Spatial patterns and dynamics of woody vegetation in an arid savanna

The spatial distribution of woody plants was studied in an arid savanna in Botswana. The study included stands of mixed species and sizes as well as monospecific even-sized stands of different size classes of the tree Acacia erioloba and the shrub Acacia mellifera. In the case of A. mellifera both dense stands on overgrazed land and more open stands were included. The analysis used all plant-to-plant distances, and individuals were represented with a realistic canopy extension. The mixed stands showed aggregated distribution of individuals, mainly caused by strong clumping of small shrubs. In A. erioloba saplings were aggregated, small trees were randomly or regularly distributed and large trees were randomly spaced. In open stands of A. mellifera aggregation increased with size of the shrubs, while in dense stands with overgrazing aggregation decreased with increasing size. The different patterns are discussed in relation to the relative importance of inter- and intraspecific competition for water and of disturbance by fire as regulatory mechanisms for total amount and spatial distribution of woody plants in this savanna.     

Termite‐induced heterogeneity in African savanna vegetation: mechanisms and patterns

Objectives: To (1) assess the strength of evidence for the role of termites in vegetation heterogeneity in African savannas, and (2) identify the mechanisms by which termites induce such heterogeneity. Location: African savannas. Methods: We conducted a review of the literature, a meta‐analysis and qualitative systems analysis to identify mechanisms to explain the observed patterns. Results: The review provided evidence for termite‐induced heterogeneity in floristic composition and vegetation patterning in savannas across Africa. Termites induced vegetation heterogeneity directly or indirectly through their nest‐building and foraging activities, associated nutrient cycling and their interaction with mammalian herbivores and fire. The literature reviewed indicated that termite mounds essentially act as islands of fertility, which are responsible for ecosystem‐level spatial heterogeneity in savannas. This was supported by the meta‐analysis, which demonstrated that mounds of Ancistrotermes, Macrotermes, Odontotermes (family Macrotermitinae), Cubitermes (family Termitinae) and Trinervitermes (Nasutitermitinae) are significantly enriched in clay (75%), carbon (16%), total nitrogen (42%), calcium (232%), potassium (306%) and magnesium (154%) compared to the surrounding savanna soil. Conclusions: Termite activity is one of the major factors that induce vegetation patterning in African savannas. The implications of this are discussed and research questions for future studies and modelling efforts are indicated.     

Soil nutrients distribution and moisture dynamics on upper catena in a semi-arid nature reserve

The distribution of soil nutrients and soil moisture dynamics on slopes of catenary landscape, were recorded in a semi-arid nature reserve located in the Eastern Transvaal, South Africa. Soil nutrients were accumulating in the slopes of the catena whereas the crests were highly leached. Open grassland bands segregating between the two soil types and their associated plant species, did not show clear stratification of nutrient deposition amongst ground levels down to 1.5 m below the surface. There was however, a tendency of soils with high nutrient contents to correlate with areas of high woody vegetation density. Patterns of soil moisture levels during two annual rainfall periods showed some segregation between soil depths, although the trends indicated that the duration of saturation and dry out of soils levels differed more between rainy seasons than between soil levels during a particular season.     

Structure and biomass along an Acacia zanzibarica woodland–savanna gradient in a former ranching area in coastal Tanzania

Questions: What factors influence the density, size and growth form of trees in secondary Acacia zanzibarica woodlands on a former humid savanna rangeland? How does tree density relate to variation in tree foliage and spines, and woody and grass biomass? Location: Tropical coastal Tanzania (former Mkwaja Ranch, now in Saadani National Park). Methods: We surveyed 97 circular plots (4‐m radius) representing a gradient from open savanna to dense woodland. Within each plot, we measured all trees and estimated the biomass of spines. Foliage biomass of tree and grass layers was estimated on three occasions, twice during the wet season and once in the dry season. Soil samples were taken from each plot and analysed for texture and nutrient content. Interrelationships among various variables were investigated using linear multiple regression and mixed effects models. Results: Tree densities were highest on more nutrient‐rich, heavy soils. Spinescence was highest on trees in open savanna. Biomass of tree foliage in the wet season was best explained by numbers of ant nests and tree live‐wood ratio. Foliage biomass in the dry season was less than half that in the wet season and best predicted by grass biomass. Variables related to biomass of the grass layer were strongly influenced by fire; living grass biomass also decreased with increasing tree density. Conclusions: A. zanzibarica is a tree with a high water demand, and the association with heavy soils is probably due to greater availability of water on these sites. Establishment of A. zanzibarica woodlands significantly reduced grazing resources at Mkwaja Ranch. Under post‐ranching conditions, however, fires and soil conditions predominate. The woodlands may, therefore, represent a transient state of woody density in a still resilient humid savanna.     

