Effects of nutrients and shade on tree‐grass interactions in an East African savanna

Abstract. Savanna trees have a multitude of positive and negative effects on understorey grass production, but little is known about how these effects interact. We report on a fertilization and shading experiment carried out in a Tanzanian tropical dry savanna around Acacia tortilis trees. In two years of study there was no difference in grass production under tree canopies or in open grassland. Fertilization, however, indicate that trees do affect the nutrient limitation of the grass layer with an N‐limited system in open grassland to a P‐limited system under the trees. The N:P ratios of grass gave a reliable indication of the nature of nutrient limitation, but only when assessed at the end of the wet season. Mid‐wet season nutrient concentrations of grasses were higher under than outside the tree canopy, suggesting that factors other than nutrients limit grass production. A shading experiment indicated that light may be such a limiting factor during the wet season when water and nutrients are sufficiently available. However, in the dry season when water is scarce, the effect of shade on plant production became positive. We conclude that whether trees increase or decrease production of the herbaceous layer depends on how positive effects (increased soil fertility) and negative effects (shade and soil water availability) interact and that these interactions may significantly change between wet and dry seasons.     

Positive versus negative environmental impacts of tree encroachment in South Africa

Woody plant encroachment in grasslands is a worldwide phenomenon. Despite many studies, the consequences of woody plant encroachment on sub-canopy vegetation and soil properties are still unclear. To better understand the impacts of trees on grassland properties we examined the following questions using a mountainous sub-tropical grassland of South Africa encroached by an indigenous tree, Acacia sieberiana as a case study: (1) Do trees increase sub-canopy herbaceous diversity, quality and biomass and soil nitrogen content? (2) Do large trees have a stronger effect than medium-sized trees on grass and soil properties? (3) Does the impact of trees change with the presence of livestock and position of trees in a catena? We studied grass and non-graminoid species diversity and biomass, grass quality and soil properties during the wet season of 2009. Nitrogen in grass leaves, soil cation exchange capacity and calcium and magnesium ion concentrations in the soil increased under tall Acacia versus open areas. Medium-sized Acacia decreased the gross energy content, digestibility and neutral detergent fibre of grasses but increased the species richness of non-graminoids. Tall and medium Acacia trees were associated with the presence of Senecio inaequidens, an indigenous species that is toxic to horses and cattle. The presence of livestock resulted in a decrease in herbaceous root biomass and an increase in soil carbon and leaf biomass of grass under Acacia. Tree position in the catena did not modify the impact of trees on the herbaceous layer and soil properties. For management of livestock we recommend retaining tall Acacia trees and partially removing medium-sized Acacia trees because the latter had negative effects on grass quality.     

Impacts of savanna trees on forage quality for a large African herbivore

Recently, cover of large trees in African savannas has rapidly declined due to elephant pressure, frequent fires and charcoal production. The reduction in large trees could have consequences for large herbivores through a change in forage quality. In Tarangire National Park, in Northern Tanzania, we studied the impact of large savanna trees on forage quality for wildebeest by collecting samples of dominant grass species in open grassland and under and around large Acacia tortilis trees. Grasses growing under trees had a much higher forage quality than grasses from the open field indicated by a more favourable leaf/stem ratio and higher protein and lower fibre concentrations. Analysing the grass leaf data with a linear programming model indicated that large savanna trees could be essential for the survival of wildebeest, the dominant herbivore in Tarangire. Due to the high fibre content and low nutrient and protein concentrations of grasses from the open field, maximum fibre intake is reached before nutrient requirements are satisfied. All requirements can only be satisfied by combining forage from open grassland with either forage from under or around tree canopies. Forage quality was also higher around dead trees than in the open field. So forage quality does not reduce immediately after trees die which explains why negative effects of reduced tree numbers probably go initially unnoticed. In conclusion our results suggest that continued destruction of large trees could affect future numbers of large herbivores in African savannas and better protection of large trees is probably necessary to sustain high animal densities in these ecosystems.     

