Log in

Effects of rain, nitrogen, fire and grazing on tree recruitment and early survival in bush-encroached savanna, South Africa

  • Published:
Plant Ecology Aims and scope Submit manuscript

Abstract

Moisture, nutrients, fire and herbivory are the principal factors governing tree–grass cover ratios of savannas. We investigated tree (Acacia mellifera) recruitment after fire and under conditions of maximum-recorded rainfall, nitrogen addition and grazing in a completely-crossed field experiment. We employed a similar garden experiment with the exception of the fire treatment. Tree germination in the field was extremely low, probably due to below-average natural rainfall in plots that only received natural rain, and insufficient watering frequency in irrigated plots. Due to low germination in the field experiment, no treatment significantly affected tree recruitment. In the garden experiment, frequent watering, nutrient control (i.e. no nitrogen addition) and grazing enhanced tree recruitment with significant interactions between rain, nitrogen and grazing. We infer that above-average rainfall years with frequent rainfall events are required for mass tree recruitment. Grass defoliation makes space and resources available for tree seedlings. Nitrogen enrichment increases the competitive ability of fast-growing grasses more than that of the N2-fixing tree component. In contrast to conventional wisdom that grazing alone causes encroachment, we suggest that there are complex interactions between the above-mentioned factors and ‘triggering’ events such as unusually high rainfall.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Subscribe and save

Springer+ Basic
EUR 32.99 /Month
  • Get 10 units per month
  • Download Article/Chapter or Ebook
  • 1 Unit = 1 Article or 1 Chapter
  • Cancel anytime
Subscribe now

Buy Now

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Similar content being viewed by others

References

  • Acocks J.P.H. (1988). Veld types of South Africa. Mem. Bot. Surv. S. Afr. 57:1–146

    Google Scholar 

  • Alef K., Nannipieri P. (1995). Methods in Applied Soil Microbiology and Biochemistry. Academic Press, London

    Google Scholar 

  • Andersson C.J. (1856). Lake Ngami: On Explorations and Discoveries, During Four Years’ Wanderings in the Wilds of South Western Africa. Hurst and Blackett, London

    Google Scholar 

  • Archer S. (1989). Have southern Texas savannas been converted to woodlands in recent history? Am. Nat. 134:545–561

    Article  Google Scholar 

  • Babalwa A.M., Witkowski E.T.F. (1997). Tolerance to soil temperatures experienced during and after the passage of fire in seeds of Acacia karroo, Acacia tortilis andChromolaena odorata. S. Afr. J. Bot. 63:421–425

    Google Scholar 

  • Bezuidenhout H. (1994). An ecological study of the major vegetation communities of the Vaalbos National Park, Northern Cape. 1. The Than-Droogeveld section. Koedoe 37:19–42

    Google Scholar 

  • Booth C.A. 1986. The effect of microtopography and woollybutt (Eragrostis eriopoda) competitiion on the survival of hopbush (Dodonaea attenuata) seedlings. In: Rangelands: a resource under siege. Proceedings of the Second International Rangeland Congress (Adelaide 1985). Cambridge University Press, Cambridge, pp. 32–33.

  • Bosch P.J.A. 1993. Geological survey: the geology of the Kimberley area. Explanation of sheet 2824. Scale 1:250 000. Department of Mineral and Energy Affairs, Republic of South Africa. Government Printer, Pretoria.

  • Bradstock R.A., Auld T.D. (1995). Soil temperatures during experimental bushfires in relation to fire intensity: consequences for legume germination and fire management in south-eastern. Aust. J. Appl. Ecol. 32:76–84

    Article  Google Scholar 

  • Brown J.R., Archer S. (1989). Woody plant invasion of grasslands: establishment of honey mesquite (Prosopis glandulosa var. glandulosa) on sites differing in herbaceous biomass and grazing history. Oecologia 80:19–26

    Article  Google Scholar 

  • Brown J.R., Archer S. (1999). Shrub invasion of grassland: recruitment is continuous and not regulated by herbaceous biomass or density. Ecology 80:2385–2396

    Article  Google Scholar 

  • Bush J.K., Van Auken O.W. (1995). Woody plant growth related to planting time and clip** of a C4 grass. Ecology 76:1603–1609

    Article  Google Scholar 

  • Cohn E.J., Van Auken O.W., Bush J.K. (1989). Competitive interactions between Cynodon dactylon and Acacia smallii seedlings at different nutrient levels. Am. Midl. Nat. 121:265–272

    Article  Google Scholar 

  • Condon R.W. 1986. Scrub invasion on semi-arid grazing lands in western New South Wales – causes and effects. In: Rangelands: A resource under siege. Proceedings of the Second International Rangeland Congress (Adelaide 1985). Cambridge University Press, Cambridge, p. 40.

