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

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

Seasonal variation in top-down and bottom-up processes in a grassland arthropod community

Both top-down and bottom-up processes are common in terrestrial ecosystems, but how these opposing forces interact and vary over time is poorly understood. We tested the variation of these processes over seasonal time in a natural temperate zone grassland, a field site characterized by strong seasonal changes in abiotic and biotic conditions. Separate factorial experiments manipulating nutrients and cursorial spiders were performed in the wet and dry seasons. We also performed a water-addition experiment during the summer (dry season) to determine the degree of water limitation during this time. In the spring, nutrient addition increased plant growth and carnivore abundance, indicating a bottom-up control process. Among herbivores, sap-feeders were significantly enhanced while grazers significantly declined resulting in no net change in herbivore abundance. In the summer, water limitation was predominant increasing plants and all herbivores while nutrient (N) effects were non-significant. Top-down processes were present only in the spring season and only impacted the guild of grazing herbivores. These results show that bottom-up limitation is present throughout the season in this grassland, although the specific limiting resource changes as the season progresses. Bottom-up processes affected all trophic levels and many different guilds, while top-down effects were limited to a select group of herbivores and did not extend to the plant trophic level. Our results show that the relative strengths of top-down and bottom-up processes can shift over relatively short periods of time in habitats with a strong seasonal component.     

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.     

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.     

Interacting effects of pollination,water and nutrients on fruit tree performance

Pollination is critical to fruit production, but the interactions of pollination with plant resources on a plant's reproductive and vegetative features are largely overlooked. We examined the influences of pollination, irrigation and fertilisation on the performance of almond, Prunus dulcis, in northern California. We used a full‐factorial design to test for the effects of pollination limitation on fruit production and foliage variables of whole trees experiencing four resource treatments: (i) normal water and nutrients, (ii) reduced water, (iii) no nutrients, and (iv) reduced water and no nutrients. In each of these combinations, we applied three pollination treatments: hand‐cross pollination, open‐pollination and pollinator exclusion. Pollination strongly affected yield even under reduced water and no nutrient applications. Hand‐cross pollination resulted in over 50% fruit set with small kernels, while open‐pollinated flowers showed over 30% fruit set with moderate‐sized kernels. Pollinator‐excluded flowers had a maximum fruit set of 5%, with big and heavy kernels. Reduced water interacted with the open‐ and hand‐cross pollination treatments, reducing yield more than in the pollinator exclusion treatment. The number of kernels negatively influenced the number of leaves, and reduced water and no nutrient applications interacted with the pollination treatments. Overall, our results indicate that the influences of pollination on fruit tree yield interact with the plant availability of nutrients and water and that excess pollination can reduce fruit quality and the production of leaves for photosynthesis. Such information is critical to understand how pollination influences fruit tree performance.     

Seasonal,interannual and decadal drivers of tree and grass productivity in an Australian tropical savanna

Tree–grass savannas are a widespread biome and are highly valued for their ecosystem services. There is a need to understand the long‐term dynamics and meteorological drivers of both tree and grass productivity separately in order to successfully manage savannas in the future. This study investigated the interannual variability (IAV) of tree and grass gross primary productivity (GPP) by combining a long‐term (15 year) eddy covariance flux record and model estimates of tree and grass GPP inferred from satellite remote sensing. On a seasonal basis, the primary drivers of tree and grass GPP were solar radiation in the wet season and soil moisture in the dry season. On an interannual basis, soil water availability had a positive effect on tree GPP and a negative effect on grass GPP. No linear trend in the tree–grass GPP ratio was observed over the 15‐year study period. However, the tree–grass GPP ratio was correlated with the modes of climate variability, namely the Southern Oscillation Index. This study has provided insight into the long‐term contributions of trees and grasses to savanna productivity, along with their respective meteorological determinants of IAV.     

Dry season watering alters the significance of climate factors influencing phenology and growth of saplings of savanna woody species in central Zambia,southern Africa

