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.     

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.     

Changes over 46 years in plant community structure in a cleared brigalow (Acacia harpophylla) forest

Plant succession theory underpins the development of strategies for the conservation and regeneration of native communities. Current theory has been based largely on space‐for‐time rather than long‐term monitoring data, which have known limitations. There is general consensus that more site‐specific studies are needed to corroborate existing hypotheses. The target vegetation is a brigalow (Acacia harpophylla, Mimosaceae) forest in one of Australia's most endangered ecosystems, which was cleared and burnt in 1963. Forty quadrats were placed systematically within each of six 20 m × 20 m permanent plots. Presence, density and per cent canopy cover data were recorded for each species at 18 times over 46 years. Brigalow dominated the original vegetation, assumed dominance soon after clearing through massive root suckering and remained dominant throughout the study. It achieved maximum density within two years when severe intraspecific competition led to self‐thinning. After approximately 30 years, vacant niches appeared. Woody understorey species were slow to recolonise. Species richness and other diversity indices increased rapidly to a maximum after 2–4 years, declined until the 30th year when they again increased. This was the pattern of the species‐rich herbaceous layer; woody species showed a steady monotonic increase. The ‘hump‐shaped’ relationship between cover (biomass) and species richness was confirmed. This example fits the inhibition model for which few examples have been described. While the long‐term successional pattern is slightly confounded by climatic variability preceding sample surveys, this space‐for‐time study not only supports a bimodal pattern of diversity over time but also indicates that the relative species richness of the herbaceous and woody layers may explain the extreme variability reported in the literature.     

Secondary succession in Acacia nilotica (L.) savanna in the Hluhluwe Game Reserve,South Africa

Analysis of aerial photographs indicates that woody plant biomass has rapidly increased in Hluhluwe Game Reserve over the last 40 years. Open Acacia nilotica savanna is being replaced by broadleaf species, especially Euclea spp. We were interested in whether this secondary successional shift was due to high numbers of seedlings establishing and growing to maturity under acacias (facilitation) or due to the release of already established, but suppressed individuals (gullivers) of the resprouting broadleaf species. We examined the recruitment patterns and size-class distributions (height, basal diameter) of important species in this savanna.Densities of euclea seedlings (<0.6 cm basal diameter) under Acacia nilotica were low (median of 0 and mean of 0.06 m²) below adult canopies and effectively zero in adjacent interspaces. No differences in numbers of other broadleaf species were found between open sites and under Acacia nilotica sites. Few large eucleas or other broadleaf species occurred under Acacia nilotica. Few Acacia nilotica recruits were found either under adults or in the open whereas Acacia karroo recruits were more common.The overall size class distribution for eucleas was dominated by individuals in the intermediate size class, suggesting that recruitment is not the dominant demographic process. We propose that the escape of intermediate sized eucleas from the fire trap has caused the increase in woody plants.Analysis of a time sequence of aerial photographs shows that invasion occurred rapidly between 1954 and 1975. A common feature for sites where woody plant invasion has taken place, was the presence of barriers to fire (especially roads). We suggest that the recent and rapid increase in woody vegetation is due to a decrease in the frequency of intense fires, rather than the recent absence of megaherbivores which allowed Acacia nilotica establishment. Few intense fires allow suppressed tree or shrub individuals, ('gullivers'), to escape the grass/fire layer and thereby become tall and fire-resistant. This release may explain the rapid rate of invasion by inherently slow growing broad-leaf species.     

Bird community composition in an actively managed savanna reserve, importance of vegetation structure and vegetation composition

The mosaic of trees, shrubs and open grassland in mesic African savannas is highly dynamic and strongly influenced by mammal herbivory and fire. We investigated the bird fauna in four different savanna habitats to help assess the impacts of vegetation change on this component of faunal diversity. Birds were censused, plant species were identified and vegetation structure was measured in four different vegetation types (Acacia nilotica woodland, Acacia nigrescens woodland, broadleaf thicket and open grassland) in the Hluhluwe-Umfolozi Park in northern KwaZulu Natal, South Africa. Multivariate ordination analyses were used to determine the relative importance of vegetation structure and floristic composition in defining bird assemblages. The bird communities of the grasslands, the acacia woodlands, and the broadleaf woodlands were clearly separated on the first axis of the detrended canonical correspondence analysis (DCCA). Canopy cover and foliage height diversity (FHD) were strongly correlated with the first axis of DCCA, possibly reflecting a secondary successional series from grassland to woodland, known as bush encroachment. Floristic composition (based on presence–absence data only) seemed to be less important for bird community composition than vegetation structure. The results indicate that changes in vegetation structure, caused by bush encroachment, could cause concomitant changes in bird community composition.     

