Long-term vegetation dynamics mediated by herbivores,weather and fire in a Juniperus-Quercus savanna

Abstract. Long-term (45-yr) basal area dynamics of dominant graminoid species were analyzed across three grazing intensity treatments (heavily grazed, moderately grazed and ungrazed) at the Texas A&M University Agricultural Research Station on the Edwards Plateau, Texas. Grazing intensity was identified as the primary influence on long-term variations in species composition. Periodic weather events, including a severe drought (1951–1956), had little direct influence on composition dynamics. However, the drought interacted with grazing intensity in the heavily grazed treatment to exacerbate directional changes caused by grazing intensity. Species response to grazing was individualistic and noisy. Three response groups were identified. Taller, more productive mid-grasses were most abundant under moderate or no grazing. Short grasses were most abundant under heavy grazing. Intermediate species were most abundant under moderate grazing and opportunistic to weather patterns. Graminoid diversity increased with the removal or reduction of grazing intensity. The moderately and ungrazed treatments appeared most resistant to short-term weather fluctuations, while the heavily grazed treatment demonstrated significant resilience when grazing intensity was reduced after over 110 yr of overgrazing. Identification of a ‘climax’ state is difficult. Significant directional change, which took nearly 20 yr, appears to continue in the ungrazed treatment after 45 yr of succession. The observed, relatively linear patterns of perennial grass composition within the herbaceous patches of this savanna were generally explained by traditional Clementsian succession. However, when dynamics of the herbaceous community are combined with the woody component of this savanna, the frequency and intensity of fire becomes more important. Across the landscape, successional changes follow several pathways. When vegetation change is influenced by several factors, a multi-scale model is necessary to demonstrate interactions and feedbacks and accurately describe successional patterns. Absence of fires, with or without grazing, leads ultimately to a Juniperus/Quercus woodland with grazing intensity primarily influencing the fuel load and hence fire intensity.     

The influence of abiotic and spatial variables on woody and herbaceous species abundances in a woodland–grassland system in the Eastern Terai of India

印度特莱东部林地-草地系统中非生物和空间变量对木本和草本物种丰度的影响 目前尚不清楚哪些环境因素决定了热带稀树草原特别是在潮湿地带的林地和草地镶嵌处的林地和草地的物种多度。基于此,本研究探究了非生物和空间变量对印度东北部的台拉河生态系统木本和草本物种分布的影响,评估了气候和非气候因素在整个景观中保持可变的树草比和空间连通性和分散性的相对重要性。在519 km²的受保护的特莱栖息地中随机建立了134个30 m × 30 m的抽样样方,并调查了每个样方的木本和草本植物的物种多度和气候,以及非气候环境因素。基于不同的地点空间连通性模型,通过构建变量检验气候和非气候环境因素对物种多度的影响。使用冗余分析和方差分解定量解析环境变量和空间结构对林地和草地物种多度的相对重要性。研究结果表明,降雨、火灾、水分胁迫、地形和土壤养分在内的环境变量对物种多度和林草比有显著的影响。空间结构显著,最佳空间模型为反距离加权模型(inverse distance-weighted model), 而且显示最大的空间扩散距离可以达到23.5 km,表明扩散限制较弱。约21%的物种多度变化能够被环境和空间因素解释。这些结果揭示了植物群落动态的决定因素,即环境因子的时空变化可能驱动物种分布和多度的随机性,并对植被镶嵌产生主导影响。     

Differential effects of goat browsing on herbaceous plant community in a two-phase mosaic

The impact of herbivores on herbaceous plant communities is usually attributed to direct consumption of plants. We hypothesized that goats affect herbaceous plants both directly (consumption by foraging) and indirectly, by changing environmental conditions through modification of woody plant structure. We assessed the effects of goats browsing on environmental conditions, landscape structure, and herbaceous plants to link the direct and indirect effects of goats on herbaceous communities. Our model system was the Mediterranean woodland in Mt. Carmel, Israel. This is a two-phase mosaic landscape, composed of herbaceous (open) and woody patches. We delineated 10 plots of 1000 m², goats were introduced to five plots and five plots remained without goats. We monitored plant species richness and composition in two adjacent patch types (woody and open) in each plot. For each patch type, in all plots, we collected data on environmental conditions. We analyzed landscape structure using landscape metrics derived from a high-resolution vegetation map. We found that goats modified the structure of woody plants and hence the landscape mosaic. This alteration was associated with changes in environmental conditions, with more light penetration and higher temperatures. The impact of goats on the herbaceous plant community depended on patch type. In open patches, goats affected the herbaceous community mostly by direct consumption, whereas in woody patches they affected the herbaceous community mainly by modification of abiotic conditions. Our results stress the importance of considering landscape and patch structure in analyzing the effect of herbivory on plant communities.     

