, Volume 22, Issue 1, pp 125–136 | Cite as

Are Termite Mounds Always Grazing Hotspots? Grazing Variability with Mound Size, Season and Geology in an African Savanna

  • Justice MuvengwiEmail author
  • Francesca Parrini
  • E. T. F. Witkowski
  • Andrew B. Davies


Foraging site selection by large herbivores is influenced by multiple factors varying across landscapes and spatial scales. Termite mounds harbour highly nutritious plants compared with the savanna matrix, making them preferred foraging patches in many savannas. However, it is unknown whether termite mounds equally influence herbivore grazing intensity across geological substrates and mound sizes. These knowledge gaps hamper our ability to draw general trans-ecosystem conclusions about the effect of termite mounds for savanna herbivores. We measured grazing intensity on mounds of three different size classes (small, medium and large) across two geologies with differing soil nutrition (granite and basalt) in Gonarezhou National Park, Zimbabwe. We recorded measurements across three seasons (hot wet, cool dry and hot dry), and at multiple distances from mounds. Grazing intensity on mounds was higher on nutrient-poor granite than nutrient-rich basalt, and termite mounds of all sizes had a significant effect on grazing on granite during the cool dry season. Grazing was highest on large mounds on both geologies throughout the year. Large mounds also had the largest spatial influence on grazing in the cool dry season, up to 8 m beyond the mound edge on granite and 2 m on basalt. When scaled up to the landscape level, mounds influenced about 15% of the granite landscape, but only about 0.5% of the basalt landscape. Our results show that the positive effects of mounds on grazing intensity were pronounced on nutrient-poor soils but negligible on nutrient-rich soils, and that the magnitude of these effects varied across seasons and with mound size.


basalt granite grazing intensity landscape large herbivore nutrient hotspots semi-arid savannas 

Supplementary material

10021_2018_257_MOESM1_ESM.docx (69 kb)
Supplementary material 1 (DOCX 69 kb)