Regeneration patterns in a Central European dry heathland: effects of burning,sod-cutting and cutting

Vegetation development in dry heathlands was studied during a six-year period following experimental burning, sod-cutting and cutting in a continental area in the southern part of the Czech Republic (Podyjí National Park). Calluna vulgaris-dominated heathlands in the area were composed of uneven-aged stands and had been unmanaged for several decades. The aim of the study was testing the applicability of Western European management systems to nature conservation in a continental area. Species composition was recorded in 8 permanent plots using 25×25 cm grids, and vegetation recovery following different treatments was analysed. Calluna recovered successfully after burning. Burning dense heathlands, which contained a large amount of woody fuel, resulted in a medium-intensity fire that destroyed moss mats and litter and exposed patches of bare ground to facilitate Calluna regeneration by seed, in addition to vegetative regrowth. On the contrary, in open heathlands with patches of herbaceous vegetation, low-intensity fires which failed to expose mineral soil were more typical. Almost all Calluna regeneration was vegetative in this case, and regrowth was slower. Heathland recovery after sod-cutting to mineral soil depended on whether or not Calluna seed germination occurred in the plot. With germination, the community developed towards heathland; without towards grassland. Cutting promoted a striking increase in grass cover, which was followed by the slow recovery of Calluna. The experiments suggest that of these management systems, burning is perhaps the most appropriate in the study area. A nature conservation management system is proposed, based on sheep grazing combined with rotational burning of restricted patches.     

Sapling survival and growth of coniferous and broad‐leaved trees in successional highland habitats in Mexico

Abstract. We assessed survival and growth of transplanted saplings of understory broad‐leaved trees (Oreopanax xalapensis, Rapaneajuergensenii, Rhamnus sharpii and Tern‐stroemia lineata) and canopy conifers (Abies guatemalensis, Pinus ayacahuite and P. pseudostrobus) into Grassland (GR), Mid‐Successional Oak Forest (MS), and Old‐Growth Oak Forest (OF) stands in the central highlands of Chiapas, Mexico. A total of 727 plants were monitored over eight years. The results suggest habitat preferences of the studied species that should be considered for their conservation oriented management. Conifers had highest survival and growth in GR, while broad‐leaved trees survived better under sparse canopy in MS, but grew tallest in full light. Saplings of all species in dense canopy sites in OF were smaller, and the survival of conifers was lower. An integrated response index (IRI) calculated for each species with survival and relative growth rates showed similar trends. Re‐introduction of the endangered A. guatemalensis is possible in open habitats if fire, trampling and grazing are suppressed. Open habitats in current man‐made landscapes in the highlands of Chiapas may limit the establishment of understory trees requiring partial shade. Inclusion of such species in restoration programs has not been usual so far; yet, local and global interest in native species for such programs is increasing. Information on the response of these kind of species in contrasting habitats may help to incorporate them in the management of high‐diversity forests that may follow pine plantations.     

Impacts of ungulates on the demography and diversity of small mammals in central Kenya

