Long-term historical and projected herbivore population dynamics in Ngorongoro crater, Tanzania.

The Ngorongoro Crater is an intact caldera with an area of approximately 310 km2 located within the Ngorongoro Conservation Area (NCA) in northern Tanzania. It is known for the abundance and diversity of its wildlife and is a UNESCO World Heritage Site and an International Biosphere Reserve. Long term records (1963-2012) on herbivore populations, vegetation and rainfall made it possible to analyze historic and project future herbivore population dynamics. NCA was established as a multiple use area in 1959. In 1974 there was a perturbation in that resident Maasai and their livestock were removed from the Ngorongoro Crater. Thus, their pasture management that was a combination of livestock grazing and fire was also removed and 'burning' stopped being a regular occurrence until it was resumed in 2001 by NCA management. The Maasai pasture management would have selected for shorter grasses and more palatable species. Vegetation mapping in 1966-1967 recorded predominately short grasslands. Subsequent vegetation mapping in the crater in 1995 determined that the grassland structure had changed such that mid and tall grasses were dominant. After removal of the Maasai pastoralists from the Ngorongoro Crater in 1974, there were significant changes in population trends for some herbivore species. Buffalo, elephant and ostrich numbers increased significantly during 1974-2012. The zebra population was stable from 1963 to 2012 whereas population numbers of five species declined substantially between 1974 and 2012 relative to their peak numbers during 1974-1976. Grant's and Thomson's gazelles, eland, kongoni, and waterbuck (wet season only) declined significantly in the Crater in both seasons after 1974. In addition, some herbivore species were consistently more abundant inside the Crater during the wet than the dry season. This pattern was most evident for the large herbivore species requiring bulk forage, i.e., buffalo, eland, and elephant. Even with a change in grassland structure, total herbivore biomass remained relatively stable from 1963 to 2012, implying that the crater has a stable carrying capacity. Analyses of rainfall indicated that there was a persistent cycle of 4.83 years for the annual component. Herbivore population size was correlated with rainfall in both the wet and dry seasons. The relationships established between the time series of historic animal counts in the wet and dry seasons and lagged wet and dry season rainfall series were used to forecast the likely future trajectories of the wet and dry season population size for each species under three alternative climate change scenarios.

The impact of a pathogenic bacterium on a social carnivore population.

1. The long-term ecological impact of pathogens on group-living, large mammal populations is largely unknown. We evaluated the impact of a pathogenic bacterium, Streptococcus equi ruminatorum, and other key ecological factors on the dynamics of the spotted hyena Crocuta crocuta population in the Ngorongoro Crater, Tanzania. 2. We compared key demographic parameters during two years when external signs of bacterial infection were prevalent ('outbreak') and periods of five years before and after the outbreak when such signs were absent or rare. We also tested for density dependence and calculated the basic reproductive rate R(0) of the bacterium. 3. During the five pre-outbreak years, the mean annual hyena mortality rate was 0.088, and annual population growth was relatively high (13.6%). During the outbreak, mortality increased by 78% to a rate of 0.156, resulting in an annual population decline of 4.3%. After the outbreak, population size increased moderately (5.1%) during the first three post-outbreak years before resuming a growth similar to pre-outbreak levels (13.9%). We found no evidence that these demographic changes were driven by density dependence or other ecological factors. 4. Most hyenas showed signs of infection when prey abundance in their territory was low. During the outbreak, mortality increased among adult males and yearlings, but not among adult females - the socially dominant group members. These results suggest that infection and mortality were modulated by factors linked to low social status and poor nutrition. During the outbreak, we estimated R(0) for the bacterium to be 2.7, indicating relatively fast transmission. 5. Our results suggest that the short-term 'top-down' impact of S. equi ruminatorum during the outbreak was driven by 'bottom-up' effects on nutritionally disadvantaged age-sex classes, whereas the longer-term post-outbreak reduction in population growth was caused by poor survival of juveniles during the outbreak and subsequent poor recruitment of breeding females. These results suggest synergistic effects of 'bottom-up' and 'top-down' processes on host population dynamics.