Kristine L. Metzger

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 ∼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 ∼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.

A disease-mediated trophic cascade in the Serengeti and its implications for ecosystem C.

The removal of rinderpest had cascading effects on herbivore populations, fire, tree density, and even ecosystem carbon in the Serengeti ecosystem of East Africa.

Inorganic nitrogen availability after severe stand-replacing fire in the Greater Yellowstone ecosystem

Understanding ecosystem processes as they relate to wildfire and vegetation dynamics is of growing importance as fire frequency and extent increase throughout the western United States. However, the effects of severe, stand-replacing wildfires are poorly understood. We studied inorganic nitrogen pools and mineralization rates after stand-replacing wildfires in the Greater Yellowstone Ecosystem, Wyoming. After fires that burned in summer 2000, soil ammonium concentration peaked in 2001 (33 mg NH4-N· kgsoil−1); soil nitrate increased subsequently (2.7 mg NO3-N·kgsoil−1 in 2003) but was still low. However, annual net ammonification rates were largely negative from 2001 to 2004, indicating ammonium depletion. Thus, although net nitrification rates were positive, annual net nitrogen mineralization (net ammonification plus net nitrification) remained low. Aboveground net primary production (ANPP) increased from 0.25 to 1.6 Mg·ha−1·yr−1 from 2001 to 2004, but variation in ANPP among stands was not related to net nitrogen mineralization rates. Across a broader temporal gradient (stand age zero to >250 yr), negative rates of net annual ammonification were especially pronounced in the first postfire year. Laboratory incubations using 15N isotope pool dilution revealed that gross production of ammonium was reduced and ammonium consumption greatly exceeded gross production during the initial postfire years. Our results suggest a microbial nitrogen sink for several years after severe, stand-replacing fire, confirming earlier hypotheses about postdisturbance succession and nutrient cycling in cold, fire-dominated coniferous forests. Postfire forests in Yellowstone seem to be highly conservative for nitrogen, and microbial immobilization of ammonium plays a key role during early succession.

Evaluating the protection of wildlife in parks: the case of African buffalo in Serengeti

Human population growth rates on the borders of protected areas in Africa are nearly double the average rural growth, suggesting that protected areas attract human settlement. Increasing human populations could be a threat to biodiversity through increases in illegal hunting. In the Serengeti ecosystem, Tanzania, there have been marked declines in black rhino (Diceros bicornis), elephant (Loxodonta africana) and African buffalo (Syncerus caffer) inside the protected area during a period when there was a reduction of protection through anti-poaching effort (1976–1996). Subsequently, protection effort has increased and has remained stable. During both periods there were major differences in population decline and recovery in different areas. The purpose of this paper is to analyse the possible causes of the spatial differences. We used a spatially structured population model to analyze the impacts of three factors—(i) hunting, (ii) food shortage and (iii) natural predation. Population changes were best explained by illegal hunting but model fit improved with the addition of predation mortality and the effect of food supply in areas where hunting was least. We used a GIS analysis to determine variation in human settlement rates and related those rates to intrinsic population changes in buffalo. Buffalo populations in close proximity to areas with higher rates of human settlement had low or negative rates of increase and were slowest to recover or failed to recover at all. The increase in human populations along the western boundary of the Serengeti ecosystem has led to negative consequences for wildlife populations, pointing to the need for enforcement of wildlife laws to mitigate these effects.

