Abstract
After rainfall and soils, fire and herbivory are two of the main determinants of savanna ecosystems. Although the interactive effects of fire and herbivores on soil and vegetation are widely acknowledged few studies have addressed these two factors in concert, and none of the studies has focused on the Kalahari sand system. We experimentally studied how annual late dry season fires and grazing affect herbaceous plant species composition, above- and belowground biomass, and soil and grass nutrient concentrations in the nutrient-poor semi-arid Kalahari system in northern Botswana. Four treatments (fire, grazing, fire + grazing, and no-fire–no-grazing) were applied for two consecutive years in the late dry season. Plant species composition was affected by treatment and year. The no-fire–no-grazing treatment was distinctly different from all the other treatments in terms of species composition. Beta diversity was lower on the fire treatment and grazing treatment, but not where fire and grazing were combined. Fire and grazing alone or in combination did not have a substantial effect on biomass, soil and plant nutrients or plant species alpha diversity. Plant nitrogen was the only element that differed between treatments, with high concentrations on all the grazed treatments in the first year and low levels on the fire-alone treatment during the second year. The results show that fire and grazing mainly affect species composition and large-scale biodiversity patterns as indicated by the no-fire–no-grazing treatment being distinctly different from other treatments, suggesting the evolutionary adaptation of this dystrophic Kalahari sand system to herbivory and fire.
Similar content being viewed by others
References
AOAC. 1995. Official methods of analysis. 16th edn. Arlington (VA): Association of Official Analytical Chemists.
Anderson MJ. 2006. Distance-based tests for homogeneity of multivariate dispersions. Biometrics 62:245–53.
Anderson MJ, Ellingsen KE, McArdle BH. 2006. Multivariate dispersion as a measure of beta diversity. Ecol Lett 9:683–93.
Archibald S, Bond WJ, Stock WD, Fairbanks DHK. 2005. Shaping the landscape: fire–grazer interactions in an African savanna. Ecol Appl 15:96–109.
Augustine DJ, McNaughton SJ. 1998. Ungulate effects on the functional species composition of plant communities: herbivore selectivity and plant tolerance. J Wildl Manag 62:1165–83.
Augustine DJ, Frank DA. 2001. Effects of migratory grazers on spatial heterogeneity of soil nitrogen properties in a grassland ecosystem. Ecology 82:3149–62.
Belsky AJ. 1987. Revegetation of natural and human-caused disturbances in the Serengeti National Park, Tanzania. Plant Ecol 70:51–60.
Ben-Shahar R, MacDonald DW. 2002. The role of soil factors and leaf protein in the utilization of mopane plants by elephants in northern Botswana. BMC Ecol 3:1–6.
Bhalotra YPR. 1987. Climate of Botswana. Part II: elements of climate. 1. Rainfall. Department of Meteorological Services. Ministry of Works, Transport and Communications, Gaborone.
Böhm W. 1979. Methods of studying root systems. New York: Springer.
Bond WJ, Van Wilgen BW. 1996. Fire and plants. Population and community biology Series 14. London: Chapman and Hall.
Brockway DG, Gatewood RG, Paris RB. 2002. Restoring fire as an ecological process in shortgrass prairie ecosystems: initial effects of prescribed burning during the dormant and growing seasons. J Environ Manage 65:135–52.
Chao A, Jost L, Chiang SC, Jian YH, Chazdon R. 2008. A two-stage probabilistic approach to multiple-community similarity indices. Biometrics 64:1178–86.
Child G. 1968. An ecological survey of northeastern Botswana. UNDP Report to the government of Botswana, F.A.O. Technical Assistance No. 2563, Rome.
Coetsee C, Bond WJ, February EC. 2010. Frequent fire affects soil nitrogen and carbon in an African savanna by changing woody cover. Oecologia 162:1027–34.
Daubenmire R. 1968. Ecology of fire in grasslands. In: Cragg JB, Ed. Advances in ecological research. New York: Academic Press. p 209–66.
DeBano LF, Neary DG, Ffolliott PF. 1998. Fire’s effects on ecosystems. New York: Wiley. pp 263–83.
De Castro EA, Kauffman JB. 1998. Ecosystem structure in the Brazilian Cerrado: a vegetation gradient of aboveground biomass, root mass and consumption by fire. J Trop Ecol 14:263–83.
DeLuca TH, Zouhar KL. 2000. Effects of selection harvest and prescribed fire on the soil nitrogen status of ponderosa pine forests. For Ecol Manage 138:263–71.
Dougill AJ, Heathwaite AL, Thomas DSG. 1998. Soil water movement and nutrient cycling in semi-arid rangeland: vegetation change and system resilience. Hydrol Process 12:443–59.