Although tree growth in southern African savannas is correlated with rainfall in the wet season, some studies have shown that tree growth is controlled more by rainfall in the dry season. If more rainfall occurred in the dry season in future climates, it would affect the growth of savanna trees, especially saplings that have shallower roots which limit access to subsoil water during the dry season when leaf flush and shoot extension occur. Recent paleobotanical evidence has revealed that there was relatively more precipitation in the dry season in eastern Africa in the Eocene than under the current climate. Saplings therefore can be expected to respond more to water addition during the dry season than mature trees that have more stored water and deeper roots that access subsoil water. Accordingly, I hypothesized that irrigation in the dry season should (i) advance the onset of the growing season, (ii) increase growth rates and (iii) alter the growth responses of saplings to climate factors. To test these hypotheses saplings of five savanna woody species were irrigated during the hot‐dry season at a site in central Zambia and their monthly and annual growth rates compared to those of conspecifics growing under control conditions. Although the responses among the species were variable, all irrigated saplings had significantly higher monthly and annual growth rates than control plants. In addition, dry season watering significantly altered the climatic determinants of sapling growth by either strengthening the role of the same climatic factors that were important under control conditions or displacing them altogether. In conclusion, more precipitation during the hot‐dry season is likely to have significant positive effects on sapling growth and consequently reduce the sapling‐tree transition periods and promote future tree population recruitment in some southern African savanna tree species.     

Selection of shade trees in forest restoration plantings should not be based on crown tree architecture alone

The planting of tree seedlings is a common restoration technique in the tropics, and using large‐crowned, fast‐growing shade species is recommended to suppress invasive grasses and accelerate forest succession. We analyzed the effectiveness of shade species in shading the forest floor during the rainy and dry seasons at young forest restoration sites, whether shade changes according to site for a given species, and whether crown architecture can predict the shade level. We measured the photosynthetically active radiation (PAR) intercepted by the tree crowns of 14 species in two 3‐year‐old restoration plantings. The ability to predict shade based on crown architecture traits was evaluated using multiple linear regressions. The interception of PAR varied according to species, site, and season for seven species and was generally higher during the rainy season. Low values of tree and first branch height and high values of trunk diameter and mean area of a leaf predicted greater light interception. For the dry season, the ability to predict PAR interception was weaker than that for the rainy season and affected by a shorter tree height and a greater crown area. The crown architecture of shade species did not completely predict their shading ability, and the preselection of shade species for forest restoration purposes based only on crown architecture traits is not effective. Therefore, it is important to consider other factors, such as how long trees retain their leaves throughout the year and the soil and management conditions of the sites undergoing restoration, during the selection of species.     

Nutrient supply and bird predation additively control insect herbivory and tree growth in two contrasting forest habitats

It has been suggested that bottom–up and top–down forces interactively control food web dynamics. While top–down effects would increase with resource availability to plants, bottom–up effects would be stronger under low predator abundance. These predictions, however, have rarely been tested at contrasting sites while keeping the dominant plant species unchanged. Furthermore, few studies have factorially manipulated both types of forces in forest communities. For two years, we evaluated the effects of fertiliser (NPK) addition and bird exclusion on tree growth, leaf traits, insect abundance, and folivory rates in a dry/warm and a wet/cold Nothofagus pumilio forest in Patagonia, Argentina. Overall, we found no interaction between nutrient supply and bird predation, although the strength of bottom–up and top–down forces differed markedly between forest sites. Treatment effects were generally weak in the wet forest, where tree growth rates and insect herbivory were low relative to the dry forest. In the dry forest, fertilisation increased sapling growth, insect abundance and folivory, whereas bird exclusion increased leaf damage and reduced tree growth. In the wet forest, fertilisation enhanced leaf nutrient contents and folivore abundance but not sapling growth, while bird exclusion had little impact on insects or trees. These results imply that factors other than nutrients and birds were important in controlling tree growth and folivore activity in the wet forest. While treatment effect sizes varied widely among feeding guilds, in general, nutrient effects on folivores were stronger than predator effects. We conclude that, within the time‐frame of this study, tree growth and herbivory were additively affected by soil nutrients and predator presence, as bird exclusion effects did not change with elevated folivore activity on fertilised trees. We also show that both top–down and bottom–up cascades were weaker in a forest site characterised by slow‐growing juvenile trees subjected to low folivore pressure.     

Non‐Native Grass Removal and Shade Increase Soil Moisture and Seedling Performance during Hawaiian Dry Forest Restoration

Invasive non‐native species can create especially problematic restoration barriers in subtropical and tropical dry forests. Native dry forests in Hawaii presently cover less than 10% of their original area. Many sites that historically supported dry forest are now completely dominated by non‐native species, particularly grasses. Within a grass‐dominated site in leeward Hawaii, we explored the mechanisms by which non‐native Pennisetum setaceum, African fountain grass, limits seedlings of native species. We planted 1,800 seedlings of five native trees, three native shrubs, and two native vines into a factorial field experiment to examine the effects of grass removal (bulldozed vs. clipped plus herbicide vs. control), shade (60% shade vs. full sun), and water (supplemental vs. ambient) on seedling survival, growth, and physiology. Both grass removal and shade independently increased survival and growth, as well as soil moisture. Seedling survival and relative growth rate were also significantly dependent on soil moisture. These results suggest that altering soil moisture may be one of the primary mechanisms by which grasses limit native seedlings. Grass removal increased foliar nitrogen content of seedlings, which resulted in an increase in leaf‐level photosynthesis and intrinsic water use efficiency. Thus in the absence of grasses, native species showed increased productivity and resource acquisition. We conclude that the combination of grass removal and shading may be an effective approach to the restoration of degraded tropical dry forests in Hawaii and other ecologically similar ecosystems.     

