Large-scale impacts of herbivores on the structural diversity of African savannas

African savannas are undergoing management intensification, and decision makers are increasingly challenged to balance the needs of large herbivore populations with the maintenance of vegetation and ecosystem diversity. Ensuring the sustainability of Africa's natural protected areas requires information on the efficacy of management decisions at large spatial scales, but often neither experimental treatments nor large-scale responses are available for analysis. Using a new airborne remote sensing system, we mapped the three-dimensional (3-D) structure of vegetation at a spatial resolution of 56 cm throughout 1640 ha of savanna after 6-, 22-, 35-, and 41-year exclusions of herbivores, as well as in unprotected areas, across Kruger National Park in South Africa. Areas in which herbivores were excluded over the short term (6 years) contained 38%–80% less bare ground compared with those that were exposed to mammalian herbivory. In the longer-term (> 22 years), the 3-D structure of woody vegetation differed significantly between protected and accessible landscapes, with up to 11-fold greater woody canopy cover in the areas without herbivores. Our maps revealed 2 scales of ecosystem response to herbivore consumption, one broadly mediated by geologic substrate and the other mediated by hillslope-scale variation in soil nutrient availability and moisture conditions. Our results are the first to quantitatively illustrate the extent to which herbivores can affect the 3-D structural diversity of vegetation across large savanna landscapes.

[1]  A. Sinclair,et al.  Protected areas as biodiversity benchmarks for human impact: agriculture and the Serengeti avifauna , 2002, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[2]  Gregory P Asner,et al.  Hyperspectral and LiDAR remote sensing of fire fuels in Hawaii Volcanoes National Park. , 2008, Ecological applications : a publication of the Ecological Society of America.

[3]  M. Fenton,et al.  ELEPHANTS, WOODLANDS AND BIODIVERSITY IN SOUTHERN AFRICA , 1997 .

[4]  R. Macarthur,et al.  On Bird Species Diversity , 1961 .

[5]  C. C. Grant,et al.  The importance of nutrient hot-spots in the conservation and management of large wild mammalian herbivores in semi-arid savannas , 2006 .

[6]  R. Ben-Shahar Woodland dynamics under the influence of elephants and fire in Northern Botswana , 1996, Vegetatio.

[7]  W. Cohen,et al.  Lidar Remote Sensing for Ecosystem Studies , 2002 .

[8]  A. J. Belsky,et al.  Spatial and temporal landscape patterns in arid and semi-arid African savannas , 1995 .

[9]  H. K. Beuchner,et al.  Vegetation Change Induced by Elephants and Fire in Murchison Falls National Park, Uganda , 1961 .

[10]  M. Rietkerk,et al.  EFFECTS OF FIRE AND HERBIVORY ON THE STABILITY OF SAVANNA ECOSYSTEMS , 2003 .

[11]  D. Rubenstein,et al.  Herbivore-initiated interaction cascades and their modulation by productivity in an African savanna , 2007, Proceedings of the National Academy of Sciences.

[12]  Roberta E. Martin,et al.  Carnegie Airborne Observatory: in-flight fusion of hyperspectral imaging and waveform light detection and ranging for three-dimensional studies of ecosystems , 2007 .

[13]  R. Naiman,et al.  Large Animals and System-Level Characteristics in River Corridors Implications for river management , 1997 .

[14]  Holly T. Dublin,et al.  Elephants and Fire as Causes of Multiple Stable States in the Serengeti-Mara Woodlands , 1990 .

[15]  R. Noss Indicators for Monitoring Biodiversity: A Hierarchical Approach , 1990 .

[16]  N. Grimm,et al.  Nutrient Vectors and Riparian Processing: A Review with Special Reference to African Semiarid Savanna Ecosystems , 2007, Ecosystems.

[17]  R. Barnes Woodland changes in Ruaha National Park (Tanzania) between 1976 and 1982 , 1985 .

[18]  R. Paine,et al.  Ungulate herbivory: Indirect effects cascade into the treetops , 2007, Proceedings of the National Academy of Sciences.

