Savanna tree density, herbivores, and the herbaceous community: bottom-up vs. top-down effects

Herbivores choose their habitats both to maximize forage intake and to minimize their risk of predation. For African savanna herbivores, the available habitats range in woody cover from open areas with few trees to dense, almost-closed woodlands. This variation in woody cover or density can have a number of consequences for herbaceous species composition, cover, and productivity, as well as for ease of predator detection and avoidance. Here, we consider two alternative possibilities: first, that tree density affects the herbaceous vegetation, with concomitant "bottom-up" effects on herbivore habitat preferences; or, second, that tree density affects predator visibility, mediating "top-down" effects of predators on herbivore habitat preferences. We sampled sites spanning a 10-fold range of tree densities in an Acacia drepanolobium-dominated savanna in Laikipia, Kenya, for variation in (1) herbaceous cover, composition, and species richness; (2) wild and domestic herbivore use; and (3) degree of visibility obstruction by the tree layer. We then used structural equation modeling to consider the potential influences that tree density may have on herbivores and herbaceous community properties. Tree density was associated with substantial variation in herbaceous species composition and richness. Cattle exhibited a fairly uniform use of the landscape, whereas wild herbivores, with the exception of elephants, exhibited a strong preference for areas of low tree density. Model results suggest that this was not a response to variation in herbaceous-community characteristics, but rather a response to the greater visibility associated with more open places. Elephants, in contrast, preferred areas with higher densities of trees, apparently because of greater forage availability. These results suggest that, for all but the largest species, top-down behavioral effects of predator avoidance on herbivores are mediated by tree density. This, in turn, appears to have cascading effects on the herbaceous vegetation. These results shed light on one of the major features of the "landscape of fear" in which African savanna herbivores exist.     

Are savannas patch-dynamic systems? A landscape model

Savannas are ecosystems characterized by the coexistence of woody species (trees and bushes) and grasses. Given that savanna characteristics are mainly formed from competition, herbivory, fire, woodcutting, and patchy soil and precipitation characteristics, we propose a spatially explicit model to examine the effects of the above-mentioned parameters on savanna vegetation dynamics in space and time. Furthermore, we investigate the effects of the above-mentioned parameters on tree–bush–grass ratios, as well as the degrees of aggregation of tree–bush–grass biomass. We parameterized our model for an arid savanna with shallow soil depth as well as a mesic one with generally deeper and more variable soil depths. Our model was able to reproduce savanna vegetation characteristics for periods of time over 2000 years with daily updated time steps. According to our results, tree biomass was higher than bush biomass in the arid savanna but bush biomass exceeded tree and grass biomass in the simulated mesic savanna. Woody biomass increased in our simulations when the soil's porosity values were increased (mesic savanna), in combination with higher precipitation. Savanna vegetation varied from open savanna to woodland and back to open savanna again. Vegetation cycles varied over ∼300-year cycles in the arid and ∼220-year cycles in the mesic-simulated savanna. Autocorrelation values indicated that there are both temporal and spatial vegetation cycles. Our model indicated cycling savanna vegetation at the landscape scale, cycles in cells, and patchiness, i.e. patch dynamics.     

Ecological consequences of forest elephant declines for Afrotropical forests

Poaching is rapidly extirpating African forest elephants (Loxodonta cyclotis) from most of their historical range, leaving vast areas of elephant‐free tropical forest. Elephants are ecological engineers that create and maintain forest habitat; thus, their loss will have large consequences for the composition and structure of Afrotropical forests. Through a comprehensive literature review, we evaluated the roles of forest elephants in seed dispersal, nutrient recycling, and herbivory and physical damage to predict the cascading ecological effects of their population declines. Loss of seed dispersal by elephants will favor tree species dispersed abiotically and by smaller dispersal agents, and tree species composition will depend on the downstream effects of changes in elephant nutrient cycling and browsing. Loss of trampling and herbivory of seedlings and saplings will result in high tree density with release from browsing pressures. Diminished seed dispersal by elephants and high stem density are likely to reduce the recruitment of large trees and thus increase homogeneity of forest structure and decrease carbon stocks. The loss of ecological services by forest elephants likely means Central African forests will be more like Neotropical forests, from which megafauna were extirpated thousands of years ago. Without intervention, as much as 96% of Central African forests will have modified species composition and structure as elephants are compressed into remaining protected areas. Stopping elephant poaching is an urgent first step to mitigating these effects, but long‐term conservation will require land‐use planning that incorporates elephant habitat into forested landscapes that are being rapidly transformed by industrial agriculture and logging.     

