Effects of bush encroachment on an assemblage of diurnal lizard species in central Namibia
Introduction
Although humans have influenced the natural environment in Namibia for thousands of years, changes in land use practices over the last century have severely altered natural ecological processes (Louw & Seely, 1982, Prins & van der Jaeugd, 1993, Hoffman, 1997, Pallett, 1997). Recent human-mediated impacts include overgrazing, fire suppression, fragmentation of historical migration routes, establishment of permanent water sources and extirpation of the largest herbivores and top carnivores (for a review, see Barnard, 1998). A consequence of these ecological changes is the conversion of savannas to dense, acacia-dominated thickets with little grass cover, a process known as “bush encroachment.” This problem is especially prevalent on commercial ranchlands in central Namibia, to the extent that bush encroachment has measurably reduced the economic productivity of the Namibian livestock industry (Quan et al., 1994).
Livestock grazing has been the predominant land use practice in Namibia for several centuries; consequently, much of the native wildlife in Namibia (and throughout much of sub-Saharan Africa) must coexist with livestock production (Kinahan, 1991, Mabutt, 1984, Archer, 1995). Nevertheless, commercial ranchlands in Namibia still harbor a diverse assemblage of native species and have substantial potential as refuges for Namibia's biota. For example, the majority of Namibia's populations of native large ungulates (over 70%) and the largest remaining population of cheetahs in the world are found on commercial ranchland, outside of the formal protected area network (Joubert & Morsbach, 1982, Marker-Kraus et al., 1996). Considerable conservation efforts and government wildlife policies are focused on these large mammalian species (reviewed by Griffin, 1998, Richardson, 1998). While these efforts complement biodiversity conservation objectives in most cases (e.g. the cheetah, see Marker-Kraus et al., 1996), some less economically measurable and/or popular components of biodiversity may not be compatible with a system designed to promote livestock production and large, huntable game (Richardson, 1998). Most of the native species that do not have direct economic value in Namibia are found outside of the formal protected area network (Barnard et al., 1998). A better understanding of interactions between native species and human impacts is an important step towards native species conservation on commercial ranchland.
Here we present results of systematic surveys of an assemblage of lizard species in open savanna and bush-encroached habitats in central Namibia. Our purpose was to examine lizard species composition, relative abundance, and microhabitat use in open savanna and bush-encroached habitats. In a review of resource partitioning among species of amphibians and reptiles, Toft (1985) found habitat to be more important than temporal or dietary dimensions in interspecific separation. Patterns in the spatial use across habitat gradients by lizard assemblages have been well documented, yet few studies have examined the impacts of ecological processes on lizard assemblage stability at the landscape scale (Case, 1994, Vitt, 1994; but see Pianka, 1996). Although lizards have little direct economic value, they are important components of Namibian savannas and may be impacted by bush encroachment. We compared lizard assemblages associated with open savanna and bush-encroached habitat and interpreted our results with respect to accumulating evidence that bush encroachment and its associated ecological impacts are reorganizing savanna ecosystems throughout southern Africa.
Section snippets
Study area
The study area is located on ranchland owned by Cheetah Conservation Fund (CCF) in central Namibia, approximately 30 km east of Otjiwarongo (16° 39′ 0″ E, 20° 28′ 12″ S). Dominant topographic relief within the study area is in the form of small (∼1–8 ha), isolated granitic outcrops (kopjes). These outcrops rise above a surrounding matrix of flatland, which has virtually no exposed rock. The Waterberg Plateau, a 4100-km2 sandstone uplift, lies on the southern periphery of the study area and is
Habitat comparison of savanna and bush-encroached plots
Compared to open savanna plots, bush-encroached plots were visually different (Fig. 1; Table 1). Bush-encroached plots were characterized by having a different assemblage of woody plant species (Table 2), greater density of woody plants (x=309/ha vs. 217/ha), greater degree of canopy cover, greater amount of downed wood, and lesser amount of perennial grass cover. Bush-encroached plots also differed strikingly from open savanna plots in that there was increased homogeneity in the spatial extent
Discussion
We observed significant differences in the lizard assemblages between savanna and bush-encroached habitat types. Both Pedioplanis undata and Lygodactylus bradfieldi were, respectively, 3.9 and 2.7 times more abundant in savanna than bush-encroached plots. The assemblage differences, however, were primarily attributable to the absence of M. varia from all plots that were formerly savanna and a dramatic increase in the abundance of its congener, M. striata, which was 7.5 times more abundant in
Acknowledgements
We are indebted to L. Marker, B. Brewer, K. O'doherty and the staff at Cheetah Conservation Fund for logistical support. We thank the Namibian Ministry of Environment and Tourism for permission to conduct this research (General Permit No. 35048). We thank E. Horan and E. Ramsey for providing assistance in the field. J.M.M. would like to particularly thank E.D. Brodie, Jr., for making travel to Africa possible. We thank C. Garrard and S. Durham for valuable assistance with statistical analyses.
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