Skip to main content
Log in

Tree-grass coexistence in a flood-disturbed, semi-arid savanna system

  • Research article
  • Published:
Landscape Ecology Aims and scope Submit manuscript

An Erratum to this article was published on 06 February 2010

Abstract

The coexistence of trees and grasses in savanna ecosystems is a contentious phenomenon. Fire and herbivory disturbances are often cited as major structuring forces that create a sustainable tree–grass relationship. However, periodic flooding of savanna patches may also enable coexistence. The aim of this study was to investigate the effects of flood-disturbance on the recruitment patterns of Acacia karroo trees in a semi-arid savanna system in South Africa. We analysed the spatial coincidence of A. karroo seedlings with tussocks of the tall spiny grass Stipagrostis namaquensis in the riverbed and related herbivory intensity to spatial position. The data showed that A. karroo seedlings were significantly positively associated with S. namaquensis (Chi-square test, \( \chi_{1}^{2} = 4 5. 20 \), n = 118, P < 0.001); A. karroo seedlings growing inside of tussocks experienced less browsing pressure than those growing in the floodplain (Kruskal–Wallis test, H = 11.90, n = 118, P < 0.01); and recruitment success of A. karroo trees was spatially discrete (K–S test, D = 0.78, n = 196, P < 0.01). We suggest that floods create an enemy-free zone, which S. namaquensis colonises and then facilitates successful A. karroo establishment. High levels of A. karroo recruitment in the riverbed may replenish the woodlands fringing the river, which appear to be sink areas for A. karroo seedlings. Thus, the interaction between disturbances at different spatial and temporal scales (flooding versus herbivory) seems to maintain the inherently unstable coexistence of tree and grass species in this ecosystem. These findings also suggested that flood disturbances alter the tree–grass relationship.

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

Access this article

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

Instant access to the full article PDF.

Fig. 1
Fig. 2
Fig. 3

Similar content being viewed by others

References

  • Statsoft Inc. (2004) Statisitca (data analysis software system), version 7. www.statsoft.com

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

    Article  Google Scholar 

  • Archibald S, Bond WJ (2003) Growing tall vs. growing wide: tree architecture and allometry of A. karroo in forest, savanna and arid environments. Oikos 102:3–14

    Article  Google Scholar 

  • Archibald S, Bond WJ, Stock WD, Fairbanks DHK (2005) Shaping the landscape: fire-grazer interactions in an African savanna. Ecol Appl 15:96–109

    Article  Google Scholar 

  • Bond WJ, Midgley GF, Woodward FI (2000) The importance of low atmospheric CO2 and fire in promoting the spread of grasslands and savannas. Glob Chang Biol 9:973982

    Google Scholar 

  • Bond WJ, Lee WG, Craine JM (2004) Plant structural defence against browsing birds: a legacy of New Zealand’s extinct moas. Oikos 104:500–508

    Article  Google Scholar 

  • Briske DD, Fuhlendorf SD, Smeins FE (2005) State-and-transition models, thresholds, and rangeland health: a synthesis of ecological concepts and perspectives. Rangeland Ecol Manag 58:1–10

    Article  Google Scholar 

  • Chhin S, Wang GG (2002) Spatial and temporal pattern of white spruce regeneration within mixed-grass prairie in the Spruce Woods Provincial Park of Manitoba. J Biogeogr 23:903–912

    Article  Google Scholar 

  • Cipollini HL, Wallace-Senft DA, Whigham DF (1994) A model of patch-dynamics, seed dispersal, and sex ration in the dioecious shrub Lendera benzoin (Lauraceae). J Ecol 82:621–633

    Article  Google Scholar 

  • Cumming GS, Barnes G, Southworth J (2008) Environmental asymmetries. In: Norberg J, Cumming GS (eds) Complexity theory for a sustainable future. Columbia University Press, New York, pp 15–46

    Google Scholar 

  • Darlington RB, Smulders TV (2001) Problems with residual analysis. Anim Behav 62:599–602

    Article  Google Scholar 

  • Dean WRJ, Milton SJ, Jeltsch F (1999) Large trees, fertile islands, and birds in arid savannah. J Arid Environ 41:61–78

    Article  Google Scholar 

  • Duncan RP (1993) Flood disturbance and the coexistence of species in a lowland podocarp forest, South Westland, New Zealand. J Ecol 81:403–416

    Article  Google Scholar 

  • Fornara DA, Du Toit JT (2008) Responses of woody sapling exposed to chronic mammalian herbivory in an African savanna. Ecoscience 15:129–135