Dougill AJ, Thomas AD. 2004. Kalahari sand soils: spatial heterogeneity, biological soil crust and land degradation. Land Degrad Dev 15:233–42.
Dougill AJ, Thomas DSG, Heathwaite AL. 1999. Environmental change in the Kalahari: integrated land degradation studies for nonequilibrium dryland environments. Ann Assoc Am Geogr 89:420–42.
DWNP. 2006. Aerial census of animals in Botswana. Internal report, Department of Wildlife and National Parks, Gaborone.
Enright NJ, Miller BP, Crawford A. 1994. Environmental correlates of vegetation patterns and species richness in the northern Grampians, Victoria. Aust J Ecol 19:159–68.
Frank DA, McNaughton SJ. 1993. Evidence for the promotion of aboveground grassland production by native large herbivores in Yellowstone National Park. Oecologia 96:157–61.
Frank DA, Kuns MM, Guido DR. 2002. Consumer control of grassland plant production. Ecology 83:602–6.
Frost PGH. 1984. The responses and survival of organisms in fire-prone environments. In: de Booysen VP, Tainton NM, Eds. Ecological studies 48: ecological effects of fire in South African ecosystems. Berlin: Springer. p 273–310.
Fuhlendorf SD, Engle DM. 2004. Application of the fire–grazing interaction to restore a shifting mosaic on tallgrass prairie. J Appl Ecol 41:604–14.
Fynn RWS, O’Connor TG. 2000. Effect of stocking rate and rainfall on rangeland dynamics and cattle performance in a semi-arid savanna, South Africa. J Appl Ecol 37:491–507.
Gao Y, Giese M, Lin S, Sattelmacher B, Zhao Y, Brueck H. 2008. Belowground net primary productivity and biomass allocation of a grassland in Inner Mongolia is affected by grazing intensity. Plant Soil 307:41–50.
Gibbs-Russel GE, Watson L, Koekemoer M, Smook L, Barker NP, Anderson HM, Dallwitz MJ. 1990. Grasses of southern Africa. Memoirs of the Botanical survey of South Africa No. 58. Leistner OA, Ed. Pretoria: National Botanic Gardens.
Gignoux J, Clobert J, Menant J. 1997. Alternative fire resistance strategies in savanna trees. Oecologia 110:575–83.
Hobbs NT, Schimel DS, Clenton EO, Ojima DS. 1991. Fire and grazing in the tallgrass prairie: contingent effects on nitrogen budgets. Ecology 72:1374–82.
Hobbs RJ, Huenneke LF. 1992. Disturbance, diversity, and invasion: implications for conservation. Coserv Biol 6:324–37.
Hobbs NT. 1996. Modification of ecosystems by ungulates. J Wildl Manag 60:695–713.
Holdo RM, Mack MC, Arnold SG. 2011. Tree canopies explain fire effects on soil nitrogen, phosphorus and carbon in a savanna ecosystem. Journal of Vegetation Science. doi:10.1111/j.1654-1103.2011.01357.x (early view on-line: 20 October 2011).
Holland EA, Detling JK. 1990. Plant response to herbivory and below ground nitrogen cycling. Ecology 71:1040–9.
Hulbert LC. 1988. Causes of fire effects in tallgrass prairie. Ecology 69:46–58.
Jensen M, Michelsen A, Gashaw M. 2001. Responses in plant, soil inorganic and microbial nutrient pools to experimental fire, ash and biomass addition in a woodland savanna. Oecologia 128:85–93.
Johnson LC, Matchett JR. 2001. Fire and grazing regulate belowground processes in tallgrass prairie. Ecology 82:3377–89.
Jost L. 2008. GST and its relatives do not measure differentiation. Mol Ecol 17(4):015–4026.
Kaye JP, Hart SC. 1998. Ecological restoration alters nitrogen transformations in a ponderosa pine bunchgrass ecosystem. Ecol Appl 8:1052–60.
Kutt AS, Woinarski JCZ. 2007. The effects of grazing and fire on vegetation and the vertebrate assemblage in a tropical savanna woodland in north-eastern Australia. J Trop Ecol 23:95–106.
Lavoie M, Starr G, Mack MC, Martin TA, Gholz HL. 2010. Effects of prescribed fire on understory vegetation, carbon pools, and soil nutrients in a longleaf pine-slash pine forest in Florida. Nat Areas J 30:82–94.
Magurran AE. 2004. Measuring biological diversity. Oxford: Blackwell.
McCune B, Rosentreter R, Ponzetti JM, Shaw DC. 2000. Epiphyte habitats in an old conifer forest in western Washington, USA. Bryologist 103:417–27.
McNaughton SJ. 1985. Ecology of a grazing ecosystem: the Serengeti. Ecol Monogr 55:259–94.
McNaughton SJ, Ruess RW, Seagle SW. 1988. Large mammals and process dynamics in African ecosystems. Bioscience 38:794–800.
