Rangeland Mismanagement in South Africa: Failure to Apply Ecological Knowledge

Chronic, heavy livestock grazing and concomitant fire suppression have caused the gradual replacement of palatable grass species by less palatable trees and woody shrubs in a rangeland degradation process termed bush encroachment in South Africa. G razing policymakers and cattle farmers alike have not appreciated the ecological role fire and native browsers play in preventing bush encroachment. Unpredictable droughts are common in South Africa but have deflected too much blame for bush encroachment away from grazing mismanagement. Bush encroachment is widespread on both black and white farms, although the contributing socioeconomic, cultural, and political forces differ. Managers at Madikwe Game Reserve have reintroduced fire and native game animals into a formerly overgrazed system in an attempt to remediate bush encroachment, with encouraging preliminary results. A bush control program is needed that educates cattle farmers about the ecological causes of bush encroachment and encourages the use of fire and native browsers as tools for sustainable grazing management.     

Defoliation depletes the carbohydrate reserves of resprouting <Emphasis Type="Italic">Acacia</Emphasis> saplings in an African savanna

Over the past century there has been a global trend towards tree expansion and densification in rangelands and savannas. This phenomenon is commonly referred to as bush encroachment. In South Africa Acacia karroo is one of the key species responsible for bush encroachment. It has been suggested that the combination of fire and browsing might limit bush encroachment by A. karroo more effectively than either browsing or fire alone. We hypothesized that these repeated disturbances progressively deplete root carbohydrates and compromise resprouting ability. This was tested by burning and then manually defoliating A. karroo once a month for 1 year. Manual defoliation did not inhibit the rapid shoot elongation after topkill of A. karroo saplings. During this initial phase, the growth of the new shoots of A. karroo was dependent more on mobilised root reserves than on photoassimilates from the new shoots. Frequent manual defoliation of resprouting A. karroo saplings prevented the replenishment of starch reserves. We suggest a mechanism for how the interaction of browsing and fire can suppress and perhaps reverse bush encroachment in African savannas. Saplings that have reduced starch reserves at the end of dry season due to browsing will struggle to resprout if they are burnt. Even if they do not die, they will be less able to escape fire damage in the next fire than if they had been able to resprout unimpeded.     

Effects of Prescribed Burning and Mechanical Bush Clearing on Ungulate Space Use in an African Savannah

Savannah ecosystems exhibit constant transitions between states dominated by trees and states dominated by a combination of trees and grasses. Transitions between these states are driven by interactions between fire and herbivory. Bush encroachment (i.e. an increase in the density of woody vegetation) is often caused by anthropogenic disturbance such as climate change, invasive plants, fire control, and livestock practices. As bush encroachment alters the dynamics between fire and herbivory, it may have significant impact on savannah ecosystems. Two of the most common measures to mitigate bush encroachment are prescribed burning and bush clearing by mechanical treatments. We studied the effects of these two mitigation measures on ungulate space use in Lapalala Wilderness, a private conservation area within the Waterberg Biosphere Reserve, northern South Africa. Burning and bush clearing affected both the overall abundance and the species composition of ungulates at particular patches, but these effects were influenced by habitat and the type of bush clearing treatment. Contrary to our expectations, most species occurred less frequently in burnt patches, and also less frequently in patches that had been bush cleared. Our results suggest that combined effects of fire and bush clearing may have positive effects on grazers and negative effects on browsers. Although our sampling design did not allow us to fully resolve interactive effects of burning and bush clearing treatments across habitats, our study highlights the fact that there are complex ecological consequences of habitat alterations in savannah ecosystems.     

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.     