δ15N values of tropical savanna and monsoon forest species reflect root specialisations and soil nitrogen status

A large number of herbaceous and woody plants from tropical woodland, savanna, and monsoon forest were analysed to determine the impact of environmental factors (nutrient and water availability, fire) and biological factors (microbial associations, systematics) on plant delta(15)N values. Foliar delta(15)N values of herbaceous and woody species were not related to growth form or phenology, but a strong relationship existed between mycorrhizal status and plant delta(15)N. In woodland and savanna, woody species with ectomycorrhizal (ECM) associations and putative N(2)-fixing species with ECM/arbuscular (AM) associations had lowest foliar delta(15)N values (1.0-0.6 per thousand ), AM species had mostly intermediate delta(15)N values (average +0.6 per thousand ), while non-mycorrhizal Proteaceae had highest delta(15)N values (+2.9 to +4.1 per thousand ). Similar differences in foliar delta(15)N were observed between AM (average 0.1 and 0.2 per thousand ) and non-mycorrhizal (average +0.8 and +0.3 per thousand ) herbaceous species in woodland and savanna. Leguminous savanna species had significantly higher leaf N contents (1.8-2.5% N) than non-fixing species (0.9-1.2% N) indicating substantial N acquisition via N(2) fixation. Monsoon forest species had similar leaf N contents (average 2.4% N) and positive delta(15)N values (+0.9 to +2.4 per thousand ). Soil nitrification and plant NO(3)(-) use was substantially higher in monsoon forest than in woodland or savanna. In the studied communities, higher soil N content and nitrification rates were associated with more positive soil delta(15)N and plant delta(15)N. In support of this notion, Ficus, a high NO(3)(-) using taxa associated with NO(3)(-) rich sites in the savanna, had the highest delta(15)N values of all AM species in the savanna. delta(15)N of xylem sap was examined as a tool for studying plant delta(15)N relations. delta(15)N of xylem sap varied seasonally and between differently aged Acacia and other savanna species. Plants from annually burnt savanna had significantly higher delta(15)N values compared to plants from less frequently burnt savanna, suggesting that foliar (15)N natural abundance could be used as marker for assessing historic fire regimes. Australian woodland and savanna species had low leaf delta(15)N and N content compared to species from equivalent African communities indicating that Australian biota are the more N depauperate. The largest differences in leaf delta(15)N occurred between the dominant ECM Australian and African savanna (miombo) species, which were depleted and enriched in (15)N, respectively. While the depleted delta(15)N of Australian ECM species are similar to those of previous reports on ECM species in natural plant communities, the (15)N-enriched delta(15)N of African ECM species represent an anomaly.     

Automatic detection of woody vegetation in repeat landscape photographs using a convolutional neural network

Repeat photography is an efficient method for documenting long-term landscape changes. So far, the usage of repeat photographs for quantitative analyses is limited to approaches based on manual classification. In this paper, we demonstrate the application of a convolutional neural network (CNN) for the automatic detection and classification of woody regrowth vegetation in repeat landscape photographs. We also tested if the classification results based on the automatic approach can be used for quantifying changes in woody vegetation cover between image pairs. The CNN was trained with 50 × 50 pixel tiles of woody vegetation and non-woody vegetation. We then tested the classifier on 17 pairs of repeat photographs to assess the model performance on unseen data. Results show that the CNN performed well in differentiating woody vegetation from non-woody vegetation (accuracy = 87.7%), but accuracy varied strongly between individual images. The very similar appearance of woody vegetation and herbaceous species in photographs made this a much more challenging task compared to the classification of vegetation as a single class (accuracy = 95.2%). In this regard, image quality was identified as one important factor influencing classification accuracy. Although the automatic classification provided good individual results on most of the 34 test photographs, change statistics based on the automatic approach deviated from actual changes. Nevertheless, the automatic approach was capable of identifying clear trends in increasing or decreasing woody vegetation in repeat photographs. Generally, the use of repeat photography in landscape monitoring represents a significant added value to other quantitative data retrieved from remote sensing and field measurements. Moreover, these photographs are able to raise awareness on landscape change among policy makers and public as well as they provide clear feedback on the effects of land management.     

