Changes in structure of savanna woodlands in northern Botswana following the impacts of elephants and fire

Fluctuations in densities of woody plant species were monitored in plots within three northern Botswana woodland types subjected to elephant damage and burning. Woodlands dominated by Baikiaea plurijuga and Colophospermum mopane sustained significant changes occurring on an annual basis, whereas Acacia erioloba plots maintained a typical structure. The structure of A. erioloba woodlands appeared to be influenced by factors other than elephants and the occurrence of fire. Woodlands dominated by C. mopane plants were subjected to obtrusive elephant damage, although the densities of tall trees remained largely unchanged. The effects of fire were most prominent in B. plurijuga woodlands. Tree densities declined consistently and plants of lower height classes, such as shrubs and seedlings increased in densities in areas subjected to a high occurrence of fire.     

Browsing and grazing in elephants: the isotope record of modern and fossil proboscideans

The diet of extant elephants (Loxodonta in Africa, Elephas in Asia) is dominated by C₃ browse although some elephants have a significant C₄ grass component in their diet. This is particularly noteworthy because high-crowned elephantid cheek teeth represent adaptation to an abrasive grazing diet and because isotopic analysis demonstrates that C₄ vegetation was the dominant diet for Elephas in Asia from 5 to 1 Ma and for both Loxodonta and Elephas in Africa between 5–1 Ma. Other proboscideans in Africa and southern Asia, except deinotheres, also had a C₄-dominated diet from about 7 Ma (when the C₄ biomass radiated in tropical and subtropical regions) until their subsequent extinction. Received: 1 July 1998 / Accepted: 16 February 1999     

The effects of fire on the frillneck lizard (Chlamydosaurus kingii) in northern Australia

Abstract A population of frillneck lizards, Chlamydosaurus kingii, was monitored by radio telemetry and mark-recapture techniques between April 1991 and April 1994, as part of a landscape-scale fire experiment, in Kakadu National Park, Northern Territory. The study aimed to investigate both the short- and longer-term effects of fire on a lizard species in a tropical savanna where fires are frequent and often annual. Frillneck lizards are able to survive fires that occur in the first few months of the dry season by remaining perched in trees. A high level of mortality (29%) occurred during late dry-season fires, along with changes in their behavioural response to fire: sheltering in either larger trees or hollow termite mounds. Food is more accessible after fires due to the removal of ground vegetation. This is reflected in greater volume and diversity of prey in stomach contents after fires. This increase is more pronounced after late dry-season fires, possibly due to increased accessibility of prey caused by more complete vegetation removal. Frillneck lizards show an overall preference for trees with a dense canopy cover located in an area with a low density of grass. Fire has an effect on this relationship. Frillneck lizards in habitat unburnt for a number of years tend to perch in trees with a smaller canopy, whereas lizards in annually burnt habitat perch in trees with a dense canopy. Volume and composition of lizard stomach contents was broadly similar among fire treatments over a 2 year period, although termites were more predominant in stomach contents of lizards in unburnt habitat. Wet-season body condition is lower in lizards from unburnt habitat, although the reason for this is unclear. These results demonstrate the importance of different fire intensities and regimes on the ecology of a lizard species in a tropical savanna.     

Soil nutrients distribution and moisture dynamics on upper catena in a semi-arid nature reserve

The distribution of soil nutrients and soil moisture dynamics on slopes of catenary landscape, were recorded in a semi-arid nature reserve located in the Eastern Transvaal, South Africa. Soil nutrients were accumulating in the slopes of the catena whereas the crests were highly leached. Open grassland bands segregating between the two soil types and their associated plant species, did not show clear stratification of nutrient deposition amongst ground levels down to 1.5 m below the surface. There was however, a tendency of soils with high nutrient contents to correlate with areas of high woody vegetation density. Patterns of soil moisture levels during two annual rainfall periods showed some segregation between soil depths, although the trends indicated that the duration of saturation and dry out of soils levels differed more between rainy seasons than between soil levels during a particular season.     

