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.

Modeling elephant-mediated cascading effects of water point closure.

Wildlife management to reduce the impact of wildlife on their habitat can be done in several ways, among which removing animals (by either culling or translocation) is most often used. There are, however, alternative ways to control wildlife densities, such as opening or closing water points. The effects of these alternatives are poorly studied. In this paper, we focus on manipulating large herbivores through the closure of water points (WPs). Removal of artificial WPs has been suggested in order to change the distribution of African elephants, which occur in high densities in national parks in Southern Africa and are thought to have a destructive effect on the vegetation. Here, we modeled the long-term effects of different scenarios of WP closure on the spatial distribution of elephants, and consequential effects on the vegetation and other herbivores in Kruger National Park, South Africa. Using a dynamic ecosystem model, SAVANNA, scenarios were evaluated that varied in availability of artificial WPs; levels of natural water; and elephant densities. Our modeling results showed that elephants can indirectly negatively affect the distributions of meso-mixed feeders, meso-browsers, and some meso-grazers under wet conditions. The closure of artificial WPs hardly had any effect during these natural wet conditions. Under dry conditions, the spatial distribution of both elephant bulls and cows changed when the availability of artificial water was severely reduced in the model. These changes in spatial distribution triggered changes in the spatial availability of woody biomass over the simulation period of 80 years, and this led to changes in the rest of the herbivore community, resulting in increased densities of all herbivores, except for giraffe and steenbok, in areas close to rivers. The spatial distributions of elephant bulls and cows showed to be less affected by the closure of WPs than most of the other herbivore species. Our study contributes to ecologically informed decisions in wildlife management. The results from this modeling exercise imply that long-term effects of this intervention strategy should always be investigated at an ecosystem scale.

The spatial scaling of habitat selection by African elephants.

1. Understanding and accurately predicting the spatial patterns of habitat use by organisms is important for ecological research, biodiversity conservation and ecosystem management. However, this understanding is complicated by the effects of spatial scale, because the scale of analysis affects the quantification of species-environment relationships. 2. We therefore assessed the influence of environmental context (i.e. the characteristics of the landscape surrounding a site), varied over a large range of scales (i.e. ambit radii around focal sites), on the analysis and prediction of habitat selection by African elephants in Kruger National Park, South Africa. 3. We focused on the spatial scaling of the elephants' response to their main resources, forage and water, and found that the quantification of habitat selection strongly depended on the scales at which environmental context was considered. Moreover, the inclusion of environmental context at characteristic scales (i.e. those at which habitat selectivity was maximized) increased the predictive capacity of habitat suitability models. 4. The elephants responded to their environment in a scale-dependent and perhaps hierarchical manner, with forage characteristics driving habitat selection at coarse spatial scales, and surface water at fine spatial scales. 5. Furthermore, the elephants exhibited sexual habitat segregation, mainly in relation to vegetation characteristics. Male elephants preferred areas with high tree cover and low herbaceous biomass, whereas this pattern was reversed for female elephants. 6. We show that the spatial distribution of elephants can be better understood and predicted when scale-dependent species-environment relationships are explicitly considered. This demonstrates the importance of considering the influence of spatial scale on the analysis of spatial patterning in ecological phenomena.

Determinants of woody cover in African savannas.

Savannas are globally important ecosystems of great significance to human economies. In these biomes, which are characterized by the co-dominance of trees and grasses, woody cover is a chief determinant of ecosystem properties. The availability of resources (water, nutrients) and disturbance regimes (fire, herbivory) are thought to be important in regulating woody cover, but perceptions differ on which of these are the primary drivers of savanna structure. Here we show, using data from 854 sites across Africa, that maximum woody cover in savannas receiving a mean annual precipitation (MAP) of less than approximately 650 mm is constrained by, and increases linearly with, MAP. These arid and semi-arid savannas may be considered 'stable' systems in which water constrains woody cover and permits grasses to coexist, while fire, herbivory and soil properties interact to reduce woody cover below the MAP-controlled upper bound. Above a MAP of approximately 650 mm, savannas are 'unstable' systems in which MAP is sufficient for woody canopy closure, and disturbances (fire, herbivory) are required for the coexistence of trees and grass. These results provide insights into the nature of African savannas and suggest that future changes in precipitation may considerably affect their distribution and dynamics.

Model-based assessment of aspen responses to elk herbivory in Rocky Mountain National Park, USA.

