• Mediterranean‐type ecosystems contain 20% of all vascular plant diversity on Earth and have been identified as being particularly threatened by future increases in drought. Of particular concern is ...the Cape Floral Region of South Africa, a global biodiversity hotspot, yet there are limited experimental data to validate predicted impacts on the flora. In a field rainout experiment, we tested whether rooting depth and degree of isohydry or anisohydry could aid in the functional classification of drought responses across diverse growth forms. • We imposed a 6‐month summer drought, for 2 yr, in a mountain fynbos shrubland. We monitored a suite of parameters, from physiological traits to morphological outcomes, in seven species comprising the three dominant growth forms (deep‐rooted proteoid shrubs, shallow‐rooted ericoid shrubs and graminoid restioids). • There was considerable variation in drought response both between and within the growth forms. The shallow‐rooted, anisohydric ericoid shrubs all suffered considerable reductions in growth and flowering and increased mortality. By contrast, the shallow‐rooted, isohydric restioids and deep‐rooted, isohydric proteoid shrubs were largely unaffected by the drought. • Rooting depth and degree of iso/anisohydry allow a first‐order functional classification of drought response pathways in this flora. Consideration of additional traits would further refine this approach.
The cool season rainfall at our study site should favour C
rather than C
grasses. There are, however, several locations where C
grasses have become dominant, suggesting that rainfall seasonality is ...not a constraint on distribution. Here, we explored the limitations on C
grass distribution in a fynbos shrubland. Using δ
C values of SOM, we determined when these grasses had established. We also looked at the role of roads as conduits for establishment and asked what impact these grasses may have on fynbos species richness. We then conducted a field experiment designed to examine the extent to which soil moisture, nutrient availability, and competition with fynbos for resources influence the establishment and growth of the grasses. Finally using aerial photography, we explored the role of changes in land use on distribution. Our results showed that the establishment is recent, that roads may be acting as conduits, and that with establishment, there is a reduction in fynbos species richness and diversity. Disturbance and removing below-ground competition for resources open the system to establishment in wetter areas. This study is the first to look at the potential for C
grasses expanding into cool season rainfall shrublands such as in Western Australia and South Africa. What is interesting about these results is that C
grasses can establish and dominate in a cool season rainfall regime. Rather than temperature of the growing season, it is competition for resources from fynbos that prevents these grasses from encroaching.
The cool season rainfall at our study site should favour C
3
rather than C
4
grasses. There are, however, several locations where C
4
grasses have become dominant, suggesting that rainfall ...seasonality is not a constraint on distribution. Here, we explored the limitations on C
4
grass distribution in a fynbos shrubland. Using δ
13
C values of SOM, we determined when these grasses had established. We also looked at the role of roads as conduits for establishment and asked what impact these grasses may have on fynbos species richness. We then conducted a field experiment designed to examine the extent to which soil moisture, nutrient availability, and competition with fynbos for resources influence the establishment and growth of the grasses. Finally using aerial photography, we explored the role of changes in land use on distribution. Our results showed that the establishment is recent, that roads may be acting as conduits, and that with establishment, there is a reduction in fynbos species richness and diversity. Disturbance and removing below-ground competition for resources open the system to establishment in wetter areas. This study is the first to look at the potential for C
4
grasses expanding into cool season rainfall shrublands such as in Western Australia and South Africa. What is interesting about these results is that C
4
grasses can establish and dominate in a cool season rainfall regime. Rather than temperature of the growing season, it is competition for resources from fynbos that prevents these grasses from encroaching.
Abstract
Woody plant encroachment (WPE) is a global trend that occurs in many biomes, including savannas, and accelerates with fire suppression. Since WPE can result in increased storage of soil ...organic carbon (SOC), fire management, which may include fire suppression, can improve ecosystem carbon (C) sequestration in savannas.
At our study site in Kruger National Park, South Africa, we used a long‐term (~70 year) fire experiment to study the drivers and consequences of changes in woody cover (trees and shrubs) on SOC sequestration. We surveyed four fire manipulation treatments, replicated at eight locations within the park: annual high‐intensity burns, triennial high (dry season) and low‐intensity (wet season) burns, and fire exclusion, to capture the range of fire management scenarios under consideration. The changes in woody cover were calculated over a period similar to the experiment's duration (~80 years) using aerial photographs (1944–2018). Soils were analysed to 30 cm depth for SOC and δ
13
C, under and away from the tree canopy to isolate local‐ and landscape‐level effects of WPE on SOC.
