Linking woody species diversity with plant available water at a landscape scale in a Brazilian savanna

Question: How is the diversity of woody species in a seasonally dry savanna related to plant available water (PAW)? Location: Savannas in central Brazil. Methods: Two‐dimensional soil resistivity profiles to 10‐m depth previously measured along three 10 m × 275 m replicate transects revealed differences in belowground water resources among and within transects: (1) driest/most heterogeneous; (2) wettest/least heterogeneous; and (3) PAW‐intermediate. All woody plants along these transects were identified to species, and height and basal circumference measured. Species diversity was evaluated for the whole transect (total diversity), 100‐m² plots (alpha‐diversity) and dissimilarity among 100‐m² plots within transects (beta‐diversity). Correlation analyses were conducted between PAW and vegetation variables at the 100‐m² scale. Results: The driest/most heterogeneous transect had the lowest total species diversity, while the wettest/least heterogeneous transect showed the lowest beta‐diversity. Floristic variation was correlated with PAW in all transects. In the most heterogeneous transect, species density was positively correlated with PAW in the 0‐400 cm soil layer. Evenness and Simpson's diversity were negatively correlated with PAW in the 700‐1000 cm soil layer. Conclusion: Woody species diversity was related to PAW at a fine spatial scale. Abundant PAW in the top 4 m of soil may favour many species and increase species total diversity. Conversely, abundant PAW at depth may result in lower evenness and total diversity, probably because the few species adapted to obtaining deep soil water can become dominant. Environmental changes altering soil water availability and partitioning in soil layers could affect the diversity of woody plants in this savanna.     

Seasonal leaf dynamics across a tree density gradient in a Brazilian savanna

Interactions between trees and grasses that influence leaf area index (LAI) have important consequences for savanna ecosystem processes through their controls on water, carbon, and energy fluxes as well as fire regimes. We measured LAI, of the groundlayer (herbaceous and woody plants <1-m tall) and shrub and tree layer (woody plants >1-m tall), in the Brazilian cerrado over a range of tree densities from open shrub savanna to closed woodland through the annual cycle. During the dry season, soil water potential was strongly and positively correlated with grass LAI, and less strongly with tree and shrub LAI. By the end of the dry season, LAI of grasses, groundlayer dicots and trees declined to 28, 60, and 68% of mean wet-season values, respectively. We compared the data to remotely sensed vegetation indices, finding that field measurements were more strongly correlated to the enhanced vegetation index (EVI, r ²=0.71) than to the normalized difference vegetation index (NDVI, r ²=0.49). Although the latter has been more widely used in quantifying leaf dynamics of tropical savannas, EVI appears better suited for this purpose. Our ground-based measurements demonstrate that groundlayer LAI declines with increasing tree density across sites, with savanna grasses being excluded at a tree LAI of approximately 3.3. LAI averaged 4.2 in nearby gallery (riparian) forest, so savanna grasses were absent, thereby greatly reducing fire risk and permitting survival of fire-sensitive forest tree species. Although edaphic conditions may partly explain the larger tree LAI of forests, relative to savanna, biological differences between savanna and forest tree species play an important role. Overall, forest tree species had 48% greater LAI than congeneric savanna trees under similar growing conditions. Savanna and forest species play distinct roles in the structure and dynamics of savanna–forest boundaries, contributing to the differences in fire regimes, microclimate, and nutrient cycling between savanna and forest ecosystems.     

Multiple Scales of Control on the Structure and Spatial Distribution of Woody Vegetation in African Savanna Watersheds

