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Termites facilitate and ungulates limit savanna tree regeneration

  • Plant-animal interactions - Original research
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Abstract

Both large herbivores and termites are key functional groups in savanna ecosystems, and in many savanna areas, large termite mounds (termitaria) are associated with distinct woody clusters. Studies on the effect of large mammals on tree regeneration are few, and the results are conflicting. Large herbivores have been found to be important seedling predators in some areas, but facilitate tree regeneration by outcompeting small mammals and reducing grass cover in other areas. Through the use of the experimental fencing of termite mounds and adjacent savanna areas in this study, we investigated how termites and large herbivores influence tree regeneration. Termite mounds had a higher number of seedlings, more species richness, more alpha diversity (OD) and lower evenness (E) than savanna plots. Large herbivores did not significantly affect overall seedling density, species richness, OD or E. Beta diversity was higher in savanna areas than on termitaria, and beta diversity decreased in savanna areas when herbivores were excluded. Herbivore exclusion increased the density of the 12 (40 %) most common seedling species, representing 79 % of all seedlings, and fenced plots had relatively taller seedlings than open plots. Thus, termites were the main determinants of tree regeneration in our study area, but large mammals regulated the most common species. Although our study confirms previous work suggesting that large herbivores affect tree regeneration, we found that termites were an even more important determinant. Termite impacts on tree regeneration deserve increased attention by savanna ecologists.

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References

  • Anderson T (2008) Plant compositional change over time increases with rainfall in Serengeti grasslands. Oikos 117:675–682

    Article  Google Scholar 

  • Asner GP, Levick SR, Kennedy-Bowdoin T, Knapp DE, Emerson R, Jaconson J, Colgan MS, Martin RE (2009) Large-scale impacts of herbivores on the structural diversity of African savannas. Proc Natl Acad Sci USA 106:4947–4952

    Article  PubMed  CAS  Google Scholar 

  • Augustine DJ, McNaughton SJ (2004) Regulation of shrub dynamics by native browsing ungulates on East African rangeland. J Appl Ecol 41:45–58

    Article  Google Scholar 

  • Bakker ES, Olff H (2003) Impact of different-sized herbivores on recruitment opportunities for subordinate herbs in grasslands. J Veg Sci 14:465–474

    Article  Google Scholar 

  • Bardgett RD, Wardle DA (2003) Herbivore-mediated linkages between aboveground and belowground communities. Ecology 84:2258–2268

    Article  Google Scholar 

  • Beentje HJ, Cheek M (2003) Flora of tropical East Africa: glossary. Balkema, Lisse

    Google Scholar 

  • Bloesch U (2002) The dynamics of thicket clumps in the Kagera savanna landscape, East Africa. Swiss Federal Institute of Technology, Zurich

    Google Scholar 

  • Bloesch U (2008) Thicket clumps: a characteristic feature of the Kagera savanna landscape, East Africa. J Veg Sci 19:31–44

    Article  Google Scholar 

  • Bond WJ (2008) What limits trees in C4 grassy biomes? Annu Rev Ecol Syst 39:641–659

    Article  Google Scholar 

  • Bond WJ, Smythe K-A, Balfour DA (2001) Acacia species turnover in space and time in an African savanna. J Biogeogr 28:117–128

    Article  Google Scholar 

  • Brody A, Palmer T, Fox-Dobbs K, Doak D (2010) Termites, vertebrate herbivores, and the fruiting success of Acacia drepanolobium. Ecology 91:399–407

    Article  PubMed  Google Scholar 

  • Bucini G, Hanan NP (2007) A continental-scale analysis of tree cover in African savannas. Glob Ecol Biogeogr 16:593–605

    Article  Google Scholar 

  • Chase JM (2010) Stochastic community assembly causes higher biodiversity in more productive environment. Science 328:1388–1391

    Article  PubMed  CAS  Google Scholar 

  • Davies AB, Parr CL, Van Rensburg BJ (2010) Termites and fire: current understanding and future research directions for improved savanna conservation. Aust Ecol 35:482–486

    Article  Google Scholar 

  • de Knegt HJ, Groen TA, van de Vijver CADM, Prins HHT, van Langevelde H (2008) Herbivores as architects of savannas: inducing and modifying spatial vegetation patterning. Oikos 117:543–554

    Article  Google Scholar 

  • Dickson TL, Foster BL (2008) The relative importance of the species pool, productivity and disturbance in regulating grassland plant species richness: a field experiment. J Ecol 96:937–946

    Article  Google Scholar 

  • Fensham RJ, Fairfax RJ, Buckley YM (2008) An experimental study of fire and moisture stress on the survivorship of savanna eucalypt seedlings. Aust J Bot 56:693–697

    Article  Google Scholar 

  • 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 91:1296–1307

    Article  PubMed  Google Scholar 

  • Gignoux J, Lahoreau G, Julliard R, Barot S (2009) Establishment and early persistence of tree seedlings in an annually burned savanna. J Ecol 97:484–495

    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, Palmer TM, Keesing F, Riginos C, Young TP (2010) Large herbivores facilitate savanna tree establishment via diverse and indirect pathways. J Anim Ecol 79:372–382