Tree cover in sub-Saharan Africa: rainfall and fire constrain forest and savanna as alternative stable states

Savannas are known as ecosystems with tree cover below climate-defined equilibrium values. However, a predictive framework for understanding constraints on tree cover is lacking. We present (a) a spatially extensive analysis of tree cover and fire distribution in sub-Saharan Africa, and (b) a model, based on empirical results, demonstrating that savanna and forest may be alternative stable states in parts of Africa, with implications for understanding savanna distributions. Tree cover does not increase continuously with rainfall, but rather is constrained to low (<50%, "savanna") or high tree cover (>75%, "forest"). Intermediate tree cover rarely occurs. Fire, which prevents trees from establishing, differentiates high and low tree cover, especially in areas with rainfall between 1000 mm and 2000 mm. Fire is less important at low rainfall (<1000 mm), where rainfall limits tree cover, and at high rainfall (>2000 mm), where fire is rare. This pattern suggests that complex interactions between climate and disturbance produce emergent alternative states in tree cover. The relationship between tree cover and fire was incorporated into a dynamic model including grass, savanna tree saplings, and savanna trees. Only recruitment from sapling to adult tree varied depending on the amount of grass in the system. Based on our empirical analysis and previous work, fires spread only at tree cover of 40% or less, producing a sigmoidal fire probability distribution as a function of grass cover and therefore a sigmoidal sapling to tree recruitment function. This model demonstrates that, given relatively conservative and empirically supported assumptions about the establishment of trees in savannas, alternative stable states for the same set of environmental conditions (i.e., model parameters) are possible via a fire feedback mechanism. Integrating alternative stable state dynamics into models of biome distributions could improve our ability to predict changes in biome distributions and in carbon storage under climate and global change scenarios.     

Disturbance, Diversity, and Invasion: Implications for Conservation

Disturbance is an important component of many ecosystems, and variations in disturbance regime can affect ecosystem and community structure and functioning. The "intermediate disturbance hypothesis" suggests that species diversity should be highest at moderate levels of disturbance. However, disturbance is also known to increase the invasibility of communities. Disturbance therefore poses an important problem for conservation management, Here, we review the effects of disturbances such as fire grazing, soil disturbance and nutrient addition on plant species diversity and invasion with particular emphasis on grassland vegetation. Individual components of the disturbance regime can have marked effects on species diversity, but it is often modifications of the existing regime that have the largest influence. Similarly, disturbance can enhance invasion of natural communities, but frequently it is the interaction between different disturbances that has the largest effect. The natural disturbance regime is now unlikely to persist within conservation areas since fragmentation and human intervention have usually modified physical and biotic conditions. Active management decisions must now be made on what disturbance regime is required and this requires decisions on what species are to be encouraged or discouraged.     

Grass feedbacks on fire stabilize savannas

Savannas commonly consist of a discontinuous cover of overstory trees and a groundcover of grasses. Savanna models have previously demonstrated that vegetation feedbacks on fire frequency can limit the density of overstory trees, thereby maintaining savannas. Positive feedbacks of either savanna trees alone or trees and grasses together on fire frequency have been shown to result in a stable savanna equilibrium. Grass feedbacks on fire frequency, in contrast, have resulted in stable equilibria in either a grassland or forest state, but not in a savanna. These results, however, were derived from a system of differential equations that assumes that fire occurrence is strictly deterministic and that vegetation losses due to fire are continuous in time. We develop an alternative formulation of the grass-fire feedback model that assumes that fires are discrete and occur stochastically in time to examine the influence of these assumptions on the predicted state of the system. We show that incorporating fire as a discrete event can produce a recurring temporal refuge in which both grass and trees co-occur in a stable, bounded savanna. In our model, tree abundance is limited without invoking demographic bottlenecks in the transition from fire-sensitive to fire-resistant life history stages. An increasing strength of grass feedback on fire results in regular, predictable fires, which suggests that the system can also be modeled using a set of difference equations. We implement this discrete system using modified Leslie/Gower difference equations and demonstrate the existence of a bounded savanna state in this model framework. Our results confirm the potential for grass feedbacks to result in stable savannas, and indicate the importance of modeling fire as a discrete event rather than as a loss rate that is continuous in time.     