Reproductive potential and seed fate of Sclerocarya birrea subsp. caffra (marula) in the low altitude savannas of South Africa

Even though Sclerocarya birrea subsp. caffra (marula) is a well-studied, keystone tree species with high ecological, commercial and cultural value, significant gaps in our understanding of its reproductive biology exist, particularly the factors limiting fruit and seed production, seed fate and the persistence of the seed bank. Therefore, a detailed quantitative assessment of these factors was conducted at five sites in the low altitude savannas of South Africa. Sites varied with respect to fire regime, large mammals, geology and rainfall. Most sites showed male-biased secondary sex ratios and the minimum fruiting stem diameter ranged between 7.1 and 15.7 cm across sites. Sites with higher levels of disturbance (fire and large browsers) had trees producing fruit at larger minimum stem diameters than sites with lower levels of disturbance. Fruit production was highly variable between individuals, within and between sites, and from year to year. Variability in fruit production across years at one site was greater than the variability across sites in one year, indicating that drivers such as weather, insect herbivory, fire and predator numbers, which vary annually, play a greater role than more constant drivers such as mammalian herbivory, soil types or long-term rainfall. No significant relationship was found between environmental variables (rainfall and temperature) and annual fruit production, indicating a trade-off between vegetative growth and reproduction between years. Since marula fruits are large and heavy, the species relies primarily on mammalian dispersal agents such as the African elephant, which have also been shown to increase the germination rate. However, rodents also appear to play a significant role in seed dispersal. Seed predation rates tend to be highest in areas of low disturbance (no fire and no large browsers). While marula has a small persistent seed bank, recruitment appears to be reliant on the current season's fruit crop. This study provides a detailed quantitative assessment of important reproductive and seed fate vital rates for future population modelling.     

The impact of elephants, Loxodonta africana, on woody vegetation through selective debarking in Samburu and Buffalo Springs National Reserves, Kenya

The impact of elephants on the woody plant community through debarking was investigated in Samburu and Buffalo Springs National Reserves, Kenya. Acacia elatior Brenan , the most abundant tree species in the riverine zone, accounted for 68% (n = 1375) of woody plants. A. tortilis (Forsskal) Hayne dominated plots away from the river. Debarking incidences were significantly higher for A. elatior than for other species indicating selective utilization. The riverine zone by virtue of having more trees of the preferred species, A. elatior, had the highest debarking incidences. Presence of very few saplings along the river is attributed to both elephant trampling and herbivory by other species. An estimated 38.5% and 22.5% of the riverine A. elatior and A. tortilis trees respectively, were bound to die within the next 4–5 years because of severe debarking, ≥75% of bark circumference. Debarking was positively correlated with stem circumference; the medium-sized trees being the worst affected by the elephants' selective debarking behaviour. Intense debarking incidences were recorded during the dry season. Through the elephants' selective debarking, the riverine habitat is bound to open up gradually, leading to considerable habitat change in the near future. Elephant impact on vegetation is less away from the river and increases with their densities.     

Water relations and growth of shrubs before and after fire in a semi-arid woodland

Summary Plant water relations and shoot growth rate of shrubs resprouting after fire or unburnt were measured in a semi-arid poplar box (Eucalyptus populnea) shrub woodland of eastern Australia. In vegetation unburnt for about 60 years, the dawn xylem water potential (_x) of the dominant shrub species was about-1.0 MPa when the soil was wet and-8.0 MPa when the soil was very dry. At any one time, the dominant shrub species,Eremophila mitchellii, E. sturtii, Geijera parviflora andCassia nemophila, were similar in _x butAcacia aneura andDodonaea viscosa were consistently higher in _x than this group when the soil was moist and lower when the soil was dry. The dominant tree species,Eucalyptus populnea andE. intertexta, appeared to have access to additional water beneath the hardpan which is located 60–80 cm below the surface. When shrubs were under extreme water stress (_x of-8 MPa), the trees had a _x of-3 to-3.6 MPa. Following a fire, both _x and leaf stomatal conductance (g _s) of resprouting shrubs were higher for about 5 years than comparable-aged unburnt vegetation, with relative differences in _x increasing with drought stress. Elongation rate of resprouts was positively linked to prefire shrub height in 3 of 4 species. However, shrubs resprouting after high intensity fires had substantially higher rates of shoot elongation than after low intensity fires which were in turn higher than for foliar expansion of unburnt shrubs. It is concluded that the growth rate of resprouting shrubs is primarily determined by physiological/ morphological factors associated with plant size but is also assisted by greater availability of water and possibly nutrients for a period after fire.     