Acacia trees as keystone species in Negev desert ecosystems

Abstract. The only trees in most of the Negev desert are 3 native Acacia species. We tested the hypothesis that they act as keystone species as a result of the improved soil conditions under their canopies. Furthermore, because many Acacia populations suffer high levels of mortality due to water stress, we tested whether trees in high mortality populations had diminished effects on plant species and soil quality under their canopies. We show that plant species diversity beneath the tree canopies is higher than in the surrounding areas. There was also a clearly identifiable suite of species with higher occurrence under the trees. Plant species composition differed significantly between high and low mortality sites. However, there was higher species diversity in high mortality sites and under trees with higher water stress. Soil nutrient content was higher under the trees than in the open areas, especially under larger trees and trees with higher water status. The results indicate that there is a combination of positive and negative effects of Acacia trees on the under‐canopy environment, which may include positive effects of higher soil nutrients and a negative influence of higher soil salinity.     

Oak canopy effects on the distribution patterns of two annual grasses: the role of competition and soil nutrients

Within the oak woodlands of California there is often a distinct shift in the botanical composition between the open grassland and the herbaceous understory beneath oak canopy. Botanical sampling at two woodland sites indicated that the annual grass Bromus diandrus was dominant under deciduous blue oak canopy, while a congener, Bromus hordeaceus, was dominant in open grassland. We examined the relative importance of congeneric competition and edaphic factors in creating these differences in species distribution in two separate field experiments that manipulated both congeneric and intraspecific competition, as well as soil type. We used the demographic measure of relative reproductive rate as an index of population growth. In general, demographic performance correctly predicted the distribution of the two annual grasses in the field. Our results indicate that reduced abundance of B. hordeaceus under canopy reflects the negative effects of competition with B. diandrus. In contrast, B. diandrus is little affected by competition from B. hordeaceus. The reduced abundance of B. diandrus in open grassland may result, in part, from its inability to adapt as well as B. hordeaceus to lower nutrient availability in soils of the open grassland.     

Combined effects between temporal heterogeneity of water supply,nutrient level,and population density on biomass of four broadly distributed herbaceous species

Temporal heterogeneity of water supply affects grassland community productivity and it can interact with nutrient level and intraspecific competition. To understand community responses, the responses of individual species to water heterogeneity must be evaluated while considering the interactions of this heterogeneity with nutrient levels and population density. We compared responses of four herbaceous species grown in monocultures to various combinations of water heterogeneity, nutrient level, and population density: two grasses (Cynodon dactylon and Lolium perenne), a forb (Artemisia princeps), and a legume (Trifolium repens). Treatment effects on shoot and root biomass were analyzed. In all four species, shoot biomass was larger under homogeneous than under heterogeneous water supply. Shoot responses of L. perenne tended to be greater at high nutrient levels. Although root biomass was also larger under homogeneous water supply, effects of water heterogeneity on root biomass were not significant in the grasses. Trifolium repens showed marked root responses, particularly at high population density. Although greater shoot and root growth under homogeneous water supply appears to be a general trend among herbaceous species, our results suggested differences among species could be found in the degree of response to water heterogeneity and its interactions with nutrient level and intraspecific competition.     

Anthropogenic effects on population structure of Acacia tortilis subsp. raddiana along a gradient of water availability in South Sinai,Egypt

Acacia trees in Sinai desert are suffering population decline. This study aims to explore this assumption and to determine the effects of grazing and urbanization on size structure of Acacia tortilis subsp. raddiana along a gradient of water availability in Wadi Feiran basin, South Sinai, Egypt. Size structure of 289 Acacia trees in thirteen isolated populations was analysed. Catchment area and lineament density as water availability indicators were calculated. Effects of grazing and urbanization on Acacia populations were evaluated. A strong positive correlation was recognized between tree height, crown diameter and trunk circumference. Acacia tree density and crown diameter are positively correlated with water availability indicators. Population structure of A. tortilis is characterized by absence of juveniles, clear reduction in numbers of small and large categories and increasing in medium categories. This pattern indicates very low recruitment and high mortality of small and large trees. The variation in water availability may have considerable effects on the structure of A. tortilis. Overgrazing, cutting and urbanization are the main causes of population decline of Acacia in Sinai desert. In addition, drought conditions and bruchid seed beetles aggravate the anthropogenic effect on recruitment of Acacia in South Sinai.     

