The Ecology of Black Cotton Glades Kari E. Veblen University of California-Davis, USA (keveblen@ucdavis.edu) Making glades Research overview Introduction My research explores the ecology of abandoned boma sites in the black cotton ecosystem. In Laikipia, old boma sites, referred to as “glades,” are recognized as large treeless patches The objective of this experiment is to test methods of creating glade-like structures—areas that are (approx. 0.5-1.0 ha) of nutrient-rich grass that dot the savanna landscape (Figures 1 and 2). nutrient rich and attractive to wildlife and livestock. I am testing the effects of three factors on glade Abandoned boma sites such as these occur throughout much of eastern and southern Africa development: in various forms (Stelfox 1986, Blackmore et al. 1990, Reid and Ellis 1995). In Turkana, • fertilization with dung or commercial fertilizer • clearing of trees to make open areas • addition of Acacia brush, which may create safe sites for the establishment and Kenya, old boma sites are the mirror image of Laikipia’s old bomas: dense Acacia patches in an otherwise treeless landscape (Reid and Ellis 1995). Glades can remain on the landscape for many decades—at least 45 years in Laikipia. Understanding how glades develop over growth of certain plant species preferred by herbivores (Figure 6). time and how plants and animals fit into their development are integral to a scientific understanding of the black cotton ecosystem as a whole. Additionally, better understanding Fertilization may increase the nutritive value of the plants in the area or encourage the glades, which attract both wild and domestic animals, may be useful to livestock and wildlife establishment of lawns of Cynodon, thereby attracting wildlife or cattle. If cleared or brush-addition managers. Here I present two components of my research: 1) description of the plants, areas attract cattle or wildlife, increased defecation and/or grazing may stimulate grass production animals, and soil associated with black cotton glades as they develop over time and 2) to produce more attractive forage. Each of these three factors or some combination of them could experimental boma-building that explores how dung addition, tree-clearing and Acacia perpetuate an escalating cycle of increased nutrients and increased herbivore use, leading to brush-addition— three major components of bomas—influence glade development. glade-like structures. Methods a) I am using a combination of 50x50 m. plots (shared with C. Riginos’ tree density experiment) and Figure 1. Quickbird satellite image (June 2003) displayed in false color infrared, with red indicating higher plant productivity. Image covers portions of Mpala, Jessel and Segera black cotton soil. Glades appear as small red patches across the landscape. Recently abandoned boma 10x10 m. plots to test the influence of dung and fertilizer addition, tree clearing and Acacia brush addition on glade development in the Mpala and Jessel black cotton soil (Figure 7). In 50x50 m. plots (with a 10 m. buffer on each side), I will compare five replicates each of the following three treatments: 1) trees cleared, 2) trees cleared with 10x10 m. patch of dung at center, and 3) no tree Description of black cotton glades Cynodon plectostachyus Pennisetum stramineum addition in inner 10x10 m. area, 2) tree clearing in full 20x20 m. area, 3) Acacia drepanolobium 100 Although the vegetation, soil and animals associated with glades in red soil have been brush addition around inner 10x10 m. area, 4) brush addition+dung addition, 5) clearing +dung 80 described (Young et al. 1995, Augustine 2003), black cotton glades have not yet been addition, 6) brush addition +clearing+dung addition, 7) fertilizer addition in inner 10x10m. + 60 formally described. Because the two ecosystems differ so dramatically, we would expect clearing, and 8) control (no manipulations). In addition to making comparisons among plots of the 40 black cotton glades to be characterized by very different communities of plants and same size, I will be able to compare similar treatments of different sizes. I am regularly sampling 20 animals, and by unique soil characteristics. I surveyed plants, animals and soil on new vegetation, dung (animal use), termite presence, and soils to test which factors are most and old glades to better understand how glades develop over time in the black cotton soil. responsible for the functional traits of glades. Methods Significance/application I surveyed five young glades (<45 years old, Figure 2b) and five old glades (>45 years Understanding how old bomas develop into glades is of significance for understanding dynamics of old, Figure 2c) on Mpala and Jessel properties. In these glades, I measured plant cover the ecosystem and is also potentially of interest to livestock and wildlife managers. The results of pin hits Introduction 0 New glades b) Glade ~7 years old clearing. In 20x20 m. plots, I will compare five replicates each of the following treatments: 1) dung Old glades Figure 4. Mean (+1 S.E.) of Cynodon and Pennisetum cover in 5 new (<45 yrs) and 5 old (>45 yrs) glades. inside and outside of the glades, using a series of pin-frame measurements (Figure 3). I this study will complement my descriptive study by lending insight into the mechanisms behind also assessed animal use of these glades by identifying