Advertisement

Landscape Ecology

, Volume 28, Issue 8, pp 1505–1516 | Cite as

Escaping the flames: large termitaria as refugia from fire in miombo woodland

  • Grant S. Joseph
  • Colleen L. Seymour
  • Graeme S. Cumming
  • Zacheus Mahlangu
  • David H. M. Cumming
Research article

Abstract

At finer scales, spatial heterogeneity can influence fire intensity and severity. To test whether Macrotermes termite mounds act as fire refugia for woody plants, we assessed effects of fire on individual plants, woody plant structure and composition in a miombo woodland in Zimbabwe, where elephants have decreased tree cover, leading to increased grass cover, fuelling greater intensity fires. We compared exposure to fire on 47 paired mound-matrix plots at three sites. Mound-based woody plants were less exposed to fire than those in matrix positions. Woody species composition differed between mound and matrix, and there were more tall trees on mounds. We assessed grass cover, elephant damage, fire damage and resprouting response for all woody plants found on 10 paired mound-matrix plots that had been equally exposed to severe late dry season fires. Grass cover was three times greater for matrix sites, where 85 % of woody species experienced heavy fire damage, compared to 29 % for mounds. Matrix species were nearly 31 times more likely than mound species to exhibit a vigorous resprouting response after fire damage, all else being equal. The distinct composition of termitaria vegetation has been attributed to edaphic factors. To this should be added the fire-retardant properties of mounds, allowing woody species that might otherwise have been excluded, to persist in a fire-prone system. Thus, spatial pattern created by termitaria is reinforced through exclusion of fire, allowing different species composition and structure. Since termitaria are important for productivity and biodiversity, the refuge effect is significant for the system.

Keywords

Chizarira National Park Elephant herbivory Macrotermes Ordinal logistic regression Resilience Resprouting Savanna Spatial heterogeneity 

Notes

Acknowledgments

This research was funded by an NRF-SADC collaborative grant and the DST-NRF Centre of Excellence at the Percy FitzPatrick Institute. The Director General of the Zimbabwe National Parks and Wildlife Management Authority (ZNPWMA) granted research permission for work within CNP through a memorandum of understanding between ZNPWMA and the Tropical Resource Ecology Programme (TREP) at the University of Zimbabwe. We gratefully acknowledge the support we received from both agencies and CNP staff in the field, and thank four anonymous reviewers for comments that strengthened the manuscript.

Supplementary material

10980_2013_9897_MOESM1_ESM.docx (19 kb)
Supplementary material 1 (DOCX 19 kb)