Asynchronous food-web pathways could buffer the response of Serengeti predators to El Niño Southern Oscillation

Understanding how entire ecosystems maintain stability in the face of climatic and human disturbance is one of the most fundamental challenges in ecology. Theory suggests that a crucial factor determining the degree of ecosystem stability is simply the degree of synchrony with which different species in ecological food webs respond to environmental stochasticity. Ecosystems in which all food-web pathways are affected similarly by external disturbance should amplify variability in top carnivore abundance over time due to population interactions, whereas ecosystems in which a large fraction of pathways are nonresponsive or even inversely responsive to external disturbance will have more constant levels of abundance at upper trophic levels. To test the mechanism underlying this hypothesis, we used over half a century of demographic data for multiple species in the Serengeti (Tanzania) ecosystem to measure the degree of synchrony to variation imposed by an external environmental driver, the El Niño Southern Oscillation (ENSO). ENSO effects were mediated largely via changes in dry-season vs. wet-season rainfall and consequent changes in vegetation availability, propagating via bottom-up effects to higher levels of the Serengeti food web to influence herbivores, predators and parasites. Some species in the Serengeti food web responded to the influence of ENSO in opposite ways, whereas other species were insensitive to variation in ENSO. Although far from conclusive, our results suggest that a diffuse mixture of herbivore responses could help buffer top carnivores, such as Serengeti lions, from variability in climate. Future global climate changes that favor some pathways over others, however, could alter the effectiveness of such processes in the future.

Anthropogenic stressors influence small mammal communities in tropical East African savanna at multiple spatial scales

Abstract Context. Protection of natural ecosystems undoubtedly safeguards ecological communities, with positive benefits for ecosystem processes and function. However, ecosystems are under threat from anthropogenic stressors that reduce the resilience both of component species and the system as a whole. Aims. To determine how anthropogenic stressors (land use and climate change) could impact the diversity and resilience of a small mammal community in the greater Serengeti ecosystem, an East African savanna comprising Serengeti National Park (SNP) and adjacent agro-ecosystems, at local (SNP) and Africa-wide geographic scales. Methods. We recorded small mammal species in 10 habitats in the greater Serengeti ecosystem, including the agro-ecosystem, over 48 years (1962–2010). We calculated richness and diversity for each habitat type, and used an index of similarity to quantify differences in the community among habitats. Species accumulation curves were also generated for each habitat type. Key results. We recorded 40 species of small mammals in the greater Serengeti ecosystem. At the local scale, restricted habitat types in SNP (each <1% of the total area) made a disproportionately large contribution to diversity. Agro-ecosystems had lower richness and were less likely to contain specialist species. At regional and Africa-wide scales, local endemics were less likely to be recorded in the agro-ecosystem (57% species loss) compared with those with regional (33% loss) or Africa-wide (31%) geographic distributions. Conclusions. At the local scale, the variety of habitats in SNP contributed to overall diversity. However, the ability to maintain this diversity in the adjacent agro-ecosystem was compromised for localised endemics compared with species with Africa-wide ranges. Land use intensification adjacent to SNP and projected changes in rainfall patterns for East Africa under global climate scenarios may compromise the future resilience of the small mammal community in this tropical savanna ecosystem. Implications. The loss of rare or specialised species from protected areas and human-modified ecosystems could be mitigated by: (1) increasing habitat complexity and maintaining specialist habitats in the agro-ecosystem; and (2) creating buffers at the boundary of protected natural ecosystems that accommodate regime shifts in response to climatic change. These measures would increase the resilience of this coupled human–natural savanna ecosystem.

Advances in wildlife ecology and the influence of Graeme Caughley

Graeme Caughley produced substantial advances in our understanding of interactions between large mammalian herbivores and the environments they occupy. The strength of his work lay in the logical approach to answering fundamental questions. While his life work contributed to our understanding of animal population dynamics, it is in the application of his research and ideas that we have greatly advanced the science of conservation biology. Two central legacies of Caughley’s lifelong work are that an understanding of basic science leads to more appropriate management, and that underlying assumptions must be explicitly stated and tested. By arguing that efficient management of ecosystems requires an understanding of the underlying mechanisms, he moved forward the application of basic research to management. Future advances in wildlife conservation must focus on three aspects: (1) the rules for stability in ecosystems, and how humans cause instability; (2) the decline in native habitats, mostly from agriculture, and how to renew and reconstruct them while expanding threatened populations; and (3) how to breed species in captivity, and then reintroduce them as a last line of defence.