Decadal changes in mean annual rainfall drive long‐term changes in bush‐encroached southern African savannas

Bush encroachment can have profound effects on the ability of savanna ecosystems to provide goods and services to society. It is therefore crucial to understand the key drivers of bush encroachment in savannas. In this study, we test whether decadal changes in mean annual rainfall significantly explain changes in the dominant patch size as well as the density of bush patches at six protected savanna sites located along a rainfall gradient in Zimbabwe. We first performed Maximal Overlap Discrete Wavelet transform within the intensity‐dominant scale theoretical framework on multi‐temporal aerial photographs and high spatial resolution satellite imagery to objectively detect changes in the dominant patch dimension as well as the intensity of bush cover over a 40‐year period at six test sites. We then pooled the data and performed regression analysis relating changes in dominant scale and intensity to decadal changes in mean annual rainfall in order to deduce a possible connection between dynamics of bush encroachment and rainfall variability. Our results indicate a significant nonlinear relationship between changes in the dominant scale and decadal changes in mean annual rainfall (R² = 0.85, F₁₃ = 35.96, P < 0.01). In contrast, the relationship between decadal changes in mean annual rainfall and changes in intensity was weak and not significant (R² = 0.29, F₁₃ = 2.69, P = 0.106). These results imply the importance of annual rainfall in explaining long‐term changes in the dominant scale of woody patches. However, mechanisms other than rainfall probably explain changes in the intensity of bush cover, and this needs further investigation.     

非洲稀树草原生态系统概况

介绍了非洲稀树草原的生态系统研究概况。涵盖了生理生态。生态系统评估,火,灌木入侵并丛生,植被动态。植被退化和恢复以及生态系统管理策略等内容。在此基础上提出了非洲稀树草原生态系统管理的研究趋势。     

Bird community composition in an actively managed savanna reserve, importance of vegetation structure and vegetation composition

The mosaic of trees, shrubs and open grassland in mesic African savannas is highly dynamic and strongly influenced by mammal herbivory and fire. We investigated the bird fauna in four different savanna habitats to help assess the impacts of vegetation change on this component of faunal diversity. Birds were censused, plant species were identified and vegetation structure was measured in four different vegetation types (Acacia nilotica woodland, Acacia nigrescens woodland, broadleaf thicket and open grassland) in the Hluhluwe-Umfolozi Park in northern KwaZulu Natal, South Africa. Multivariate ordination analyses were used to determine the relative importance of vegetation structure and floristic composition in defining bird assemblages. The bird communities of the grasslands, the acacia woodlands, and the broadleaf woodlands were clearly separated on the first axis of the detrended canonical correspondence analysis (DCCA). Canopy cover and foliage height diversity (FHD) were strongly correlated with the first axis of DCCA, possibly reflecting a secondary successional series from grassland to woodland, known as bush encroachment. Floristic composition (based on presence–absence data only) seemed to be less important for bird community composition than vegetation structure. The results indicate that changes in vegetation structure, caused by bush encroachment, could cause concomitant changes in bird community composition.     

Seventy years of continuous encroachment substantially increases ‘blue carbon’ capacity as mangroves replace intertidal salt marshes

Shifts in ecosystem structure have been observed over recent decades as woody plants encroach upon grasslands and wetlands globally. The migration of mangrove forests into salt marsh ecosystems is one such shift which could have important implications for global ‘blue carbon’ stocks. To date, attempts to quantify changes in ecosystem function are essentially constrained to climate‐mediated pulses (30 years or less) of encroachment occurring at the thermal limits of mangroves. In this study, we track the continuous, lateral encroachment of mangroves into two south‐eastern Australian salt marshes over a period of 70 years and quantify corresponding changes in biomass and belowground C stores. Substantial increases in biomass and belowground C stores have resulted as mangroves replaced salt marsh at both marine and estuarine sites. After 30 years, aboveground biomass was significantly higher than salt marsh, with biomass continuing to increase with mangrove age. Biomass increased at the mesohaline river site by 130 ± 18 Mg biomass km^?2 yr^?1 (mean ± SE), a 2.5 times higher rate than the marine embayment site (52 ± 10 Mg biomass km^?2 yr^?1), suggesting local constraints on biomass production. At both sites, and across all vegetation categories, belowground C considerably outweighed aboveground biomass stocks, with belowground C stocks increasing at up to 230 ± 62 Mg C km^?2 yr^?1 (± SE) as mangrove forests developed. Over the past 70 years, we estimate mangrove encroachment may have already enhanced intertidal biomass by up to 283 097 Mg and belowground C stocks by over 500 000 Mg in the state of New South Wales alone. Under changing climatic conditions and rising sea levels, global blue carbon storage may be enhanced as mangrove encroachment becomes more widespread, thereby countering global warming.     

