causing bush encroachment by investigating the scale dependency of patterns and processes in savannas? Location: An arid savanna in the Khomas Hochland, Namibia. Methods: Patterns of bush, grass, and soil nutrient distribution were surveyed on several scales along a rainfall gradient, with emphasis on intraspecific interactions within the dominant woody species, Acacia reficiens. Results: Savannas can be interpreted as patch‐dynamic systems where landscapes are composed of many patches (a few ha in size) in different states of transition between grassy and woody dominance. Conclusions: In arid savannas, this patchiness is driven both by rainfall that is highly variable in space and time and by inter‐tree competition. Within the paradigm of patch‐dynamic savannas, bush encroachment is part of a cyclical succession between open savanna and woody dominance. The conversion from a patch of open savanna to a bush‐encroached area is initiated by the spatial and temporal overlap of several (localized) rainfall events sufficient for Acacia germination and establishment. With time, growth and self‐thinning will transform the bush‐encroached area into a mature Acacia stand and eventually into open savanna again. Patchiness is sustained due to the local rarity (and patchiness) of rainfall sufficient for germination of woody plants as well as by plant‐soil interactions.     

Influence of conspecific and heterospecific adults on riparian tree species establishment during encroachment of a humid palm savanna

Woody plant encroachment of savanna ecosystems has been related to altered disturbance regimes, mainly fire suppression and herbivore exclusion. In contrast, neighbourhood interactions among resident and colonising woody species have received little attention, despite their likely influence on the pattern and rate of tree establishment. We examined how resident palm trees (Butia yatay) and established adults of two riparian forest tree species (Allophylus edulis and Sebastiania commersoniana) influenced seed arrival and seedling performance of the latter two species in a humid savanna of east-central Argentina. Seed traps and seedlings of both riparian species were placed in herbaceous openings, and beneath palm, conspecific and heterospecific adult trees in two unburned savanna patches, and were monitored for 2 years. Only seeds of the bird-dispersed Allophylus arrived in palm microsites, yet survival of Allophylus seedlings near adult palms was limited by animal damage through trampling and burrowing, a non-trophic mechanism of apparent competition. Seeds of both riparian species dispersed into conspecific microsites, although adult trees selectively reduced growth of conspecific seedlings, a pattern consistent with the “escape hypothesis”. Further, survival of Sebastiania increased in the moister Allophylus microsites, suggesting a one-way facilitative interaction between woody colonisers. Our results indicate that dispersal facilitation by resident savanna trees may be critical to riparian species invasion after fire suppression. Distance-dependent effects of conspecific and heterospecific adult trees could contribute to shape the subsequent dynamics of woody seedling establishment. Overall, we show that indirect interactions can play a prominent role in savanna encroachment by non-resident woody species.     

Predicting the Effects of Woody Encroachment on Mammal Communities,Grazing Biomass and Fire Frequency in African Savannas

With grasslands and savannas covering 20% of the world’s land surface, accounting for 30–35% of worldwide Net Primary Productivity and supporting hundreds of millions of people, predicting changes in tree/grass systems is priority. Inappropriate land management and rising atmospheric CO₂ levels result in increased woody cover in savannas. Although woody encroachment occurs world-wide, Africa’s tourism and livestock grazing industries may be particularly vulnerable. Forecasts of responses of African wildlife and available grazing biomass to increases in woody cover are thus urgently needed. These predictions are hard to make due to non-linear responses and poorly understood feedback mechanisms between woody cover and other ecological responders, problems further amplified by the lack of long-term and large-scale datasets. We propose that a space-for-time analysis along an existing woody cover gradient overcomes some of these forecasting problems. Here we show, using an existing woody cover gradient (0–65%) across the Kruger National Park, South Africa, that increased woody cover is associated with (i) changed herbivore assemblage composition, (ii) reduced grass biomass, and (iii) reduced fire frequency. Furthermore, although increased woody cover is associated with reduced livestock production, we found indigenous herbivore biomass (excluding elephants) remains unchanged between 20–65% woody cover. This is due to a significant reorganization in the herbivore assemblage composition, mostly as a result of meso-grazers being substituted by browsers at increasing woody cover. Our results suggest that woody encroachment will have cascading consequences for Africa’s grazing systems, fire regimes and iconic wildlife. These effects will pose challenges and require adaptation of livelihoods and industries dependent on conditions currently prevailing.     

