African primates are naturally infected with over 40 different simian immunodeficiency viruses (SIVs), two of which have crossed the species barrier and generated human immunodeficiency virus types 1 and 2 (HIV-1 and HIV-2)1,2. Unlike the human viruses, however, SIVs do not generally cause acquired immunodeficiency syndrome (AIDS) in their natural hosts3. Here we show that SIVcpz, the immediate precursor of HIV-1, is pathogenic in free-ranging chimpanzees. By following 94 members of two habituated chimpanzee communities in Gombe National Park, Tanzania, for over 9 years, we found a 10- to 16-fold higher age-corrected death hazard for SIVcpz-infected (n = 17) compared to uninfected (n = 77) chimpanzees. We also found that SIVcpz-infected females were less likely to give birth and had a higher infant mortality rate than uninfected females. Immunohistochemistry and in situ hybridization of post-mortem spleen and lymph node samples from three infected and two uninfected chimpanzees revealed significant CD4+ T-cell depletion in all infected individuals, with evidence of high viral replication and extensive follicular dendritic cell virus trapping in one of them. One female, who died within 3 years of acquiring SIVcpz, had histopathological findings consistent with end-stage AIDS. These results indicate that SIVcpz, like HIV-1, is associated with progressive CD4+ T-cell loss, lymphatic tissue destruction and premature death. These findings challenge the prevailing view that all natural SIV infections are non-pathogenic and suggest that SIVcpz has a substantial negative impact on the health, reproduction and lifespan of chimpanzees in the wild.

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Territorial behavior is expected to buffer populations against short-term environmental perturbations, but we have found that group living in African lions causes a complex response to long-term ecological change. Despite numerous gradual changes in prey availability and vegetative cover, regional populations of Serengeti lions remained stable for 10- to 20-year periods and only shifted to new equilibria in sudden leaps. Although gradually improving environmental conditions provided sufficient resources to permit the subdivision of preexisting territories, regional lion populations did not expand until short-term conditions supplied enough prey to generate large cohorts of surviving young. The results of a simulation model show that the observed pattern of "saltatory equilibria" results from the lions' grouping behavior.

The conservation of migratory species can be problematic because of their requirements for large protected areas. We investigated this issue by examining the annual movements of the migratory wildebeest, Connochaetes taurinus, in the 25 000 km 2 Serengeti-Mara Ecosystem of Tanzania and Kenya. We used Global Positioning System telemetry to track eight wildebeest during 1999-2000 in relation to protected area status in different parts of the ecosystem. The collared wildebeest spent 90% of their time within well-protected core areas. However, two sections of the wildebeest migration route -the Ikoma Open Area and the Mara Group Ranches -currently receive limited protection and are threatened by poaching or agriculture. Comparison of current wildebeest migration routes to those recorded during 1971-73 indicates that the western buffer zones appear to be used more extensively than in the past. This tentative conclusion has important repercussions for management and needs further study. The current development of community-run Wildlife Management Areas as additional buffer zones around the Serengeti represents an important step in the conservation of this UNESCO World Heritage Site. This study demonstrates that detailed knowledge of movement of migratory species is required to plan effective conservation action.

Theoretical ecology is largely founded on the principle of mass action, in which uncoordinated populations of predators and prey move in a random and well-mixed fashion across a featureless landscape. The conceptual core of this body of theory is the functional response, predicting the rate of prey consumption by individual predators as a function of predator and/or prey densities. This assumption is seriously violated in many ecosystems in which predators and/or prey form social groups. Here we develop a new set of group-dependent functional responses to consider the ecological implications of sociality and apply the model to the Serengeti ecosystem. All of the prey species typically captured by Serengeti lions (Panthera leo) are gregarious, exhibiting nonlinear relationships between prey-group density and population density. The observed patterns of group formation profoundly reduce food intake rates below the levels expected under random mixing, having as strong an impact on intake rates as the seasonal migratory behaviour of the herbivores. A dynamical system model parameterized for the Serengeti ecosystem (using wildebeest (Connochaetes taurinus) as a well-studied example) shows that grouping strongly stabilizes interactions between lions and wildebeest. Our results suggest that social groups rather than individuals are the basic building blocks around which predator-prey interactions should be modelled and that group formation may provide the underlying stability of many ecosystems.

Summary1. Within the rapidly developing field of movement ecology, much attention has been given to studying the movement of individuals within a subset of their population's occupied range. Our understanding of the effects of landscape heterogeneity on animal movement is still fairly limited as it requires studying the movement of multiple individuals across a variety of environmental conditions. Gaining deeper understanding of the environmental drivers of movement is a crucial component of predictive models of population spread and habitat selection and may help inform management and conservation. 2. In Ontario, woodland caribou (Rangifer tarandus caribou) occur along a wide geographical gradient ranging from the boreal forest to the Hudson Bay floodplains. We used highresolution GPS data, collected from 114 individuals across a 450 000 km 2 area in northern Ontario, to link movement behaviour to underlying local environmental variables associated with habitat permeability, predation risk and forage availability. 3. We show that a great deal of observed variability in movement patterns across space and time can be attributed to local environmental conditions, with residual individual differences that may reflect spatial population structure. 4. We discuss our results in the context of current knowledge of movement and caribou ecology and highlight potential applications of our approach to the study of wide-ranging animals.

Habitat quality is typically inferred by assuming a direct relationship between consumer density and resource abundance, although it has been suggested that consumer fitness may be a more accurate measure of habitat quality. We examined density vs. fitness-based measures of habitat quality for lions in the Serengeti National Park, Tanzania. A 40-year average of female reproductive success (yearling cubs per female) was best explained by proximity to river confluences, whereas patterns of productivity (yearling cubs per km(2)) and adult female density (individuals per km(2)) were associated with more general measures of habitat quality and areas of shelter in poor habitat. This suggests that density may not accurately distinguish between high-quality 'source' areas and low-quality sites that merely provide refuges for effectively non-reproductive individuals. Our results indicate that density may be a misleading indicator of real estate value, particularly for populations that do not conform to an ideal free distribution.

Population size and trends of large carnivores are difficult to determine, but are often needed to inform conservation actions. Direct counts maintained over long time periods are extremely difficult to achieve. Indices of population sizes can be used to estimate large carnivore abundances, but are often case-, species-and sitespecific. Here, we test the general applicability of track-based indices to estimate large carnivore abundance. We surveyed 15 306.4 km of roads associated with 339 transects across a wide geographical scale, large range of densities and variable substrates for tracks of African large carnivores. A combined model for all carnivore species on sandy soils serves as a robust approach to predict large carnivore densities. Thus, indices based on track counts can provide useful estimates of carnivore abundance. We found consistent relationships between track densities and the actual carnivore densities, having taken account of substrate.