Bush encroachment under three contrasting land-use practices in a mesic South African savanna

This study determined the effects of land-use practice had on the rate and extent of bush encroachment in a mesic savanna in KwaZulu-Natal, South Africa. Changes in woody cover were measured for 1 km² sites in areas under communal, commercial and conservation land-use systems for the period between 1937 and 2000. Land users from each area were interviewed to gain the histories of each area and to determine how the changes in woody cover had impacted them and whether anything was being done to counteract the spread of trees and shrubs on their land. Bush encroachment occurred across all three of the land-use types in the 67-year period between 1937 and 2000. The results showed that land-use practice had enormous impacts on the process of bush encroachment. The communal site showed a decrease in grass (21%) and tree (5%) cover and an increase in shrub cover (13%). At the commercial site, there was a considerable decrease in grass cover (46%) and moderate increase in shrub cover (10%) and a massive increase in tree cover (36%). The area under conservation showed a substantial decrease in grass cover (47%), a slight decrease in shrub cover (19%) and a massive increase in tree cover (66%). The perceived causes of these changes were fairly similar amongst the different land users. The changes were mostly not perceived to be a problem for the communal land users. The main advantages mentioned were increased woody resources for building and firewood and increased browse availability. The commercial and conservation land users perceived the changes to have significant negative connotations including the loss of grazing land and biodiversity and secondary invasion of encroached areas by alien plant species. Despite these perceptions, very little has been done to combat bush encroachment in the commercial and conservation land use systems.     

Savanna tree–grass competition is modified by substrate type and herbivory

Question: Woody plant and grass interactions in savannas have frequently been studied from the perspective of the response of one growth form on the other but seldom evaluated as two‐way interactions. What causes woody plant encroachment in semi‐arid savannas and what are the competitive responses of tree seedlings and grasses on rocky and sandy substrates? Methods: In this greenhouse study, we investigated the influence of substrate and grazing on responses to interspecific competition by tree seedlings and grasses. We measured competitive/facilitative responses on biomass and nutrient status of tree seedlings and grasses grown together. Results: Interspecific competition suppressed growth of trees and grasses. Tree seedlings and uncut grass accumulated double the biomass when grown without competition relative to when they competed. Competitive responses varied on different substrates. Grass biomass on rocky substrate showed no response to tree competition, but appeared to be facilitated by trees on sandy substrate. Grass clipping resulted in higher tree seedling biomass on rocky substrate, but not on sandy substrate. There was a positive response of grass nutrient status to competition from tree seedlings. Conclusion: Selective grass herbivory in the absence of browsing or suppression of shade‐intolerant grasses by trees are commonly cited reasons behind bush encroachment in savannas. We show that grazing may confer a competitive advantage to tree seedlings and promote bush encroachment more readily on rocky substrates. This may be due to the imposed sharing of the soil depth niche on rocky substrates, whereas possible niche separation on sandy substrates minimizes the advantage conferred by reduced competition.     

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.     

Footnote on Breeding of Paradise Flycatcher

Bird community diversity was assessed in semi-arid savanna in the Northern Cape province of South Africa and related to vegetation diversity and foliage height diversity. As this is an area in which shrub encroachment is common, the effects of shrub or bush encroachment on bird species diversity were specifically addressed. An experiment in which the dominant encroaching species, Acacia mellifera, was removed from ten 1 ha plots was used and compared with ten controls. A strong positive association with foliage height diversity was shown. No effect of experimental removal of the dominant encroacher, A. mellifera, on bird species diversity or species richness was found. However, four species increased in relative abundance in cut plots and two species decreased in relative abundance in these plots.     