Factors controlling savanna woody vegetation structure vary at multiple spatial and temporal scales, and as a consequence, unraveling their combined effects has proven to be a classic challenge in savanna ecology. We used airborne LiDAR (light detection and ranging) to map three-dimensional woody vegetation structure throughout four savanna watersheds, each contrasting in geologic substrate and climate, in Kruger National Park, South Africa. By comparison of the four watersheds, we found that geologic substrate had a stronger effect than climate in determining watershed-scale differences in vegetation structural properties, including cover, height and crown density. Generalized Linear Models were used to assess the spatial distribution of woody vegetation structural properties, including cover, height and crown density, in relation to mapped hydrologic, topographic and fire history traits. For each substrate and climate combination, models incorporating topography, hydrology and fire history explained up to 30% of the remaining variation in woody canopy structure, but inclusion of a spatial autocovariate term further improved model performance. Both crown density and the cover of shorter woody canopies were determined more by unknown factors likely to be changing on smaller spatial scales, such as soil texture, herbivore abundance or fire behavior, than by our mapped regional-scale changes in topography and hydrology. We also detected patterns in spatial covariance at distances up to 50–450 m, depending on watershed and structural metric. Our results suggest that large-scale environmental factors play a smaller role than is often attributed to them in determining woody vegetation structure in southern African savannas. This highlights the need for more spatially-explicit, wide-area analyses using high resolution remote sensing techniques.     

Net woody vegetation increase confined to seasonally inundated lowlands in an Australian tropical savanna,Victoria River District,Northern Territory

Abstract Georeferenced digital aerial photographs were used to assess changes in overstorey vegetation cover since 1948 in the Victoria River District, Northern Territory, Australia, across a range of lowland tropical savanna habitats and with explicit consideration of known and variable site‐specific grazing and fire management histories. Vegetation surveys at corresponding locations on the ground identified five distinct woody vegetation communities defined primarily by water drainage and secondarily by soil characteristics. Air‐photo analyses revealed that, contrary to popular perceptions and in contrast to results from other habitats, there has been no generalized net increase in overstorey woody vegetation cover across the full range of lowland savanna habitats. Rather, different habitats exhibited distinctly different vegetation change mechanisms: low‐lying seasonally inundated ‘wet’ habitats have experienced woody vegetation increase since 1948, whereas well‐drained ‘dry’ habitats have experienced overstorey vegetation stability or loss. In almost every instance woody vegetation increase could be attributed to the invasion or proliferation of a single species, Melaleuca minutifolia F.Muell. The extent of M. minutifolia increase was unrelated to historical grazing/fire regime. Demographic analyses for this species revealed that recruitment was often episodic and that synchronized recruitment events occurred uniformly across the full range of historical management treatments, most likely as a consequence of favourable climatic conditions in years with an extended wet season. Heavy grazing facilitated juvenile survival and/or recruitment, most likely by reducing grassy fuel loads and eliminating landscape fire. We conclude that while there has been no generalized net increase in overstorey woody vegetation cover in lowland environments, savanna dynamics are complex, and multiple change mechanisms have occurred simultaneously in different habitats, some of which have been significantly transformed since 1948. Where net woody vegetation increase has occurred it is primarily a natural consequence of episodic M. minutifolia establishment in climatically favourable years, but the extent and magnitude of this effect is likely mediated by fire/grazing regime.     

Soil seed bank dynamics of fire-prone wooded grassland, woodland and dry forest ecosystems in Ethiopia

In order to reveal the role of soil seed banks in vegetation recovery after fire in savanna, the spatial distribution and temporal changes in the soil seed banks of regularly burning savanna in Gambella, western Ethiopia, was studied. The seedling emergence technique was employed to determine the species composition and density of the soil seed bank of six sites ranging in fire severity from wooded grassland with frequent fires over woodland with intermediate fire frequency to forest with absence of fires. Species composition and density of seeds in the soil were compared between seasons, depths and sites with different types of standing vegetation. Fourteen plant species were recorded in the soil seed bank from the grassland and woodland sites and 6 from the dry forests; 60 % of the taxa in the soil seed bank were annuals and 40 % were perennials. The soil seed banks were largely dominated by graminoids and 48–97 % of the soil seed bank in the grasslands and woodlands was of a single grass species, Hyparrhenia confinis , which was absent from the dry forests. The soil seed pools ranged from less than 100 to 4700 seeds per m² depending upon the season. The soil seed bank of graminoids was nearly empty after the onset of the rainy season whereas seeds of broadleaved herbs and woody species able to germinate were still found after this time. Floristic composition, representation of life forms and density of seeds in soil did not correspond closely with that of the standing vegetation, but within graminoids there was a strong similarity between the soil seed bank and the standing vegetation. The current fire regime of Ethiopian savanna woodlands appears to maintain the dominance of graminoids over broadleaved herbs and woody plants both as seeds in the soil and in the standing vegetation.     