    Article  PubMed  Google Scholar 

  • Hillebrand H, Gruner D, Borer E, Bracken M, Cleland E, Elser J, Harpole W, Ngai J, Seabloom E, Shurin J (2007) Consumer versus resource control of producer diversity depends on ecosystem type and producer community structure. Proc Natl Acad Sci USA 104:10904–10909

    Article  PubMed  CAS  Google Scholar 

  • Hoffmann WA (1996) The effects of fire and cover on seedling establishment in a neotropical savanna. J Ecol 84:383–393

    Article  Google Scholar 

  • Holdo RM, Holt RD, Fryxell JM (2009) Grazers, browsers, and fire influence the extent and spatial pattern of tree cover in Serengeti. Ecol Appl 19:95–109

    Article  PubMed  Google Scholar 

  • Huston M (1979) General hypothesis of species-diversity. Am Nat 113:81–101

    Article  Google Scholar 

  • Jones JA (1990) Termites, soil fertility and carbon cycling in dry tropical Africa: a hypothesis. J Trop Ecol 6:291–305

    Article  Google Scholar 

  • 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 206:47–60

    Article  Google Scholar 

  • Kondoh M (2001) Unifying the relationships of species richness to productivity and disturbance. Proc R Soc Lond B 268:269–271

    Article  CAS  Google Scholar 

  • Kvålseth TO (1991) Note on biological diversity, evenness, and homogeneity measures. Oikos 62:123–127

    Article  Google Scholar 

  • Levick S, Rogers K (2009) Patch and species specific responses of savanna woody vegetation to browser exclusion. Biol Conserv 141:489–498

    Article  Google Scholar 

  • Levick SR, Asner GP, Kennedy-Bowdoin T, Knapp DE (2010a) The spatial extent of termite influences on herbivore browsing in an African savanna. Biol Conserv 143:2462–2467

    Article  Google Scholar 

  • Levick SR, Asner GP, Kennedy-Bowdoin T, Knapp DE (2010b) The relative influence of fire and herbivory on savanna three-dimensional vegetation structure. Biol Conserv 142:1693–1700

    Article  Google Scholar 

  • Loveridge JP, Moe SR (2004) Termitaria as browsing hotspots for African megaherbivores in miombo woodland. J Trop Ecol 20:337–343

    Article  Google Scholar 

  • Midgley JJ, Lawes MJ, Chamaillé-Jammes S (2010) Savanna woody plant dynamics: the role of fire and herbivory, separately and synergistically. Aust J Bot 58:1–11

    Article  Google Scholar 

  • Mills AJ, Milewski A, Fey MV, Groengroeft A, Petersen A (2009) Fungus culturing, nutrient mining and geophagy: a geochemical investigation of Macrotermes and Trinervitermes mounds in southern Africa. J Zool 278:24–35

    Article  Google Scholar 

  • Mobæk R, Narmo AK, Moe SR (2005) Termitaria are focal feeding sites for large ungulates in Lake Mburo National Park, Uganda. J Zool 267:97–102

    Article  Google Scholar 

  • Moe SR, Mobæk R, Narmo AK (2009a) Mound building termites contribute to savanna vegetation heterogeneity. Plant Ecol 202:31–40

    Article  Google Scholar 

  • Moe SR, Rutina LP, Hytteborn H, du Toit JT (2009b) What controls woodland regeneration after elephants have killed the big trees? J Appl Ecol 46:223–230

    Article  Google Scholar 

  • Oksanen J, Blanchet FG, Kindt R, Legendre P, O’Hara RB, Simpson GL, Solymos P, Stevens MHH, Wagner H (2010) Vegan: community ecology package. R package version 1.17-4. Internet: http://CRAN.R-project.org/package=vegan. Accessed 2 Oct 2010

  • Okullo P, Moe SR (2012a) Termite activity, not grazing, is the main determinant of spatial variation in savanna herbaceous vegetation. J Ecol 100:232–241

    Article  Google Scholar 

  • Okullo P, Moe SR (2012b) Large herbivores maintain termite-caused differences in herbaceous species diversity patterns. Ecology 93:2095–2103

    Google Scholar 

  • Olff H, Ritchie M (1998) Effects of herbivores on grassland plant diversity. Trends Ecol Evol 13:261–265

    Article  PubMed  CAS  Google Scholar 

  • Osem Y, Perevolotsky A, Kigel J (2004) Site productivity and plant size explain the response of annual species to grazing exclusion in a Mediterranean semi-arid rangeland. J Ecol 92:297–309

    Article  Google Scholar 

  • Petty AM, Werner PA, Lehmann CER, Riley JE, Banfai DS, Elliott LP (2007) Savanna responses to feral buffalo in Kakadu national park, Australia. Ecol Monogr 77:441–463

    Article  Google Scholar 

  • Porensky LM, Veblen KE (2012) Grasses and browsers reinforce landscape heterogeneity by excluding trees from ecosystem hotspots. Oecologia 168:749–759