灌丛化对黄土高原草地植物群落结构和地上生物量的影响

中国北方草地普遍出现灌丛化现象,灌丛化改变植物群落结构、植物多样性和生产力,直接影响着草地生态保护与可持续利用.该研究以黄土高原灌丛化草地为研究对象,通过植被调查,分析比较不同坡向的灌丛斑块与禾草斑块植物群落结构(物种组成、优势种及物种多样性)和地上生物量的差异.结果发现:(1)灌丛化草地不同坡向对物种多样性及地上生物...     

<Emphasis Type="Italic">Molinia caerulea</Emphasis> responses to N and P fertilisation in a dry heathland ecosystem (NW-Germany)

In the present study we analysed whether airborne N pollution may constitute one important driver for the encroachment of Molinia caerulea in dry heathland ecosystems. Based on full-factorial field experiments (in 2006 and 2008) and complementary greenhouse experiments (in 2008), we quantified growth responses of Molinia caerulea to N and P fertilisation (50 kg N ha⁻¹ year⁻¹, 20 kg P ha⁻¹ year⁻¹). Aboveground biomass production of Molinia caerulea was limited by P in 2006, but by N in both experiments in 2008. In the greenhouse experiment, N addition caused a sixfold increase of the biomass of vegetative tillers, and in all experiments the biomass and numbers of flowering tillers showed a significant increase due to fertilisation. Our experiments indicated that growth of Molinia caerulea was primarily limited by N, but in dry heaths the kind of nutrient limitation may be mediated by other factors such as water availability during the vegetative period. Shifts in biomass allocation patterns resulting from N fertilisation showed that Molinia caerulea encroachment in dry heaths is not only attributable to increased leaf biomass, but also due to higher investments in reproductive tissue that allow for increased seed production and thus accelerated encroachment of seedlings in places where the dwarf shrub canopy has been opened after disturbance.     

Spatial distribution of <Emphasis Type="Italic">Vachellia karroo</Emphasis> in Zimbabwean savannas (southern Africa) under a changing climate

Climate change projections in southern Africa show a drier and a warmer future climate. It is not yet clear how these changes are going to affect the suitable habitat of bush encroacher woody species in southern African savannas. Maximum Entropy niche modelling technique was used to test the extent to which climate change is likely to affect the suitable habitat of Vachellia karroo in Zimbabwe based on six Global Climate Models (GCMs) from Coupled Model Intercomparison Project Phase 5 (CMIP5) and two Representative Concentration Pathways (RCPs) for the 2070s. An overlay analysis was then performed in a Geographic Information System based on the current and future bioclimatically suitable areas for the respective GCMs and RCPs. This was done to determine the potential effect of climate change on the focal species. Results show that temperature related variables are more important in explaining the spatial distribution of V. karroo than precipitation related variables. In addition, results indicate an overall increase in the modelled suitable habitat for V. karroo by the 2070s across the GCMs and RCPs considered in this study. Specifically, the suitable habitat of V. Karroo is projected to increase by a maximum of 57,594 km² signifying a 69% increase from the current suitable habitat (83,674 km²). The suitable areas are projected to increase in eastern, western and south eastern parts of Zimbabwe. These results imply that improved understanding of the response of woody species to a changing climate is important for managing bush encroachment in savanna ecosystems.     