  • Dart P., Umali-Garcia M. and Almendras A. 1991. Role of symbiotic associations in nutrition of tropical acacias. In: Turnbull J.W. (ed.), Advances in Tropical Acacia Research, ACIAR Proceedings No. 35, pp. 13–19.

  • Davis M.A., Wrage K.J., Reich P.B. (1998). Competition between tree seedlings and herbaceous vegetation: support for a theory of resource supply and demand. J. Ecol. 86:652–661

    Article  Google Scholar 

  • Dean W.R.J., Macdonald I.A.W. (1994). Historical changes in stocking rates of domestic livestock as a measure of semi-arid and arid rangeland degradation in the Cape Province, South Africa. J. Arid Environ. 26:281–298

    Article  Google Scholar 

  • Donaldson C.H. (1966). Control of Blackthorn in the Molopo area with special reference to fire. Proc. Grassland Soc. South. Afr. 1:57–62

    Google Scholar 

  • Du Toit P.F. (1972a). The goat in a bush-grass community. Proc. Grassland Soc. South. Afr. 7:44–50

    Google Scholar 

  • Du Toit P.F. (1972b). Acacia karroo intrusion: the effect of burning and sparing. Proc. Grassland Soc. South. Afr. 7:23–27

    Google Scholar 

  • Dye P.J., Spear P.T. (1982). The effects of bush clearing and rainfall variability on grass yield and composition in south-west Zimbabwe. Zimb. J. Agric. Res. 20:103–118

    Google Scholar 

  • Ernst W.H.O. (1988). Seed and seedling ecology of Brachystegia spiciformis, a predominant tree component in miombo woodlands in South Central Africa. For. Ecol. Manage. 25:195–210

    Article  Google Scholar 

  • Fowler N. (1982). Competition and coexistence in a North Carolina grassland. III. Mixtures of component species. J. Ecol. 70:77–92

    Article  Google Scholar 

  • Frost P., Medina E., Menaut J.C., Solbrig O., Swift M. and Walker B. (eds), 1985. Responses of savannas to stress and disturbance. In: Biol. Int. 10: 1–82.

  • Higgins S.I., Bond W.J., Trollope W.S.W. (2000). Fire, resprouting and variability: a recipe for grass-tree coexistence in savanna. J. Ecol. 88:213–229

    Article  Google Scholar 

  • Jeltsch F., Milton S.J., Dean W.R.J., Van Rooyen N. (1996). Tree spacing and coexistence in sermiarid savannas. J. Ecol. 84:583–595

    Article  Google Scholar 

  • Kanz W.A. 2001. Seed and seedling dynamics of certain Acacia species as affected by herbivory, grass competition, fire, and grazing system. M.Sc., Thesis, University of Natal, Pietermaritzburg, South Africa.

  • Kelk D.M., Donaldson C.H. (1970). An investigation of veld problems in the Molopo area: II. Some practical implications or the experimental findings. Proc. Grassland Soc. South. Afr. 5:58–61

    Google Scholar 

  • Knoop W.T., Walker B.H. (1985). Interactions of woody and herbaceous vegetation in a southern African savanna. J. Ecol. 73:235–253

    Article  Google Scholar 

  • Krebs C.J. (1989). Ecological Methodology. Harper & Row, New York

    Google Scholar 

  • Kreuter U.P., Archer S.R. and Scifres C.J. 1999. Bio-economic basis for woody plant management. In: Proceedings of the Sixth International Rangeland Congress, Vol. 2. VI International Rangeland Congress Inc., Queensland, Australia, pp. 842–943.