McNaughton SJ, Banyikwa FF, McNaughton MM. 1998. Root biomass and productivity in a grazing ecosystem: the Serengeti. Ecology 79:587–92.
Midgley JJ, Lawes MJ, Chamaillé-Jammes S. 2010. Savanna woody plant dynamics: the role of fire and herbivory, separately and synergistically. Turner Review No. 19. Aust J Bot 58:1–11.
Milchunas DG, Sala OE, Lauenroth WK. 1988. A generalized model of the effects of grazing by large herbivores on grassland community structure. Am Nat 132:87–106.
Milchunas DG, Lauenroth WK, Chapman PL, Kazempour MK. 1989. Effects of grazing, topography, and precipitation on the structure of a semi-arid grassland. Plant Ecol 80:11–23.
Mills AJ, Fey MV. 2005. Interactive response of herbivores, soils and vegetation to annual burning in a South African savanna. Austral Ecol 30:435–44.
Moe SR, Wegge P. 1997. The effects of cutting and burning on grass quality and axis deer (Axis axis) use of grassland in lowland Nepal. J Trop Ecol 13:279–92.
Moe SR, Wegge P. 2008. Effects of deposition of deer dung on nutrient redistribution and on soil and plant nutrients on intensively grazed grasslands in lowland Nepal. Ecol Res 23:227–34.
Nelson DW, Sommers LE. 1996. Total carbon, organic carbon, and organic matter. In: Sparks DL, Ed. Methods of soil analysis. Part 3. Chemical methods. SSSA Book Series: 5. Madison. Wisconsin. pp 961–1010.
Oba G, Stenseth NC, Lusigi WJ. 2000. New perspectives on sustainable grazing management in arid zones of sub-saharan Africa. Bioscience 50:35–51.
O’Connor TG. 1991. Influence of rainfall and grazing on the compositional change of the herbaceous layer of a sandveld savanna. J Grassl Soc Southern Afr 8:103–9.
Oksanen J, Blanchet FG, Kindt R, Legendre P, Minchin PR, O’Hara RB, Simpson GL, Solymos P, Henry M, Stevens H, Wagner H. 2012. Vegan: community ecology package. R package version 2.0-3. http://vegan.r-forge.r-project.org.
Okullo P, Moe SR. 2012. Large herbivores maintain termite-caused differences in herbaceous species diversity patterns. Ecology 93:2095–103.
Olff H, Ritchie ME. 1998. Effects of herbivores on grassland plant diversity. Trends Ecol Evol 13:261–5.
Pickett STA, White PS. 1985. Patch dynamics: a synthesis. In: Pickett STA, White PS, Eds. The ecology of natural disturbance and patch dynamics. San Diego: Academic Express. p 371–84.
Ritchie ME, David T, Knops JMH. 1998. Herbivore effects on plant and nitrogen dynamics in Oak savanna. Ecology 79:165–77.
Ruess RW, McNaughton SJ. 1987. Grazing and the dynamics of nutrient and energy regulated microbial processes in the Serengeti grasslands. Oikos 49:101–10.
Ruthven DCIII, Gallagher JF, Synatzske DR. 2000. Effect of fire and grazing on forbs in the Western South Texas plains. Southwestern Nat 45:89–94.
Sankaran M, Hanan NP, Scholes RJ, Ratnam J, Augustine DJ, Cade BS, Gignoux J, Higgins SI, Le Roux X, Ludwig F, Ardo J, Banyikwa F, Bronn A, Bucini G, Caylor KK, Coughenour MB, Diouf A, Ekaya W, Feral CJ, February EC, Frost PGH, Hiernaux P, Hrabar H, Metzger KL, Prins HHT, Ringrose S, Sea W, Tews J, Worden J, Zambatis N. 2005. Determinants of woody cover in African savannas. Nature 438:846–9.
Singer FJ, Schoenecker KA. 2003. Do ungulates accelerate or decelerate nitrogen cycling? For Ecol Manage 181:189–204.
Skarpe C. 1986. Plant community structure in relation to grazing and environmental changes along a north-south transect in the western Kalahari. Plant Ecol 68:3–18.
Skarpe C. 1992. Dynamics of savanna ecosystems. J Veg Sci 3:293–300.
Skarpe C, Aarrestad PA, Andreassen HP, Dhillion SS, Dimakatso T, Du Toit JT, Halley DJ, Hytteborn H, Makhabu S, Mari M, Marokane W, Masunga G, Modise D, Moe SR, Mojaphoko R, Mosugelo D, Motsumi S, Neo-Mahupeleng G, Ramotadima M, Rutina L, Sechele L, Sejoe TB, Stokke S, Swenson JE, Taolo C, Vandewalle M, Wegge P. 2004. The return of the giants: ecological effects of an increasing elephant population. Ambio 33:276–82.