    Article  Google Scholar 

  • Fukami T (2001) Sequence effects of disturbance on community structure. Oikos 92:215–224

    Article  Google Scholar 

  • Gagnon PR, Platt WJ, Moser EB (2007) Response of a native bamboo [Arundinaria gigantean (Walt.) Muhl.] in a wind-disturbed forest. Forest Ecol Manag 241:288–294

    Article  Google Scholar 

  • Gillson L (2004) Evidence of hierarchical patch dynamics in an east African savanna? Landscape Ecol 19:883–894

    Article  Google Scholar 

  • Goheen JR, Keesing F, Allan BF, Ogada D, Ostfeld RS (2004) Net effects of large mammals on Acacia seedling survival in an African savanna. Ecology 85:1555–1561

    Article  Google Scholar 

  • Goheen JR, Young TP, Keesing F, Palmer TM (2007) Consequences of herbivory by native ungulates for the reproduction of a savanna tree. J Ecol 95:129–138

    Article  Google Scholar 

  • Gregory SV, Swanson FJ, McKee WA, Cummins KW (1991) An ecosystem perspective of riparian zones. Bioscience 41:540–551

    Article  Google Scholar 

  • Grubb PJ (1992) A positive distrust in simplicity: lessons from plant defence and from competition among plants and among animals. Ecology 80:585–610

    Article  Google Scholar 

  • Higgins SI, Bond WJ, Trollope WSW (2000) Fire, resprouting and variability: a recipe for grass-tree coexistence in savannah. J Ecol 88:213–229

    Article  Google Scholar 

  • Hobbs RJ, Huenneke LF (1992) Disturbance, diversity and invasion: implications for conservation. Conserv Biol 6:324–337

    Article  Google Scholar 

  • Holl KD (2002) Effect of shrubs on tree seedling establishment in an abandoned tropical pasture. J Ecol 90:179–187

    Article  Google Scholar 

  • House JI, Archer S, Breshears DD, Scholes RJ, NCEAS Tree-Grass Interaction Participants (2003) Conundrums in mixed woody-herbaceous plant systems. J Biogeogr 30:1763–1777

    Article  Google Scholar 

  • Jeffries MJ, Lawton JH (1984) Enemy free space and the structure of ecological communities. Biol J Linn Soc 23:269–286

    Article  Google Scholar 

  • Jeltsch F, Weber GE, Grimm V (2000) Ecological buffering mechanisms in savannahs: a unifying theory of tree-grass coexistence. Plant Ecol 161:161–171

    Article  Google Scholar 

  • Jurena PN, Archer S (2003) Woody plant establishment and spatial heterogeneity in grasslands. Ecology 84:907–919

    Article  Google Scholar 

  • Levin SA, Paine RT (1974) Disturbance, patch formation, and community structure. Proc Nat Acad Sci USA 71:2744–2747

    Article  PubMed  CAS  Google Scholar 

  • Medina E, Silva JF (1990) Savannahs of northern South America: a steady state regulated by water-fire interactions on a background of low nutrient availability. J Biogeogr 17:403–413

    Article  Google Scholar 

  • Milewski AV, Madden D (2006) Interactions between large African browsers and thorny Acacia on a wildlife ranch in Kenya. Afr J Ecol 44:515–522

    Article  Google Scholar 

  • Milton SJ, Dean WRJ, Kerley GIH (1992) Tierberg Karoo Research Centre: history, physical environment, flora and fauna. T Roy Soc S Afr 48:15–46

    Google Scholar 

  • Mucina L, Jurgens N, le Roux A, Rutherford MC, Schmiedel U, Esler KJ, Powrie LW, Desmet PG, Milton SJ (2006) Succulent Karoo Biome. In: Mucina L, Rutherford MC (eds) The Vegetation of South Africa, Lesotho and Swaziland, Strelitzia 19. South African National Biodiversity Institute, Pretoria, pp 221–299

    Google Scholar 

  • Naiman RJ, Decamps H (1997) The ecology of interfaces: riparian zones. Ann Rev Ecol Syst 28:621–658

    Article  Google Scholar 

  • O’Connor TG (1995) Acacia karroo invasion of grassland: environmental and biotic effects influencing seedling emergence and establishment. Oecologia 103:214–223

    Article  Google Scholar 

  • Paige KN (1992) Overcompensation in response to mammalian herbivory: from mutualistic to antagonistic interactions. Ecology 73:2076–2085