Effects of four decades of fire manipulation on woody vegetation structure in Savanna

The amount of carbon stored in savannas represents a significant uncertainty in global carbon budgets, primarily because fire causes actual biomass to differ from potential biomass. We analyzed the structural response of woody plants to long-term experimental burning in savannas. The experiment uses a randomized block design to examine fire exclusion and the season and frequency of burn in 192 7-ha experimental plots located in four different savanna ecosystems. Although previous studies would lead us to expect tree density to respond to the fire regime, our results, obtained from four different savanna ecosystems, suggest that the density of woody individuals was unresponsive to fire. The relative dominance of small trees was, however, highly responsive to fire regime. The observed shift in the structure of tree populations has potentially large impacts on the carbon balance. However, the response of tree biomass to fire of the different savannas studied were different, making it difficult to generalize about the extent to which fire can be used to manipulate carbon sequestration in savannas. This study provides evidence that savannas are demographically resilient to fire, but structurally responsive.     

Fire and forest landscapes in the Georgia Piedmont: an assessment of spatial modeling assumptions

Landscape simulation models are widely used to study the behavior of ecological systems. As computing power has increased, these models have become more complex and incorporated more realistic spatial representations of landscape patterns and ecological processes. The goal of this research was to examine the sensitivity of simulated landscape patterns to fundamental spatial modeling assumptions. The LANDIS simulator was parameterized for forests of the Georgia Piedmont and used to model landscape-scale community dynamics at fire return intervals from 20 to 100 years. A base scenario incorporating localized seed dispersal along with landform-related variation in species establishment rates and disturbance regimes was contrasted with three alternative scenarios. The uniform habitat scenario applied the same set of species establishment coefficients across all landforms. The uniform dispersal scenario removed the effects of seed source abundance and pattern on species establishment. The uniform disturbance scenario assumed identical disturbance regimes on all landforms.At the shortest fire return intervals, fire severities were low and the stand age distribution was dominated by older forests. At longer fire return intervals, fire severities were high and the stand age distribution was skewed toward younger forests. Species composition generally followed a gradient from fire-resistant species at short fire return intervals to fire-sensitive species at longer fire return intervals. However, some species exhibited bimodal distributions with high abundances at both short and long fire return intervals. Landscape responses to fire were similar in the uniform habitat scenario and the base scenario. Communities were less sensitive to fire return interval and had more fire-sensitive species in the uniform dispersal scenario than in the base scenario. Species composition in the uniform disturbance scenario was similar to the base scenario for the longest fire-intervals, but was more sensitive to changes in the fire regime at shorter fire return intervals. In models of Piedmont forest landscapes, accurate spatial representations of dispersal and fire regime heterogeneity are essential for predicting landscape-scale species composition under changing fire regimes. In contrast, the precise spatial representation of species–habitat relationships may be considerably less important.     

Megafaunal Extinctions: The Conservation Message from 11,000 Years B.P.

Abstract: At the end of the Pleistocene, the Americas, northern Eurasia: and Australia experienced a vast decline in large mammal diversity, while Africa and tropical Asia were hardly affected. The elimination of the megaberbivores (animals weighing >1000 kg, probably by human predation, removed the vegetation impact of these species. The resultant reduction in habitat mosaic diversity and in forage quality probably precipitated the extinctions of lesser large mammalian species. Surviving megaherbivores in the form of elephants and rhinoceroses are currently being exterminated from many African conservation areas. African savanna ecosystems could prove more resistant to species losses than north temperate ecosystems, because geomorphic factors plus low and erratic rainfall enhance spatial heterogeneity and vegetation quality independently of large herbivore impact Nevertheless, the history of the Hluhluwe Game Reserve in South Africa suggests that certain African ecosystems may become susceptible to an inexorable decline in populations of some large herbivores following the extermination of elephants. If elephants and rhinoceroses cannot be conserved active habitat manipulation will be needed to retain a diverse fauna of large mammals in such regions.     