Soil type, microsite, and herbivory influence growth and survival of Schinus molle (Peruvian pepper tree) invading semi-arid African savanna

Naturalization of Schinus molle (Anacardiaceae) has been observed in semi arid savanna of the Northern Cape Province of South Africa. However, with high dispersal ability, the species is expected to achieve greater densities and invade more widely. The study involved a field manipulation experiment over 14 months using a factorial block design to examine transplanted seedlings in different savanna environments. The experiments examine the effects of soil type (sandy and clay), microsite, and herbivores on seedling performance (establishment, growth and survival). Seedlings were grown in a greenhouse and individually transplanted into four treatment groups: in open grassland, under tree canopies, and with and without cages to exclude large herbivores (cattle and game). The same experiment was repeated in two different soil types: coarse sand and fine-textured clay soil. Results suggest that protection provided by canopies of large indigenous Acacia trees facilitates S. molle invasion into semi-arid savanna. In the field, S. molle seedlings performed considerably better beneath canopies of indigenous Acacia trees than in open areas regardless of soil type. Whether exposed or protected from large herbivores, no seedlings planted in open grassland survived the first winter. Although, seedlings grew better and had higher survival rates beneath tree canopies than in the open sites, exposure to large herbivores significantly decreased heights and canopy areas of seedlings compared with those protected from large herbivores. The effect was greater on clay soil than on sandy soil. The results suggest that low temperature (frost), and possibly inter-specific competition with grasses, may limit S. molle seedling establishment, survival and growth away from tree canopies in semi arid savannas. Low soil nutrient status and browsing may also delay growth and development of this species. The invasive potential of S. molle is thus greatest on fertile soils where sub-canopy microsites are present and browsing mammals are absent.     

Influence of tree cover on herbaceous layer development and carbon and water fluxes in a Portuguese cork-oak woodland

Facilitation and competition between different vegetation layers may have a large impact on small-scale vegetation development. We propose that this should not only influence overall herbaceous layer yield but also species distribution and understory longevity, and hence the ecosystems carbon uptake capacity especially during spring. We analyzed the effects of trees on microclimate and soil properties (water and nitrate content) as well as the development of an herbaceous community layer regarding species composition, aboveground biomass and net water and carbon fluxes in a cork-oak woodland in Portugal, between April and November 2011.The presence of trees caused a significant reduction in photosynthetic active radiation of 35 mol m⁻² d⁻¹ and in soil temperature of 5 °C from April to October. At the same time differences in species composition between experimental plots located in open areas and directly below trees could be observed: species composition and abundance of functional groups became increasingly different between locations from mid April onwards. During late spring drought adapted native forbs had significantly higher cover and biomass in the open area while cover and biomass of grasses and nitrogen fixing forbs was highest under the trees. Further, evapotranspiration and net carbon exchange decreased significantly stronger under the tree crowns compared to the open during late spring and the die back of herbaceous plants occurred earlier and faster under trees. This was most likely caused by interspecific competition for water between trees and herbaceous plants, despite the more favorable microclimate conditions under the trees during the onset of summer drought.     

Below-ground competition between trees and grasses may overwhelm the facilitative effects of hydraulic lift

Under large East African Acacia trees, which were known to show hydraulic lift, we experimentally tested whether tree roots facilitate grass production or compete with grasses for below‐ground resources. Prevention of tree–grass interactions through root trenching led to increased soil water content indicating that trees took up more water from the topsoil than they exuded via hydraulic lift. Biomass was higher in trenched plots compared to controls probably because of reduced competition for water. Stable isotope analyses of plant and source water showed that grasses which competed with trees used a greater proportion of deep water compared with grasses in trenched plots. Grasses therefore used hydraulically lifted water provided by trees, or took up deep soil water directly by growing deeper roots when competition with trees occurred. We conclude that any facilitative effect of hydraulic lift for neighbouring species may easily be overwhelmed by water competition in (semi‐) arid regions.     