    Article  Google Scholar 

  • Pettit NE, Naiman RJ (2006) Flood-deposited wood creates regeneration niches for riparian vegetation on a semi-arid South African river. J Veg Sci 17:615–624

    Article  Google Scholar 

  • Pinay G, Ruffinoni C, Fabre A (1995) Nitrogen cycling in two riparian forest soils under different geomorphic conditions. Biogeochemistry 30:9–29

    Article  CAS  Google Scholar 

  • Pulliam HR (1988) Sources, sinks and population regulation. Am Nat 132:652–661

    Article  Google Scholar 

  • Rooke T, Bergstrom R, Skarpe C, Danell K (2004) Morphological responses of woody species to simulated twig-browsing in Botswana. J Trop Ecol 20:281–289

    Article  Google Scholar 

  • Rubin F, Palmer AR (1996) The physical environment and major plant communities of the Karoo National Park, South Africa. Koedoe 39:25–52

    Google Scholar 

  • 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, Forst 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 438:846–849

    Article  PubMed  CAS  Google Scholar 

  • Scholes RJ, Archer SR (1997) Tree-grass interactions in savannahs. Ann Rev Ecol Syst 28:517–544

    Article  Google Scholar 

  • Shreve F (1931) Physical conditions in sun and shade. Ecology 12:96–104

    Article  Google Scholar 

  • Simmons MT, Archer SR, Ansley RJ, Teague WR (2007) Grass effects of tree (Prosopis glandulosa) growth in an arid savannah. J Arid Environ 69:212–227

    Article  Google Scholar 

  • Smit GN (2005) Tree thinning as an option to increase herbaceous yield of an encroached semi-arid savannah in South Africa. BioMed Cent Ecol 5:4

    Google Scholar 

  • Sweet SS (1980) Allometric inference in biology. Integr Comp Biol 20:643–652

    Article  Google Scholar 

  • Tewksbury JJ, Lloyd JD (2001) Positive interactions under nurse plants: spatial scale, stress gradients and benefactor size. Oecologia 127:425–434

    Article  Google Scholar 

  • Tiver F, Andrew MH (1997) Relative effects of herbivory by sheep, rabbits, goats, and kangaroos on recruitment and regeneration of shrubs and trees in eastern South Australia. J Appl Ecol 34:903–914

    Article  Google Scholar 

  • Van Auken OW (2000) Shrub invasions of North American semiarid grasslands. Annu Rev Ecol Syst 31:197–215

    Article  Google Scholar 

  • Van Langevelde FC, van de Vijver AD, 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 84:337–350

    Article  Google Scholar 

  • Walter H (1971) Ecology of tropical and subtropical vegetation. Oliver & Boyd, Edinburgh

    Google Scholar 

  • Wu J, Levin SA (1994) A spatial patch dynamic modelling approach to pattern and process in an annual grassland. Ecol Monogr 64:447–464

    Article  Google Scholar 

  • Wu J, Loucks OL (1995) From balance of nature to hierarchical patch dynamics: a paradigm shift in ecology. Q Rev Biol 70:439–466

    Article  Google Scholar 

  • Young TP (1987) Increased thorn length in Acacia depranolobium- an induced response to browsing. Oecologia 71:436–438

    Article  Google Scholar 

  • Young TP, Lindsay WK (1988) Role of even-age population structure in the disappearance of Acacia xanthophloea woodlands. Afr J Ecol 26:69–72

    Article  Google Scholar 

Download references

Acknowledgements

We thank the Percy FitzPatrick Institute of African Ornithology for providing funding for the project. The study was stimulated by discussions with Dr. Shayne Jacobs, Conservation Ecology and Entomology, University of Stellenbosch. We also thank three anonymous reviewers for greatly improving this manuscript.

Author information

Authors and Affiliations

Authors

Corresponding author

Correspondence to Matthew F. Child.

Additional information

An erratum to this article can be found at http://dx.doi.org/10.1007/s10980-010-9455-4

Electronic supplementary material

Below is the link to the electronic supplementary material.

Supplementary material 1 (DOCX 2141 kb)

Rights and permissions

Reprints and permissions

About this article

Cite this article

Child, M.F., Milton, S.J., Dean, R.W.J. et al. Tree-grass coexistence in a flood-disturbed, semi-arid savanna system. Landscape Ecol 25, 315–326 (2010). https://doi.org/10.1007/s10980-009-9409-x

Download citation

  • Received:

  • Accepted:

  • Published:

  • Issue Date:

  • DOI: https://doi.org/10.1007/s10980-009-9409-x

Keywords

Navigation