The impacts of ungulates on small mammals in an East African savanna habitat were investigated by monitoring the population and community responses of small mammals on replicated 4-ha plots from which ungulates had been excluded. The dominant small mammal in this habitat is the pouched mouse, Saccostomusmearnsi, a medium-sized murid rodent. Eight other small mammal species, including Arvicanthis sp., Mus sp., Mastomys sp., Dendromus sp., Crocidura sp., and, rarely, Tatera sp., Aethomys sp., and Acomys sp., were also captured. The dominant ungulates are elephant (Loxodonta africana), giraffe (Giraffa camelopardalis), Grevy's and common zebra (Equus grevyi and E. burchelli), buffalo (Syncerus cafer), eland (Taurotragus oryx), Grant's gazelle (Gazella granti), and domestic cattle. Within 1 year, S. mearnsi populations had responded dramatically to the exclusion of large mammals by a two-fold increase in density, a difference that was maintained through pronounced seasonal fluctuations in the second year. Though individual pouched mice showed no significant differences in their use of space with and without ungulates, male S. mearnsi maintained significantly higher body weights in the absence of ungulates, indicating that habitat quality had increased. One other species, Mastomys sp., also increased in the absence of ungulates. Overall, the small mammal community maintained relatively constant species diversity on the plots to which ungulates did not have access. On the plots to which ungulates did have access, on the other hand, there was a rapid 75% decrease in diversity in the control plots during one trapping session. Ungulates are most likely affecting small mammals through their effects on food quality, since there were no detectable differences in their exposure to predators, as determined by vegetative cover, in the absence of ungulates. These results demonstrate that ungulates can have strong and rapid impacts on small mammal abundance and diversity in East African savannas, an interaction which has not previously been given serious consideration. Received: 28 December 1997 / Accepted: 11 May 1998     

Preliminary observations on the role of Acacia gum chemistry in Acacia utilization by Galago senegalensis in Kenya

A census and preliminary observations on the ranging and feeding behavior of Galago senegalensis were made in a savannah-woodland site in Kenya. Population densities in three slightly different habitats were all approximately 1.5 animals/ha. Ranging patterns suggested the existence of territorial behavior. The diet of the galagos was exclusively insects and gums of two species of Acacia. The gum of A. drepanolobium appeared to be preferred to that of A. xanthophloea. The chemistry of the gums suggests that this preference is not due simply to the total level of phenolics or to avoidance of condensed tannins but may relate to the presence of compounds (e. g., flavonoids) having nutritional or hormonelike action.     

Carbon-cycling changes during regeneration of a decinduous broadleaf forest after clear-cutting II. Aboveground net production

Growth and death rates of aboveground plant parts were measured in a mature forest and four different-aged deciduous broadleaf forests regeneratede after clear-cutting, with special reference to rates for woody parts (stems and branches) of different diameters (ø) in rerms of the pipe model theory (Shinozaki et al., 1964). The total biomass increment of woody parts of trees higher than 1.3 m varied within a range of 2.1-4.6 ton ha^?1 yr^?1, the increase beingdue largely to the growth of canopy trees exposed to direct sunlight. Biomass increments of small (ø<1 cm) and medium (1≤ø<5 cm) woody parts were negligibly small except in the youngest forest, and changes in aboveground woody biomass with forest age after clear-cutting mainly resulted from accumulation of large (5 cm<ø) woody parts of canopy trees. Biomass loss of trees due to death and grazing increased with forest age from 4.0 to 8.3 ton ha^?1 yr^?1. Recovery of leaf and small wood falls was observed at the early stage of regeneration, while large wood falls increased during regeneration. Flower and fruit fall was markedly higher in the mature forest than in the other four forest types. Mortality of woody parts became higher with forest age and was 20, 5.0 and 0.46% yr^?1 for small, medium and large parts, respectively, at the mature stage. Aboveground net production of the forest was in therange 7.6-13.3 ton ha^?1 yr^?1 with the undergrowth vegetation lower than 1.3 m being 0.4-1.4 ton ha^?1 yr^?1. Production recovered rapidly at an early stage of regeneration and was highest in mature forest.     

广东黑石顶自然保护区森林次生演替过程中群落的种间联结性分析

 采用2×2列联表，通过方差分析、X2检验、Ochiai指数对黑石顶森林次生演替过程中各阶段群落优势种的种间联结性进行了定量分析。研究结果表明：群落的总体种间关联性在演替恢复初期表现为无相关性，针阔叶混交林阶段为总体的正关联，阳生性常绿阔叶林阶段呈总体负关联，中生性常绿阔叶林阶段总体呈显著正关联。在演替初期主要种对间尚未形成一定的种间关系，演替至混交林和阳生性常绿阔叶林阶段，种间关系开始形成，但大多表现为直接的竞争关系，中生性常绿阔叶林阶段形成了复杂多样的种间关系，表现为同层树种间的种间结合或竞争和不同层种间的结合关系。     

Composition of grazed and cleared temperate grassy woodlands in eastern Australia: patterns in space and inferences in time