The largest increases in woody cover occurred with fire exclusion. We found that plots with higher increases in woody cover also had higher SOC. However, trees were not the only contributor to SOC gains, sustained high inputs of C
4
‐derived C (grasses), even under canopies in fire suppression plots, contributed significantly to SOC. We observed little difference in SOC sequestration between cooler triennial (wet season) burns and fire suppression.
Synthesis
. Grass input to soil organic carbon (SOC) remained high across the full range of woody cover created by varying burning regimes. The total SOC stocks stored from tree input only matched grass‐derived SOC stocks after almost 70 years of fire exclusion. Our results point to C
4
grasses as a resilient contributor to SOC under altered fire regimes and further challenge the assumption that increasing tree cover, either through afforestation schemes or fire suppression, will result in large gains in C sequestration in savanna soils, even after 70 years.
Savanna is a species-rich biome, that includes many modern mammal lineages and C₄ grass (Poaceae) species. The greater productivity and grazing pressure associated with savannas is likely ...attributable to the foliar traits of the grasses they support. Thus, it is important to understand the complex relationships between the abiotic environment, foliar attributes and the floristic composition of savanna grasses, and the supported grazer densities. We sampled 37 grass communities in the Kruger National Park (South Africa) across three soil types and along a rainfall gradient and found that these communities lack strong phylogenetic structure. We then measured specific leaf area and leaf tensile strength for 384 individuals representing 66 species and found that both traits were strongly phylogenetically structured and associated with both rainfall and soil type. Finally, we found that grazer densities in the Park are correlated with the foliar traits of the associated grass communities, but the resolution of our data do not allow for a thorough analysis of this association. Our results demonstrate the complex interactions between climate, soils and grazer densities relative to C₄ grass functional traits.
Here we describe the fine root distribution of trees and grasses relative to soil nitrogen and water profiles. The primary objective is to improve our understanding of edaphic processes influencing ...the relative abundance of trees and grasses in savanna systems. We do this at both a mesic (737
mm MAP) site on sandy-loam soils and at an arid (547
mm MAP) site on clay rich soils in the Kruger National Park in South Africa. The proportion of tree and grass fine roots at each soil depth were estimated using the
δ
13C values of fine roots and the
δ
13C end members of the fine roots of the dominant trees and grasses at our study sites. Changes in soil nitrogen concentrations with depth were indexed using total soil nitrogen concentrations and soil
δ
15N values. Soil water content was measured at different depths using capacitance probes. We show that most tree and grass roots are located in the upper layers of the soil and that both tree and grass roots are present at the bottom of the profile. We demonstrate that root density is positively related to the distribution of soil nitrogen and negatively related to soil moisture. We attribute the negative correlation with soil moisture to evaporation from the soil surface and uptake by roots. Our data is a snapshot of a dynamic process, here the picture it provides is potentially misleading. To understand whether roots in this system are primarily foraging for water or for nitrogen future studies need to include a dynamic component.
Both resource and disturbance controls have been invoked to explain tree persistence among grasses in savannas. Here we determine the extent to which competition for available resources restricts the ...rooting depth of both grasses and trees, and how this may influence nutrient cycling under an infrequently burned savanna near Darwin, Australia. We sampled fine roots <2 mm in diameter from 24 soil pits under perennial as well as annual grasses and three levels of canopy cover. The relative proportion of C3 (trees) and C4 (grasses) derived carbon in a sample was determined using mass balance calculations. Our results show that regardless of the type of grass both tree and grass roots are concentrated in the top 20 cm of the soil. While trees have greater root production and contribute more fine root biomass grass roots contribute a disproportional amount of nitrogen and carbon to the soil relative to total root biomass. We postulate that grasses maintain soil nutrient pools and provide biomass for regular fires that prevent forest trees from establishing while savanna trees, are important for increasing soil N content, cycling and mineralization rates. We put forward our ideas as a hypothesis of resource‐regulated tree–grass coexistence in tropical savannas.
Conifer populations appear disproportionately threatened by global change. Most examples are, however, drawn from the northern hemisphere and long-term rates of population decline are not well ...documented as historical data are often lacking. We use a large and long-term (1931-2013) repeat photography dataset together with environmental data and fire records to account for the decline of the critically endangered Widdringtonia cedarbergensis. Eighty-seven historical and repeat photo-pairs were analysed to establish 20th century changes in W. cedarbergensis demography. A generalized linear mixed-effects model was fitted to determine the relative importance of environmental factors and fire-return interval on mortality for the species.