遮荫对青桐幼苗生长性状与化学计量特征的影响

营养元素含量及化学计量比可反映植物器官营养元素的分配及互作关系,亦可反映其营养利用效率及生长环境的养分限制状况.以青桐(Firmiana platanifolia)幼苗为材料,分析4种不同光强(全光照,50％、75％和95％遮荫)对青桐幼苗生长,C、N、P、K含量及其化学计量特征的影响.结果 表明:遮荫对青桐幼苗的生长...     

The influence of tree species on canopy soil nutrient status in a tropical lowland wet forest in Costa Rica

The canopy is host to a large percentage of the flora and fauna in tropical wet forests and is distinct from the forest floor in plant richness, soil type and microclimate. In this study, we examined the influence of tree species and season on soil nutrient cycling processes in canopy soils of four tree species common to Costa Rican wet forests. We also compared the canopy soils to the associated forest floor mineral soils. Both tree species and season had strong effects on canopy soil nutrients and processes. Canopy soils from trees with high litter lignin concentrations had higher net N-mineralization rates and higher dissolved inorganic N concentrations than those with low lignin concentrations. During the dry season, net N-immobilization occurred and dissolved organic and inorganic N and available P concentrations were significantly higher than during the wet season. Overall, canopy soils had higher N levels and higher fungi + bacteria richness than forest floor mineral soils. The differences in canopy soil properties observed among tree species indicates that these species have distinct N cycles that reflect differences in both soil origin and biological controls.     

Foliar demand and resource economy of nutrients in dry tropical forest species

Important phenological activities in seasonally dry tropical forest species occur within the hot‐dry period when soil water is limiting, while the subsequent wet period is utilized for carbon accumulation. Leaf emergence and leaf area expansion in most of these tree species precedes the rainy season when the weather is very dry and hot and the soil cannot support nutrient uptake by the plants. The nutrient requirement for leaf expansion during the dry summer period, however, is substantial in these species. We tested the hypothesis that the nutrients withdrawn from the senescing leaves support the emergence and expansion of leaves in dry tropical woody species to a significant extent. We examined the leaf traits (with parameters such as leaf life span, leaf nutrient content and retranslocation of nutrients during senescence) in eight selected tree species in northern India. The concentrations of N, P and K declined in the senescing foliage while those of Na and Ca increased. Time series observations on foliar nutrients indicated a substantial amount of nutrient resorption before senescence and a ‘tight nutrient budgeting’. The resorbed N‐mass could potentially support 50 to 100% and 46 to 80% of the leaf growth in terms of area and weight, respectively, across the eight species studied. Corresponding values for P were 29 to 100% and 20 to 91%, for K 29 to 100% and 20 to 57%, for Na 3 to 100% and 1 to 54%, and for Ca 0 to 32% and 0 to 30%. The species differed significantly with respect to their efficiency in nutrient resorption. Such interspecific differences in leaf nutrient economy enhance the conservative utilization of soil nutrients by the dry forest community. This reflects an adaptational strategy of the species growing on seasonally dry, nutrient‐poor soils as they tend to depend more or less on efficient internal cycling and, thus, utilize the retranslocated nutrients for the production of new foliage biomass in summer when the availability of soil moisture and nutrients is severely limited.     

Modelling ungulate dependence on higher quality forage under large trees in African savannahs

