Abstract
Aims
While patterns of variation in woody plant aboveground traits related to disturbance and resource availability in savanna ecosystems are fairly well understood, dimensions of variation in belowground traits remain poorly understood. We investigate how sapling coarse root (>2 mm diameter) traits and belowground storage organs vary with respect to differing disturbance regimes (i.e. fire or herbivore dominated) and soil fertility in Southern African savannas.
Methods
We examined how sapling rooting strategies (type of rooting system, storage organs, root allocation) as well as root total non-structural carbohydrate concentrations [TNC], root N and C:N of 69 common woody savanna and forest species at 16 sites vary with respect to differing disturbance regimes and soil fertility.
Results
We found that root [TNC] and the occurrence of storage organs were highest while root N was lowest for species growing at fire-driven sites, compared to herbivore-driven sites and competition-controlled forest species. Allocation to belowground biomass was higher at fire-dominated sites but root volume:stem basal area did not differ between disturbance regimes. None of the measured traits were found to differ between nutrient rich clayey soils and nutrient poor sandy soils.
Conclusions
Our results suggest that disturbance related controls are important drivers of savanna belowground traits.
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References
Archibald S, Bond W (2003) Growing tall vs growing wide: tree architecture and allometry of Acacia karroo in forest, savanna, and arid environments. Oikos 102:3–14
Archibald S, Roy DP, Van Wilgen BW, Scholes RJ (2009) What limits fire? An examination of drivers of burnt area in southern Africa. Glob Chang Biol 15:613–630
Bhattachan A, Tatlhego M, Dintwe K, O'Donnell F, Caylor KK, Okin GS, Perrot DO, Ringrose S, D'odorico P (2012) Evaluating Ecohydrological theories of Woody root distribution in the Kalahari. PLoS One 7:e33996
Bond WJ (2005) Large parts of the world are brown or black: a different view on the ‘green world’ hypothesis. J Veg Sci 16:261–266
Bond WJ (2008) What limits trees in C4 grasslands and savannas? Annu Rev Ecol Evol Syst 39:641–659
Bond W, Keeley J (2005) Fire as a global “herbivore”: the ecology and evolution of flammable ecosystems. Trends Ecol Evol 20:387–394
Bond WJ, Midgley JJ (2001) Ecology of sprouting in woody plants: the persistence niche. Trends Ecol Evol 16:45–51
Boone RD, Grigal DF, Sollins P, Ahrens RJ, Armstrong DE (1999) Soil sampling, preparation, archiving, and quality control standard soil methods for long-term ecological research. Oxford University Press, New York, pp 3–28
Burrows GE, Hornby SK, Waters DA, Bellairs SM, Prior LD, Bowman DMJS (2008) Leaf axil anatomy and bud reserves in 21 Myrtaceae species from northern Australia. Int J Plant Sci 169:1174–1186
Charles-Dominique T, Beckett H, Midgley GF, Bond WJ (2015a) Bud protection: a key trait for species sorting in a forest–savanna mosaic. New Phytol 207:1052–1060
Charles-Dominique T, Midgley GF, Bond WJ (2015b) An index for assessing effectiveness of plant structural defences against mammal browsing. Plant Ecol 216:433–1440
Charles-Dominique T, Staver AC, Midgley GF, Bond WJ (2015c) Functional differentiation of biomes in an African savanna/forest mosaic. S Afr J Bot 101:82–90
Clarke PJ, Lawes MJ, Midgley JJ, Lamont BB, Ojeda F, Burrows GE, Enright NJ, Knox KJE (2012) Resprouting as a key functional trait: how buds, protection and resources drive persistence after fire. New Phytol 197:19–35
Cramer MD, van Cauter A, Bond WJ (2010) Growth of N2-fixing African savanna Acacia species is constrained by below-ground competition with grass. J Ecol 98:156–167
Dantas VL, Pausas JG (2013) The lanky and the corky: fire-escape strategies in savanna woody species. J Ecol 101:1265–1272