  1. Anderson DR, Link WA, Johnson DH, Burnham KP, Press A. 2001. Suggestions for presenting the results of data analyses invited paper: the results of data analyses suggestions. J Wildl Manag 65:373–8.
  2. Anderson JM, Ingram JSI. 1993. Tropical soil biology and fertility: a handbook of methods. second. Wallingford: CABI.Google Scholar
  3. Asner GP, Levick SR, Kennedy-Bowdoin T, Knapp DE, Emerson R, Jacobson J, Colgan MS, Martin RE. 2009. Large-scale impacts of herbivores on the structural diversity of African savannas. Proc Natl Acad Sci USA 106:4947–52.CrossRefGoogle Scholar
  4. Barton K. 2016. MuMIn: multi-model inference. R package Version 1156,
  5. Bates D. 2007. nlme: Linear and Nonlinear Mixed Effects Models. R package version 31-128,
  6. Benjamin Y, Yekutieli D. 2001. The control of the false discovery rate in multiple testing under dependency. Ann Stat 29:1165–88.CrossRefGoogle Scholar
  7. Bottinelli N, Jouquet P, Capowiez Y, Podwojewski P, Grimaldi M, Peng X. 2015. Why is the influence of soil macrofauna on soil structure only considered by soil ecologists? Soil Tillage Res 146:118–24.CrossRefGoogle Scholar
  8. Cromsigt J, Olff H. 2006. Resource partitioning among savanna grazers mediated by local heterogeneity: an experimental approach. Ecology 87:1532–41.CrossRefGoogle Scholar
  9. Cromsigt JPGM, te Beest M. 2014. Restoration of a megaherbivore: landscape-level impacts of white rhinoceros in Kruger National Park, South Africa. J Ecol 102:566–75.CrossRefGoogle Scholar
  10. Crutzen P, Andreae MO. 1990. Biomass burning in the tropics: impact on atmospheric chemistry and biogeochemical cycles. Science 250:1669–78.CrossRefGoogle Scholar
  11. Davies AB, Baldeck CA, Asner GP. 2016a. Termite mounds alter the spatial distribution of African savanna tree species. J Biogeogr 43:301–13.CrossRefGoogle Scholar
  12. Davies AB, Levick SR, Robertson MP, van Rensburg BJ, Asner GP, Parr CL. 2016b. Termite mounds differ in their importance for herbivores across savanna types, seasons and spatial scales. Oikos 125:726–34.CrossRefGoogle Scholar
  13. Davies AB, Robertson MP, Levick SR, Asner GP, van Rensburg BJ, Parr CL. 2014. Variable effects of termite mounds on African savanna grass communities across a rainfall gradient. Journal of Vegetation Science 25:1405–16.
  14. Fox J, Weisberg S, Adler D, Bates D, Baud-Bovy G, Ellison S, Firth D, Friendly M, Gorjanc G, Graves S, Heiberger R, Laboissiere R, Monette G, Murdoch D, Nilsson H, Ogle D, Ripley B, Venables W, Winsemius D, Zeileis A. 2015. CAR: companion to applied regression. R package version 21-2,
  15. Fryxell JM. 1991. Forage quality and aggregation by large herbivores. Am Nat 138:478–98.CrossRefGoogle Scholar
  16. Gandiwa E, Chikorowondo G, Muvengwi J. 2011. Structure and composition of Androstachys johnsonii woodland across various strata in Gonarezhou National Park, southeast Zimbabwe. Trop Conserv Sci 4:218–29.CrossRefGoogle Scholar
  17. Genz A, Bretz F, Miwa T, Mi X, Leisch F, Scheipl F, Bornkamp B, Maechler M, Hothorn T. 2016. Multivariate normal and t distributions. R package version 10-5:
  18. Gosling CM, Cromsigt JPGM, Mpanza N, Olff H. 2012. Effects of erosion from mounds of different termite genera on distinct functional grassland types in an African Savannah. Ecosystems 15:128–39.CrossRefGoogle Scholar
  19. Grant CC, Scholes MC. 2006. The importance of nutrient hot-spots in the conservation and management of large wild mammalian herbivores in semi-arid savannas. Biol Conserv 130:426–37.CrossRefGoogle Scholar
  20. Heady HF. 1949. Methods of determining utilization of range forage. J Range Manag 2:53–63.CrossRefGoogle Scholar
  21. Holt J, Coventry R. 1990. Nutrient cycling in Australian savannas. Journal of Biogeography 17:427–32.
  22. Hothorn T, Bretz F, Westfall P, Heiberger RM, Schuetzenmeister A, Scheibe S. 2016. Simultaneous inference in general parametric models. R package version 14-4:
  23. Jones CG, Lawton JH, Shachak M. 1994. Organisms as ecosystem engineers. Oikos 69:373–86.CrossRefGoogle Scholar
  24. Joseph GS, Seymour CL, Cumming GS, Cumming DHM, Mahlangu Z. 2013. Termite mounds as islands: woody plant assemblages relative to termitarium size and soil properties. J Veg Sci 24:702–11.CrossRefGoogle Scholar
  25. Kartzinel TR, Chen PA, Coverdale TC, Erickson DL, Kress WJ, Kuzmina ML, Rubenstein DI, Wang W, Pringle RM. 2015. DNA metabarcoding illuminates dietary niche partitioning by African large herbivores. Proc Natl Acad Sci: 6.Google Scholar
  26. Knox NM, Skidmore AK, Prins HHT, Asner GP, van der Werff HMA, de Boer WF, van der Waal C, de Knegt HJ, Kohi EM, Slotow R, Grant RC. 2011. Dry season mapping of savanna forage quality, using the hyperspectral Carnegie Airborne Observatory sensor. Remote Sens Environ 115:1478–88.
  27. Konaté S, Le Roux X, Tessier D, Lepage M. 1999. Influence of large termitaria on soil characteristics, soil water regime, and tree leaf shedding pattern in a West African savanna. Plant Soil 206:47–60.CrossRefGoogle Scholar
  28. Kruckeberg AR. 1986. An essay: the stimulus of unusual geologies for plant speciation. Syst Bot 11:455–63.
  29. Lagendijk DDG, Davies AB, Eggleton P, Slotow R. 2016. No evidence for an elephant-termite feedback loop in Sand Forest, South Africa. Biol Conserv 203:125–33.
  30. Lepage M, Abbadie L, Mariotti A. 1993. Food habits of sympatric termite species (Isoptera, Macrotermitinae) as determined by stable carbon isotope analysis in a Guinean savanna (Lamto, Cote d’Ivoire). J Trop Ecol 9:303–11.CrossRefGoogle Scholar