The consequences of poaching and anthropogenic change for forest elephants

Poaching has devastated forest elephant populations (Loxodonta cyclotis), and their habitat is dramatically changing. The long‐term effects of poaching and other anthropogenic threats have been well studied in savannah elephants (Loxodonta africana), but the impacts of these changes for Central Africa's forest elephants have not been discussed. We examined potential repercussions of these threats and the related consequences for forest elephants in Central Africa by summarizing the lessons learned from savannah elephants and small forest elephant populations in West Africa. Forest elephant social organization is less known than the social organization of savannah elephants, but the close evolutionary history of these species suggests that they will respond to anthropogenic threats in broadly similar ways. The loss of older, experienced individuals in an elephant population disrupts ecological, social, and population parameters. Severe reduction of elephant abundance within Central Africa's forests can alter plant communities and ecosystem functions. Poaching, habitat alterations, and human population increase are probably compressing forest elephants into protected areas and increasing human–elephant conflict, which negatively affects their conservation. We encourage conservationists to look beyond documenting forest elephant population decline and address the causes of these declines when developing conversation strategies. We suggest assessing the effectiveness of the existing protected‐area networks for landscape connectivity in light of current industrial and infrastructure development. Longitudinal assessments of the effects of landscape changes on forest elephant sociality and behavior are also needed. Finally, lessons learned from West African elephant population loss and habitat fragmentation should be used to inform strategies for land‐use planning and managing human–elephant interactions.     

Are trade-offs in plant resprouting manifested in community seed banks?

Trade-offs in allocation to resprouting vs. seedling regeneration in plants are predicted to occur along fire disturbance gradients. Increased resprouting ability should be generally favored in plant communities with a high probability of crown fire return. Hence, communities dominated by resprouters are predicted to have smaller seed banks than those dominated by species killed by fire. We tested whether there were trait shifts in resprouting ability among growth forms (short-lived herbaceous vs. ground-dwelling perennials vs. shrubs) and among communities (rocky outcrop vs. sclerophyll forest) with contrasting probabilities of crown fire return. Resprouting was more common in ground-dwelling perennials and in the sclerophyll forest community with a high probability of crown fire. Soil seed banks were sampled in rocky outcrop and sclerophyll forest communities in recently burned (18 months) and long-since-burned (12 years) locations at interspersed replicated sites. Collected seed banks were treated with orthogonal treatments of fire stimuli or no stimuli, and seedling emergence was measured in controlled conditions. Seed bank composition reflected the pattern of extant vegetation, with resprouting species being more common in the community with a higher probability of crown fire. Overall, however, resprouting species were poorly represented in the seed bank compared to those species killed by fire. Predicted shifts in allocation to seed production were strongly manifested in community seed banks across the disturbance gradient. Fewer species, seedlings, and seedlings per adult emerged from seed banks in the sclerophyll forest. This suggests that the dominance of resprouting species influences recruitment at the community scale. Community patterns in the seed bank also reflected predicted trade-offs with plant size and growth rate. Short-lived species that are killed by fire dominated the seed bank on rocky outcrops, while longer-lived resprouting species were found in low abundance. Life history trade-offs in persistence and regeneration strongly contribute to coexistence patterns between and within communities with contrasting probabilities of fire return.     

A model of surface fire,climate and forest pattern in the Sierra Nevada,California