Dietze MC, Sala A, Carbone MS, Czimczik CI, Mantooth JA, Richardson AD, Vargas R (2014) Nonstructural carbon in Woody plants. Annu Rev Plant Biol 65:667–687
Dybzinski R, Farrior C, Wolf A, Reich PB, Pacala SW (2011) Evolutionarily stable strategy carbon allocation to foliage, wood, and fine roots in trees competing for light and nitrogen: an analytically tractable, individual-based model and quantitative comparisons to data. Am Nat 177:153–166
Fornara DA, DuToit JD (2007) Browsing lawns? Responses of Acacia nigrescens to ungulate browsing in an African savanna. Ecology 88:200–209
Fornara DA, DuToit JT (2008) Community-level interactions between ungulate browsers and woody plants in an African savanna dominated by palatable-spinescent Acacia trees. J Arid Environ 72:534–545
Gignoux J, Clobert J, Menaut J (1997) Alternative fire resistance strategies in savanna trees. Oecologia 110:576–583
Grady JM, Hoffmann WA (2012) Caught in a fire trap: recurring fire creates stable size equilibria in woody resprouters. Ecology 93:2052–2060
Haissig BE, DICKSON RE (1979) Starch measurement in plant tissue using enzymatic hydrolysis. Physiol Plant 47:151–157
Hansen J, Møller I (1975) Percolation of starch and soluble carbohydrates from plant tissue for quantitative determination with anthrone. Anal Biochem 68:87–94
Higgins SI, Bond WJ, Trollope WS (2000) Fire, resprouting and variability: a recipe for grass–tree coexistence in savanna. J Ecol 88:113–229
Hoffmann W, Orthen B, Do Nascimento P (2003) Comparative fire ecology of tropical savanna and forest trees. Funct Ecol 17:720–726
Hoffmann W, Orthen B, Franco A (2004) Constraints to seedling success of savanna and forest trees across the savanna-forest boundary. Oecologia 140:252–260
Janos DP, Scott J, Bowman DMJS (2008) Temporal and spatial variation of fine roots in a northern Australian Eucalyptus tetrodonta savanna. J Trop Ecol 24:177–188
Kellman M (1979) Soil enrichment by neotropical savanna trees. J Ecol 67:565–577
Kruger L, Charles-Dominique T, Bond W, Midgley J, Balfour D, Mkhwanazi A (2017) Woody Plant traits and life-history strategies across disturbance gradients and biome boundaries in the Hluhluwe-iMfolozi park. In: Cromsigt J, Archibald S, Owen-Smith N (eds) Conserving Africa's mega-diversity in the Anthropocene: the Hluhluwe-iMfolozi park story. Cambridge University Press, Cambridge, p 406
Kulmatiski A, Beard KH, Verweij RJ, February EC (2010) A depth-controlled tracer technique measures vertical, horizontal and temporal patterns of water use by trees and grasses in a subtropical savanna. New Phytol 188:199–209
Laliberté E (2016) Below-ground frontiers in trait-based plant ecology. New Phytol 213:1597–1603
Lambers H, Raven J, Shaver G, Smith S (2008) Plant nutrient-acquisition strategies change with soil age. Trends Ecol Evol 23:95–103
Mokany K, Raison R, Prokushkin A (2006) Critical analysis of root: shoot ratios in terrestrial biomes. Glob Chang Biol 12:84–96
Mordelet P, Menaut JC, Mariotti A (1997) Tree and grass rooting patterns in an African humid savanna. J Veg Sci 8:65–70
Mugasha WA, Eid T, Bollandsås OM, Malimbwi RE, Chamshama SAO, Zahabu E, Katani JZ (2013) Allometric models for prediction of above- and belowground biomass of trees in the miombo woodlands of Tanzania. Forest Ecol Manag 310:87–101
O'Donnell FC, Caylor KK, Bhattachan A, Dintwe K, D'odorico P, Okin GS (2015) A quantitative description of the interspecies diversity of belowground structure in savanna woody plants. Ecosphere 6:1–15
Plante PM, Rivest D, Vézina A, Vanasse A (2014) Root distribution of different mature tree species growing on contrasting textured soils in temperate windbreaks. Plant Soil 380:429–439
R Development Core Team (2016) R: A language and environment for statistical computing [Computer software] Vienna: R Foundation for Statistical Computing