From an initial total of 1313 live trees in historical photographs, 74% had died and only 44 (3.4%) had recruited in the repeat photographs, leaving 387 live individuals. Juveniles (mature adults) had decreased (increased) from 27% (73%) to 8% (92%) over the intervening period. Our model demonstrates that mortality is related to greater fire frequency, higher temperatures, lower elevations, less rocky habitats and aspect (i.e. east-facing slopes had the least mortality).
Our results show that W. cedarbergensis populations have declined significantly over the recorded period, with a pronounced decline in the last 30 years. Individuals that established in open habitats at lower, hotter elevations and experienced a greater fire frequency appear to be more vulnerable to mortality than individuals growing within protected, rocky environments at higher, cooler locations with less frequent fires. Climate models predict increasing temperatures for our study area (and likely increases in wildfires). If these predictions are realised, further declines in the species can be expected. Urgent management interventions, including seedling out-planting in fire-protected high elevation sites, reducing fire frequency in higher elevation populations, and assisted migration, should be considered.
In this study, we explored how rainfall manipulation influenced competitive interactions between grasses and juvenile trees (small nonreproductive trees capable of resprouting) in savanna. To do ...this, we manipulated rainfall amount in the field using an incomplete factorial experiment that determined the effects of rainfall reduction, no manipulation, rainfall addition, and competition between grasses and trees on grass and tree growth. As response variables, we focused on several measures of tree growth and Disc Pasture Meter settling height as an estimate of grass aboveground biomass. We conducted the study over four years, at two sites in the Kruger National Park, South Africa. Our results show that rainfall manipulation did not have substantial effects on any of the measures of tree growth we considered. However, trees at plots where grasses had been removed grew on average 15 cm more in height and 1.3-1.7 times more in basal area per year than those in plots with grasses. Grass biomass was not influenced by the presence of trees but was significantly and positively influenced by rainfall addition. These findings were not fundamentally influenced by soil type or by prevailing precipitation, suggesting applicability of our results to a wide range of savannas. Our results suggest that, in savannas, increasing rainfall serves to increase the competitive pressure exerted by grasses on trees. The implication is that recruitment into the adult tree stage from the juvenile stage is most likely in drought years when there is little competition from grass for resources and grass fuel loads are low.
Stable isotopes in water (δ2H and δ18O) are important indicators of hydrological and ecological pattern and process. δ2H and δ18O of water are incorporated into geological and biological systems in a ...predictable manner and have been used extensively as tracers in hydrological, ecological and forensic studies. Physical processes result in spatial variation of δ2H, δ18O in water across the landscape (so-called “isoscapes”) and provide the basis for hydrological, ecological, archaeological and forensic studies. Southern Africa is a globally important meeting point for ocean and climate systems, biological diversity and human societies, yet there is little information on the spatial variability of δ2H and δ18O in water across this important region. Here we present the first ground water and tap water isoscapes for southern Africa. We compare and contrast these two water resources, and consider how well global models of precipitation isotopes capture isotopic variation across South Africa. Ground water and tap water samples were collected from across South Africa, analysed for δ2H and δ18O, and used to generate interpolated δ2H, δ18O and deuterium-excess (d=δ2H – 8*δ18O) isoscapes. We found coherent spatial structure in δ2H, δ18O and d of ground water and tap water that could be predicted by a geostatistical model based on simple environmental parameters (elevation, mean annual precipitation, precipitation minus potential evaporation, distance to coast and modeled isotope ratio of precipitation). This spatial structure resulted in considerable differences in isotopic composition of water in many of the major wildlife reserves in South Africa, indicating a good potential for wildlife forensics in this region. δ2H and δ18O of ground water, and to a lesser extent tap water, reflected the δ2H and δ18O of long-term weighted annual precipitation at the two GNIP stations in South Africa. However, large discrepancies between modelled isotopic composition of precipitation and our ground water and tap water isoscapes, particularly at higher elevations, highlighted uncertainty in the accuracy of modelled precipitation isoscapes for this region. Increased spatial sampling of precipitation, especially for high elevation regions, and temporal sampling of ground and tap water would considerably aid isotopic studies in this region.
•We present the first ground and tap water isoscapes (δ2H, δ18O) for South Africa.•Coherent spatial structure indicates good potential for wildlife forensics.•We compare ground and tap water with measured and modelled precipitation.•Ground water reflected long-term measured precipitation isotope ratios.•We highlight uncertainty in precipitation isoscapes, particularly at high elevation.
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