References

  1. Anderson GD, Walker BH (1974) Vegetation composition and elephant damage in the Sengwa Wildlife Research Area, Rhodesia. J S Afr Wildl Manage Assoc 4:1–14Google Scholar
  2. Belsky AJ (1994) Influences of trees on savanna productivity: tests of shade, nutrients, and tree-grass competition. Ecology 75(4):922–932CrossRefGoogle Scholar
  3. Bloesch U (2008) Thicket clumps: a characteristic feature of the Kagera savanna landscape, East Africa. J Veg Sci 19(1):31–44CrossRefGoogle Scholar
  4. Bond WJ (2008) What limits trees in C4 grasslands and savannas? Annu Rev Ecol Evol Syst 39:641–659CrossRefGoogle Scholar
  5. Bond WJ, Midgley GF (2000) A proposed CO2-controlled mechanism of woody plant invasion in grasslands and savannas. Glob Change Biol 6:865–869CrossRefGoogle Scholar
  6. Bond WJ, van Wilgen BW (1996) Fire and Plants. Chapman and Hall, LondonCrossRefGoogle Scholar
  7. Bond WJ, Midgley GF, Woodward FI (2003) What controls South African vegetation: climate or fire? S Afr J Bot 69(1):79–91Google Scholar
  8. Brody AK, Palmer TM, Fox-Dobbs K, Doak DF (2010) Termites, vertebrate herbivores, and the fruiting success of Acacia drepanolobium. Ecology 91(2):399–407PubMedCrossRefGoogle Scholar
  9. Clarke PJ (2002) Habitat islands in fire-prone vegetation: do landscape features influence community composition? J Biogeogr 29(5–6):677–684CrossRefGoogle Scholar
  10. Clarke KR, Gorley RN (2001) Plymouth routines in multivariate ecological research (PRIMER). Primer-E LtdGoogle Scholar
  11. Cumming DHM (1981) Elephants and woodlands in Chizarira National Park. In: Jewell PA, Hold S, Hart D (eds) Problems in management of locally abundant wild animals. Academic Press, New YorkGoogle Scholar
  12. Dangerfield JM, McCarthy TS, Ellery WN (1998) The mound-building termite Macrotermes michaelseni as an ecosystem engineer. J Trop Ecol 14(4):507–520CrossRefGoogle Scholar
  13. Dunham KM, Mackie CS, Musemburi OC et al (2006) Aerial survey of elephants and other large herbivores in the Sebungwe Region, Zimbabwe:2006. WWF-SAROPO Occasional Paper Number 12, Harare, ZimbabweGoogle Scholar
  14. Farina A (2006) Principles and methods in landscape ecology: towards a science of landscape. Springer, DordrechtGoogle Scholar
  15. Fleming PA, Loveridge JP (2003) Miombo woodland termite mounds: resource islands for small vertebrates? J Zool 259(2):161–168CrossRefGoogle Scholar
  16. Fox-Dobbs K, Doak DF, Brody AK, Palmer TM (2010) Termites create spatial structure and govern ecosystem function by affecting N2 fixation in an East African savanna. Ecology 91(5):1296–1307PubMedCrossRefGoogle Scholar
  17. Frost PGH (1996) The ecology of miombo woodlands. In: Campbell BM (ed) The miombo in transition: woodlands and welfare in Africa. Centre for International Forestry Research (CIFOR), Bogor, Indonesia, pp. 11–57Google Scholar
  18. Furley PA, Rees RM, Ryan CM, Saiz G (2008) Savanna burning and the assessment of long-term fire experiments with particular reference to Zimbabwe. Prog Phys Geog 32:611–634CrossRefGoogle Scholar
  19. Gignoux J, Clobert J, Menaut J-C (1997) Alternative fire resistance strategies in savanna trees. Oecologia 110:576–583CrossRefGoogle Scholar
  20. Glover PE, Trump EC, Wateridge LED (1964) Termitaria and vegetation patterns on the loita plains of Kenya. J Ecol 52(2):367–377CrossRefGoogle Scholar
  21. Guisan A, Harrell FE (2000) Ordinal response regression models in ecology. J Veg Sci 11(5):617–626CrossRefGoogle Scholar
  22. Harrell FE (2009) Design: design package. R package version 2.3-0. http://CRAN.R-project.org/package=Design
  23. Harrell FEJ, contributions from many other users (2010) Hmisc: Harrell Miscellaneous. R package version 3.8-3. http://CRAN.R-project.org/package=Hmisc