A resource ratio model of the effects of changes in CO<Subscript>2</Subscript> on woody plant invasion

It is expected that elevated CO₂ levels may have an important positive effect on the dominance of woody plants over grasses in savannas and grasslands. I propose that these changes in the relative abundance of trees and shrubs over grasses may be explained by Tilman’s resource ratio models. This change will occur because C₃ trees will have higher net photosynthetic rates than C₄ grasses which predominate in savannas. This will cause trees to have higher growth rates than grasses. An additional factor in trees and shrubs with carbon-based defences (such as tannins or other polyphenols) is that they may be better defended and, thus, lose less material to herbivory. Consequently, trees and shrubs should invade savannas and grasslands because their R* values will be lower. I compare this model to another less parsimonious model based on fire and carbon storage and allocation. Although these models are not necessarily mutually exclusive, the resource ratio model may be differentiated from the fire-carbon model on the basis of the presence of fire and/or the sensitivity to elevations in global CO₂ levels.     

A lepidopteran (<Emphasis Type="Italic">Imbrasia belina</Emphasis>) might influence tree-grass balance of <Emphasis Type="Italic">Colophospermum mopane</Emphasis> savanna

Traditional explanations of tree-grass coexistence in African savannas are based on competition between these growth forms or demographic bottlenecks of trees maintained by fire or mammalian browsers. Perturbation of their “balance” may result in an alternate system state of woody encroachment. Invertebrate herbivory has never been offered as an explanation. We developed a consumer-resource model which illustrated that annual irruptions of a lepidopteran (Imbrasia belina), known as mopane worm, can determine the tree-grass balance of semi-arid Colophospermum mopane savanna in southern Africa. Model performance was sensitive to the abundance, hence mortality, of mopane worms, owing to their complete defoliation of tree leaf biomass resulting in altered competitive relations between trees and grasses. Invertebrate herbivores have been recognized in other systems as agents for effecting a state change of host tree populations; this modeling study offers a first indication of such a role for the well-researched tree-grass relations of African savannas.     

Shifts in functional traits elevate risk of fire‐driven tree dieback in tropical savanna and forest biomes

Numerous predictions indicate rising CO₂ will accelerate the expansion of forests into savannas. Although encroaching forests can sequester carbon over the short term, increased fires and drought‐fire interactions could offset carbon gains, which may be amplified by the shift toward forest plant communities more susceptible to fire‐driven dieback. We quantify how bark thickness determines the ability of individual tree species to tolerate fire and subsequently determine the fire sensitivity of ecosystem carbon across 180 plots in savannas and forests throughout the 2.2‐million km² Cerrado region in Brazil. We find that not accounting for variation in bark thickness across tree species underestimated carbon losses in forests by ~50%, totaling 0.22 PgC across the Cerrado region. The lower bark thicknesses of plant species in forests decreased fire tolerance to such an extent that a third of carbon gains during forest encroachment may be at risk of dieback if burned. These results illustrate that consideration of trait‐based differences in fire tolerance is critical for determining the climate‐carbon‐fire feedback in tropical savanna and forest biomes.     