  31. Loveridge JP, Moe SR. 2004. Termitaria as browsing hotspots for African megaherbivores in miombo woodland. J Trop Ecol 20:337–43.CrossRefGoogle Scholar
  32. Mando A, Stroosnijder L, Brussaard L. 1996. Effects of termites on infiltration into crusted soil. Geoderma 74:107–13.CrossRefGoogle Scholar
  33. Mobæk R, Narmo AK, Moe SR. 2005. Termitaria are focal feeding sites for large ungulates in Lake Mburo National Park, Uganda. J Zool Lond 267:97–102.
  34. Moe SR, Mobæk R, Narmo AK. 2009. Mound building termites contribute to savanna vegetation heterogeneity. Plant Ecol 202:31–40.CrossRefGoogle Scholar
  35. Muvengwi J. 2016. Relationships between termite (Macrotermes) mound distribution, plant diversity and large mammalian herbivory patterns in Gonarezhou National Park, Zimbabwe.Google Scholar
  36. Muvengwi J, Davies AB, Parrini F, Witkowski ETF. 2018a. Contrasting termite diversity and assemblages on granitic and basaltic African savanna landscapes. Insectes Soc 65:25–35.CrossRefGoogle Scholar
  37. Muvengwi J, Davies AB, Parrini F, Witkowski ETF. 2018b. Geology drives the spatial patterning and structure of termite mounds in an African savanna. Ecosphere 9:e02148.
  38. Muvengwi J, Mbiba M, Nyenda T. 2013. Termite mounds may not be foraging hotspots for mega-herbivores in a nutrient-rich matrix. J Trop Ecol 29:551–8.
  39. Muvengwi J, Ndagurwa HGT, Nyenda T, Mbiba M. 2016. Nutrient dynamics and plant assemblages of Macrotermes falciger mounds in a savanna ecosystem. Acta Oecol 76:13–21.
  40. Muvengwi J, Ndagurwa HGT, Nyenda T, Mlambo I. 2014. Termitaria as preferred browsing patches for black rhinoceros (Diceros bicornis) in Chipinge Safari Area, Zimbabwe. J Trop Ecol 30:591–8.
  41. Muvengwi J, Witkowski ETF, Parrini F, Davies AB. 2017. Termite mounds vary in their importance as sources of vegetation heterogeneity across savanna landscapes. J Veg Sci 28:1008–17.CrossRefGoogle Scholar
  42. Naiman R, Braack L, Grant C, Kemp A, Du Toit J, Venter F. 2003. Interactions between species and ecosystem characteristics. In: Du Toit JT, Rogers KH, Biggs HC, Eds. The Kruger experience, ecology and management of savanna heterogeneity. Washington: Long Island Press. p 221–41.Google Scholar
  43. Okalebo JR, Gathma KW, Woomer PL. 2002. Laboratory methods of soil and plant analysis: a working manual. second. Soil Science Society of East Africa, Nairobi.Google Scholar
  44. Pretorius Y, de Boer FW, van der Waal C, de Knegt HJ, Grant RC, Knox NM, Kohi EM, Mwakiwa E, Page BR, Peel MJS, Skidmore AK, Slotow R, van Wieren SE, Prins HHT. 2011. Soil nutrient status determines how elephant utilize trees and shape environments. J Anim Ecol 80:875–83.CrossRefGoogle Scholar
  45. Pringle RM, Doak DF, Brody AK, Jocqué R, Palmer TM. 2010. Spatial pattern enhances ecosystem functioning in an African savanna. PLoS Biol 8:e1000377.CrossRefGoogle Scholar
  46. Pringle RM, Prior KM, Palmer TM, Young TP, Goheen JR. 2016. Large herbivores promote habitat specialization and beta diversity of African savanna trees. Ecology 97:2640–57.CrossRefGoogle Scholar
  47. Scholes RJ, Bond WJ, Eckhardt HC. 2003. Vegetation dynamics in the Kruger ecosystem. In: du Toit JT, Rogers KH, Biggs HC, Eds. The Kruger experience: ecology and management of savanna heterogeneity. Washington: Island Press. p 242–62.Google Scholar
  48. Seymour CL, Milewski AV, Mills AJ, Joseph GS, Cumming GS, Cumming DHM, Mahlangu Z. 2014. Do the large termite mounds of Macrotermes concentrate micronutrients in addition to macronutrients in nutrient-poor African savannas? Soil Biol Biochem 68:95–105.CrossRefGoogle Scholar
  49. Sibbesen E. 1978. An investigation of the anion-exchange resin method for soil phosphate extraction. Plant Soil 50:305–21.CrossRefGoogle Scholar
  50. Turner JS. 2006. Termites as mediators of the water economy of arid savanna ecosystems. In: D’Odorico P, Porporato A, Eds. Dryland ecohydrology. New York: Springer. p 303–13.CrossRefGoogle Scholar
  51. Van der Plas F, Howison R, Reinders J, Fokkema W, Olff H. 2013. Functional traits of trees on and off termite mounds: Understanding the origin of biotically-driven heterogeneity in savannas. J Veg Sci 24:227–38.CrossRefGoogle Scholar
  52. Venter FJ. 1990. A classification of land use for management planning in the Kruger National Park.Google Scholar
  53. Venter FJ, Scholes RJ, Eckhardt HC. 2003. The abiotic template and its associated vegetation pattern. In: Du Toit JT, Biggs HC, Rogers KH, Eds. The Kruger experience: ecology and management of savanna heterogeneity. Washington: Island Press. p 81–129.Google Scholar
  54. Wallace LL, Turner M, Romme WH, O’Neill R, Wu Y. 1995. Scale of heterogeneity of forage production and winter foraging by elk and bison. Landsc Ecol 10:75–83.CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media, LLC, part of Springer Nature 2018

Authors and Affiliations

  • Justice Muvengwi
    • 1
    • 2
    Email author
  • Francesca Parrini
    • 3
  • E. T. F. Witkowski
    • 1
  • Andrew B. Davies
    • 1
    • 4
  1. 1.Restoration and Conservation Biology Research Group, School of Animal, Plant and Environmental SciencesUniversity of the WitwatersrandJohannesburgSouth Africa
  2. 2.Department of Natural ResourcesBindura University of Science EducationBinduraZimbabwe
  3. 3.Centre for African Ecology, School of Animal, Plant and Environmental SciencesUniversity of the WitwatersrandJohannesburgSouth Africa
  4. 4.Department of Global EcologyCarnegie Institution for ScienceStanfordUSA

Personalised recommendations