Smith AMS, Wooster MJ, Drake NA, Dipotso FM, Falkowski MJ, Hudak AT. 2005. Testing the potential of multi-spectral remote sensing for retrospectively estimating fire severity in African Savannahs. Remote Sens Environ 97:92–115.
Soltanpour PN, Johnson GW, Workman SM, Jones JB, Miller RO. 1996. Inductively coupled plasma emission spectrometry and inductively coupled plasma-mass spectrometry. In: Sparks DL, Ed., Methods of soil analysis. Part 3. Chemical methods. SSSA Book Series: 5. Madison. Wisconsin. pp 91–139.
Sullivan S, Rohde R. 2002. On non-equilibrium in arid and semi-arid grazing systems. J Biogeogr 29:1595–618.
Sweet RJ. 1982. Bush control with fire in Acacia nigrescens/Combretum apiculatum savanna in Botswana. Proc Ann Congr Grassl Soc Southern Afr 17:25–8.
Taolo CL. 2003. Population ecology, seasonal movement and habitat use of the African buffalo (Syncerus caffer) in Chobe National Park, Botswana. PhD thesis. Norwegian University of Science and Technology, Trondheim.
ter Braak CJF. 1986. Canonical correspondence analysis: a new eigenvector technique for multivariate direct gradient analysis. Ecology 67:1167–79.
Tongway DJ, Hodgkinson KC. 1992. The effects of fire on the soil in a degraded semi arid woodland. III. Nutrient pool sizes, biological activity and herbage response. Soil fertility and plant nutrition. Aust J Soil Res 30:17–26.
Trollope WSW. 1984. Fire in savanna. In: Booysen de VP, Tainton NM, Ed., Ecological effects of fire in South African ecosystems. Ecological Studies 48. Berlin: Springer. pp 149–175.
Trollope WSW, Trollope LA, Harnett DC. 2002. Fire behaviour a key factor in the fire ecology of African grasslands and savannas. In: Viegas DX, Ed. Forest fire research and wildland fire safety. Rotterdam: Millpress. p 1–15.
Van de Vijver CADM, Poot P, Prins HHT. 1999. Causes of increased nutrient concentrations in post fire regrowth in an East African savanna. Plant Soil 214:173–85.
Van Langevelde F, Van de Vijver CADM, Kumar L, Van de Koppel J, De Ridder N, Van Andel J, Skidmore AK, Hearne JW, Stroosnijder L, Bond WJ, Prins HHT, Rietkerk M. 2003. Effects of fire and herbivory on the stability of savanna ecosystems. Ecology 84:337–50.
Van Oudtshoorn F. 1992. Guide to grasses of South Africa. Pretoria: Briza Publications.
Wan S, Hui D, Luo Y. 2001. Fire effects on nitrogen pools and dynamics in terrestrial ecosystems: a meta-analysis. Ecol Appl 11:1349–65.
Wang X, Yan Y, Cao Y. 2011. Impact of historic grazing on steppe soils on the northern Tibetan Plateau. Plant and Soil:1-11: online first Accepted 28 October 2011: doi:10.1007/s11104-011-1053-y.
Weber G, Moloney EK, Jeltsch F. 2000. Simulated long-term vegetation response to alternative stocking strategies in savanna rangelands. Plant Ecol 150:77–96.
Winter SL, Fuhlendorf SD, Goad CL, Davis CA, Hickman KR, Leslie DM Jr. 2012. Restoration of the fire–grazing interaction in Artemisia filifolia shrubland. J Appl Ecol 49:242–50.
Acknowledgments
The study was part of the Botswana-Norway Institutional Cooperation and Capacity building Project (BONIC) jointly funded by the Department of Wildlife and National Parks (DWNP) of Botswana and Norwegian Agency for Development (NORAD). We thank the Department of Agricultural Research in Sebele for providing a laboratory for chemical analysis and a number of DWNP officers for helping in the field. We also appreciate comments from RJ. Naiman and three anonymous reviewers which improved the quality of the manuscript.
Author information
Authors and Affiliations
Corresponding author
Additional information
Author Contributions
GSM designed the study, performed research, analyzed data and wrote the manuscript; SRM contributed to data analysis, interpretation and write-up; BP designed and performed research.
Electronic supplementary material
Below is the link to the electronic supplementary material.
Rights and permissions
About this article
Cite this article
Masunga, G.S., Moe, S.R. & Pelekekae, B. Fire and Grazing Change Herbaceous Species Composition and Reduce Beta Diversity in the Kalahari Sand System. Ecosystems 16, 252–268 (2013). https://doi.org/10.1007/s10021-012-9611-6
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s10021-012-9611-6