Abstract
Here we aim to demonstrate that in arid environments the competitive balance between species can be determined by niche separation with either nitrogen or water as the relevant niche axis. To do this we sampled roots <2 mm in diameter for 5 soil pits equidistant between two coexisting species, a shrub and a grass. Using stable carbon and nitrogen isotope ratios of fine roots we determine both photosynthetic pathway and rooting depth. We also examine the distribution of soil moisture and nitrogen relative to root biomass. Our results for root biomass and stable isotope ratios of fine roots demonstrate both niche separation and competition for resources. Root biomass is highest at the top of the profile where soil nitrogen is highest and soil moisture is lowest. We conclude that while there is competition for resources in the middle of the profile, competition is mitigated by photosynthetic pathway. The facultative CAM shrub grows whenever the soil at the surface is wet enough. The C4 photosynthetic pathway of the grass is more nitrogen and water use efficient making it better adapted to the low nitrogen in the middle of the profile and low summer rainfall.
Similar content being viewed by others
References
Belsky AJ (1994) Influences of trees on savanna productivity: Tests of shade, nutrients and tree grass competition. Ecology 75:922–932
Coetsee C, Bond WJ, February EC (2010) Frequent fire affects soil nitrogen and carbon in an African savanna by changing woody cover. Oecologia 162:1027–1034
Cowling RM, Esler KJ, Midgley GF, Honig MA (1994) Plant functional diversity, species-diversity and climate in arid and semi arid Southern Africa. J Arid Environ 27:141–158
Craine JM (2009) Resource Strategies of Wild Plants. Princeton University Press, Princeton
Cramer MD, Chimphango SBM, Van-Cauter A, Waldram MS, Bond WJ (2007) Grass competition induces N2 fixation in some species of African Acacia. J Ecol 95:1123–1133
Cushman JC, Borland AM (2002) Induction of crassulacean acid metabolism by water limitation. Plant Cell Environ 25:297–312
Desmet PG, Cowling RM (1999) Biodiversity, habitat and range-size aspects of a flora from a winter-rainfall desert in north-western Namaqualand, South Africa. pp 23–33. Kluwer Academic Publ.
Dickie IA, Schnitzer SA, Reich PB, Hobbie SE (2007) Is oak establishment in old-fields and savanna openings context dependent? J Ecol 95:309–320
February EC, Higgins SI (2010) The distribution of tree and grass roots in savannas in relation to soil nitrogen and water. S Afr J Bot 76:517–523
Francis ML, Fey MV, Prinsloo HP, Ellis F, Mills AJ, Medinski TV (2007) Soils of namaqualand: compensations for aridity. J Arid Environ 70:588–603
Griffiths H (1992) Carbon isotope discrimination and the integration of carbon assimilation pathways in terrestrial CAM plants. Plant Cell Environ 15:1051–1062
Hogberg P (1997) Tansley review No 95 - N-15 natural abundance in soil-plant systems. New Phytol 137:179–203
Kitayama K, Iwamoto K (2001) Patterns of natural 15 N abundance in the leaf-to-soil continuum of tropical rain forests differing in N availability on Mount Kinabalu, Borneo. Plant Soil 229:203–212
Midgley GF, van der Heyden F (1999) Form and function in perennial plants. In: Dean WRJ, Milton SJ (eds) The Karoo: ecological patterns and processes. Cambridge University Press, Cambridge, pp 91–106
Mooney HA, Troughton JH, Berry JA (1977) Carbon isotope measurements of succulent plants in southern Africa. Oecologia 30:295–305
Mordelet P, Menaut JC, Mariotti A (1997) Tree and grass rooting patterns in an African humid savanna. J Veg Sci 8:65–70
Mucina L, Rutherford MC Eds. (2006) The vegetation of South Africa, Lesotho and Swaziland. South African National Biodiversity Institute Pretoria.