Sleeping Parties and Nest Distribution of Chimpanzees in the Savanna Woodland, Ugalla, Tanzania

We conducted ecological studies of chimpanzees (Pan troglodytes) in the Ugalla area, western Tanzania. Ugalla is one of the driest habitats of chimpanzees and the Ugalla River is the eastern boundary of chimpanzee distribution. Most of Ugalla is occupied by savanna woodlands dominated by deciduous trees of Brachystegia and Julbernardia. Chimpanzees tended not to make nests in riverine forests in plains, but in small patchy forests dominated by Monopetalanthus richardsiae and valley forests dominated by Julbernardia unijugata on slopes in mountainous areas. We estimated population density of chimpanzees to be 7–9 × 10⁻² individuals/km² based on nest censuses, suggesting that 2–3 × 10² individuals inhabited the 3352 km² area of Ugalla. The size of the largest nest cluster (n=23) suggests that 1 unit group (community) comprised 30–35 individuals. In the daytime, chimpanzees formed small feeding parties (mean 2.0 individuals), but larger ones in the evening (mean 4.8 individuals and 5.2 individuals based on fresh nest clusters). The pattern might reduce the predation risk from large nocturnal carnivores such as lions and leopards. The sleeping sites may function as both a safe sleeping site and a meeting point for chimpanzees with a huge home range that may have difficulty in finding other members of their unit group.     

Brood raiding and the population dynamics of founding and incipient colonies of the fire ant, Solenopsis invicta

Abstract. 1. One of the first activities of minim workers in incipient fire ant nests is mutual brood raiding, the amalgamation of nests through the reciprocal stealing of brood and defection of workers.
2. Discrete mating flights created cohorts of incipient colonies. About 25% of founding nests survived the claustral period of 16–48 days (depending on season). Early incipient colony mortality was 5–6% per day. Over 60% of this mortality was accounted for through brood raiding. Most colonies raided when each cohort first became active, and many raided more than once. Raid size (number of nests, duration, distance) tended to increase during the summer as colonies grew through raiding. After the raiding period, nest mortality rate dropped 3–10-fold. Only 1–3% of founding nests were still alive at this time.
3. Queens from losing or failed nests tended to abandon their nests and attempted to enter successful ones, often following raiding trails to do so. This emigration was at least as successful as non-emigration in ultimately achieving the status of reproductrix of a successful nest (about 4%).
4. Brood raiding is a dominant process in early population dynamics, probably accounting for most of the early nest mortality. Its effect is to change the venue and unit of competition from nest-against-nest to a shifting aggregation of queens, workers and brood involving entire local populations. Nest thinning is thus very rapid, and the boost to the size of winning nests very large, allowing raiding colonies to win the competition for territory, and to achieve the early colony maturity so important to this, and other, weedy species. The importance of winning brood raids may also have driven increased minim production through the evolution of pleometrosis.     

Reactions of the fauna on the bark of trees to the frequency of fires in a North American savanna

Summary The arthropod communities living on the bark of the oak species Quercus macrocarpa and Q. ellipsoidalis were investigated in a North American oak savanna. Differences were found in the community structure of the arthropods living on the bark of these two tree species, although they have the same fissured bark type. In the North American oak savanna ecosystem the most important disturbance factor is fire, which maintains species richness. Highest numbers of species and specimens were found at moderately disturbed sites. Three main ecological groups of arthropods living on the bark of trees can be distinguished in relation to the degree of disturbance: (1) Inhabitants of bark of trees restricted to undisturbed sites: they do not occur in fire-disturbed areas; (2) Inhabitants of bark of trees adapted to a moderate degree of disturbance: many species occur in high numbers only in moderately disturbed areas; and (3) Specialist inhabitants of bark of trees in heavily disturbed areas. The number of specimens of these species increases per trunk with the frequency of disturbance.     

Tree structure,regeneration and survival of seedlings and sprouts in highmontane forests of the Garhwal Himalayas,India

The tree structure of some timberline and sub-alpine forests of the Garhwal Himalayas was studied with reference to the regeneration status and survival of tree seedlings and sprouts. Nine tree and shrub species were chosen. The dominating tree species wereAbies pindrow andQuercus semicarpifolia. Sprouts were found to have a higher survival rate than seedlings, but both suffered from winter conditions. Girth class measurements showed that trees between 1.6 and 2.5 m were best represented.     