Micro-site effects of trees and shrubs in dry savannas

Abstract. The physiognomy of dry savannas is described as a combination of discontinuous woody perennials and a continuous grassland matrix. Interactions between these two components are of vital importance for the persistence of a savanna landscape. Earlier savanna models have emphasized competitive interactions for water between the two components. Recent studies have argued that small-scale facilitating interactions between woody perennials and the herbaceous understorey are also important. This phenomenon has been given little theoretical consideration in the savanna literature, but it has been an important topic in agroforestry and arid-grassland ecology. This paper reviews some of the evidence for micro-site effects of trees and shrubs, and attempts to integrate their interactions with the surrounding open grassland. Woody perennials modify the microclimate by interception of solar radiation and rainfall. Their root systems extract nutrients horizontally and vertically, which are concentrated in the sub-canopy soil from litter decomposition and root turnover. Legumes are abundant in dry savannas, and may have symbiotic relationships with Rhizobium bacteria. This symbosis increases the availability of nitrogen in the soil. Isolated trees and shrubs initiate feedback mechanisms in their interactions with other organisms, and contribute to an uneven distribution of water and nutrients in dry savanna. This influences the species composition, and community diversity. Small-scale facilitating interaction between the woody and herbaceous components and competitive interaction on larger scales, are complementary processes which together explain a dynamic coexistence.     

Conundrums in mixed woody–herbaceous plant systems

Aims To identify approaches to improve our understanding of, and predictive capability for, mixed tree–grass systems. Elucidation of the interactions, dynamics and determinants, and identification of robust generalizations that can be broadly applied to tree–grass systems would benefit ecological theory, modelling and land management. Methods A series of workshops brought together scientific expertise to review theory, data availability, modelling approaches and key questions. Location Ecosystems characterized by mixtures of herbaceous and woody plant life‐forms, often termed ‘savannas’, range from open grasslands with few woody plants, to woodlands or forests with a grass layer. These ecosystems represent a substantial portion of the terrestrial biosphere, an important wildlife habitat, and a major resource for provision of livestock, fuel wood and other products. Results Although many concepts and principles developed for grassland and forest systems are relevant to these dual life‐form communities, the novel, complex, nonlinear behaviour of mixed tree–grass systems cannot be accounted for by simply studying or modelling woody and herbaceous components independently. A more robust understanding requires addressing three fundamental conundrums: (1) The ‘treeness’ conundrum. What controls the relative abundance of woody and herbaceous plants for a given set of conditions at given site? (2) The coexistence conundrum. How do the life‐forms interact with each other? Is a given woody–herbaceous ratio dynamically stable and persistent under a particular set of conditions? (3) The net primary productivity (NPP) conundrum. How does NPP of the woody vegetation, the herbaceous vegetation, and the total ecosystem (woody + herbaceous) change with changes in the tree–grass ratio? Tests of the theory and conceptual models of determinants of mixed woody–herbaceous systems have been largely site‐ or region‐specific and have seldom been broadly or quantitatively evaluated. Cross‐site syntheses based on data and modelling are required to address the conundrums and identify emerging patterns, yet, there are very few data sets for which either biomass or NPP have been quantified for both the woody and the herbaceous components of tree–grass systems. Furthermore, there are few cross‐site comparisons spanning the diverse array of woody–herbaceous mixtures. Hence, initial synthesis studies should focus on compiling and standardizing a global data base which could be (1) explored to ascertain if robust generalizations and consistent patterns exist; and (2) used to evaluate the performance of savanna simulation models over a range of woody–herbaceous mixtures. Savanna structure and productivity are the result of complex and dynamic interactions between climate, soils and disturbances, notably fire and herbivory. Such factors are difficult to isolate or experimentally manipulate in order to evaluate their impacts at spatial and temporal scales appropriate for assessing ecosystem dynamics. These factors can, however, be evaluated with simulation models. Existing savanna models vary markedly with respect to their conceptual approach, their data requirements and the extent to which they incorporate mechanistic processes. Model intercomparisons can elucidate those approaches most suitable for various research questions and management applications. Conclusion Theoretical and conceptual advances could be achieved by considering a broad continuum of grass–shrub–tree combinations using data meta‐analysis techniques and modelling.     