    Article  PubMed  Google Scholar 

  • Pringle RM, Doak DF, Brody AK, Jocquè R, Palmer TM (2010) Spatial pattern enhances ecosystem functioning in an African savanna. PLoS Biol 8:e1000377. doi:10.1371/journal.pbio.1000377

    Article  PubMed  Google Scholar 

  • Prins HHT, van der Jeugd HP (1993) Herbivore population crashes and woodland structure in East-Africa. J Ecol 81:305–314

    Article  Google Scholar 

  • Proulx M, Mazumder A (1998) Reversal of grazing impact on plant species richness in nutrient-poor vs. nutrient-rich ecosystems. Ecology 79:2581–2592

    Article  Google Scholar 

  • R Development Core Team (2010) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. ISBN 3-900051-07-0, URL http://www.R-project.org

  • Rannestad OT, Danielsen T, Moe SR, Stokke S (2006) Adjacent pastoral areas support higher densities of wild ungulates during the wet season than the Lake Mburo National Park in Uganda. J Trop Ecol 22:675–683

    Article  Google Scholar 

  • Reed DN, Anderson TM, Dempewolf J, Metzger K, Serneels S (2009) The spatial distribution of vegetation types in the Serengeti ecosystem: the influence of rainfall and topographic relief on vegetation patch characteristics. J Biogeogr 36:770–782

    Article  Google Scholar 

  • Renard D, Birk JJ, Glaser B, Iriarte J, Grisard G, Karl J, McKey D (2012) Origin of mound-field landscapes: a multi-proxy approach combining contemporary vegetation, carbon stable isotopes and phytoliths. Plant Soil 341:337–353

    Article  Google Scholar 

  • Riginos C (2009) Grass competition suppresses savanna tree growth across multiple demographic stages. Ecology 90:335–340

    Article  PubMed  Google Scholar 

  • Riginos C, Young TP (2007) Positive and negative effects of grass, cattle, and wild herbivores on Acacia saplings in an East African savanna. Oecologia 153:985–995

    Article  PubMed  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, Frost 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 savannas. Annu Rev Ecol Syst 28:517–544

    Article  Google Scholar 

  • Sharam G, Sinclair ARE, Turkington R (2006) Establishment of broadleaved thickets in Serengeti, Tanzania: the influence of fire, browsers, grass competition, and elephants. Biotropica 38:599–605

    Article  Google Scholar 

  • Shaw MT, Keesing F, Ostfeld RS (2002) Herbivory on Acacia seedlings in an East African savanna. Oikos 99:385–392

    Article  Google Scholar 

  • Sileshi G, Arshad M, Konaté S, Nkunika P (2010) Termite induced heterogeneity in African savanna vegetation: mechanisms and patterns. J Veg Sci 21:923–937

    Article  Google Scholar 

  • Smith B, Wilson JB (1996) A consumer’s guide to evenness indices. Oikos 76:70–82

    Article  Google Scholar 

  • Spain AV, McIvor JG (1988) The nature of herbaceous vegetion associated with termitaria in North Eastern Australia. J Ecol 76:181–191

    Article  Google Scholar 

  • Staver C, Levin SA (2012) Integrating theoretical climate and fire effects on Savanna and forest systems. Am Nat 180:211–224

    Article  PubMed  Google Scholar 

  • Staver C, Bond WJ, Stock WD, van Rensburg SJ, Waldram MS (2009) Browsing and fire interact to suppress tree density in an African savanna. Ecol Appl 19:1909–1919

    Article  PubMed  Google Scholar 

  • Traoré S, Lepage M (2008) Effects of controlled livestock grazing and annual prescribed fire on epigeal termite mounds in a savannah woodland in Burkina Faso. Insect Soc 55:183–189

    Article  Google Scholar 

  • Traoré S, Nygård R, Guinko S, Lepage M (2008) Impact of Macrotermes termitaria as a source of heterogeneity on tree diversity and structure in a Sudanian savannah under controlled grazing and annual prescribed fire (Burkina Faso). For Ecol Manag 255:2337–2346

    Article  Google Scholar 

  • Williams PR (2009) Contrasting demographics of tropical savanna and temperate forest eucalypts provide insight into how savannas and forests function. A case study using Corymbia clarksoniana from north-eastern Australia. Aust Ecol 34:120–131

    Article  Google Scholar 

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Acknowledgments

We thank James Nkwatsibwe, Patrict Bodo, Antonia Nyamukuru and Protase Rwaburindore for field assistance. We also thank Rose Kentalu who ensured that the meals were always ready, and the Chief Park warden, Adonia Bintora, for help and support in providing field work permits. We also thank Jon E. Swenson for reading a former draft and for his useful comments. The experiments comply with the current laws of the country (Uganda) in which the experiments were performed.

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Correspondence to Ole-Gunnar Støen.

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Communicated by Peter Clarke.

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Støen, OG., Okullo, P., Eid, T. et al. Termites facilitate and ungulates limit savanna tree regeneration. Oecologia 172, 1085–1093 (2013). https://doi.org/10.1007/s00442-012-2561-z

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