  24. Harrell FE, Margolis PA, Gove S et al (1998) Development of a clinical prediction model for an ordinal outcome: the World Health Organization multicentre study of clinical signs and etiological agents of pneumonia, sepsis and meningitis in young infants. Stat Med 17:909–944PubMedCrossRefGoogle Scholar
  25. Higgins SI, Bond WJ, Trollope WSW (2000) Fire, resprouting and variability: a recipe for grass-tree coexistence in savanna. J Ecol 88:213–229CrossRefGoogle Scholar
  26. Hoffmann WA, Bazzaz FA, Chatterton NJ, Harrison PA, Jackson RB (2000) Elevated CO2 enhances resprouting of a tropical savanna tree. Oecologia 123:312–317CrossRefGoogle Scholar
  27. Huntley BJ (1982) Southern African Savannas. In: Huntley BJ, Walker BH (eds) Ecology of Tropical Savannas. Springer, Berlin, pp 101–119CrossRefGoogle Scholar
  28. Jones CG, Lawton JH, Shachak M (1994) Organisms as ecosystem engineers. Oikos 69:373–386CrossRefGoogle Scholar
  29. Joseph GS, Cumming GS, Cumming DHM, Mahlangu Z, Altwegg R, Seymour CL (2011) Large termitaria act as refugia for tall trees, deadwood and cavity-using birds in a miombo woodland. Lanscape Ecol 26(3):439–448CrossRefGoogle Scholar
  30. Joseph GS, Seymour CL, Cumming GS, Cumming DHM, Mahlangu Z (2012) Termite mounds as islands: woody plant assemblages relative to termitarium size and soil properties. J Veg Sci. doi: 10.1111/j.1654-1103.2012.01489.x Google Scholar
  31. 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 206:47–60CrossRefGoogle Scholar
  32. Krebs CJ (1999) Ecological methodology Benjamin Cummings. Menlo Park, CaliforniaGoogle Scholar
  33. Lawes MJ, Adie H, Russell-Smith J, Murphy B, Midgley J (2011) How do small savanna trees avoid stem mortality by fire? The roles of stem diameter, height and bark thickness. Ecosphere 2(42):1–13Google Scholar
  34. Laws RM (1970) Elephants as agents of habitat and landscape change in East Africa. Oikos 21:1–15CrossRefGoogle Scholar
  35. Lee KE, Wood TG (1971) Termites and soils. Academic Press, LondonGoogle Scholar
  36. Levick SR, Asner GP, Kennedy-Bowdoin T, Knapp DE (2009) The relative influence of fire and herbivory on savanna three-dimensional vegetation structure. Biol Conserv 142:1693–1700CrossRefGoogle Scholar
  37. Levick SR, Asner GP, Kennedy-Bowdoin T, Knapp DE (2010) The spatial extent of termite influences on herbivore browsing in an African savanna. Biol Conserv 143(11):2462–2467CrossRefGoogle Scholar
  38. Lineham S (1965) Rainfall in Rhodesia. In: Collins MO (ed) Rhodesia: its natural resources and economic development. Collins, Salisbury, pp 26–27Google Scholar
  39. Loveridge JP, Moe SR (2004) Termitaria as browsing hotspots for African megaherbivores in miombo woodland. J Trop Ecol 20(3):337–343CrossRefGoogle Scholar
  40. Malaisse F (1978) High Termitaria. In: Werger MJA (ed) Biogeography and ecology of southern Africa. Junk, The Hague, pp 1279–1300CrossRefGoogle Scholar
  41. Mapaure I, Campbell BM (2002) Changes in miombo woodland cover in and around Sengwa Wildlife Research Area, Zimbabwe, in relation to elephants and fire. Afr J Ecol 40:212–219CrossRefGoogle Scholar
  42. Moe S, Mobæk R, Narmo A (2009) Mound building termites contribute to savanna vegetation heterogeneity. Plant Ecol 202(1):31–40CrossRefGoogle Scholar
  43. Morison CGT, Hoyle AC, Hope-Simpson JF (1948) Tropical soil-vegetation catenas and mosaics: a study in the south-western part of the Anglo-Egyptian Sudan. J Ecol 36:1–84CrossRefGoogle Scholar
  44. Morvan N, Burel F, Baudry J et al (1995) Landscape and fire in Britanny heathlands. Landscape Urban Plan 31:81–88CrossRefGoogle Scholar