[19]  G. Asner,et al.  Spectral unmixing of vegetation, soil and dry carbon cover in arid regions: Comparing multispectral and hyperspectral observations , 2002 .

[20]  W. Bond,et al.  Ecology of sprouting in woody plants: the persistence niche. , 2001, Trends in ecology & evolution.

[21]  S. Milton,et al.  Large trees, fertile islands, and birds in arid savanna , 1999 .

[22]  S. Moe,et al.  The Return of the Giants: Ecological Effects of an Increasing Elephant Population , 2004, Ambio.

[23]  G. Asner,et al.  Grazing gradient detection with airborne imaging spectroscopy on a semi-arid rangeland , 2003 .

[24]  H. Biggs,et al.  A new policy for the management of the Kruger National Park's elephant population , 1999 .

[25]  Gregory P. Asner,et al.  Ecosystem structure along bioclimatic gradients in Hawai'i from imaging spectroscopy , 2005 .

[26]  Ignas M. A. Heitkönig,et al.  Trees improve grass quality for herbivores in African savannas , 2007 .

[27]  Kelly K. Caylor,et al.  Determinants of woody cover in African savannas , 2005, Nature.

[28]  A. Watkinson,et al.  Dynamics of shrub encroachment in an African savanna: relative influences of fire, herbivory, rainfall and density dependence , 2001 .

[29]  R. Slotow,et al.  The utilization of large savanna trees by elephant in southern Kruger National Park , 2008, Journal of Tropical Ecology.

[30]  S. Moe,et al.  Vegetation changes during a 36‐year period in northern Chobe National Park, Botswana , 2002 .

[31]  Lalit Kumar,et al.  Spatial Heterogeneity and Irreversible Vegetation Change in Semiarid Grazing Systems , 2002, The American Naturalist.

[32]  Robert J. Scholes,et al.  Trends in savanna structure and composition along an aridity gradient in the Kalahari , 2002 .

[33]  Todd M. Scanlon,et al.  Determining land surface fractional cover from NDVI and rainfall time series for a savanna ecosystem , 2002 .

[34]  S. Levick,et al.  Patch and species specific responses of savanna woody vegetation to browser exclusion , 2008 .

[35]  L. Prior,et al.  The Kruger Experience. Ecology and Management of Savanna Heterogeneity , 2005 .

[36]  H. Olff,et al.  Changes in the woody component of an East African savanna during 25 years , 1999, Journal of Tropical Ecology.

[37]  Monica G. Turner,et al.  Landscape Heterogeneity and Ungulate Dynamics: What Spatial Scales are Important? , 1997 .

[38]  M. Fenton,et al.  Bats and the Loss of Tree Canopy in African Woodlands , 1998 .

[39]  D. Western A half a century of habitat change in Amboseli National Park, Kenya , 2007 .

[40]  R. Scholes,et al.  An African Savanna: Synthesis of the Nylsvley Study. , 1993 .

[41]  R. Biggs,et al.  A biodiversity intactness index , 2005, Nature.

[42]  K. Wiegand,et al.  Multi-scale patterns and bush encroachment in an arid savanna with a shallow soil layer , 2005 .

[43]  C. Canham,et al.  Forest Gaps and Isolated Savanna Trees , 1994 .

[44]  W. Cohen,et al.  Lidar Remote Sensing of the Canopy Structure and Biophysical Properties of Douglas-Fir Western Hemlock Forests , 1999 .

[45]  Charles Breen,et al.  The Kruger Experience: Ecology and Management of Savanna Heterogeneity , 2004, Environmental Conservation.

[46]  K. Rogers,et al.  A Framework for Exploring the Determinants of Savanna and Grassland Distribution , 2006 .

[47]  F. James Rohlf,et al.  Biometry: The Principles and Practice of Statistics in Biological Research , 1969 .

[48]  H. Biggs,et al.  Long-term changes in the woody vegetation of the Kruger National Park, with special reference to the effects of elephants and fire , 1998 .

[49]  S. McNaughton,et al.  FEEDBACKS BETWEEN SOIL NUTRIENTS AND LARGE HERBIVORES IN A MANAGED SAVANNA ECOSYSTEM , 2003 .