A spatially explicit forest gap model was developed for the Sierra Nevada, California, and is the first of its kind because it integrates climate, fire and forest pattern. The model simulates a forest stand as a grid of 15×15 m forest plots and simulates the growth of individual trees within each plot. Fuel inputs are generated from each individual tree according to tree size and species. Fuel moisture varies both temporally and spatially with the local site water balance and forest condition, thus linking climate with the fire regime. Fires occur as a function of the simulated fuel loads and fuel moisture, and the burnable area is simulated as a result of the spatially heterogeneous fuel bed conditions. We demonstrate the model’s ability to couple the fire regime to both climate and forest pattern. In addition, we use the model to investigate the importance of climate and forest pattern as controls on the fire regime. Comparison of model results with independent data indicate that the model performs well in several areas. Patterns of fuel accumulation, climatic control of fire frequency and the influence of fuel loads on the spatial extent of fires in the model are particularly well-supported by data. This model can be used to examine the complex interactions among climate, fire and forest pattern across a wide range of environmental conditions and vegetation types. Our results suggest that, in the Sierra Nevada, fuel moisture can exert an important control on fire frequency and this control is especially pronounced at sites where most of the annual precipitation is in the form of snow. Fuel loads, on the other hand, may limit the spatial extent of fire, especially at elevations below 1500 m. Above this elevation, fuel moisture may play an increasingly important role in limiting the area burned.     

Coexistence between People and Elephants in African Savannas

Abstract: The decline in the range and numbers of elephants as a result of expanding human activity in Africa is recognized as one of the continent's more serious conservation problems. Understanding the relationship between human settlement patterns and elephant abundance is fundamental to predicting the viability of elephant populations. The prevailing model of human-elephant interaction predicts a negative linear relationship between rising human density and declining elephant density at a coarse (national or subcontinental) scale. Using observed elephant densities and human population data, we tested this prediction in a study area of 15,000 km² in northwestern Zimbabwe. The results did not fit a linear model. Elephant and human coexistence occurs at various levels of human density, up to a threshold of human density beyond which elephant populations disappear. This threshold seems to be related to a particular stage in the process of agriculturally transformed land becoming spatially dominant over the natural woodland that constitutes elephant habitat. Within the contexts of conservation and sustainable development in African savannas, investigating spatial relationships between elephant and human abundance should be a priority topic for future research.     

Understanding the effects of rainfall on elephant-vegetation interactions around waterholes

The distribution of surface water affects herbivore-vegetation interactions in arid and semi-arid regions. Limited access to surface water typically results in the emergence of vegetation gradients around natural and artificial water sources. In particular, African elephants can create large-scale gradients of woody vegetation. Understanding the dynamics of these gradients is of particular importance for the conservation of other, less mobile herbivores that depend on woody vegetation in areas close to water. While rainfall is known to be a key determinant of herbivore-vegetation interactions in dry areas, we only have limited understanding on how it impacts woody vegetation gradients around waterholes. To address this problem, we developed a deterministic simulation model that describes the interplay of rainfall, elephants and woody vegetation in the vicinity of waterholes. The model is based on elephant telemetry data and the ecological conditions in Etosha National Park (ENP), Namibia. We found that decreasing amounts of rainfall led to an increased degradation of woody vegetation, which was particularly severe in areas close to water. Based on this result we conclude that low rainfall was an important driver of recently observed patterns of vegetation degradation in ENP. More generally, rainfall appears to be a key factor that determines elephant-vegetation interactions and thus dynamics of woody vegetation gradients around waterholes. Using long-term rainfall data from ENP, we also demonstrate that an increase in the number of water sources during periods of low rainfall can mitigate the destructive impact of elephants in areas close to water. However, more research is required to assess the sustainability and effectiveness of rainfall-adapted strategies of artificial water provisioning in more detail. In particular it is important to investigate potential effects on elephant population dynamics.     

A spatially explicit model to predict future landscape composition of aspen woodlands under various management scenarios

Quaking aspen (Populus tremuloides) is declining across the western United States. Aspen habitats are among the most diverse plant communities in this region and loss of these habitats can result in shifts in biodiversity, productivity, and hydrology across a range of spatial scales. Western aspen occurs on the majority of sites seral to conifer species, and long-term maintenance of these aspen woodlands requires periodic fire. Over the past century, fire intervals, extents, and intensities have been insufficient to regenerate aspen stands at historic rates; however the effects of various fire regimes and management scenarios on aspen vegetation dynamics at broad spatial and temporal scales are unexplored. Here we use field data, remotely sensed data, and fire atlas information to develop a spatially explicit landscape simulation model to assess the effects of current and historic wildfire regimes and prescribed burning programs on landscape vegetation composition across two mountain ranges in the Owyhee Plateau, Idaho. Model outputs depict the future structural makeup and species composition of the landscape at selected time steps under simulated management scenarios. We found that under current fire regimes and in the absence of management activities, loss of seral aspen stands will continue to occur over the next two centuries. However, a return to historic fire regimes (burning 12–14% of the modeled landscape per decade) would maintain the majority of aspen stands in early and mid seral woodland stages and minimizes the loss of aspen. A fire rotation of 70–80 years was estimated for the historic fire regime while the current fire regime resulted in a fire rotation of 340–450 years, underscoring the fact that fire is currently lacking in the system. Implementation of prescribed burning programs, treating aspen and young conifer woodlands according to historic fire occurrence probabilities, are predicted to prevent conifer dominance and loss of aspen stands.     