Successional patterns of woody plants in catchment areas in a semi-arid region

Patterns of woody plants dispersal in a semi-arid nature reserve situated in Eastern Transvaal, South Africa, revealed that trees have spread from core areas and converted previously open grasslands to densely vegetated woodlands. These patterns were found in catchment areas of the gently undulating terrain which characterizes the region.Two plant communities dominated by Acacia senegal-Acacia tortilis and Euclea divinorum-Acacia nilotica were distinguished. Analysis of nearest-neighbour distances, dispersal patterns of seedlings and mature woody plants identified successional processes. These were manifested through intra and interspecific competition among the dominant tree species. Within each plant community, a transition of relative abundance was occurring namely, A. senegal became dominant in areas previously dominated by A. tortilis while E. divinorum was replacing previous A. nilotica dominance.Comparative assessment of the two plant communities was facilitated by the summary of competition and seedling dispersal indices in a multivariate analysis. Results indicated that tree species had characteristic dispersal strategies. Identification the patterns of woody plants establishment could advance the evaluation of successional dynamics and management of savannas in areas prone to bush encroachment.     

Interacting effects of grass height and herbivores on the establishment of an encroaching savanna shrub

Shrub encroachment is a widely observed problem in Southern African savannas. Although the effects of herbivory and grass height on woody species recruitment have been studied individually, little information exists about how these factors interact. In this study seeds and seedlings of the encroaching shrub Dichrostachys cinerea were planted in clipped and unclipped grass plots, with and without large herbivores present. Seed germination, seedling survival and seedling predation were monitored for 8 months. Germination started earlier in plots where herbivores were excluded. Overall, the earlier the seeds germinated, the longer the seedlings survived. Clipping positively affected the number of germinated seeds, seedling growth and survival but effects varied among herbivore exclusion treatments and sites. Invertebrates caused the majority of the seedling damage. We conclude the recruitment of D. cinerea is influenced by the interplay of grass height and herbivory. In this study, the presence of large herbivores early in the wet season, and the absence of simulated grazing later on, affected the regeneration of D. cinerea negatively. However, differences in effects among sites suggest that the mechanisms found here may work differently in other habitats.      

Phenology of nine common woody species in semi-arid,deciduous Kalahari Sand vegetation

The phenological patterns of 9 woody species that are common components of Kalahari Sand communities were observed over a period of 32 months. The observations were made on plants adjacent to the soil moisture study sites which have been described earlier (Childes & Walker 1987). A pre-rain flush was noted in the 2 shrub/small tree species Ochna pulchra and Terminalia sericea. Leaf fall in all species was related to a decrease in soil moisture and minimum temperature. The rooting habits and other adaptations of the species resulted in a sequential leaf fall in the communities. Open woodland and scrub species such as Pterocarpus angolensis showed a generally synchronised pre-rain flowering, while the closed woodland species e.g. Baikiaea plurijuga had protracted asynchronous flowering periods. Fruit ripening generally occurred in the dry season when conditions were ideal for fruit and seed dispersal.     

Patch and soil characteristics of pastures with water‐spreading banks and woody encroachment in semi‐arid Australia

Water‐spreading banks are used in semi‐arid areas such as the Cobar pediplain in western New South Wales, Australia to encourage pasture growth, often after removal of woody encroachment. We studied the arrangement of bare inter‐patches and vegetated patches, and associated surface soil variables, in three pastures following installation of water‐spreading banks (2, 15, 38 years ago) and in an area of woody encroachment near Cobar. The aims of the study were as follows: (i) to determine the number and percent area of inter‐patches and vegetated patches, and associated surface soil variables at the three pasture sites and at the woody encroachment site and (ii) by inference, explore effects of establishing water‐spreading banks and pasture following removal of woody encroachment on these factors, to understand the role of water‐spreading banks as a management tool. The percent area of inter‐patches in pasture with 38‐year‐old water‐spreading banks was much lower, and the percent area of medium‐vegetated patches (but not of well‐vegetated patches) was substantially higher, than in the woody encroachment. Differences in soil carbon and nitrogen between the sites were related to their percent areas of inter‐patches and vegetated patches. The results suggest that the mosaic of bare inter‐patches and vegetated patches changes over time after clearing of woody encroachment and establishment of pasture with water‐spreading banks, from many large inter‐patches to a few small inter‐patches, and from small to large medium‐vegetated patches. Water‐spreading banks are a useful management tool in these landscapes because of their benefits for landscape function, that is, bare areas become less connected, the percent area of moderately vegetated patches increases, and soil carbon builds up with time following their installation.     