O’Connor TG, Roux PW (1995) Vegetation changes (1949–71) in a semi-arid, grassy dwarf shrubland in the Karoo, South Africa: influence of rainfall variabilty and grazing by sheep. J Appl Ecol 32:612–626
Pardo LH, Nadelhoffer KJP (2010) Using nitrogen isotope ratios to assess terrestrial ecosystems at regional and global scales. In: West JB, Bowen GJ, Dawson TE, Tu KP (eds) Isoscapes: understanding movement, pattern, and process on Earth through isotope mapping. Springer-Verlag, New York, pp 221–249
Pärtell M, Wilson SD (2002) Root dynamics and spatial pattern in prairie and forest. Ecology 83:1199–1203
Post WM, Pastor J, Zinke PJ, Stangenberger AG (1985) Global patterns of soil nitrogen storage. Nature 317:613–616
Roux PW (1966) Die uitwerking van seisonreenval en beweiding op gemengde karooveld. Prc Grass Soc Sthn Afr 1:131–134
Roux PW, Vorster M (1983) Vegetation change in the Karoo. Prc Grass Soc Sthn Afr 18:18–24
Rubio G, Walk T, Ge ZY, Yan XL, Liao H, Lynch JP (2001) Root gravitropism and below-ground competition among neighbouring plants: A modelling approach. Ann Bot 88:929–940
Rundel PW, Esler KJ, Cowling RM (1999) Ecological and phylogenetic patterns of carbon isotope discrimination in the winter rainfall flora of the Richtersveld, South Africa. Plant Ecol 142:133–148
Sage RF, Pearcy RW (1987) The nitrogen use efficiency of C3 and C4 plants. ll. Leaf nitrogen effects on the gas exchange characteristics of Chenopodium album (L.) and Amaranthus retroflexus (L.). Plant Physiol 84:959–963
Schenk HJ, Jackson RB (2002) Rooting depths, lateral root spreads and below-ground/above ground allometries of plants in water-limited ecosystems. J Ecol 90:480–494
Scholes RJ, Archer SR (1997) Tree-grass interactions in savannas. Annu Rev Ecol Syst 28:517–544
Scholes RJ, Walker BH (eds) (1993) An African Savanna. Synthesis of the Nylsvley study. Cambridge University Press, Cambridge
Shiponeni NN (2007) Spatio-temporal distribution of grass and shrubs at the ecotone between an arid grassland and succulent shrubland: ecological interactions and influences of soils. PhD. Department of Botany. University of Cape Town
Vitousek PM, Shearer G, Kohl DM (1989) Foliar 15 N natural abundance in Hawaiin rainforest: patterns and possible mechanisms. Oecologia 78:383–388
Vogel JC, Fuls A, Ellis RP (1978) The Geographical distribution of Krantz Grasses in South Africa. S Afr J Sci 74:209–215
Walker BH, Noy-Meir I (1982) Aspects of the stability and resilience of savanna ecosystems. In: Huntley BJ (ed) Ecology of tropical savannas. Springer-Verlag, Berlin, pp 556–590
Walter H (1970) Vegetationszonen und Klima. Gustav Fischer, Verlag
Walter H (1971) Ecology of tropical and subtropical vegetation. Oliver and Boyd, Edinburgh
Watkeys MK (1999) Soils of the Arid south-western zone of Africa. In The Karoo: Ecological Patterns and Processes. Eds. W R J Dean and M S.J. Cambridge University Press
Westoby M (1980) Elements of a theory of vegetation dynamics in arid rangelands. Isr J Bot 28:169–194
Winter K, Holtum JAM (2002) How closely do the δ13 C values of crassulacean acid metabolism plants reflect the proportion of CO2 fixed during day and night? Plant Physiol 129:1843–1851
Acknowledgements
Funding from BIOTA Southern Africa sponsored by the German Federal Ministry of Education and Research under promotion number 01 LC 0024A is acknowledged. February acknowledges funding from both the South Africa NRF as well as the Mellon Foundation. Mr Christo Visser very kindly allowed us to conduct this research on his farm. Corli Coetsee and Stephan Woodbourne made suggestions on improving the manuscript.
Author information
Authors and Affiliations
Corresponding author
Additional information
Responsible Editor: Tibor Kalapos.
Rights and permissions
About this article
Cite this article
February, E.C., Allsopp, N., Shabane, T. et al. Coexistence of a C4 grass and a leaf succulent shrub in an arid ecosystem. The relationship between rooting depth, water and nitrogen. Plant Soil 349, 253–260 (2011). https://doi.org/10.1007/s11104-011-0867-y
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s11104-011-0867-y