Strong positive effects of termites on savanna bird abundance and diversity are amplified by large herbivore exclusion

Vast areas of the African savanna landscapes are characterized by tree‐covered Macrotermes termite mounds embedded within a relatively open savanna matrix. In concert with termites, large herbivores are important determinants of savanna woody vegetation cover. The relative cover of woody species has considerable effects on savanna function. Despite the potentially important ecological relationships between termite mounds, woody plants, large herbivores, and birds, these associations have previously received surprisingly little attention. We experimentally studied the effects of termites and large herbivores on the avian community in Lake Mburo National Park, Uganda, where woody vegetation is essentially limited to termite mounds. Our experiment comprised of four treatments in nine replicates; unfenced termite mounds, fenced mounds (excluding large mammals), unfenced adjacent savanna, and fenced savanna. We recorded species identity, abundance, and behavior of all birds observed on these plots over a two‐month period, from late dry until wet season. Birds used termite mounds almost exclusively, with only 3.5% of observations occurring in the treeless intermound savanna matrix. Mean abundance and species richness of birds doubled on fenced (large herbivores excluded) compared to unfenced mounds. Feeding behavior increased when large mammals were excluded from mounds, both in absolute number of observed individuals, and relative to other behaviors. This study documents the fundamental positive impact of Macrotermes termites on bird abundance and diversity in an African savanna. Birds play crucial functional roles in savanna ecosystems, for example, by dispersing fruits or regulating herbivorous insect populations. Thus, the role of birds in savanna dynamics depends on the distribution and abundance of termite mounds.     

Floristic relationships of the woody component in rocky outcrops savanna areas in Central Brazil

Rocky outcrops usually have a peculiar vegetation which differs from that of the surrounding vegetation matrix. In Brazil, a savanna type (cerrado rupestre) is particularly characterized by rocky outcrops and is one of eleven vegetation types which occur throughout Cerrado biome. It has been scarcely studied in detail. Therefore, the aim of this study was to investigate the phytogeographical patterns of the woody species in ten areas of the rocky outcrop savanna in the Brazilian Central Plateau. The difference in spatial distribution of the species was assessed through indicator species and CCA ordination analyses. Ten indicator species showed significantly different distribution. The first axis of the ordination showed association with the edaphic gradient, based on maximum temperature, pH and physical properties of the soils (eigenvalue 0.63). The second axis showed an eigenvalue 0.46, associated with poor rainfall and land declivity. It appears that the woody flora of the rocky outcrop savanna areas of the Brazilian Central Plateau is relatively homogeneous, with low β diversity. Apparently, the size of the woody species populations is what distinguishes the ten investigated areas. Therefore, initiatives and actions for the Cerrado biome conservation must consider not only the size of the protected areas but also that of the populations to be protected, especially the woody species on rocky outcrops.     

Nigerian Man and Biosphere research plots in Guinea savanna: floristics and structure of the vegetation

A study was made of the vegetation of three plots used in the Nigerian Man and Biosphere Research Programme for Savanna Studies. The aim is to provide detailed information on the vegetation of the plots which may possibly contribute to an understanding of factors influencing savanna structure and relative abundances of forbs, grasses and woody species. Results show that there are differences in the species composition of the plots. There are more forb species than grass species in each plot. The woody basal areas and crown areas of the plots do not depend entirely on the density but also on the size of the woody species. There are differences in herbage yield in the plots that could be attributed to differences in soil properties, species composition and level of human and animal activities between the plots.     

The influence of savanna trees on nutrient,water and light availability and the understorey vegetation

