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Interactions among keystone species: effects of Odontotermes and ungulates on savanna biodiversity Alison K. Brody, University of Vermont; Todd M. Palmer, Univ. of Florida; Dan Doak, University of California, Santa Cruz, USA. Preliminary Results

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Interactions among keystone species: effects of Odontotermes and ungulates on savanna biodiversity

Alison K. Brody, University of Vermont; Todd M. Palmer, Univ. of Florida; Dan Doak, University of California, Santa Cruz, USA

Preliminary Results

Important soil characteristics are strongly positively correlated with proximity to termite mounds (Fig. 2), and invertebrates are 3x more abundant on termite mounds than off (Fig. 3).


Understanding the links between species and the structure and function of ecological communities has long been a primary goal of ecologists. As human modification of the biosphere intensifies, understanding these linkages has become a fundamental conservation concern as well (Chapin et al. 1997, Walker & Noy-Meir 1982). Species differ in the extent to which they impact the communities they inhabit. In particular, species that create, modify, or maintain physical habitats (“physical ecosystem engineers” sensu Jones et al. 1994), may strongly influence local and regional biodiversity and species interactions (Gutterman et al. 1990, Folgarait 1998, Williams et al. 1986, Doak and Loso 2003, Castilla et al. 2004).

Perhaps because their presence across landscapes is so widespread and conspicuous (Fig. 1), the importance of termites to the structure and function of African savanna communities is almost cliché. Despite this, few hard data exist on the magnitude of and mechanisms for community-wide effects of termites in these unique and threatened ecosystems. Here, we propose manipulative experiments to assess the influence of an abundant termite on a widespread model ecosystem, the “black cotton savanna” of East Africa and how these effects are mediated by interactions with another commonly identified “keystone” guild, grazing ungulates (Sinclair & Arcese 1995, Jackson 1997, Mduma et al. 1999, Larter et al. 2000, Gordon et al. 2004).

Fig. 2. Mean % organic C at 5-m intervals radiating from mound centers (N=4 mounds). Vertical line = mound edge.



Fig. 3. a) Mean number of insects caught in pitfall traps on mounds vs. > 10 m from mound edges.

Fig. 1. An Odontotermes mound in fore-ground. Distinguishable from matrix by green vegetation with a distinct margin and absence of trees and shrubs.

  • Ongoing Experiments

  • We will quantify patterns by sampling plants and invertebrates on and off termite mounds. We will remove termites from some mounds to measure how termite activity per se produces the patterns observed.

  • 2) We will conduct an experiment in which termites, native ungulates and cattle are allowed or excluded to partition each of their effects on plant and invertebrate communities.

  • 3) We will examine the patterns of plant and invertebrate diversity on and off mounds across a range of cattle grazing intensity to examine how cattle and termites interact in producing patterns observed.


Hypothesis I: Termite mound building activity leads to predictable patterns of density and diversity of plant, invertebrate and vertebrate community by creating a mosaic of soil fertility and soil structure. The landscape-scale effect of these islands of fertility is to enhance and structure biodiversity.

Hypothesis II: High nutrient levels of termite mound soils are maintained and/or enhanced by preferential use of mounds by large herbivores that deposit nutrients in dung and urine. This positive feedback loop contributes to nutrient heterogeneity both directly through deposition, and indirectly through enhancement of nitrogen mineralization rates.

Hypothesis III: High levels of cattle grazing will reduce termite populations and at high numbers cattle function as exporters of nutrients from mounds. These negative effects, in turn, cascade throughout the plant, invertebrate, and vertebrate communities to ultimately impact the diversity and stability of savanna ecosystems.

Scientific Significance

Termite mounds appear to be a major organizing feature of the black-cotton ecosystem (Young et al. 1998, Palmer 2003) and play important roles in other semi-arid savannas as well (Scholes & Walker 1993). One of the most visible and striking characteristics of termites is the degree to which their subterranean mounds are associated with above-ground heterogeneity in plant communities. In East Africa, habitat heterogeneity has consequences for plants and animals at all levels of ecological organization (Sinclair & Arcese 1995). Understanding the forces that generate and maintain this patchiness has direct bearing on our ability to manage these systems (Walker & Menaut 1988, Sinclair 1996).

Broader Significance

A primary goal of the project is to train both U.S. and Kenyan researchers and students and to interact with the broader, Kenyan public and surrounding land-owners and ranch managers. The project will support a Post-doc, one African Master’s student, provide partial support for one U.S. graduate student, employ three full-time technicians (two in Kenya and one in the US) and provide part-time support to at least 2 additional Kenyan field assistants.


We thank Dr. Nick Georgiadis, Director, and the staff at Mpala Research Center for supporting this work and making it all possible. We give special thanks to John Lemboi and Wilson Ndurito for help in the field. The project is sponsored by NSF DEB-0519223.

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