Spatial partitioning of the soil water resource between grass and shrub components in a West African humid savanna

Most savanna water balance models assume water partitioning between grasses and shrubs in a two-layer hypothesis, but this hypothesis has not been tested for humid savanna environments. Spatial partitioning of soil water between grasses and shrubs was investigated in a West African humid savanna by comparing the isotopic composition (oxygen-18 and deuterium) of soil water and plant stem water during rainy and dry conditions. Both grass and shrub species acquire most of their water from the top soil layer during both rainy and dry periods. A shift of water uptake pattern towards deeper horizons was observed only at the end of the dry season after shrub defoliation. The mean depth of water uptake, as determined by the isotopic signature of stem water, was consistent with grass and shrub root profiles and with changes in soil water content profiles as surveyed by a neutron probe. This provides evidence for potentially strong competition between shrubs and grasses for soil water in these humid savannas. Limited nutrient availability may explain these competitive interactions. These results enhance our understanding of shrub-grass interactions, and will contribute to models of ecosystem functioning in humid savannas.     

Net changes in regional woody vegetation cover and carbon storage in Texas Drylands, 1937–1999

Although local increases in woody plant cover have been documented in arid and semiarid ecosystems worldwide, there have been few long‐term, large‐scale analyses of changes in woody plant cover and aboveground carbon (C) stocks. We used historical aerial photography, contemporary Landsat satellite data, field observations, and image analysis techniques to assess spatially specific changes in woody vegetation cover and aboveground C stocks between 1937 and 1999 in a 400‐km² region of northern Texas, USA. Changes in land cover were then related to topo‐edaphic setting and historical land‐use practices. Mechanical or chemical brush management occurred over much of the region in the 1940–1950s. Rangelands not targeted for brush management experienced woody cover increases of up to 500% in 63 years. Areas managed with herbicides, mechanical treatments or fire exhibited a wide range of woody cover changes relative to 1937 (?75% to + 280%), depending on soil type and time since last management action. At the integrated regional scale, there was a net 30% increase in woody plant cover over the 63‐year period. Regional increases were greatest in riparian corridors (33%) and shallow clay uplands (26%) and least on upland clay loams (15%). Allometric relationships between canopy cover and aboveground biomass were used to estimate net aboveground C storage changes in upland (nonriparian) portions of regional landscapes. Carbon stocks increased from 380 g C m^?2 in 1937 to 500 g C m^?2 in 1999, a 32% net increase across the 400 km² region over the 63‐year period. These plant C storage change estimates are highly conservative in that they did not include the substantial increases in woody plant cover observed within riparian landscape elements. Results are discussed in terms of implications for ‘carbon accounting’ and the global C cycle.     

A preliminary report on the vegetation zonation of palsas in the Arctic National Wildlife Refuge,northern Alaska,USA

We measured vegetation patterns on palsas with reference to topographic characteristics on the Arctic National Wildlife Refuge, northern Alaska, to obtain benchmark data because of the changes expected from global warming. Vegetation was examined in 60 plots of area 50 cm × 50 cm by five environmental factors: water content in the peat and duff layers, groundwater level, slope angle, depth to frozen surface, and presence of pellets and feces. Three palsas were selected for the survey, and the heights were fewer than 50 cm from the groundwater surface. Based on TWINSPAN and canonical correspondence analysis, we confirmed that clear patterns of vegetation zonation had developed within a 60-cm difference in water level. Vaccinium vitis-idaea occurred well on the top areas of palsas, while Carex aquatilis was established on the bottom areas. Sphagnum spp. were established on intermediate locations between V. vitis-idaea and C. aquatilis. The prime determinant of the vegetation zonation seems to be water content in peat and duff layers rather than water level, although the five factors that we examined interact intricately with each other.     

Assessment of species composition change in savanna vegetation by means of woody plants' size class distributions and local information

A new method is presented for assessment of compositional changes in disturbed savanna systems and for identification of species susceptible to local extinction. Such information is urgently needed as a basis for management plans that aim at ensuring conservation and sustainable use of savanna vegetation. Size class distributions (SCDs) of trees and shrubs from a fire-disturbed savanna in Senegal are analysed and compared to observations provided by local people on declining species. Many species, particularly large trees, have few young individuals and therefore flat SCDs that deviate from the reverse-J shape SCD characterizing species with abundant and constant rejuvenation. Based on comparisons of SCDs with local information and historical records, it is concluded that most of the flat distributions analysed are effects of declining population sizes caused by human impact, particularly fire. Results from the analyses certify that a change in species composition from a vegetation characterized by large trees to one dominated by shrubs takes place. Whereas most of the declining species are locally appreciated, this is not the case for the few species with good rejuvenation. Therefore, if the observed trend continues, it will have severe implications for the local communities.     

