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Colwell, R.K., C.X. Mao and J. Chang. 2004. Interpolating, extrapolating, and comparing incidence-based species accumulation curves. Ecology 85: 2717–2727. Chang J. Interpolating

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In the present study, we examined the effect of shrub encroachment in temperate semi-natural grasslands on plant species diversity. We tested the hypothesis that an initial shrub encroachment leads to enhanced habitat heterogeneity and thereby to a higher diversity. In a calcareous grassland near Göttingen (Germany) we analysed the effect of shrub encroachment on the species richness in 30, 100 m2 plots each with 10 subplots of 1m2 size. The 30 main plots belonged to six different shrub encroachment classes. A descriptive correlative gradient analysis of shrub invaded grasslands and their species number of flowering plants was performed. Within the 30 plots of different shrub encroachment a total of 203 plant species were recorded. The mean α1-diversity (level of subplots), mean α2-diversity (level of plots) as well as the α3-diversity (level of shrub encroachment classes) have their highest values at medium shrub invaded sites. This finding is in line with our hypothesis of a hump-back relation between shrub encroachment and species richness, and can be explained by the increased habitat heterogeneity. However, Detrended Correspondence Analysis (DCA) emphasized the importance of the present vegetation composition for species richness. The dominance of highly competitive, clonal-growing grass species is accompanied by low diversity swards with a lower facilitation of shrub establishment. Species accumulation curves highlight the benefit of shrub encroachment for γ-(landscape-) diversity. This result emphasises the importance of habitat heterogeneity for biodiversity and, therefore, nature conservation.

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Species richness in an assemblage is frequently used as a measure of biological diversity. However, observed species richness is strongly dependent on sample size. If more samples are collected, then more species are observed. Non-parametric species richness estimators, such as the jackknife 1 and 2 and the Chao 1 and 2, are indicated in the literature as potential solutions to the problem of dependence of observed species richness on sampling effort. These methods are intended to estimate the total species richness in an area or assemblage with small sampling effort. Non-parametric estimators are based on the number of species observed, and the number of rare species in a sample, i.e., that occurred in one and/or two sampling units, or with one and/or two individuals. High estimates are produced when samples contain large proportions of rare species. Using a range of real datasets, I show that estimates produced by non-parametric methods are generally dependent on observed species richness. An implicit assumption of these non-parametric techniques is that the rare species curve should present high values at small sample sizes and decreasing values as sampling effort is increased. This assumption was observed in only one out of eight datasets presented. Instead, the rare species curve generally flattens off around a constant value as sampling effort increases. I conclude that non-parametric estimators are not reliable to estimate species richness in an assemblage when the rare species curve does not show a decreasing trend. Comments are made on the possibilities of using non-parametric estimators in the comparison of species assemblages.

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.K., C.X. Mao and J. Chang. 2004. Interpolating, extrapolating, and comparing incidence-based species accumulation curves. Ecology 85: 2717–2727. Chang J. Interpolating

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.E. Ellingsen. 2003. The species-accumulation curve and estimation of species richness. J. Animal Ecol. 72:888–897 Ellingsen K.E. The species-accumulation curve and estimation of species

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Ecol. Lett. 2001 4 379 391 Hui, C. 2008. On species-area and species accumulation curves: A

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Moreno, C.E. and G. Halffter. 2001. On the measure of sampling effort used in species accumulation curves. J. App. Ecol. 38: 487–490. Halffter G. On the measure of sampling effort

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.O. 1973 . Diversity and evenness: a unifying notation and its consequences . Ecology 54 : 427 – 432 . Hui , C. 2008 . On species-area and species accumulation

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