Abstract. A regional vegetation survey of the temperate grassy woodlands (temperate savanna) in Australia was designed to assess the effects of clearing and grazing on the composition of vegetation remnants and the adjacent pasture matrix. Vegetation was sampled across a range of habitats using 77 0.1024‐ha quadrats; the relative abundance of species was recorded. Classification analysis clustered the sites into three main groups that corresponded to intensity of grazing/clearing followed by groups based on underlying lithology (basalt, metasediment, granites). Using Canonical Correspondence Analysis, exogenous disturbance and environmental variables were related to the relative abundance of species; grazing intensity had the highest eigenvalue (0.27) followed by tree canopy cover (0.25), lithology (0.18), altitude (0.17) and slope (0.10). Based on two‐dimensional ordination scores, six species response groups were defined relating to intensity of pastoralism and nutrient status of the landscape. Abundance and dominance of native shrubs, sub‐shrubs, twiners and geophytes were strongly associated with areas of less‐intense pastoralism on low‐nutrient soils. The strongest effects on species richness were grazing followed by canopy cover. Continuously grazed sites had lower native species richness across all growth forms except native grasses. There was no indication that intermediate grazing intensities enhanced forb richness as a result of competitive release. Species richness for all native plants was lowest where trees were absent especially under grazed conditions. Canopy cover in ungrazed sites appeared to promote the co‐existence of shrubs with the herbaceous layer. Predicted declines in forb richness in treeless, ungrazed, sites were not detected. The lack of a disturbance‐mediated enhancement of the herbaceous layer was attributed to habitat heterogeneity at 0.1 ha sampling scale.     

黑石顶自然保护区森林次生演替过程中群落主要种的种间协变分析

采用2×2列联表,通过Pearson积矩相关系数和Speaman秩相关系数对黑石顶森林次生演替过程中各阶段群落优势种的种间协变进行了定量分析。结果表明,在演替初期的裸地恢复阶段,显著种间协变的种对较多,有14对(正协变9对负协变5对),在针阔混交林阶段,显著种间协变的种对较少,仅有5种。全为正协变,表现为主要种间对群落环境的相同要求,并形成了主要种间的竞争格局,在阳生性常绿阔叶林阶段。显著种间协变的种对最少,只有4对,其中3对为负协变,这是主要种之间对群落环境相同要求的激烈竞争结果,在中生性常绿阔叶林阶段,显著种间协变的种对最多有20对,19对为正协变,主要表现为群落上层种和中下层种间形成了大量的正协变,但群落上层主要种间无显著的种间协变。     

The Community Dynamics of the Forest Secondary Succession in Heishiding Natural Reserve of Guangdong Province

Studying on the community structure, species composition and species diversities of the secondary succession of lower subtropical forest in Heishiding Natural Reserve, Guangdong Province, the following results were obtained. In 2 ~4 years of restored stage the seedlings of pioneer species, heliophytes and mesophytes occur simultaneously. In 10 years of restored stage pioneer species are superior as heliophytes maintain stable and mesophytes are being decreased. In the stage of mixed forest pioneer species are on the decreasing and heliopytes are on the increasing trend. In the stage of evergreen broad-leaved forest dominated by heliophytes pioneer species are being declined, heliophytes are dominant and mesophytes are being increased. In the stage of evergTeen broad-leaved forest dominated by mesophytes heliophytes are being declined and mesophytes are superior. The vertical structure, horizontal structure, the structure of species composition determine stability of communities in secondary succession process. The vertical structure of community is building up in the stage of restoration. The community is composed of the stands under the third class. The forest canopy is not stratified. The individual density is stable. From restored stage to mixed forest the vertical structure of community is diversified. There are the third and forth class stands in community. The tree layer is divided into three sub-layers. The individual density declines rapidly because of self-thinning. In the mixed forest the vertical stmcture is relatively stable. The abundance of the fifth class stands and area of breast height reach the first peak in the succession serial. The individual density increases rapidly. In stage of evergreen broad-leaved forest dominated by heliophytes, the vertical structure changes largely. The abundance of the fifth class and area of breast height decline rapidly, but the individual density changes slowly. In the stage of evergreen broad-leaved forest dominated by mesophytes, the vertical structure of community is stable and the composition of every stand class becomes more stable. The area of breast height is the biggest in the succession serial. When succession develops, the coverage of communities increases gradually but the turnover rate of species declines. The species composition tends to be stabilized. The diversity of species and community evenness increase gradually thus, the ecological dominance declines gradually.     