In African savannahs, large trees improve grass quality, particularly in dry and nutrient poor areas. Enhanced below-canopy grass nutrients, such as nitrogen and phosphorus contents should therefore attract and benefit grazers. To predict whether ungulates really need these forage quality islands we focused on four grazer species, i.e., zebra, buffalo, wildebeest, and warthog, differing in body size and digestive system. We confronted literature estimations of their feeding requirements with forage availability and quality, observed in three South African savannah systems, through linear modelling. The model predicted the proportion of below-canopy grass that grazers should include in their diet to meet their nutritional requirements.During the wet season, the model predicted that all animals could satisfy their daily nutrient requirements when feeding on a combination of below- and outside-canopy grasses. However, wildebeest, having relatively high nutrient demands, could meet their nutrient requirements only by feeding almost exclusively below canopies.During the dry season, all animals could gain almost twice as much digestible protein when feeding on below – compared to outside-canopy forage. Nonetheless, only warthogs could satisfy their nutrient requirements – when feeding almost exclusively on below-canopy grasses. The other ungulate species could not meet their phosphorus demands by feeding at either site without exceeding their maximum fibre intake, indicating the unfavourable conditions during the dry season.We conclude that grazing ungulates, particularly warthog, zebra, and buffalo, actually depend on the available below-canopy grass resources. Our model therefore helps to quantify the importance of higher quality forage patches beneath savannah trees. The composition of grazer communities depending on below-canopy grasses can be anticipated if grazer food requirements and the abundance of large trees in savannahs are known. The model suggests that the conservation of large single-standing trees in savannahs is crucial for maintenance of locally grazing herbivores.     

Effects of host shading on consumption and growth of the geometrid Epirrita autumnata: interactive roles of water, primary and secondary compounds

Shading is assumed to reduce allocation to plant phenolics and to defense in general. We here report the results of experimental shading of individual branches or whole canopies in mountain birch on foliar chemistry and on the growth and consumption of a geometrid, Epirritaautumnata. Branch‐wide shading tended to have at least as strong effects on both leaf chemistry and herbivore performance as canopy‐wide shading, indicating local responses of the host to shading. Responses to shading varied among the key leaf traits. Leaf water content was higher and toughness lower in shaded than in non‐shaded leaves. Leaf sugars were lower and protein‐bound and free amino acids higher in shaded than in control leaves. Sucrose and galactose were at high levels in unshaded branches adjacent to shaded ones, suggesting that partial shading enhanced translocation of sugars within canopies. Total phenolics and soluble proanthocyanidins were low in both shading treatments. Of the other phenolic groups, concentrations of gallotannins and cell‐wall‐bound proanthocyanidins did not differ between shaded and non‐shaded leaves. Epirrita larvae grew better in both types of shading treatments compared to either unshaded control trees or to unshaded branches in the branch‐shading trees. By far the most important correlate of larval growth was the amount of water consumed with leaf mass (r=0.94). When variance in water intake was standardized (also largely eliminating parallel variation in proteins), fructose and glucose still had significant positive correlations and proanthocyanidins negative with larval growth on control but not on shade leaves. Concentrations of several phenolic compounds correlated negatively with intake of dry matter and especially water, and different phenolics were important in shaded (gallotannins) and in control (flavonoids) leaves. Our findings strongly suggest that the effects of putatively defensive leaf traits on insect consumption and growth interact with nutritive leaf traits, particularly with water.     

Herbaceous production in South India—limiting factors and implications for large herbivores

This study’s goal was to better understand the growth pattern and limitations of the herbaceous production that supports South India’s rich large herbivore grazer assemblage. We conducted a fully factorial nitrogen and water (three levels each) treatment field experiment in the herbivore rich South Indian Western Ghats region to determine the seasonal pattern and the extent to which nitrogen and water availability limit herbaceous production. Graminoid production was found to be nitrogen limited. Despite low rainfall, additional water did not significantly increase overall biomass production nor extend growth in the dry season. Accumulated standing biomass was highest in the late wet season (November) and lowest in the dry season (May). Leaf nitrogen was highest in the early wet season (June) and lowest in the late dry season (March). Grazing had a positive effect on grass production by extending the growing season. Biomass production and graminoid leaf nitrogen concentration levels in the study area were similar to other tropical areas in the world. Also similar to other tropical large herbivore areas, the dry season poses an annual challenge for large herbivores in the study area —particularly the smaller bodied species—to satisfy their nutrient requirements.     