Vegetation management shapes arthropod and bird communities in an African savanna

Habitat heterogeneity is a key driver of the diversity and distribution of species. African savannas are experiencing changes in their vegetation structure causing shifts towards increased woody plant cover, which results in vegetation structure homogenization. Given the impact that increasing woody plant cover has on patterns of animal use, resource managers across Africa are implementing habitat management practices that are intended to reduce woody plant cover. To understand the ecological implications of various habitat management practices on arthropod and bird communities, we leveraged large‐scale tree clearing and subsequent mowing in an African savanna to understand how changes in both the herbaceous layer and woody plant cover (i.e., structural heterogeneity) may shape arthropod and bird communities at the local scale. We focused on four replicated treatments: (1) annual summer mow, (2) annual winter mow, (3) >5 years since last mow (rest), and (4) an adjacent unmanipulated savanna to act as a control. We found that the mowing treatments significantly influenced vegetation structure both with respect to tree density and herbaceous layer. Both arthropod and bird community composition varied across treatments. Grass biomass was the best predictor of arthropod richness and abundance, with arthropods selecting for areas with high biomass. Insectivorous bird richness and abundance was driven by tree density (i.e., perching locations) and not arthropod abundance. Our results suggest that vegetation management practices contribute to habitat heterogeneity at the landscape scale and increase bird species richness through species turnover. However, we caution that if a single vegetation management practice dominates the landscape, it is plausible that it could lead to the simplification of the avian community.     

Use and conservation of woody vegetation by the Batemi of Ngorongoro District,Tanzania

The Batemi, agropastoralists of north-central Tanzania, inhabit a landscape of savanna and scrub woodland. We identified 61 species of woody plants used by the Batemi for 19 types of items. The reports indicated 39 species are used for construction, 19 species for cultural artifacts, 23 for firewood and 6 live trees for fences, boundaries and shade. The Batemi have extensive knowledge of species habitat and distinctive management strategies for woody vegetation, particularly for the well-forested irrigation channels and spring sources. To date, a total of 90 species of woody plants have been identified as useful to the Batemi. Compared to the total number of species found within the study area, the Batemi use 79% of woody plant species in the area.     

Change in the vegetation mosaic of central Arizona USA between 1940 and 1989

The landscape of central Arizona U.S.A. is characterized by a patchy distribution of three major vegetation types: chaparral, woodland and grassland. Disturbance is common in the landscape, primarily livestock grazing, fire and conversion (i.e., removal of woody plants to increase forage for livestock). The purposes of this research were to determine changes in the landscape mosaic of central Arizona between 1940 and 1989 and to predict future changes. Using aerial photographs from 1940, 1968 and 1989 and digital overlays followed by transition matrix analysis, we found that chaparral and its adjacent grassland had changed less than woodland and its grassland. However, both had nearly equal projected stabilization times. Moreover, disturbance increased time for stabilization and some results were scale-dependent.     

Self‐thinning and Tree Competition in Savannas

This paper examines the feasibility of applying self‐thinning concepts to savannas and how competition with herbaceous vegetation may modify self‐thinning patterns among woody plants in these ecosystems. Competition among woody plants has seldom been invoked as a major explanation for the persistence of herbaceous vegetation in mixed tree–grass ecosystems. On the contrary, the primary resource‐based explanations for tree–grass coexistence are based on tree–grass competition (niche‐separation) that assumes that trees are inferior competitors unless deeper rooting depths provide them exclusive access to water. Alternative nonresource‐based hypotheses postulate that trees are the better competitors, but that tree populations are suppressed by mortality related to fire, herbivores, and other disturbances. If self‐thinning of woody plants can be detected in savannas, stronger evidence for resource‐limitation and competitive interactions among woody plants would suggest that the primary models of savannas need to be adjusted. We present data from savanna sites in South Africa to suggest that self‐thinning among woody plants can be detected in low‐disturbance situations, while also showing signs that juvenile trees, more so than adults, are suppressed when growing with herbaceous vegetation in these ecosystems. This finding we suggest is evidence for size‐asymmetric competition in savannas.     

