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Diversity of the Herbaceous Layer in Mixed Bushveld

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Diversity of the Herbaceous Layer in Mixed Bushveld J. Range Manage. 52:519–524 September 1999 Diversity of the herbaceous layer in mixed bushveld WERNER G. DÖRGELOH Author is a wildlife biologist, Applied Natural Sciences, Technikon SA, Private Bag X6, Florida, 1710, South Africa. e-mail: [email protected] Abstract Resumen The diversity of the herbaceous layer in the Nylsvley Nature Se investigó la diversidad del estrato herbáceo en la Reserve (Mixed Bushveld) was investigated to enhance the Reserva Natural Nylsvley (Mixed Bushveld) para aumentar understanding of savanna ecosystems and to serve as a base- el entendimiento de los ecosistemas de savana , que sirva de line for future monitoring to facilitate management. Species referencia para futuras comparaciones y facilitar el manejo. composition and density, dry weight contribution per species, La composición y densidad de las especies, la contribución and grass density was measured with a small-quadrat del peso seco por especie y la densidad de pasto fueron medi- method. A total of 73 grass species were recorded. Plant com- das con el método del cuadrante pequeño. Se registraron un munities were compared in terms of percentage composition total de 73 especies de pastos. Las comunidades de plantas se and percentage dry weight per ecological group, species compararon en términos de porcentaje de composición y peso diversity and grass density with general linear modeling. The seco por grupo ecológico, diversidad de especies y densidad herbaceous layer of most plant communities consisted pre- de pastos la comparacion se hizó mediante modelos lineales dominantly of increaser I species (increasing in under-utilized generales. El estrato herbáceo de la mayoría de las comu- areas), with increaser I and decreaser species (decreasing nidades de plantas consistió principalmente de especies incre- with under or over-utilization) producing the highest bio- soras I (que aumentan en áreas subutilizadas), las especies mass. The dominance of increaser III species (becoming dom- incresoras I y decresoras (que disminuyen con sub o sobre- inant in heavily over-grazed areas) in the Sporobolus ioclados utilización) produjeron la mayor cantidad de biomasa. La -Acacia tortilis Savanna indicates previous over-utilization of dominancia de especies incresoras III (dominantes en áreas this plant community. A generally high species diversity fuertemente sobreutlizadas) en la savana de Isporobolus (Simpson’s index 1-D >0.72) in the reserve is influenced by iocladosy-Acacia tortilis indican una sobreutilización previa environmental factors and is a reflection of previous low ani- de esta comunidad. La alta diversidad de especies (Indice de mal stocking densities and a rotational burning regime. Grass Simpson 1-D>0.72) de la reserva esta influenciada por fac- density varied from 53.07 to 219.13 grasses/m-2. A negative tores ambientales y también es un reflejo de la baja densidad de carga animal anterior y un régimen rotacional de quemas. correlation (r = –0.6654) between grass density and species -2 density supports the principle that species diversity is La densidad de pasto vario de 53.07 a 219.13 pastos m . Una reduced in over-grazed areas. The high diversity of the correlación negativa (r = –0.6654) entre los pastos y la densi- herbaceous layer in the Nylsvley Nature Reserve may serve as dad de especies refuerza el principio de que la diversidad de a benchmark for comparing range diversity over time and especies es reducida en áreas sobreutilizadas. La alta diversi- space within this vegetation type. dad del estrato herbáceo en la reserva natural de Nylsvley puede servir como una referencia para comparar el rango de diversidad a través del tiempo y el espacio dentro de este tipo Key Words: species composition, species density, species de vegetación. diversity, percentage dry weight, grass density, Simpson’s index, bootstrapping 1991). The high species diversity in the Nylsvley Nature The herbaceous layer composition is changing continuously Reserve is attributable to the diversity of habitats, the climatic, in space and time due to a combination of factors, such as geomorphological and biogeographical history of the region, grazing, fire, and rainfall which differ in intensity, duration, and the long period of uninterrupted, evolutionary develop- and timing. O’Connor (1991) found that rainfall variability ment (Scholes and Walker 1993). over 1 or 2 years could induce substantial changes in compo- When only a sample of species in the community is sition. Composition is further affected by grazing. Tuft sur- obtained, it is necessary to distinguish between numerical vival of grass species decreased by approximately 48 % under species richness, that is number of species per specified num- rainfall interception and grazing experiments (O’Connor ber of individuals or biomass, and species density, that is number of species per specified collection area (Magurran 1991). Species evenness or equitability refers to how equally Research was funded by the Foundation for Research