  • Lay D.W. (1965). Effects of periodic clip** on yield of some common browse species. J. Range Manage. 18:181–184

    Article  Google Scholar 

  • Low A.B., Rebelo A.T.G. (eds). (1998). Vegetation of South Africa, Lesotho and Swaziland. Department of Environmental Affairs and Tourism, Pretoria, South Africa

    Google Scholar 

  • Machlis L., Torrey J.G. (1956). Plants in Action: A Laboratory Manual of Plant Physiology. WH Freeman & Company Inc., San Francisco

    Google Scholar 

  • Milton S.J. 1980. Studies on Australian Acacias in the south western Cape, South Africa. M.Sc., Thesis, University of Cape Town, Cape Town, South Africa.

  • Moshe D., Bailey C.L. and Scholes R.J. 2000. The effect of elevated atmospheric carbon dioxide on selected savanna plants. In: Seydack A.H.W., Vermeulen W.J. and Vermeulen C. (eds), Proceedings: Natural forests and savanna woodlands symposium II: towards sustainable management based on scientific understanding of natural forests and woodlands. Department of Water Affairs and Forestry, Knysna, South Africa, pp. 142–144.

  • O’Connor T.G. (1995). Acacia karroo invasion of grassland: environmental and biotic effects influencing seedling emergence and establishment. Oecologia 103:214–223

    Article  Google Scholar 

  • Palgrave K.C. (1977). Trees of Southern Africa. Struik, Cape Town

    Google Scholar 

  • Perkins J.S., Thomas D.S.G. (1993). Spreading deserts or spatially confined impacts? Land degradation and cattle ranching in the Kalahari desert of Botswana. Land Degrad. Rehabil. 4:179–194

    Article  Google Scholar 

  • Pitt M.D., Newman R.F., Youwe P.L., Wikeem B.M. and Quinton D.A. (1998). Using a grazing pressure index to predict cattle damage of regenerating tree seedlings. J. Range Manage. 51:417–422

    Article  Google Scholar 

  • Rohner C., Ward D. (1997). Chemical and mechanical defense against herbivory in two sympatric species of desert Acacia. J. Veg. Sci. 8:717–726

    Article  Google Scholar 

  • Salihi D.O., Norton B.E. (1987). Survival of perennial grass seedlings under intensive grazing in semi-arid rangelands. J. Appl. Ecol. 24:145–151

    Article  Google Scholar 

  • Scholes R.J., Walker B.H. (1993). An African Savanna: Synthesis of the Nylsvley Study. Cambridge University Press, Cambridge

    Google Scholar 

  • Schultka W., Cornelius R. (1997). Vegetation structure of a heavily grazed range in northern Kenya: tree and shrub canopy. J. Arid Environ. 36:290–306

    Article  Google Scholar 

  • Schultz A.M., Launchbauch J.L., Biswell H.H. (1955). Relationship between grass density and brush seedling survival. Ecology 36:226–238

    Article  Google Scholar 

  • Scott J.D. (1967). Bush encroachment in South Africa. S. Afr. J. Sci. 63:311–314

    Google Scholar 

  • Siegel S., Castellan N.J. (1989). Nonparametric Statistics for the Behavioural Sciences. McGraw-Hill Inc., New York

    Google Scholar 

  • Skarpe C. (1990). Structure of the woody vegetation in disturbed and undisturbed arid savanna, Botswana. Vegetatio 87:11–18

    Article  Google Scholar 

  • Smit G.N., Rethman N.F.G. (1992). Inter-related floristic changes associated with different long-term grazing treatments in Sourish Mixed Bushveld. J. Grassland Soc. S. Afr. 5:42–47

    Google Scholar 

  • Stuart-Hill G.C., Tainton N.M. (1988). Browse and herbage production in the Eastern Cape Thornveld in response to tree size and defoliation frequency. J. Grassland Soc. S. Afr. 5:42–47

    Google Scholar 

  • Stuart-Hill G.C., Tainton N.M. (1989). The competitive interaction between Acacia karroo and the herbaceous layer and how this is influenced by defoliation. J. Appl. Ecol. 26:285–298

    Article  Google Scholar 

  • Teague W.R., Smit G.N. (1992). Relations between woody and herbaceous components and the effects of bush-clearing in southern African savannas. J. Grassland Soc. S. Afr. 9:60–71

    Google Scholar 

  • Tilman D. (1987). Secondary succession and the pattern of plant dominance along experimental nitrogen gradients. Ecol. Monogr. 57:189–214