In an East African savanna herbaceous layer productivity and species composition were studied around Acacia tortilis trees of three different age classes, as well as around dead trees and in open grassland patches. The effects of trees on nutrient, light and water availability were measured to obtain an insight into which resources determine changes in productivity and composition of the herbaceous layer. Soil nutrient availability increased with tree age and size and was lowest in open grassland and highest under dead trees. The lower N:P ratios of grasses from open grassland compared to grasses from under trees suggested that productivity in open grassland was limited by nitrogen, while under trees the limiting nutrient was probably P. N:P ratios of grasses growing under bushes and small trees were intermediate between large trees and open grassland indicating that the understorey of Acacia trees seemed to change gradually from a N-limited to a P-limited vegetation. Soil moisture contents were lower under than those outside of canopies of large Acacia trees suggesting that water competition between trees and grasses was important. Species composition of the herbaceous layer under Acacia trees was completely different from the vegetation in open grassland. Also the vegetation under bushes of Acacia tortilis was different from both open grassland and the understorey of large trees. The main factor causing differences in species composition was probably nutrient availability because species compositions were similar for stands of similar soil nutrient concentrations even when light and water availability was different. Changes in species composition did not result in differences in above-ground biomass, which was remarkably similar under different sized trees and in open grassland. The only exception was around dead trees where herbaceous plant production was 60% higher than under living trees. The results suggest that herbaceous layer productivity did not increase under trees by a higher soil nutrient availability, probably because grass production was limited by competition for water. This was consistent with the high plant production around dead trees because when trees die, water competition disappears but the high soil nutrient availability remains. Hence, in addition to tree soil nutrient enrichment, below-ground competition for water appears to be an important process regulating tree-grass interactions in semi-arid savanna.     

Vegetation,fire and soil feedbacks of dynamic boundaries between rainforest,savanna and grassland

At fine spatial scales, savanna‐rainforest‐grassland boundary dynamics are thought to be mediated by the interplay between fire, vegetation and soil feedbacks. These processes were investigated by quantifying tree species composition, the light environment, quantities and flammability of fuels, bark thickness, and soil conditions across stable and dynamic rainforest boundaries that adjoin grassland and eucalypt savanna in the highlands of the Bunya Mountains, southeast Queensland, Australia. The size class distribution of savanna and rainforest stems was indicative of the encroachment of rainforest species into savanna and grassland. Increasing dominance of rainforest trees corresponds to an increase in woody canopy cover, the dominance of litter fuels (woody debris and leaf), and decline in grass occurrence. There is marked difference in litter and grass fuel flammability and this result is largely an influence of strongly dissimilar fuel bulk densities. Relative bark thickness, a measure of stem fire resistance, was found to be generally greater in savanna species when compared to that of rainforest species, with notable exceptions being the conifers Araucaria bidwillii and Araucaria cunninghamii. A transect study of soil nutrients across one dynamic rainforest – grassland boundary indicated the mass of carbon and nitrogen, but not phosphorus, increased across the successional gradient. Soil carbon turnover time is shortest in stable rainforest, intermediate in dynamic rainforest and longest in grassland highlighting nutrient cycling differentiation. We conclude that the general absence of fire in the Bunya Mountains, due to a divergence from traditional Aboriginal burning practices, has allowed for the encroachment of fire‐sensitive rainforest species into the flammable biomes of this landscape. Rainforest invasion is likely to have reduced fire risk via changes to fuel composition and microclimatic conditions, and this feedback will be reinforced by altered nutrient cycling. The mechanics of the feedbacks here identified are discussed in terms of landscape change theory.     

Do feral buffalo (Bubalus bubalis) explain the increase of woody cover in savannas of Kakadu National Park,Australia?

Aim To study changes in woody vegetation in both floodplains and eucalypt savanna over a 40‐year period using multi‐temporal spatial analysis of variation in density of a large introduced herbivore, the Asian water buffalo (Bubalus bubalis). Feral buffalo built up to high densities in the study area until c. 1985, after which a control programme almost eliminated the animals. From 1990, low densities of managed buffalo were maintained inside an enclosure. We compared trends in woody vegetation when buffalo were high‐density feral, low‐density managed or absent. Location The study area was located in and around a 116‐km² buffalo enclosure inside Kakadu National Park, in monsoonal northern Australia. Methods We analysed sequences of digitized and geo‐rectified aerial photographs, acquired in 1964, 1975, 1984, 1991 and 2004, to chart changes in woody cover on the floodplain and in the savanna. On the floodplain we assessed whether trees were present at these times at 14,568 points, and buffalo density was estimated from the density of animal tracks. In the savanna we estimated woody cover at pre‐selected sites. Generalized linear modelling was used to analyse changes in woody vegetation, using elevation and presence of woody vegetation in neighbouring points on the floodplain, and buffalo regime and initial woody cover in the savanna. Results Changes in animal track density reflected park‐wide historical estimates of buffalo numbers. Tree cover increased in both floodplain and savanna, but this was only weakly related to buffalo density. The best predictor of whether a floodplain cell converted from treeless to woody, or the converse, was the woodiness of neighbouring vegetation. There was slightly less thickening with high buffalo densities. In savanna, low densities of managed buffalo were weakly associated with increases in tree cover relative to either high densities of feral buffalo or no buffalo. Main conclusions Our study indicates that buffalo are not a major driver of floodplain and eucalypt savanna dynamics. Rather, the observed increase in woody cover in both savanna and flood plains concords with regional trends and may be related to increased atmospheric CO₂, increasing rainfall and changing fire regimes during the study period.     