N biogeochemistry and cycling in two well-drained savannas: a comparison between the Orinoco Basin (Llanos,Venezuela) and Ivory Coast (western Africa)

When a savanna burns, a decline in the input of organic matter and nutrients to the soil occurs. However, the existence of recurrent fires is a natural condition and N depletion by fire is not incompatible with the existence of savannas per se. Consequently, savanna vegetations have evolved under fires, implying a near to steady-state N budget. In some Australian and African ecosystems, N fixation appears to be insufficient to replace losses inducing soil-N depletion, whereas in neotropical Llanos and western African savannas, the N-fixation and precipitation seem to be enough to maintain production despite fires. This review presents information about well-drained savannas, namely Trachypogon savannas in Orinoco Llanos and Andropogonae savannas in Ivory Coast. The sites present similarities in climate and fire regime differing in soil parent material. A report on N budgets is presented. The budget was positive, since losses seem to be balanced by inputs in rainfall and biological fixation. Uncertainties in flux measures indicate that more work on those aspects are required. Results indicate that the estimated amounts of dinitrogen fixation and deposition seem to be enough to maintain plant production. Data support the hypothesis that the vegetation in the natural savanna has evolved under fire constraint.     

Longleaf pine (Pinus palustris) and hardwood dynamics in a fire-maintained ecosystem: A simulation approach

Longleaf pine (Pinus palustris) savannas of the southeastern U.S. represent an archetype of a fire dependent ecosystem. They are known to have very short fire return intervals (∼1-3 years) that perpetuate understory plant diversity (up to 50 species m⁻²), support pine recruitment, and suppress fire sensitive hardwoods. Understanding the relationships that regulate longleaf and southern hardwoods is especially critical. With decreased fire frequency, insufficient intensity, or lack of underground competition, a woody mid-story rapidly develops, dominated by fire sensitive trees and shrubs that in-turn suppress more fire dependent species (including pine seedlings). This may occur in forest gaps, where pine-needle abundance is diminished, reducing fire spread potential. The interactions between longleaf pine, hardwoods, forest fuels, and fire frequency are complex and difficult to understand spatially. The objective of this study was to develop a spatially explicit longleaf pine-hardwood stochastic simulation model (LLM), incorporating tree demography, plant competition, and fuel and fire characteristics. Data from two longleaf pine study sites were used to develop and evaluate the model with the goal to incorporate simple site-specific calibration parameters for model versatility. Specific model components included pine seed masting, hardwood clonal sprouting, response to fire (re-sprouting, mortality), and tree density driven competition effects. LLM spatial outputs were consistent with observed forest gap dynamics associated with pine seedling establishment and hardwood encroachment. Changes in fire frequency (i.e., fire probability = 0.35-0.05) illustrated a shift in community structure from longleaf pine dominated to a hardwood dominated community. This approach to assessing model response may be useful in characterizing longleaf ecosystem resilience, especially at intermediate fire frequencies (e.g., 0.15) where the community may be sensitive to small changes in the fire regime. Height distributions and population densities were similar to in situ findings (field and LIDAR data) for both study sites. Height distributions output by the LLM illustrated fluctuations in population structure. The LLM was especially useful in determining knowledge gaps associated with fuel and fire heterogeneity, plant-plant interactions, population structure and its temporal fluctuations, and hardwood demography. This is the first known modeling work to simulate interactions between longleaf pine and hardwoods and provides a foundation for further studies on fire and forest management, especially in relation to ecological forestry practices, restoration, and site-specific applications.     