Vegetation dynamics in a Quercus‐Juniperus savanna: An isotopic assessment

Abstract. Woody plants are increasing in many grassland and savanna ecosystems around the world. As a case in point, the Edwards Plateau of Texas, USA, is a vast region (93 000 km²) in which rapid woody encroachment appears to be occurring. The native vegetation (prior to the Anglo‐European settlement 150–200 yr ago) and the biogeochemical consequences of woody encroachment in this region, however, are poorly understood. To assess these matters we measured plant and soil δ¹³C, soil organic C and soil N content from grasslands and two important woody patch types (mature Quercus virginiana clusters and Juniperus ashei woodlands) in this region. Soil δ¹³C values showed that relative productivity of C₃ species has increased in grassland and both woody habitats in recent times. δ¹³C of SOC in grasslands and Q. virginiana clusters increased with depth from the litter layer to 30 cm (grasslands =?21 to ?13‰Q. virginiana clusters =?27 to ?17‰) and were significantly different between habitats at all depths, indicating that Q. virginiana has been a long‐term component of the landscape. In J. ashei woodlands, soil δ¹³C values (at 20–30 cm depth) near the woodland edge (‐13‰) converged with those of an adjacent grassland (‐13‰) while those from the woodland interior (‐15‰) remained distinct, indicating that the woodland has been present for many years but has recently expanded. Concentrations and densities of SOC and total N were generally greater in woody patches than in grasslands. However, differences in the amount of SOC and N stored beneath the two woody patch types indicates that C and N sequestration potentials are species dependent.     

Exotic herbivores and fire energy drive standing herbaceous biomass but do not alter compositional patterns in a semiarid savanna ecosystem

Questions

Fire regime alterations are pushing open ecosystems worldwide past tipping points where alternative steady states characterized by woody dominance prevail. This reduces the frequency and intensity of surface fires, further limiting their effectiveness for controlling cover of woody plants. In addition, grazing pressure (exotic or native grazers) can reinforce woody encroachment by potentially reducing fine-fuel loads. We investigated the effects of different fire energies on the herbaceous plant community, together with mammalian wildlife herbivory (exotic and native combined) exclusion, to inform best management practices.

Location

Texas semi-arid savanna, southern Great Plains, USA.

Methods

We conducted an experiment in which we manipulated fire intensity and herbivore access to herbaceous biomass in a split-plot design. We altered fire energy via fuel addition rather than applying fire under different environmental conditions to control for differences in standing biomass and composition attributable to differential plant physiological status and fire season.

Results

High-energy fire did not reduce herbaceous biomass or alter plant community composition, although it did increase among-plot variability in composition and forb biomass relative to low-energy fire and non-burned controls. Grazing pressure from native and non-native mammalian herbivores reduced above-ground herbaceous biomass regardless of fire treatments, but did not alter community composition.

Conclusions

Managers seeking to apply high-intensity prescribed fire to reduce woody encroachment will not negatively impact herbaceous plant productivity or alter community composition. However, they should be cognizant that repeated fires necessary for greatly reducing woody plants in heavily invaded areas might be difficult to accomplish due to fine-fuel reduction from wild herbivores. High fencing to restrict access by wildlife herbivores or culling might be necessary to build fuels sufficient to conduct high-intensity burns for woody-plant reduction.     

Woody overstorey effects on soil carbon and nitrogen pools in South African savanna

Abstract Woody plant encroachment in savannas may alter carbon (C) and nitrogen (N) pools over the long‐term, which could have regional or global biogeochemical implications given the widespread encroachment observed in the vast savanna biome. Soil and litter %C and %N were surveyed across four soil types in two encroached, semi‐arid savanna landscapes in northern South Africa. Litter at sampling points with a woody component had a higher %N and lower C : N ratio than litter at solely herbaceous points. Severely encroached areas had lower C : N ratios throughout the soil profile than less encroached areas. Soil %C and %N were highly influenced by soil texture but were also influenced by the presence of a woody overstorey, which increased surface soil %C on three soil types but decreased it on the most heavily encroached soil type. Soil C sequestration may initially increase with bush encroachment but then decline if bush densities become so high as to inhibit understorey grass growth.