Raz-Yaseef N, Koteen L, Baldocchi DD (2013) Coarse root distribution of a semi-arid oak savanna estimated with ground penetrating radar. J Geophys Res Biogeosci 118:135–147
Ryan CM, Williams M (2011) How does fire intensity and frequency affect miombo woodland tree populations and biomass? Ecol Appl 21:48–60
Sala OE, Gherardi LA, Reichmann L, Jobbagy E, Peters D (2012) Legacies of precipitation fluctuations on primary production: theory and data synthesis. Philos Trans R Soc Lond Ser B Biol Sci 367:3135–3144
Schenk HJ, Jackson RB (2002) The global biogeography of roots. Ecol Monogr 72:311–328
Schutz AEN, Bond WJ, Cramer MD (2009) Juggling carbon: allocation patterns of a dominant tree in a fire-prone savanna. Oecologia 160:235–246
Schutz AEN, Bond WJ, Cramer MD (2010) Defoliation depletes the carbohydrate reserves of resprouting Acacia saplings in an African savanna. Plant Ecol 212:2047–2055
Staver AC (2018) Prediction and scale in savanna ecosystems. New Phytol 219:52–57
Staver AC, Bond WJ (2014) Is there a ‘browse trap’? Dynamics of herbivore impacts on trees and grasses in an African savanna. J Ecol 102:595–602
Staver AC, Bond WJ, Cramer MD, Wakeling JL (2012) Top-down determinants of niche structure and adaptation among African acacias. Ecol Lett 15:673–679
Staver AC, Botha J, Hedin LO (2017) Soils and fire jointly determine vegetation structure in an African savanna. New Phytol 216:1151–1160
Strong WL, La Roi GH (1985) Root density-soil relationships in selected boreal forests of Central Alberta, Canada. For Ecol Manag 12:233–251
Tjoelker TMG, Craine JM, Wedin D, Reich PB, Tilman D (2005) Linking leaf and root trait syndromes among 39 grassland and savannah species. New Phytol 167:493–508
Tomlinson KW, Sterck FJ, Bongers F, da Silva DA, Barbosa ERM, Ward D, Bakker FT, van Kaauwen M, Prins HHT, de Bie S, van Langevelde F (2012) Biomass partitioning and root morphology of savanna trees across a water gradient. J Ecol 100:1113–1121
Trollope WSW, Tainton NM (1986) Effect of fire intensity on the grass and bush components of the eastern cape thornveld. J Grassland Soc Southern Afr 3:37–42
Walkley A (1947) A critical examination of a rapid method for determining organic carbon in soils-effect of variations in digestion conditions and of inorganic soil constituents. Soil Sci 63:251–264
Wigley BJ, Cramer MD, Bond WJ (2009) Sapling survival in a frequently burnt savanna: mobilisation of carbon reserves in Acacia karroo. Plant Ecol 203:1–11
Wigley BJ, Bond WJ, Fritz H, Coetsee C (2015) Mammal browsers and rainfall affect Acacia leaf nutrient content, defense, and growth in south African savannas. Biotropica 47:190–200
Wigley BJ, Slingsby JA, Díaz S, Bond WJ, Fritz H, Coetsee C (2016) Leaf traits of African woody savanna species across climate and soil fertility gradients: evidence for conservative versus acquisitive resource-use strategies. J Ecol 104:1357–1369
Wigley BJ, Fritz H, Coetsee C (2018) Defence strategies in African savanna trees. Oecologia 187:797–809
Zhou Y, Boutton TW, Ben Wu X, Wright CL, Dion AL (2018) Rooting strategies in a subtropical savanna: a landscape-scale three-dimensional assessment. Oecologia 186:1127–1135
Acknowledgements
We are grateful to SANParks, Ezemvelo KZN Wildlife, the Zimbabwe Parks and Wildlife Authority and several anonymous private landowners for their support. Lab analyses for this study were partially supported by the Institute of Dendrology, Polish Academy of Sciences, Kórnik, Poland. We also want to thank the reviewers for their thoughtful inputs and comments on an earlier draft.
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Wigley, B.J., Staver, A.C., Zytkowiak, R. et al. Root trait variation in African savannas. Plant Soil 441, 555–565 (2019). https://doi.org/10.1007/s11104-019-04145-3
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DOI: https://doi.org/10.1007/s11104-019-04145-3