Using plant functional types to compare vegetation structure of alien-invaded and uninvaded Acacia nilotica savannas

Four plant functional types (PFTs) were used to compare the vegetation structure of an alien-invaded Acacia nilotica savanna with one of negligible invasions. Heights, canopy covers and species richness of three native PFTs (woody plants, grasses and herbs) and one alien PFT (woody plants) were measured in 14, 1-m² quadrats sampled in a stratified-random pattern in a 400-m² plot demarcated in each savanna. In the uninvaded plot, mean heights of native PFTs were stratified. In the invaded plot, the mean height of aliens extended into the native woody stratum with the lower range of native woody PFT heights reduced to the grass stratum. Discriminant analysis of canopy covers and species richness of the four PFTs revealed significant differences in composition between plots with the alien PFT being the most important variable correlated with these differences. Univariate analysis confirmed the dominance of alien woody plants in the invaded plot but also showed significant reductions in the canopy covers and species richness of native herbs and grasses compared to those in the uninvaded plot. These results suggest that PFTs can rapidly measure small-scale, spatial differences in the physiognomy, composition and species richness of A. nilotica savannas when invaded by alien woody plants.     

Mound building termites contribute to savanna vegetation heterogeneity

With biomass densities comparable to large ungulates and megaherbivores, termites play a key functional role in many tropical savanna ecosystems. This study focuses on vegetated termite mounds (termitaria) constructed by the Termitidae species Macrotermes herus. We studied how resource rich termitaria affect graminoid herbs (Poaceae and Cyperaceae), forbs and woody species composition and diversity. The density of termitaria explained 89% of the variation in dense thickets in the area. Fire tolerant Acacia species dominated the open savanna while fire sensitive species like Grewia spp. and the succulent Euphorbia candelabrum were restricted to termite mounds. Termitaria plots had four times the mean number of woody species and supported three times as many forb species as the adjacent savanna. For woody species, both the Shannon–Wiener index and the Shannon evenness index were higher on temitaria than on the savanna. There were no differences for graminoid herbs, except for the Shannon evenness index which was higher on termitaria. Our results indicate that graminoid herb richness peaks at lower productivity levels than trees and forbs in savanna ecosystems, as also recently found in temperate areas.     

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.     

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.     

Regeneration of monsoon rain forest in northern Australia: the dormant seed bank

Abstract. As part of a wider study examining regeneration pathways in monsoon rain forest vegetation in northern Australia, the dormant component of the soil seed bank was assessed by storing soil samples for over six dry season months, before watering in shade-house trials. Six soil samples were collected from each of 34 sites broadly representative of the range of regional monsoon rain forest vegetation. Four floristic seed bank groups were derived through TWINSPAN classification. Mean group densities of germinants ranged from 25–144/m². Dormant seed banks were least dense, and most sparsely distributed, in sandy soils. Seed bank samples were dominated by woody pioneer monsoon rain forest species, especially figs; exotic weeds and savanna taxa (e.g. Poaceae) were relatively more common at seasonally dry sites. Dormant seed banks comprised species mostly present in the standing vegetation, although a small number of germinants represented species not growing at half the sites. Regeneration of woody pioneers from dormant seed banks is least likely to be of importance on infertile, seasonally dry sites.     