Geochemical and mineralogical characteristics of elephant geophagic soils in Udawalawe National Park,Sri Lanka

Geophagy or deliberate ingestion of soils was observed among Asian elephants (Elephas maximus) in the Udawalwe National Park, Sri Lanka, for several years. The geochemical and mineralogical composition of the clayey soil layers which are purposefully selected and eaten by elephants in the park were studied, in order to identify the possible reasons for elephant geophagy. The concentrations of major and trace elements were determined by means of X-ray fluorescence spectrometry in 21 soil samples from eight geophagic sites and six soil samples collected from four non-geophagic sites. The mineralogical composition of selected soil samples was investigated using X-ray diffractometry (XRD). These geochemical analyses revealed that geophagic soils in the study areas are deeply weathered and that most of the elements are leached from the soil layers under extreme weathering conditions. The XRD data showed that the soils of the area consisted mainly quartz, feldspar, and the clay minerals kaolinite, Fe-rich illite, and smectite. Although no significant geochemical differences were identified between geophagic and non-geophagic soils, a clear difference was observed in their clay mineralogical content. Soils eaten by elephants are richer in kaolinite and illite than non-geophagic soils, which contain a higher amount of smectite. It is suggested that elephants in Udawalawe National Park ingest soils mainly not to supplement the mineral contents of their forage but to detoxify unpalatable compounds in their diet.     

Reassessment of the Use of Fire as a Management Tool in Deciduous Forests of Eastern North America

Prescribed burning is increasingly being used in the deciduous forests of eastern North America. Recent work suggests that historical fire frequency has been overestimated east of the prairie–woodland transition zone, and its introduction could potentially reduce forest herb and shrub diversity. Fire‐history recreations derived from sedimentary charcoal, tree fire scars, and estimates of Native American burning suggest point‐return times ranging from 5–10 years to centuries and millennia. Actual return times were probably longer because such records suffer from selective sampling, small sample sizes, and a probable publication bias toward frequent fire. Archeological evidence shows the environmental effect of fire could be severe in the immediate neighborhood of a Native American village. Population density appears to have been low through most of the Holocene, however, and villages were strongly clustered at a regional scale. Thus, it appears that the majority of forests of the eastern United States were little affected by burning before European settlement. Use of prescribed burning assumes that most forest species are tolerant of fire and that burning will have only a minimal effect on diversity. However, common adaptations such as serotiny, epicormic sprouting, resprouting from rhizomes, and smoke‐cued germination are unknown across most of the deciduous region. Experimental studies of burning show vegetation responses similar to other forms of disturbance that remove stems and litter and do not necessarily imply adaptation to fire. The general lack of adaptation could potentially cause a reduction in diversity if burning were introduced. These observations suggest a need for a fine‐grained examination of fire history with systematic sampling in which all subregions, landscape positions, and community types are represented. Responses to burning need to be examined in noncommercial and nonwoody species in rigorous manipulative experiments. Until such information is available, it seems prudent to limit the use of prescribed burning east of the prairie–woodland transition zone. Reevaluación del Uso de Fuego como Herramienta de Manejo en Bosques Deciduos de América del Norte     

Molecular genetic and behavioral analysis of social organization in the Asian elephant (Elephas maximus)

We report on the genetic evaluation and behavioral study of social organization in the Asian elephant (Elephas maximus). Although Asian elephants and African elephants (Loxodonta africana) were previously thought to have similar social organizations, our results demonstrate a substantial difference in the complexity and structure of Asian elephant social groupings from that described for African savanna elephants. Photographic cataloging of individuals, radio telemetry, and behavioral observations in Ruhuna National Park, Sri Lanka, enabled us to assign associated females and young to four groups with overlapping ranges. Genetic sampling of individuals from the four groups in Ruhuna National Park and three other groups in surrounding areas, conducted through PCR amplification and sequencing of mitochondrial DNA from dung, supported the matriarchal nature of female groups and the lack of inter-group transfer of females. Behaviorally and genetically, the identified social groups were best described as ”family groups”. We did not find any evidence for the existence of social groups of higher complexity than family groups. Received: 25 March 2000 / Received in revised form: 28 March 2000 / Accepted: 1 April 2000