In Rocky Mountain National Park (RMNP), aspen (Populus tremuloides Michx.) has been observed to be declining on elk (Cervus elaphus nelsoni) winter range for many decades. To support elk management decisions, the SAVANNA ecosystem model was adapted to explore interactions between elk herbivory and aspen dynamics. The simulated probability of successful vegetative regeneration for senescent aspen stands declines sharply when elk densities reach levels of 3-5 elk/km2, depending on model assumptions for the seasonal duration of elk foraging activities. For aspen stands with a substantial component of younger trees, the simulated regeneration probability declines more continuously with increasing elk density, dropping below 50% from densities at 8-14 elk/km2. At the landscape scale, simulated aspen regeneration probability under a scenario of extensive seasonal use was little affected by elk population level, when this level was above 300-600 elk (25%-50% current population) over the ca. 107 km2 winter range. This was because elk distribution was highly aggregated, so that a high density of elk occupied certain areas, even at low population levels overall. At approximately current elk population levels (1000-1200 elk), only 35%-45% of senescent aspen stands are simulated as having at least a 90% probability of regeneration, nearly all of them located on the periphery of the winter range. Successful management for aspen persistence on core winter range will likely require some combination of elk population reduction, management of elk distribution, and fencing to protect aspen suckers from elk browsing.

An ecosystem approach to population management of ungulates.

Harvest objectives for wild ungulates have traditionally been based on population models that do not consider ecosystem effects of ungulate herbivory, nor interactions between native and domestic ungulate species. There is a need for ecosystem models to allow wildlife managers to evaluate potential ecosystem effects of management scenarios. The utility of the SAVANNA simulation model for estimating elk population objectives within an ecosystem context was demonstrated for North Park, Colorado, USA. Effects of different elk population levels were evaluated for range condition, elk and cattle forage, elk and cattle condition, forage and condition of mule deer and moose, plant production, and plant community composition. Analyses were based on 30-year simulation runs using variable, historical weather. Another set of analyses utilized stochastic weather patterns. For management scenarios using the historical climate pattern, increasing elk populations caused biomass reductions of palatable plant species, particularly on areas of high winter density, where mean leaf biomass of palatable shrubs declined from 26.97 g/m2 at 0 elk to 20.82 g/m2 at 4000 elk (3.73 elk/km2), a 23% decline. At population levels of 5000 elk (4.68 elk/km2) or greater, elk body condition declined sharply following a severe winter. The availability of palatable browse on critical winter range was likely the limiting factor. However, when random climate patterns were simulated for the same scenarios, the threshold level for density-dependent effects varied with climate, ranging from 2000 to 10,000 elk. We suggest that elk population levels from 4000 to 5000 animals represent a conservative population objective for the North Park elk herd. Also, increasing elk population levels appears to intensify intraspecific competition among elk, far more than interspecific competition with cattle. Resolution of elk-cattle conflicts is likely to be facilitated by managing elk distribution, rather than overall elk population levels.

Changes induced by elk browsing in the aboveground biomass production and distribution of willow (Salix monticola Bebb): their relationships with plant water, carbon, and nitrogen dynamics.

Willows are dominant woody plants of many high-elevation riparian areas of the western USA, and constitute an important food resource for various ungulates, which tend to concentrate in riparian areas. The response of willow to browsing was analyzed in the elk winter range of Rocky Mountain National Park, by considering the effect of elk browsing on Salix monticola Bebb, one of the most common willow species in this area. Unbrowsed and browsed treatments were established during the 1997 growing season (May to October), using eight long-term exclosures built in the fall of 1994. Plants in the browsed treatment were in the areas open to browsing, but they were protected from browsing by small exclosures during the experimental period. Winter browsing by elk induced the following measured responses in plant morphology and development: (1) higher shoot biomass production but similar leaf biomass and leaf area per plant, (2) a lower number of and bigger shoots, (3) a lower number of and bigger leaves, and (4) flower inhibition. In addition, we infer that browsing induces (5) lower belowground allocation and (6) a more negative N balance but a higher soil N uptake. We conclude that elk browsing negatively affects willow even though willow compensate for aboveground biomass removal. Continuous browsing produces long-term changes in willow morphology which constrain plant growth and development. High browsing utilization, as occurred in this experiment, could therefore reduce the competitive ability and survivorship of willow, in particular under drier environmental conditions.