The irreversible cattle-driven transformation of a seasonally flooded Australian savanna

Aim Anecdotal historical and photographic evidence suggests that woody vegetation is increasing dramatically in some northern Australian savanna habitats. Vegetation change in savannas has important implications for pastoral land‐use, conservation management, and landscape‐scale carbon storage, and informs theoretical debates about ecosystem function. This study seeks to determine the nature, extent and cause(s) of woody vegetation change in a seasonally flooded alluvial savanna habitat. Location The study area is located within the seasonally inundated alluvial zone of the tidal portion of the Victoria River, Northern Territory, Australia. The study area has been grazed by domestic stock since c. 1900, prior to which the area was inhabited and more likely regularly burnt by Aboriginal people for thousands of years. Methods Digital georeferenced aerial photographic coverages were used to examine and quantify woody vegetation change between 1948 and 1993. Transect surveys of woody and herbaceous vegetation were carried out to ground‐truth air‐photo results and determine the nature and causes of observed vegetation changes. Results There has been a dramatic increase in woody vegetation cover throughout the study area. Vegetation change patterns are roughly uniform across the full range of edaphic habitat variation and are unrelated to the depositional age of fluvial sediments. Two woody species, Eucalyptus microtheca and Excoecaria parvifolia, are predominantly responsible for observed increases. Demographic analyses reveal that woody invasions have been episodic and indicate that in most locations peak woody species establishment occurred in the mid‐1970s. Grasses are almost absent in a majority of habitats within the study area. Instead, large areas are covered by scalded soil, dense invasive weed populations, and unpalatable forbs and sedges. What grasses do occur are predominantly of very low value for grazing. The condition of the herbaceous layer renders most of the study area almost completely non‐flammable; what fires do burn are small and of low intensity. Main conclusions Multiple working hypotheses explaining observed patterns of woody vegetation increase were considered and rejected in turn. The only hypothesis consistent with the evidence is as follows: (1) observed changes are a direct consequence of extreme overgrazing by cattle, most likely when stocking rates peaked in the mid‐1970s; (2) prolonged heavy grazing effected the complete transformation of much of the herbaceous vegetation to a new state that is not flammable; and (3) in the absence of regular fire mortality, woody vegetation increased rapidly. The relatively treeless system that existed in 1948 was apparently stable and resilient to moderate grazing levels, and perhaps also to episodic heavy grazing events. However, grazing intensity in excess of a sustainable threshold has forced a transition that is irreversible in the foreseeable future. Stable‐state transitions such as this one inform debates at the heart of ecological theory, such as the nature of stability, resilience, equilibrium and carrying capacity in dynamic savanna ecosystems.     

Spatial variation in vegetation structure coupled to plant available water determined by two-dimensional soil resistivity profiling in a Brazilian savanna

Tropical savannas commonly exhibit large spatial heterogeneity in vegetation structure. Fine-scale patterns of soil moisture, particularly in the deeper soil layers, have not been well investigated as factors possibly influencing vegetation patterns in savannas. Here we investigate the role of soil water availability and heterogeneity related to vegetation structure in an area of the Brazilian savanna (Cerrado). Our objective was to determine whether horizontal spatial variations of soil water are coupled with patterns of vegetation structure across tens of meters. We applied a novel methodological approach to convert soil electrical resistivity measurements along three 275-m transects to volumetric water content and then to estimates of plant available water (PAW). Structural attributes of the woody vegetation, including plant position, height, basal circumference, crown dimensions, and leaf area index, were surveyed within twenty-two 100-m² plots along the same transects, where no obvious vegetation gradients had been apparent. Spatial heterogeneity was evaluated through measurements of spatial autocorrelation in both PAW and vegetation structure. Comparisons with null models suggest that plants were randomly distributed over the transect with the greatest mean PAW and lowest PAW heterogeneity, and clustered in the driest and most heterogeneous transect. Plant density was positively related with PAW in the top 4 m of soil. The density-dependent vegetation attributes that are related to plot biomass, such as sum of tree heights per plot, exhibited spatial variation patterns that were remarkably similar to spatial variation of PAW in the top 4 m of soil. For PAW below 4 m depth, mean vegetation attributes, such as mean height, were negatively correlated with PAW, suggesting greater water uptake from the deep soil by plants of larger stature. These results are consistent with PAW heterogeneity being an important structuring factor in the plant distribution at the scale of tens of meters in this ecosystem.     