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.     

Interactions of Acacia raddiana with herbaceous vegetation change with intensity of abiotic stress

Theoretical models predict that the relative importance of facilitation and competition may vary inversely across gradients of abiotic stress. However, these predictions have not been thoroughly tested in the field, especially in arid and semi-arid environments. In this study, we evaluated how the net effect of Acacia tortilis subsp. raddiana (syn. A. raddiana) trees on the herbaceous species varies across a gradient of water stress in arid Tunisian ecosystems. Our results show that the influence of trees on the herbaceous stratum is beneficial. An Acacia tree improves the richness of herbaceous species around it by two to three times. The positive effect of Acacia raddiana on species composition is characterised in particular by a better development of some species of high pastoral value such as Cenchrusciliaris, Cynodon dactylon, Eragrostis papposa, Sisymbrium irio and Chenopodium album. We fitted the relationship between seasonal rainfall and the relative neighbour effect index, which varied across this gradient, to a linear model. Our results show that herbaceous plant cover, plant biomass, plant density and diversity are higher under a tree canopy and this positive interaction still increase with higher abiotic stress conditions.     

Effect of woody vegetation clearing on nutrient and carbon relations of semi-arid dystrophic savanna

Throughout the savanna biome, woody vegetation is cleared to increase productivity of herbaceous pasture. While clearing can result in increased pasture production of semi-arid dystrophic savannas in the short term, it is uncertain whether production is sustained in the long term. There is insufficient knowledge of how clearing affects soil nutrient and organic carbon (SOC) stocks. Using cleared-uncleared site pairs, we evaluated techniques for time-integrated assessment of nutrient and carbon relations in Australian savanna. Short-term in situ resin incubation showed that soil at cleared sites had a higher time-integrated availability of ammonium and nitrate, indicating that nitrogen (N) may turn over faster and/or is taken up slower at cleared sites than uncleared savanna. Nitrate and ammonium availability was approximately 2-fold higher in spring than in summer, likely due to greater uptake and/or loss of nitrate during summer rains. Nitrate was a prominent N source for evergreen trees, especially before summer rain, pointing to a role of trees as permanent N sinks. Stable isotope signatures of soil and vegetation indicate that N input occurs via N₂ fixing microbiotic crusts and Acacia species. 30 years after clearing, SOC contained more C₄ grass-derived carbon than uncleared savanna, but this shift in C source was not associated with the net C gain often observed in grasslands. Interactions between altered nutrient and C relations and composition of the understorey should be assessed in context of introduced buffelgrass (Cenchrus ciliaris) which had higher macronutrient concentrations than native grasses. Heterogeneity of the studied soils highlights the need for replication at several spatial scales to infer long-term dynamics with space-for-time chronosequences. We conclude that the techniques presented here are useful for gaining knowledge of the biogeochemical processes governing savannas and the systems that result from clearing.     

The role of microsite sunlight environment on growth,architecture, and resource allocation in dominant Acacia tree seedlings,in Serengeti,East Africa

Seedling establishment is a critical life history stage for savanna tree recruitment due to variability in resource availability. While tree–grass competition for water is recognized as an important driver of tree seedling mortality, the importance of sunlight exposure on tree seedling performance has received little attention in savanna ecosystems despite variable seedling light environments caused by heterogeneity in biomass of the grass canopy. We studied the seasonal sunlight micro-environment for two dominant East African tree species (Acacia?=?Vachellia) robusta (Burch) and A. tortilis (Forssk) under natural field conditions. In the Serengeti National Park, Tanzania, A. robusta trees occur in tall grasslands of the north (shady) and A. tortilis in the southern short grasslands (less shaded). We also designed a greenhouse experiment to quantify sunlight effects on seedling growth, architecture, and resource allocation traits. In the field, A. robusta seedlings were associated with lower understorey sunlight during the wet season compared to A. tortilis, with this trend switching during the dry season. In the greenhouse experiments, under low sunlight (25% radiation), A. robusta gained height faster than A. tortilis and self-shading among canopy leaves was evident in A. tortilis but not A. robusta. Biomass allocation to leaves, stems, and roots differed between species under different light environments suggesting phenotypic plasticity in response to variable light availability. Our study suggests that microsite light variability should be incorporated in models of the spatial and temporal variability of savanna tree recruitment.