Large herbivores influence the composition and diversity of shrub-steppe communities in the Rocky Mountains, USA

It is widely believed that wild and domestic herbivores have modified the structure and composition of arid and semi-arid plant communities of western North America, but these beliefs have rarely been tested in long-term, well-replicated studies. We examined the effects of removing large herbivores from semi-arid shrublands for 40–50 years using 17 fenced exclosures in western Colorado, USA. Shrub cover was greater (F=5.87, P=0.0020) and cover (F=3.01, P=0.0601) and frequency (F=3.89, P=0.0211) of forbs was less inside the exclosures (protected) relative to grazed plots. However, we found no significant effects (minimum P=0.18) of protection from grazing on cover or frequency of grasses, biotic crusts, or bare soil. Although mean species richness and diversity were similar between treatments, protected areas had much higher dominance by fewer species, primarily sagebrush. Exclusion of herbivores changed the relationship between species richness and evenness. Consistent with theoretical expectations, species evenness was positively correlated with richness in protected plots (r ²=0.54). However, contrary to theory, evenness and richness were inversely related in grazed plots (r ² _adjacent=0.72, r ² _distant=0.84). We suggest that these differences resulted because grazing acts as a stressor promoting facilitative relationships between plant species that might compete for resources in the absence of grazing. We conclude that exclusion of grazing in the sites we studied caused minor changes in cover and diversity of herbaceous plants, but caused a clear increase in the cover of shrubs. Importantly, the exclusion of ungulates changed the relationship between evenness and richness.     

The role of protective understorey in the regeneration system of a heavily browsed woodland

The paper tests an apparent relationship between young understorey of Ilex aquifolium and the presence of seedlings of the canopy species Fagus sylvatica and Quercus species, in heavily browsed woodland in the New Forest, southern England. A small block of woodland was searched and mapped, and the information collected subject to graphical and statistical analyses.Regeneration, though of low frequency, is present and shows non-random patterns of distribution. Older seedlings appear to be located in or near areas of understorey, or near fallen branches, logs, etc. Principal components analysis of woodland characteristics suggests six main habitat types are available to seedlings, and predictions are made about likely regeneration patterns associated with these habitats, according to a model based on the hypothesis of a protective understorey and heavy browsing pressure. Chi-square tests of actual seedling occurrences versus habitat types demonstrate marked associations of regeneration with non-trampled areas, including young Ilex understorey but also other forms of protective barrier. The association is more marked for larger seedlings.The implications for a wider understanding of regeneration processes and community dynamics in heavily browsed forest are discussed. It is concluded that regeneration theories appropriate to such forests are required, since the dominant gap-phase theories are not successful in predicting regeneration patterns when browsing is a major factor.     

Sampling techniques and the use of Tang's procedure in insect population dynamics studies

Some calculations were performed usingTang 's method as an aid in planning experiments for studying the population dynamics of the Jeffrey pine beetle. The population dynamics studies were aimed at detecting the importance of specific effects, e. g., tree diameter, tree height. TheTang procedure is a method of estimating the sample size required to detect effects of a given magnitude with analysis of variance tests. Using this procedure some sample calculations were performed which indicated the sample size needed, and the efficacy of different strategies of improving the results, e. g., increasing the number of trees sampled versus increasing the area of the tree sampled. The statistical parameters used in the calculations were estimated from some preliminary sampling data. Use of this procedure is recommended in insect population studies as a method of optimally planning experiments, and as a method of making precise conclusions about the significance of specific effects.     

The importance of low atmospheric CO2 and fire in promoting the spread of grasslands and savannas