Rangeland degradation is poised to cause Africa's first recorded avian extinction

Rangeland degradation by livestock threatens several restricted-range species, but is largely overlooked by conservation biologists. The Sidamo lark Heteromirafra sidamoensis , confined to the Liben Plain grassland in southern Ethiopia, is critically endangered by bush encroachment, permanent settlement and agricultural conversion. Its global range was previously estimated at 760 km², but in 2007–2008 available habitat covered<35 km². Density estimates from multi-model inference analysis of distance transect data provided a global population estimate of 90–256 adults (possibly with a serious sex-ratio bias towards males). Logistic regression models of habitat selection showed that males preferentially occurred in areas of grassland with greater cover of medium-length grass (5–15 cm), less cover of bare ground and fewer bushes. Habitat transects extending outward from its core range revealed massive and rapid bush encroachment, corroborating information from semi-structured interviews. The survival of both local Borana pastoralism and this species – mainland Africa's likeliest first avian extinction – depends on restoring seasonal patterns of grazing, resisting agricultural conversion of grasslands, reversing fire suppression policies and clearing bush.     

Winners and losers: tropical forest tree seedling survival across a West African forest–savanna transition

Forest encroachment into savanna is occurring at an unprecedented rate across tropical Africa, leading to a loss of valuable savanna habitat. One of the first stages of forest encroachment is the establishment of tree seedlings at the forest–savanna transition. This study examines the demographic bottleneck in the seedlings of five species of tropical forest pioneer trees in a forest–savanna transition zone in West Africa. Five species of tropical pioneer forest tree seedlings were planted in savanna, mixed/transition, and forest vegetation types and grown for 12 months, during which time fire occurred in the area. We examined seedling survival rates, height, and stem diameter before and after fire; and seedling biomass and starch allocation patterns after fire. Seedling survival rates were significantly affected by fire, drought, and vegetation type. Seedlings that preferentially allocated more resources to increasing root and leaf starch (starch storage helps recovery from fire) survived better in savanna environments (frequently burnt), while seedlings that allocated more resources to growth and resource‐capture traits (height, the number of leaves, stem diameter, specific leaf area, specific root length, root‐to‐shoot ratio) survived better in mixed/transition and forest environments. Larger (taller with a greater stem diameter) seedlings survived burning better than smaller seedlings. However, larger seedlings survived better than smaller ones even in the absence of fire. Bombax buonopozense was the forest species that survived best in the savanna environment, likely as a result of increased access to light allowing greater investment in belowground starch storage capacity and therefore a greater ability to cope with fire. Synthesis: Forest pioneer tree species survived best through fire and drought in the savanna compared to the other two vegetation types. This was likely a result of the open‐canopied savanna providing greater access to light, thereby releasing seedlings from light limitation and enabling them to make and store more starch. Fire can be used as a management tool for controlling forest encroachment into savanna as it significantly affects seedling survival. However, if rainfall increases as a result of global change factors, encroachment may be more difficult to control as seedling survival ostensibly increases when the pressure of drought is lifted. We propose B. buonopozense as an indicator species for forest encroachment into savanna in West African forest–savanna transitions.     

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.     

The impact of black wattle encroachment of indigenous grasslands on soil carbon,Eastern Cape,South Africa

Black wattle (Acacia mearnsii, De Wild.) is a fast growing tree species introduced into South Africa in the nineteenth century for commercial purposes. While being an important source of timber and firewood for local communities, black wattle is an aggressive invasive species and has pervasive adverse environmental impacts in South Africa. Little is known about the effects of black wattle encroachment on soil carbon, therefore the aim of this study was to investigate the impact of black wattle encroachment of natural grassland on soil carbon stocks and dynamics. Focussing on two sites in the Eastern Cape, South Africa, the study analysed carbon stocks in soil and litter on a chronosequence of black wattle stands of varying ages (up to >50 years) and compared these with adjacent native grassland. The study found that woody encroachment of grassland at one site had an insignificant effect on soil and litter carbon stocks. The second site showed a clear decline in combined soil and litter carbon stocks following wattle encroachment. The lowest stock was in the oldest wattle stand, meaning that carbon stocks are still declining after 50 years of encroachment. The results from the two sites demonstrate the importance of considering changes in soil carbon when evaluating ecosystem effects of invasive 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.