Soil phosphorus does not keep pace with soil carbon and nitrogen accumulation following woody encroachment

Soil carbon, nitrogen, and phosphorus cycles are strongly interlinked and controlled through biological processes, and the phosphorus cycle is further controlled through geochemical processes. In dryland ecosystems, woody encroachment often modifies soil carbon, nitrogen, and phosphorus stores, although it remains unknown if these three elements change proportionally in response to this vegetation change. We evaluated proportional changes and spatial patterns of soil organic carbon (SOC), total nitrogen (TN), and total phosphorus (TP) concentrations following woody encroachment by taking spatially explicit soil cores to a depth of 1.2 m across a subtropical savanna landscape which has undergone encroachment by Prosopis glandulosa (an N₂ fixer) and other woody species during the past century in southern Texas, USA. SOC and TN were coupled with respect to increasing magnitudes and spatial patterns throughout the soil profile following woody encroachment, while TP increased slower than SOC and TN in topmost surface soils (0–5 cm) but faster in subsurface soils (15–120 cm). Spatial patterns of TP strongly resembled those of vegetation cover throughout the soil profile, but differed from those of SOC and TN, especially in subsurface soils. The encroachment of woody species dominated by N₂‐fixing trees into this P‐limited ecosystem resulted in the accumulation of proportionally less soil P compared to C and N in surface soils; however, proportionally more P accrued in deeper portions of the soil profile beneath woody patches where alkaline soil pH and high carbonate concentrations would favor precipitation of P as relatively insoluble calcium phosphates. This imbalanced relationship highlights that the relative importance of biotic vs. abiotic mechanisms controlling C and N vs. P accumulation following vegetation change may vary with depth. Our findings suggest that efforts to incorporate effects of land cover changes into coupled climate–biogeochemical models should attempt to represent C‐N‐P imbalances that may arise following vegetation change.     

Can stable carbon isotopes (δ13C) in soil carbon be used to describe the dynamics of Eucalyptus savanna–rainforest boundaries in the Australian monsoon tropics?

The history of isolated patches of monsoon rainforest within large tracts of Eucalyptus savanna is poorly understood because of the scarcity of reliable palaeoecological records in the Australian monsoon tropics. Elsewhere in the world, the ratio of the stable isotopes ¹³C to ¹²C (δ¹³C) in soil organic matter has shed light on the dynamics of rainforest–savanna boundaries because tropical grasses with the C4 photosynthetic pathway have a distinct δ¹³C signature (–17 to –9‰) compared with that of woody plants with the C3 photosynthetic pathway (–32 to –22‰). In order to determine the magnitude of the variation in δ¹³C, unreplicated soil profiles were sampled beneath different vegetation types on three boundaries between Eucalyptus savanna and rainforest that were both growing on Tertiary age laterite parent material. Replicated (n = 3) soil profiles, which were also derived from Tertiary age laterite, were sampled from beneath: (i) dense stands of African grasses within a frequently burnt Eucalyptus savanna; and within the same long unburnt Eucalyptus savanna, (ii) patches of African and natives grasses and (iii) clumps of Acacia trees. The strongly negative δ¹³C values of soil organic matter derived from the frequently burnt and long unburnt grassy understoreys in the Eucalyptus savannas showed that a considerable amount of the soil carbon was derived from C3 (woody) species despite the presence of a ground layer dominated by C4 grasses. However, a feature of these data was the considerable variability among the three ‘replicate’ profiles. The surface soil samples from beneath three clumps of Acacia trees in the unburnt Eucalyptus savanna had much less variable δ¹³C values and were similar to two of the three monsoon rainforests sampled. The pattern of δ¹³C values from unreplicated soil profiles from different vegetation types across three rainforest boundaries was also very variable and not always obviously related the known disturbance history of the extant vegetation. Given the considerable variability within and between vegetation types with contrasting disturbance histories, it is concluded that the use of carbon stable isotopes to advance understanding of the dynamics of rainforest and Eucalyptus savanna boundaries will require further development, such as determination of the ¹⁴C age and δ¹³C values of different soil carbon fractions.