     

Balance of Competitive and Facilitative Effects of Exotic Trees on a Native Patagonian Grass

The balance between facilitation and competition in plants changes with species characteristics and environmental conditions. Facilitative effects are common in natural ecosystems, particularly in stressful environments or years. Contrarily, in artificial associations of plants, such as agroforestry systems, some authors have suggested that even when facilitative effects may occur, net balance of tree effects on grasses is usually negative, particularly in dry environments. The aim of this study was to determine the net effect of the exotic ponderosa pine on the native grass Festuca pallescens (St. Ives) Parodi in agroforestry systems in Patagonia. Soil water content, plant water status, and relative growth were measured in the grass growing in different treatments (determined by tree cover level) during two growing seasons with contrasting climatic conditions. Facilitative effects of trees over grass water status were recorded only when water availability was high. A net negative effect was detected on dates when soil water content was very low and evaporative demand was high. The strength of these negative effects depended on tree density and climatic conditions, being higher in treatments with lower tree canopy cover. These results indicate that the positive effect of trees could only be expected under relatively low stress conditions. However, relative growth of grasses was always similar in plants growing in forested plots than in open grassland. Differences in biomass allocation for grasses growing in shade and open habitats may reconcile these contrary results. Our results highlight the importance of the physiology of a species (relative drought and shade tolerance) in determining the response of a plant to a particular interacting species.     

Trees improve grass quality for herbivores in African savannas

The tree–grass interactions of African savannas are mainly determined by varying rainfall patterns and soil fertility. Large savanna trees are known to modify soil nutrient conditions, but whether this has an impact on the quality of herbaceous vegetation is unclear. However, if this were the case, then the removal of trees might also affect the structure and quality of the grass layer. We studied the impact of large nitrogen- and non-nitrogen fixing trees on the sub-canopy (SC) grass layer in low- and high-rainfall areas of differing soil fertility in eastern and southern Africa. We compared the structure and nutrient levels of SC grasses with those outside the canopy. Grass leaf nitrogen and phosphorus contents beneath tree canopies were elevated at all study sites and were up to 25% higher than those outside the canopy in the site of lowest rainfall and soil fertility. Grass leaf fibre and organic matter (OM) contents were slightly enhanced beneath tree canopies. At the site of highest rainfall and soil fertility, grasses beneath the canopy had significantly lower ratios of stem:leaf biomass and dead:living leaf material. Grass species composition differed significantly, with the highly nutritious Panicum spp. being most abundant underneath tree crowns. In the two drier study sites, soil nitrogen and OM contents were enhanced by 30% beneath trees. N-fixation capacity of trees did not contribute to the improved quality of grass under the canopy. We conclude that trees improve grass quality, especially in dry savannas. In otherwise nutrient-poor savanna grasslands, the greater abundance of high-quality grass species with higher contents of N and P and favourable grass structure beneath trees could attract grazing ungulates. As these benefits may be lost with tree clearance, trees should be protected in low fertility savannas and their benefits for grazing wildlife recognised in conservation strategies.     