    Article  Google Scholar 

  • Trollope W.S.W. (1974). Role of fire in preventing bush encroachment in the Eastern Cape. Proc. Grassland Soc. South. Afr. 9:67–72

    Google Scholar 

  • Trollope W.S.W. (1980). Controlling bush encroachment with fire in the savanna areas of South Africa. Proceedings of the Grassland Society of Southern Africa 15:173–177

    Google Scholar 

  • Trollope W.S.W. (1982). Ecological effects of fire in South African savannas. In: Huntley B.J., Walker BH. (eds) Ecological Studies 42: Ecology of Tropical Savannas. Springer-Verlag, New York, pp. 292–306

    Google Scholar 

  • Van Auken O.W., Gese E.M., Connors K. (1985). Fertilization response of early and late successional species: Acacia smalliiand Celtis laevigata. Bot. Gaz. 146:564–569

    Article  Google Scholar 

  • Van der Schijff H.P. (1964). ‘n Hervaluasie van die probleem van bosindringing in Suid-Afrika. Tydskrif van Natuurwetenskappe 4:67–80

    Google Scholar 

  • Van Vegten J.A. (1983). Thornbush invasion in a savanna ecosystem in eastern Botswana. Vegetatio 56:3–7

    Article  Google Scholar 

  • Walker B.H., Knoop W.T. (1987). The response of the herbaceous layer in a dystrophic Burkea africana savanna to increased levels of nitrogen, phosphate and potassium. J. Grassland Soc. South. Afr. 4:31–34

    Google Scholar 

  • Walker B.H., Ludwig D., Holling C.S., Peterman R.M. (1981). Stability of semi-arid savanna grazing systems. J. Ecol. 69:473–498

    Article  Google Scholar 

  • Walter H. (1971). Ecology of Tropical and Subtropical Vegetation. Oliver & Boyd, Edinburgh

    Google Scholar 

  • Ward D. 2000. Rangeland vegetation overgrazing and recovery–Namibia. Final Report to US Agency for International Development, Tel Aviv, Israel.

  • Ward D. 2004. The effects of grazing on plant biodiversity in arid ecosystems. In: Shachak M., Pickett S.T.A., Gosz J. R.and Perevolotsky A. (eds), Biodiversity in drylands: towards a unified framework. Oxford University Press, Oxford, pp. 219–235

    Google Scholar 

  • Wiegand K., Jeltsch F., Ward D. (1999). Analysis of the population dynamics of Acacia trees in the Negev desert, Israel with a spatially-explicit computer simulation model. Ecol. Model. 117:203–224

    Article  Google Scholar 

  • Wiegand K., Ward D., Thulke H., Jeltsch F. (2000). From snapshot information to long-term population dynamics of Acacias by a simulation model. Plant Ecol. 150:97–114

    Article  Google Scholar 

  • Wiegand K., Ward D. and Saltz D. 2005. Multi-scale patterns and bush encroachment in an arid savanna with a shallow soil layer. J. Veg. Sci. 16: 311–320.

    Article  Google Scholar 

  • Wilson J.B. (1988). Shoot competition and root competition. J. Appl. Ecol. 25:279–296

    Article  Google Scholar 

  • Wilson T.B., Witkowski E.T.F. (1998). Water requirements for germination and early seedling establishment in four African savanna woody plant species. J. Arid Environ. 38:541–550

    Article  Google Scholar 

Download references

Acknowledgements

We are especially indebted to Errol and Barbara Tegg for their hospitality and many kindnesses during this study and without whom the work at Pniel Estates would not have been possible. We thank the Pniel community, and in particular Solomon Mojanaga, Ben Stemmer, Paul Swarts and George April for extensive help with setting up and running the field experiment. This study was funded by the University of Stellenbosch and the National Research Foundation of South Africa (to DW). We further acknowledge Kynoch Fertilizers and The House of Irrigation for donations.

Author information

Authors and Affiliations

Authors

Corresponding author

Correspondence to D. Ward.

Rights and permissions

Reprints and permissions

About this article

Cite this article

Kraaij, T., Ward, D. Effects of rain, nitrogen, fire and grazing on tree recruitment and early survival in bush-encroached savanna, South Africa. Plant Ecol 186, 235–246 (2006). https://doi.org/10.1007/s11258-006-9125-4

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s11258-006-9125-4

Keywords

Navigation