  45. Mueller-Dombois D, Ellenberg H (1974) Aims and methods of vegetation ecology. Wiley, New YorkGoogle Scholar
  46. Mushove PT, Makoni JT (1993) Coppicing ability of Colophospermum mopane. In: Pierce GD, Gumbo DJ (eds) Proceedings of the international symposium on the ecology and management of indigenous forests in southern Africa, Victoria Falls, Zimbabwe 1993. Zimbabwe Forestry Commission and SAREC (Harare)Google Scholar
  47. Nano CEM, Clarke PJ (2011) How do drought and fire influence the patterns of resprouting in Australian deserts? Plant Ecol 12:2095–2110CrossRefGoogle Scholar
  48. Nefabas LL, Gambiza J (2007) Fire-tolerance mechanisms of common woody plant species in a semiarid savanna in south-western Zimbabwe. Afr J Ecol 45:550–556CrossRefGoogle Scholar
  49. Nzunda EF, Lawes MJ (2011) Costs of resprouting are traded off against reproduction in subtropical coastal dune forest trees. Plant Ecol 12:1991–2001CrossRefGoogle Scholar
  50. Pringle RM, Doak DF, Brody AK, Jocqué R, Palmer TM (2010) Spatial pattern enhances ecosystem functioning in an African savanna. PLoS Biol 8(5):e1000377PubMedCrossRefGoogle Scholar
  51. Romme VH (1982) Fire and landscape diversity in subalpine forests of Yellowstone National Park. Ecol Monogr 52:199–221CrossRefGoogle Scholar
  52. Sankaran M, Hanan NP, Scholes RJ et al (2005) Determinants of woody cover in African savannas. Nature 438(8):846–849PubMedCrossRefGoogle Scholar
  53. Seymour CL (2008) Grass, rainfall and herbivores as determinants of Acacia erioloba (Meyer) recruitment in an African savanna. Plant Ecol 197:131–138CrossRefGoogle Scholar
  54. Sileshi GW, Arshad MA, Konaté S, Nkunika POY (2010) Termite-induced heterogeneity in African savanna vegetation: mechanisms and patterns. J Veg Sci 21(5):923–937CrossRefGoogle Scholar
  55. Starfield AM, Cumming DHM, Taylor RD (1993) A frame-based paradigm for dynamic ecosystem models. AI Appl 7:1–13Google Scholar
  56. Staver AC, Bond WJ, Stock WD, Waldram SJ, Waldram MS (2009) Browsing and fire interact to suppress tree density in an African savanna. Ecol Appl 19(7):1909–1919PubMedCrossRefGoogle Scholar
  57. Staver AC, Archibald S, Levin SA (2011) The global extent and determinants of savanna and forest as alternative biome states. Science 334(6053):230–232PubMedCrossRefGoogle Scholar
  58. Thomson PJ (1974) The role of elephants, and fire and other agents in the decline of Brachystegia boehmi woodland. J S Afr Wildl Manage Assoc 5:11–18Google Scholar
  59. Torrance JD (1965) The temperature of Rhodesia. In: Collins MO (ed) Rhodesia: its natural resources and economic development. Salisbury, Rhodesia, pp 28–29Google Scholar
  60. Turner MG (1989) Landscape ecology: the effect of pattern on process. Ann Rev Ecol Syst 20:171–197CrossRefGoogle Scholar
  61. van Wilgen BW, Trollope WSW, Biggs HC, Potgieter ALF, Brockett BH (2003) Fire as a driver of ecosystem variability. In: Du Toit J, Rogers KH, Biggs HC (eds) The Kruger experience: ecology and management of savanna heterogeneity. Island Press, New York, pp. 149–170Google Scholar

Copyright information

© Springer Science+Business Media Dordrecht 2013

Authors and Affiliations

  • Grant S. Joseph
    • 1
  • Colleen L. Seymour
    • 1
    • 2
  • Graeme S. Cumming
    • 1
  • Zacheus Mahlangu
    • 3
  • David H. M. Cumming
    • 1
    • 3
  1. 1.Percy FitzPatrick Institute of African Ornithology, DST/NRF Centre of ExcellenceUniversity of Cape TownRondeboschSouth Africa
  2. 2.Applied Biodiversity Research Division, South African National Biodiversity InstituteKirstenbosch Research CentreClaremontSouth Africa
  3. 3.Tropical Resource Ecology Programme, Department of Biological SciencesUniversity of ZimbabweHarareZimbabwe

Personalised recommendations