Savanna tree influence on understory vegetation and soil nutrients in northwestern Kenya

Abstract. Contrary to observations and models in which trees and herbaceous plants are viewed as competitors, we found that trees in an African savanna have positive impacts on herbaceous biomass production and composition, and on soil nutrient status. In the Turkana District of northwestern Kenya, we investigated vegetation and soil gradients along equi-angular transects radiating from the boles of individual Acacia tortilis trees. Total herbaceous biomass averaged 260 ± 17(se) g/m² at the bole and declined to 95 ± 8 g/m² in the tree interspaces. Soil organic carbon and total nitrogen concentrations were greatest (0.72 % and 0.083 %, respectively) in shallow soils near the bole and declined rapidly toward the interspaces and with increasing depth. Transects were also established between tree pairs to assess effects of differential canopy proximities. Grass production averaged 220 ± 21 g / m² below overlapping canopies, 150 ± 15 g / m² under individual canopies, and 95 ± 8 g / m² in interstitial areas. Detrended correspondence analysis revealed that shifts in species composition were correlated with distance from tree bole out to the edge of the canopy. Species response, in terms of relative cover, to increasing distance from the bole, seemed to fall into five general classes: 1) greatest at the bole, 2) increasing with distance from the bole, 3) greatest in the mid canopy zone, 4) least at the bole and 5) no response. Trees did not influence herbaceous compositionbeyondtree canopies. It is assumed that shade cast by the tree canopy with subsequent reductions of understory water stress and temperature and increased nutrient concentrations may be the most important factors affecting understory soil and vegetation.     

Abundance of trees and grasses in a woodland savanna in relation to environmental factors

The relationships of plant species associations and underlying environmental factors in a woodland savanna in South Africa were investigated. 40 plots were included with 25 tree and 17 grass species dominating the arboreal and ground cover. Correspondence Analysis described the relationships between soil moisture retention, soil nutrients and the abundance of trees and grasses. Dry matter indices represented the accumulated effects of rainfall, fire and grazing of the herbaceous layer. Variations in the abundance of plants corresponded to well-defined gradients of soil nutrients. The distribution of grass and tree species along the ordination axes indicated that soils with high water retention capacity and high nutrient contents provided a suitable substrate for many of the tree species sampled. However, grass species abundance was high in plots with porous soils and poor nutrient availability.     

Effect of competition on stable carbon isotope ratios of two tussock grass species

Summary Previous studies have shown that plant carbon isotope composition varies when plants experience differences in water and nutrient availability. However, none have addressed the effect of root interactions, including competition for these soil resources, on carbon isotope ratios. We studied the effect of interspecific root interactions on the productivity and carbon isotope ratios of two Great Basin tussock grass species (Agropyron desertorum and Pseudoroegneria spicata). We compared grasses grown in mixture with sagebrush (Artemisia tridentara) to grasses in similar mixtures but where root interactions with sagebrush were limited by fiberglass partitions. During both years of the study, tussocks growing in competition with sagebrush produced tissue with more negative ¹³C values than grasses experiencing limited root interaction with sagebrush. The magnitude of this difference (0.5 to 0.9%) is similar to that found in other studies when soil fertility and moisture availability were altered.     

Effects and mechanism of plant litter on grassland ecosystem: A review

Plant litter is dead, above and below ground; organic material i.e. leaves barks, needles, twigs and roots. Plant litter plays a key role in nutrient cycling and community organization in grassland ecosystems. Litter can have important consequences on recruitment of plant species through modification of biological, physical, and chemical features of microenvironment. Plant litter offers a major input of organic matter to the soil which modifies soil chemistry, hence impacts nutrient cycling. At early stages of litter decomposition, a particular amount of carbon is transporting to the soil nutrient pool. In terrestrial ecosystems, plant litter regulating biogeochemical cycles, maintain soil fertility, nutrient availability, and therefore influence plant growth, diversity, composition, structure, and productivity. Litter can also impact plant above net plant productivity and below net plant productivity in grassland ecosystem. Plant litter accumulation and decomposition can impact plant species composition and community structure through temperature, light and nutrient availability. The effects of plant litter on vegetation may be negative, positive or neutral due vegetation variability, study duration, habitat, latitude, quantity and quality of litter. These diverse effects of plant litter on grassland ecosystem might be due to, management practice type, management intensity, climate type, timing, precipitation and soil nutrient pool etc. Current review attempts to describe prominent effects of plant litter on vegetation, seed germination, soil fertility, Productivity, species composition, community structure and mechanism in grassland ecosystem.