Zonation of submerged macrophyte vegetation in Lake Kariba,Zimbabwe and its ecological interpretation

The submerged vegetation of Lake Kariba is described in relation to degree of slope (lake morphometry), depth and light transparency. The direct gradient analysis technique — canonical correspondence analysis and the TWINSPAN classification programs were used to analyse the data set. The western end of the lake with low transparency has a low species diversity (with Vallisneria aethiopica dominating). Species diversity increases with increased transparency in the other parts of the lake. The classification revealed monospecific communities for all species as well as mixed communities with Lagarosiphon as the associate species with the broadest distribution. The ordination revealed a first axis strongly related to depth and transparency gradients and the second axis related to slope. Vallisneria aethiopica has a growth form adapted to grow in shallow areas subjected to wave action. Potamogeton octandrus also growing in shallow areas is restricted to sheltered areas. Lagarosiphon ilicifolius precluded from the shallow exposed areas grows at medium depth but has a growth form adapted to tap light at the surface. Najas pectinata and Ceratophyllum demersum are adapted to grow in deep water.Abbreviations CCA Canonical correspondence analysis     

Form and function of grass ring patterns in arid grasslands: the role of abiotic controls

Ring-shaped growth patterns commonly occur in resource-limited arid and semi-arid environments. The spatial distribution, geometry, and scale of vegetation growth patterns result from interactions between biotic and abiotic processes, and, in turn, affect the spatial patterns of soil moisture, sediment transport, and nutrient dynamics in aridland ecosystems. Even though grass ring patterns are observed worldwide, a comprehensive understanding of the biotic and abiotic processes that lead to the formation, growth and breakup of these rings is still lacking. Our studies on patterns of infiltration and soil properties of blue grama (Bouteloua gracilis) grass rings in the northern Chihuahuan desert indicate that ring patterns result from the interaction between clonal growth mechanisms and abiotic factors such as hydrological and aeolian processes. These processes result in a negative feedback between sediment deposition and vegetation growth inside the bunch grass, which leads to grass die back at the center of the grass clump. We summarize these interactions in a simple theoretical and conceptual model that integrates key biotic and abiotic processes in ring formation, growth and decline.     

Nearest-neighbour analysis and the prevelance of woody plant competition in South African savannas

Inter- and intra-specific competition has long been regarded as asignificant determinant of the structure and function of woody plantcommunitiesin African savannas. The most widely used approach to detect the presence orabsence of competition has been the use of nearest-neighbour methods. Althoughseveral studies have been published reporting the presence of competition inAcacia dominated communities, less than 20 plots have beensampled across all these studies. Results from broad-leaved communities arevariable, and also based on a small sample number. Consequently, this studysought to assess the prevalence of competition from a large number of savannasites (45), and to identify abiotic and biotic factors characterising siteswithcompetition relative to those without. Using the nearest-neighbour method onlyfour sites (9.3%) indicated the presence of inter-specific competition; twowereAcacia dominated communities (18.2% ofAcacia sites) and two were broad-leaved communities(7.7%).These four sites had a significantly higher mean annual rainfall and fewerwoodyspecies than sites without inter-specific competition. There were alsosignificant relationships between the inter-specific competition index and thedominance index and the number of species at a site. In terms of intra-specificcompetition only ten of the 31 sites tested revealed a significant correlationbetween nearest-neighbour distance and summed canopy volume of the twoneighbours. Seven were for Acacia species (55.6% of theAcacia species comparisons) and three were forbroad-leavedspecies (21.4% of the broad-leaved species comparisons). Sites lacking evidenceof intra-specific competition were at a significantly lower slope position andhad a smaller proportion of small stems. The lower prevalence of competition inSouth African savannas detected in this study using the same method as previousstudies is discussed, and the appropriateness of nearest-neighbour analysis isexamined.     

Changes in soil properties and vegetation following livestock grazing exclusion in degraded arid environments of South Tunisia

In this study, characteristics of vegetation and soil properties under continued grazing and exclusion of livestock for 6 and 12 years were examined in a degraded Stipa tenacissima steppe in South Tunisia.     

Natural 15N abundance of vegetation and soil in the Kapalga savanna,Australia

Abstract The natural abundance of the stable isotope ¹⁵N was measured in different vegetation components and in the soil of a northern Australian savanna. Most of the vegetation was found to be ¹⁵N-depleted compared to atmospheric N₂. Herbaceous legumes, perennial grasses, tree legumes, non-legume trees and annual grasses exhibited mean δ¹⁵N of ? 1.7, ? 0.8, ? 0.7, 0.0 and + 0.3‰, respectively. These results are in good agreement with previous studies. Legumes exhibit slightly negative values, indicating that they are likely to be nitrogen-fixing plants. Non-legume plants have a δ¹⁵N close to zero, which could equally result from non-symbiotic fixation, soil organic matter mineralization, or fresh root litter mineralization. In contrast, soil organic matter was ¹⁵N-enriched. Values of δ¹⁵N increased with depth and were + 2.5, + 5.2 and +6.1‰ in the 0–10, 10–20 and 20–40cm layers, respectively. Soil organic matter δ¹⁵N shows a typical profile of mature soils.     

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.