The distribution and abundance of trees can be strongly affected by disturbance such as fire. In mixed tree/grass ecosystems, recurrent grass‐fuelled fires can strongly suppress tree saplings and therefore control tree dominance. We propose that changes in atmospheric [CO₂] could influence tree cover in such metastable ecosystems by altering their postburn recovery rates relative to flammable herbaceous growth forms such as grasses. Slow sapling recovery rates at low [CO₂] would favour the spread of grasses and a reduction of tree cover. To test the possible importance of [CO₂]/fire interactions, we first used a Dynamic Global Vegetation Model (DGVM) to simulate biomass in grassy ecosystems in South Africa with and without fire. The results indicate that fire has a major effect under higher rainfall conditions suggesting an important role for fire/[CO₂] interactions. We then used a demographic model of the effects of fire on mesic savanna trees to test the importance of grass/tree differences in postburn recovery rates. We adjusted grass and tree growth in the model according to the DGVM output of net primary production at different [CO₂] relative to current conditions. The simulations predicted elimination of trees at [CO₂] typical of the last glacial period (180 ppm) because tree growth rate is too slow (15 years) to grow to a fire‐proof size of ca. 3 m. Simulated grass growth would produce an adequate fuel load for a burn in only 2 years. Simulations of preindustrial [CO₂] (270 ppm) predict occurrence of trees but at low densities. The greatest increase in trees occurs from preindustrial to current [CO₂] (360 ppm). The simulations are consistent with palaeo‐records which indicate that trees disappeared from sites that are currently savannas in South Africa in the last glacial. Savanna trees reappeared in the Holocene. There has also been a large increase in trees over the last 50–100 years. We suggest that slow tree recovery after fire, rather than differential photosynthetic efficiencies in C₃ and C₄ plants, might have been the significant factor in the Late Tertiary spread of flammable grasslands under low [CO₂] because open, high light environments would have been a prerequisite for the spread of C₄ grasses. Our simulations suggest further that low [CO₂] could have been a significant factor in the reduction of trees during glacial times, because of their slower regrowth after disturbance, with fire favouring the spread of grasses.     

Mulch decomposition under agroforestry conditions in a sub-humid tropical savanna processes and influence of perennial plants

For the effective use of mulch materials in tropical agriculture and agroforestry knowledge of the speed of decomposition and nutrient release is of primary importance. The transfer of these informations from one site to another requires comparability of the processes of decomposition and their intensity at the two sites. In a litterbag experiment the decomposition and release of main nutrients from leaves and branches of Cajanus cajan (L.) Millsp. were investigated with regard to the underlying physical and biological processes during an 81 days period. To test the influence of perennial plants on the decomposition process, the study was conducted on an agricultural field in 1.1 m, 6.9 m and 14.9 m distance from a tree and hedge band. During the first 11 days leaching was high, especially for N and P (about 50% lost) and K (75–80% lost). After the 11th day consumption of the mulch material by the soil fauna was the dominating process of decomposition. During this phase the perennial plants significantly retarded the decomposition of Cajanus branches, but not leaves, probably by their influence on termite activity. Ca release was also retarded in leaves. After about 6–7 weeks, more than 90% of all main nutrients except Ca had been released from the samples. To minimize nutrient losses from nutrient-rich mulch materials, they should be applied repeatedly in small quantities according to the nutrient demand of the crop.     

Variation in the composition and structure of tropical savannas as a function of rainfall and soil texture along a large-scale climatic gradient in the Northern Territory, Australia

Abstract. Variation in structural and compositional attributes of tropical savannas are described in relation to variation in annual rainfall and soil texture along a subcontinental-scale gradient of rainfall in the wet-dry tropics of the Northern Territory, Australia. Rainfall varies along the gradient from over 1500 mm p.a. in the Darwin region ( c . 12° S) to less than 500 mm in the Tennant Creek region ( c . 18° S). Soils are patchy, and sands, loams and clays may occur in all major districts within the region. We utilized a large data set (1657 quadrats ° 291 woody species; with numerous measured and derived sample variables) covering an area of 0.5 million km². Correlations between floristic composition of woody species and environmental variables were assessed using DCA ordination and vector fitting of environmental variables. Vectors of annual rainfall and soil texture were highly correlated with variation in species composition. Multiple regression analyses incorporating linear and quadratic components of mean annual rainfall and topsoil clay content were performed on three structural attributes (tree height, tree cover, tree basal area) and two compositional attributes (woody species richness, deciduous tree species richness). Tree height declined with decreasing rainfall; cover, basal area, woody species richness and deciduous species richness all declined with decreasing rainfall and increasing soil clay content. Regression models accounted for between 17% and 45% of the variation in the data sets. Variation in other factors such as soil depth, landscape position and recent land-use practices (for which there were no data on an individual quadrat basis) are likely to have contributed to the large residual variation in the data set.