Development and partly by Technikon SA. Environmental Affairs Northern Province gave permission to abundances (number of individuals, biomass and cover) are conduct the study in the Nylsvley Nature Reserves. distributed among species (Goldsmith et al. 1986, Ludwig and Manuscript accepted 29 Nov. 1998. Journal of Range Management 52(5), September 1999 519 Reynolds 1988, Morrison et al. 1992). diversity and grass density between lowed where, at each site, 20 quadrats Diversity indices combine both species plant communities. (0.25 m2) were placed 5 m apart on a richness and evenness into a single single line transect (Dörgeloh 1997), value (Ludwig and Reynolds 1988, assuming independence between sam- Magurran 1991) and are therefore more Materials and Methods pling points. suitable for analysis of community structure. These are also called hetero- The study was conducted in the Grass density geneity indices (Ludwig and Reynolds Nylsvley Nature Reserve (24°39'S Each rooted herbaceous plant within a 1988). The advantage of diversity 28°42'E), South Africa, situated in 0.25 m2 square was counted. Density indices is that no assumptions about dis- Mixed Bushveld on the border between estimates were based on a convenient tribution of individuals among species a moist and dry savanna (Acocks 1988). morphological unit. In this case each have to be made (Morrison et al. 1992), The rainy season in summer (maximum rooted aerial shoot was treated as a unit. and they are therefore referred to as mean daily temperature of 29.3°C) is It is almost impossible to determine the non-parametric indices (Magurran followed by dry, cold winters (minimum density of grass species which spread 1991). Although the greatest problem mean daily temperature of 6.1°C). In the vegetatively (Goldsmith et al. 1986). with diversity indices is interpretation of 1993/1994 and 1994/1995 rainy sea- Therefore rooted nodes of stolons of rhi- their values, they are widely applied in sons, a rainfall of 480.5 mm and 512.4 community analyses (Ludwig and zomatic grasses and parts of tufts, that mm, respectively was lower than the had split and were at least 2 centimetres Reynolds 1988, Magurran 1991, long-term rainfall of 623 mm (standard Morrison et al. 1992). These indices from the nearest other plant of the same deviation of 134 mm) (Scholes and merely reflect community structure by species, were counted as individuals. Walker 1993). Vegetation surveys were changing with species richness, equi- This was applied to all monocarpic conducted in 8 plant communities, iden- tability and sometimes density (annuals and biennials) and polycarpic tified from a vegetation map constructed (Morrison et al. 1992). Simpson’s index (perennials) species which usually have (D) is commonly used (Magurran 1991) by Coetzee et al. (1976), Frost (1987), extensive below ground vegetative and is based on probability theory and Scholes and Walker (1993): growth systems (Causton 1988). (Goldsmith et al. 1986). It measures the 1. Rhus leptodictya-Combretum apic- probability that 2 individuals selected at ulatum Variation (lithosols underlain by Species composition and density random from a sample belong to differ- rock) Each rooted grass species within a ent species. Simpson’s index is most 2. Cymbopogon plurinodis-Com- 0.25 m2 square was identified according sensitive to changes in common species bretum apiculatum Variation (lithosols to Gibbs Russell et al. (1991) and Van (Magurran 1991, Morrison et al. 1992). underlain by rock) Oudtshoorn (1992). Forbs as a group Abundance or density of grasses is 3. Eragrostis nindensis-Digitaria (≈15 species) was treated as a single defined as number of individuals of a monodactyla Variation (lithosols under- species. Species density is the most particular species per unit area lain by rock) commonly used measure of species rich- (Goldsmith et al. 1986, Causton 1988, 4. Sporobolus ioclados-Acacia tortilis ness (number of species) (Magurran Goldsmith 1991) which is related to bio- Savanna (fine-textured illuvial soils with 1991) and was calculated from the num- mass. For the same height, grass quanti- high clay contents) ber of species per 15 m2 for each plant ty varies according to the density 5. Nyl River and floodplain (alluvial community. (Voisin 1988). soils) 6. Aristida bipartita-Setaria sphacela- In the late 1970s and 1980s alpha Biomass contribution per species diversity (number of species in a defined ta Savanna variation (vertisols and mol- The relative biomass contribution of area) was measured only in the lisols) grass species was estimated with the Eragrostis pallens–Burkea africana 7. Aristida bipartita-Setaria sphacela- dry-weight rank method, as developed savanna as part of the Savanna Biome ta Grassland variation (vertisols and by 't Mannetje and Haydock (1963) and Programme. An alpha diversity of 319 mollisols) plant species was high by global stan- 8. Eragrostis pallens-Burkea africana described by Kelly and McNeill (1980), dards of 80 to 100 species 0.1 ha-1. Savanna (well-drained, residual sandy Barnes et al. (1982), Ben-Shahar (1991) Disturbed areas with fertile soils within soils) and Shackleton (1992).
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