Exotic plant invasions over 40 years of old field successions: community patterns and associations

While exotic plant species often come to dominate disturbed communities, long-term patterns of invasion are poorly known. Here we present data from 40 yr of continuous vegetation sampling, documenting the temporal distribution of exotic plant species in old field succession. The relative cover of exotic species decreased with time since abandonment, with significant declines occurring ≥20 yr post-abandonment. The number of exotic species per plot also declined with time since abandonment while field-scale richness of exotics did not change. This suggests displacement occurring at small spatial scales. Life history types changed from short-lived herbaceous species to long-lived woody species for both native and exotic plant species. However, shrubs and lianas dominated woody cover of exotic plants while trees dominated native woody cover. The species richness of exotic and native species was positively correlated at most times. In abandoned hay fields, however, the proportion of exotic plant cover per plot was inversely related to total species richness. This relationship suggests that it is not the presence, but the abundance of exotic species that may cause a reduction in community diversity. While the development of closed-canopy forest appears to limit most introduced plant species, several shade-adapted exotic species are increasing within the fields. These invasions may cause a reversal of the patterns seen in the first 40 yr of succession and may result in further impacts on community structure.     

Development of vegetation in dune slack wetlands of Cape Cod National Seashore (Massachusetts,USA)

Interdunal seasonal wetlands, known as dune slack wetlands, were mapped and their vegetation surveyed across a large region of dunes within the Cape Cod National Seashore. Wetland sizes and ages were estimated from digitized, georeferenced aerial photographs available from 1938, 1947, 1960, 1986, 1994, and 2001, and from LIDAR elevation data. A total of 346 sites were found, covering an area of ∼45.4 ha., in which 97 species of vascular plants were identified. Vegetation structure and composition exhibited a distinct sequence of development with age, following a pattern of succession from herbaceous, graminoid-dominated communities to shrub- and tree-dominated communities. Floristic variables were not related to proximity to the coast and although wetland size appeared to have some bearing on species richness, the correlation was statistically weak. Soil organic matter determined for a subset of 60 wetlands was positively correlated with age and woody cover but showed no relationship with water depth. The results suggest that vegetation development is primarily driven by the internal mechanisms of succession. Notwithstanding, any changes in the environment that alter the process of succession will collectively influence these wetlands. In addition, stabilization of the dunes resulting in a reduction in the formation of new wetlands, may translate into permanent loss of early and mid-successional dune slack communities. Maintenance of these communities will depend on succession being periodically reset by disturbance or active management.     

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.     

Effects of large herbivores on wood pasture dynamics in a European wetland system

Whether self-regulating large herbivores play a key role in the development of wood-pasture landscapes remains a crucial unanswered question for both ecological theory and nature conservation. We describe and analyse how a ‘partly self-regulating’ population of cattle, horses and red deer affected the development of the woody vegetation in the Oostvaardersplassen nature reserve (Netherlands). Using aerial photographs from 1980 to 2011, we analysed the development of shrubs and trees. Before the large herbivores were introduced in the Oostvaardersplassen in 1983, the woody vegetation increased and vegetation type significantly affected the number of establishments. Cover of woody species increased further from 1983 to 1996, not only by canopy expansion but also by new establishments. After 1996, cover of the woody vegetation decreased from 30% to <1% in 2011 and no new establishments were seen on the photographs. Survival of Sambucus nigra and Salix spp. increased with increasing distance to grassland, which is the preferred foraging habitat of the herbivores. These results support the hypothesis of Associational Palatability. In addition, our results show that the relative decline in cover of S. nigra and Salix spp. over a certain period was negatively correlated with the cover of S. nigra in the beginning of this period, presenting some evidence for the Associational Resistance and Aggregational Resistance hypothesis. Our research shows aspects necessary for the woodland–grassland cycle, such as a strong decline of woody vegetation at high numbers of large herbivores and regeneration of shrubs and trees at low densities. Thorny shrubs, which are important for the cycle, have not yet established in the grasslands. It seems that a temporary decline in herbivore numbers is necessary to create a window of opportunity for the establishment of these woody species.     