The effect of clearing on plant composition in mulga (Acacia aneura) dry forest,Australia

Despite high deforestation rates in dry forests there are few assessments of the impacts of this transformation on biodiversity. This study addresses the impact of clearing on plant species composition and diversity in mulga (Acacia aneura) forests of Queensland, Australia using data collected from either side of 58 clearing boundaries, incorporating data on site and management conditions. The physical and climatic conditions of the study sites were narrowly constrained but had greater influence on plant composition and richness than clearing per se. In general, the abundance of perennial forbs and shrubs was higher in cleared forest compared with adjacent intact forest and there was no significant difference in abundance of other plant life forms. Perennial forb richness diminished with the age of clearings, particularly where acid sub‐soils had been exposed and shrubs proliferated. Where deforestation in the mulga dry forests is not accompanied by the establishment of exotic pasture it has a relatively benign effect on plant diversity, mulga itself remains in high density as small trees and the integrity of the forest could be maintained with restoration.     

Structure and composition of Acacia xanthophloea woodland in Lake Nakuru National Park, Kenya

The woody vegetation of Lake Nakuru National Park occurs along rivers, lakeshores and flood plains. Four different sites within the Acacia xanthophloea woodlands were selected for the study. Vegetation structure was not significantly different in the four woodlands used for the study, but these sites differed in the relative density of Acacia trees. Regeneration of A. xanthophloea differed in each site, with the highest regeneration rates found in the nonfenced plots where browsing took place.     

Forest Regeneration in a Chronosequence of Tropical Abandoned Pastures: Implications for Restoration Ecology

During the mid‐1900s, most of the island of Puerto Rico was deforested, but a shift in the economy from agriculture to small industry beginning in the 1950s resulted in the abandonment of agricultural lands and recovery of secondary forest. This unique history provides an excellent opportunity to study secondary forest succession and suggest strategies for tropical forest restoration. To determine the pattern of secondary succession, we describe the woody vegetation in 71 abandoned pastures and forest sites in four regions of Puerto Rico. The density, basal area, aboveground biomass, and species richness of the secondary forest sites were similar to those of the old growth forest sites (>80 yr) after approximately 40 years. The dominant species that colonized recently abandoned pastures occurred over a broad elevational range and are widespread in the neotropics. The species richness of Puerto Rican secondary forests recovered rapidly, but the species composition was quite different in comparison with old growth forest sites, suggesting that enrichment planting will be necessary to restore the original composition. Exotic species were some of the most abundant species in the secondary forest, but their long‐term impact depended on life history characteristics of each species. These data demonstrate that one restoration strategy for tropical forest in abandoned pastures is simply to protect the areas from fire, and allow natural regeneration to produce secondary forest. This strategy will be most effective if remnant forest (i.e., seed sources) still exist in the landscape and soils have not been highly degraded. Patterns of forest recovery also suggest strategies for accelerating natural recovery by planting a suite of generalist species that are common in recently abandoned pastures in Puerto Rico and throughout much of the neotropics.     

Seedling establishment of Acacia tortilis and Hyphaene compressa in the Turkwel riverine forest, Kenya

This study reports on the spatial and temporal patterns of seedling establishment in the Turkwel riverine forest, Kenya. Seedlings of the dominant tree species Acacia tortilis and Hyphaene compressa were mapped and monitored to assess the underlying causes of seedling recruitment and mortality. The broad‐scale distribution of A. tortilis was not correlated with any environmental variables, while H. compressa seedlings were confined to flood‐exposed sites in the arid downstream section of the floodplain. One year of monitoring showed that seedling recruitment of A. tortilis was evoked by prolonged rainfall, while seedling mortality was caused by desiccation, browsing and trampling. In contrast, seedling recruitment and mortality of H. compressa was largely unaffected by rainfall and livestock, probably due to the high moisture requirements of seeds and the tolerance of seedlings to disturbance. There were no effects of soils, light, or seedling density on the establishment of A. tortilis and H. compressa seedlings. This study demonstrates the importance of parallel mapping and monitoring of riverine seedlings in order to understand patterns and processes of forest regeneration in arid and semi‐arid floodplains.