Abiotic constraints on the establishment of Quercus seedlings in grassland

High evaporative demand and periodic drought characterize the growing season in midwestern grasslands relative to deciduous forests of the eastern US, and predicted climatic changes suggest that these climatic extremes may be exacerbated. Despite this less than optimal environment for tree seedling establishment, deciduous trees have expanded into adjacent tallgrass prairie within the last century leading to a dramatic land cover change. In order to determine the role of light and temperature on seedling establishment, we assessed carbon and water relations and aboveground growth of first‐year Quercus macrocarpa seedlings exposed to one of three conditions: (1) intact tallgrass prairie communities (control), (2) aboveground herbaceous biomass removed (grass removal), and (3) shade plus biomass removal to reduce light (PFD) to levels typical of the grassland‐forest ecotone (shade). In the 2000 growing season, precipitation was 35% below the long‐term average, which had a significant negative effect on oak seedling carbon gain at midseason (photosynthesis declined to 10% of maximum rates). However, net photosynthesis and stomatal conductance in the shade treatment was ca. 2.5 and 1.5 times greater, respectively, than in control treatment seedlings during this drought. During this period, leaf and air temperatures in control seedlings were similar whereas leaf temperatures in the shade treatment remained below air temperature. A late‐season recovery period, coincident with decreased air temperatures, resulted in increased net photosynthesis for all seedlings. Increased photosynthetic rates and water relations in shaded seedlings compared to seedlings in full sun suggest that, at least in dry years, high light and temperature may negatively impact oak seedling performance. However, high survival rates for all seedlings indicate that Q. macrocarpa seedlings are capable of tolerating both present‐day and future climatic extremes. Unless historic fire regimes are restored, forest expansion and land cover change are likely to continue.     

Increased growth of young citrus trees under reduced radiation load in a semi-arid climate

This study investigated the effects of radiation heat-load reduction by shading on the growth and development of citrus trees in a warm subtropical region. The experiment was conducted from mid-June until late October when daily maximal air temperature averaged 29.3 degrees C. Two-year-old de-fruited Murcott tangor (Citrus reticulata BlancoxCitrus sinensis (L.) Osb.) trees were grown under 30% or 60% shade tunnels, or 60% flat shade (providing midday shade only), using highly reflective aluminized nets. Non-shaded trees were used as the control. Shading reduced direct more than diffuse radiation. Daily radiation was reduced by 35% for the 30% Tunnel and 60% Flat treatments, and by 55% for the 60% Tunnel. Two days of intensive measurement showed that shading increased average sunlit leaf conductance by 44% and photosynthesis by 29%. Shading did not significantly influence root and stem dry weight growth, but it increased the increment in leaf dry weight during the three month period by an average of 28% relative to the control, while final tree height in the 30% Tunnel treatment exceeded the control by 35%. Shoot to root and shoot mass ratios increased and root mass ratio decreased due to shading because of the increase in leaf dry weight. Shading increased starch concentration in leaves while the shadiest treatment, 60% Tunnel, decreased starch concentration in the roots. Carbon isotope ratio (delta(13)C) of exposed leaves that developed under shading was significantly reduced by 1.9 per thousand in the 60% Tunnel, indicating that shading increased CO(2) concentrations at the chloroplasts (C(c)), as would be expected from increased conductance. Substomatal CO(2) concentrations, C(i), computed from leaf net CO(2) assimilation rate and conductance values, also indicate that shading increases internal CO(2) concentrations. Based on tree dry mass, tree height, and total carbohydrates fractions, the 30% Tunnel and the 60% Flat were the optimal shade treatments.     

Tree dieback and regeneration in secondary Acacia zanzibarica woodlands on an abandoned cattle ranch in coastal Tanzania

Questions: Bush encroachment is a major problem when African savanna ecosystems are used for cattle ranching. How do secondary woodlands develop after ranching is abandoned? What are the patterns and rates of tree mortality and regeneration? Location: Mkwaja Ranch (now part of Saadani National Park) in coastal Tanzania. Methods: Ninety‐seven circular plots (4‐m radius) were set in secondary Acacia zanzibarica woodland along a gradient of tree density. Variables relating to tree and grass layers and soil characteristics of plots were recorded. Seedlings were counted twice in the wet seasons, and resprouts once. Tree flowering and pod production were assessed during the fruiting season, while survival of trees initially present was recorded after 12 and 32 months. Interrelationships among variables were investigated using multiple linear regression, binary logistic regression and mixed effects models. Results: After 32 months, over one‐third of trees in plots had died. Most died after fire, especially on heavy soils, and mortality was significantly related to the tree live biomass ratio and soil conditions. Seed production was very low, especially in denser stands. Numbers of seedlings correlated with soil and grass variables but not with seed rain. Half of trees killed above‐ground produced coppice shoots from the base; in contrast, root suckering was independent of topkill. By the end of the study, no seedlings survived and no resprouts emerged above the grass layer. Conclusions: A. zanzibarica woodlands at Mkwaja Ranch were able to develop because of ranching, and can only persist under intensive grazing. The woodlands do not represent a successional stage towards forest and will probably revert to predominantly grassland vegetation within 10–20 years unless grazing pressure from wild ungulates increases considerably and/or fire regimes change.