and counting different piles of glade development. Furthermore, my results may aid interested land managers in creating glade- animal dung found inside two 20x20 meter blocks inside each glade and along two 100m- like structures to provide nutrient-rich grazing areas for cattle or wildlife. long (and 4m-wide) transects located at 100m and at 200m away from each glade. Finally, I collected soil (at 0-30cm depth) inside and outside each glade and had the soils analyzed for nitrogen, phosphorus, potassium, calcium, magnesium, and sand/silt/clay content. I tested the data for differences in vegetation, dung, and soil inside and outside 120 Clearing only of new versus old glades. INSIDE OUTSIDE Brush only NEW GLADES dung density (x1000) 100 80 60 OLD 40 20 Glade > 45 years old • New glades in the black cotton soil were dominated by the mat-forming Cynodon plectostachyus (Star grass), followed by dominance by the taller bunch grass, Dung only Pennisetum stramineum, in old glades (Figure 4). Dung + clear • Cynodon plectostachyus almost never occurred outside of glades. Dung + clear 100 OLD GLADES dung density (x1000) c) 0 120 Clearing only Results and conclusions 80 60 40 20 • Bracharia, Lintonia, Themeda, and Mezianum grass species, which dominate outside of glades, were almost never found inside glades. • Total dung density of all herbivores was higher inside new glades than outside (Figure 5). Zebra dung density was highest outside of glades, while Grant’s gazelle Brush + dung Figure 3. Ten-point pin-frame used to assess plant cover. Observers count how many times each plant species is hit by a vertical pin (maximum 10 hits for each species). dung was highest inside glades. Elephants appear to strongly prefer new glades, 0 None Brush, clear, dung All herbivores Cattle+ buffalo Zebra Grant's gazelle Elephant Figure 5. Herbivore dung count data inside and outside of new (<45 yrs) and old (>45 yrs) glades. Figure 2. Successional sequence of abandoned black cotton boma vegetation: (a) a newly abandoned boma, (b) a new glade (~7 years) covered with Star grass, Cynodon plectostachyus, and (c) an old glade (>45 years) covered with Pennisetum stramineum. References: Augustine, D. J. 2003. Long-term, livestock-mediated redistribution of nitrogen and phosphorus in an East African savanna. Journal of Applied Ecology 40:137-149. Blackmore, A. C., M. T. Mentis, and R. J. Scholes. 1990. The origin and extent of nutrient-enriched patches within a nutrientpoor savanna in South Africa. Journal of Biogeography 17:463-470. Reid, R. S., and J. E. Ellis. 1995. Impacts of pastoralists on woodlands in South Turkana, Kenya: livestock-mediated tree recruitment. Ecological Applications 5:978-992. Stelfox, J. B. 1986. Effects of livestock enclosures (bomas) on the vegetation of the Athi Plains, Kenya. African Journal of Ecology 24:41-45. Young, T. P., B. D. Okello, D. Kinyua, and T. M. Palmer. 1998. KLEE: a long-term multi-species herbivore exclusion experiment in Laikipia, Kenya. African Journal of Range and Forage Science 14:94-102. Young, T. P., N. Patridge, and A. Macrae. 1995. Long-term glades in acacia bushland and their edge effects in Laikipia, Kenya. Ecological Applications 5:97-108. and cattle show a slight preference for new glades. None • Relative to non-glade soils, glade soils were high in phosphorus and potassium, but low in magnesium, calcium, sodium and clay content. Nitrogen was high in all Fert + clear glades, but highest in new glades. Because cattle and wildlife are important players in the black cotton ecosystem and also appear to use glades especially heavily, herbivore activity may influence the rate at which =dung =N fertilizer =Acacia brush =trees/uncleared glades make the transition from Cynodon- to P.stramineum-dominance. For example: • A plant that has been damaged by herbivory may be at a competitive disadvantage for resources such as water or light. In such a case, heavier grazing of Cynodon would lead to a faster takeover by P. stramineum. • Herbivores may export nutrients out of glades when they graze on high-nutrient Figure 6. Experimental Acacia drepanolobium brush addition to simulate boma fence. glade grasses. If Cynodon competes better in high-nutrient environments, then Figure 7. Schematic diagram of treatments for glade-making experiment testing the effects of dung and fertilizer addition, brush addition, and tree clearing on glade development. Large plots are 50x50m., and small plots are 20x20m. Small focal plots included within larger plots are 10x10m. herbivores exporting nutrients may speed the transition from Cynodon to P. stramineum. I am using the KLEE experiment (Young et al. 1998) and conducting additional experiments on Mpala, Jessel and Segera properties to try to understand how the effects of herbivores influence the nature of competition and interactions between the two glade grass species. Acknowledgements: Many thanks are owed to the following for their help in the field: Frederick Erii, John Lochukuya, Jackson Ekadeli, Patrick Etelej, Corinna Riginos, James Ekiru, Jake Goheen, Ali Hassan, Dan Kelly, John Lemboi and Simon Lima. Thank you to Truman Young for advice, support and field assistance. Thank you to Mpala Research Centre, Mpala Ranch and Peter Jessel for permission to use field sites and for logistical support. Funded by NSF and U. California-Davis Jastro Shields grant.