Changes in miombo woodland cover in and around Sengwa Wildlife Research Area, Zimbabwe, in relation to elephants and fire

One of the consequences of impacts of elephants and fire on woodlands is a change in woody cover, which often results in major challenges for wildlife managers. Changes in miombo woodland cover in and around Sengwa Wildlife Research Area (SWRA) between 1958 and 1996 were quantified by analyzing aerial photographs. Woody cover in SWRA decreazed from 95.2% in 1958 to 68.2% in 1996, with a lowest mean of 62.9% in 1983. The annual absolute rate of woody cover change in SWRA increazed from ?1.1% per annum between 1958 and 1964 to a recovery of 1.6% per annum between 1993 and 1996, while the annual relative rate increazed from ?1.1% per annum between 1958 and 1964 to 3.3% per annum between 1993 and 1996. There was a strong negative correlation between elephant densities and woody cover in SWRA, suggesting that loss of woody cover was mainly due to elephants. Woodland recovery after 1983 was due to reductions in elephant populations through legal and illegal off‐take and reductions in fire frequency. Surrounding areas experienced less woody cover losses than SWRA, mainly due to tree removal by locals whose densities increazed after the eradication of tsetse fly in the 1970s.     

Forest expansion and grassland contraction within a Eucalyptus savanna matrix between 1941 and 1994 at Litchfield National Park in the Australian monsoon tropics

• 1 Analysis of digitized aerial photographs taken in 1941 and 1994, using image processing and geographical information system technology, enabled the quantification of change in the coverage of forest and grassland patches that occur within a Eucalyptus savanna matrix in a subcoastal region of the Australian monsoon tropics. The 3058 ha study area was orientated along a low escarpment that separated a sandstone plateau from lowlands that comprised 58% and 42% of the area, respectively.
• 2 In the 53‐year period, humans modified less than 1% of the study area, primarily for road building, and primarily in savanna areas. More than 85% of the study area at both sample times was covered by savanna. However, over the same period, the forest coverage increased from 5.03% to 9.91% of the study area and coverage of grassland decreased from 6.70% to 2.47%. The aerial photography also showed that tree density in the savanna had increased, although this was not assessed quantitatively.
• 3 There was an increase in the number of forest patches from 116 to 142. The number of grassland patches decreased (particularly those > 1 ha) from 87 to 59, although the size class distribution of forest and grassland patches was statistically similar for both sample times.
• 4 A 50‐m GIS buffer was used to distinguish creek‐lines environments from surrounding catchments. Using this criterion, 14% of the study area was classified as plateau creek‐lines and 9% lowland creek‐lines. Although the expansion of forest and loss of grassland varied significantly amongst catchment and creek‐lines on the plateau and lowlands, the 1941 rank order of coverage of each vegetation type was maintained in these four landscape categories in 1994. In both years the greatest extent of forest and grassland occurred on the lowland catchments, despite their accounting for only one‐third of the study area.
• 5 Transition matrices for vegetation change among the four landscape categories demonstrated that, unlike the other vegetation types, grasslands, particularly on the plateau, had a low probability of remaining unchanged during the study period.
• 6 The cause(s) of the overall increase of woody biomass across the topographic and edaphic gradient remains unclear but may be related to a period of increased rainfall since the 1970s, as well as to the cessation of Aboriginal landscape burning at the beginning of the study period.

     

Effects of Herbivory,Fire and N<Subscript>2</Subscript>-fixation on Nutrient Limitation in a Humid African Savanna

The quantities and spatial distribution of nutrients in savanna ecosystems are affected by many factors, of which fire, herbivory and symbiotic N₂-fixation are particularly important. We measured soil nitrogen (N) pools and the relative abundance of N and phosphorus (P) in herbaceous vegetation in five vegetation types in a humid savanna in Tanzania. We also performed a factorial fertilization experiment to investigate which nutrients most limit herbaceous production. N pools in the top 10 cm of soil were low at sites where fires were frequent, and higher in areas with woody legume encroachment, or high herbivore excretion. Biomass production was co-limited by N and P at sites that were frequently burnt or heavily grazed by native herbivores. In contrast, aboveground production was limited by N in areas receiving large amounts of excreta from livestock. N₂-fixation by woody legumes did not lead to P-limitation, but did increase the availability of N relative to P. We conclude that the effects of fire, herbivory and N₂-fixation upon soil N pools and N:P-stoichiometry in savanna ecosystems are, to a large extent, predictable. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Author Contributions  P.C., H.O.V. and P.E. designed the study and wrote the paper. P.C. and T.K. performed the research and analyzed the data.     

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.