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Aim To investigate the species-area relationship (SAR) of plants on very small islands, to examine the effect of other factors on species richness, and to check for a possible Small Island Effect (SIE).Location The study used data on the floral composition of 86 very small islands (all < 0.050 km 2 ) of the Aegean archipelago (Greece).Methods We used standard techniques for linear and nonlinear regression in order to check several models of the SAR, and stepwise multiple regression to check for the effects of factors other than area on species richness ('habitat diversity', elevation, and distance from nearest large island), as well as the performance of the Choros model. We also checked for the SAR of certain taxonomic and ecological plant groups that are of special importance in eastern Mediterranean islands, such as halophytes, therophytes, Leguminosae and Gramineae. We used one-way anova to check for differences in richness between grazed and non-grazed islands, and we explored possible effects of nesting seabirds on the islands' flora. ResultsArea explained a small percentage of total species richness variance in all cases. The linearized power model of the SAR provided the best fit for the total species list and several subgroups of species, while the semi-log model provided better fits for grazed islands, grasses and therophytes. None of the nonlinear models explained more variance. The slope of the SAR was very high, mainly due to the contribution of non-grazed islands. No significant SIE could be detected. The Choros model explained more variance than all SARs, although a large amount of variance of species richness still remained unexplained. Elevation was found to be the only important factor, other than area, to influence species richness. Habitat diversity did not seem important, although there were serious methodological problems in properly defining it, especially for plants. Grazing was an important factor influencing the flora of small islands. Grazed islands were richer than non-grazed, but the response of their species richness to area was particularly low, indicating decreased floral heterogeneity among islands. We did not detect any important effects of the presence of nesting seabird colonies.Main conclusions Species richness on small islands may behave idiosyncratically, but this does not always lead to a typical SIE. Plants of Aegean islets conform to the classical Arrhenius model of the SAR, a result mainly due to the contribution of non-grazed islands. At the same time, the factors examined explain a small portion of total variance in species richness, indicating the possible contribution of other, non-standard factors, or even of stochastic effects. The proper definition of habitat diversity as pertaining to the taxon examined in each case is a recurrent problem in such studies. Nevertheless, the combined effect of area and a proxy for environmental heterogeneity is once again superior to area alone in explaining species richness.
Aim To estimate species turnover of plants on 32 small islands within a 20-year period and to assess possible changes in community composition and properties, such as species richness and factors affecting it, nestedness, species co-occurrence and overall community similarity. Additionally, to assess the possible effects of grazing, gull colonies and fire on turnover values.Location Thirty-two islets in the eastern Aegean Sea (Greece).Methods Complete sampling of plants was performed in 1974 and in 1990-94 (mostly in 1994, which was used as the reference year). Species turnover rates were estimated using both per island and per species approaches. Multiple regression was used to evaluate factors affecting species richness. Chi-square tests were applied to compare community composition among sampling periods. The effects of various factors on turnover rates and species richness were examined using one-way anova and ancova. Mann-Whitney tests were applied in order to check for differences between frequencies of occurrence of extinct, immigrant and persisting species. Community nestedness was calculated using bitmatnest and the C-score index for co-occurrence was estimated using EcoSim7. Species similarities among islands in each of the 1974 and 1994 data sets were assessed using Jaccard's index and the two similarity matrices were compared using a Mantel test. ResultsOf 391 species recorded on the islets, 334 were present in 1974, 301 in 1994 and 244 were common to both these periods. Species richness in the 1974 and 1994 data sets was significantly correlated with elevation and area, but not with distance from the nearest large island. Richness was positively affected by grazing, but not by fire or gull colonies. The slopes of species-area and species-elevation regressions were almost identical in 1974 and 1994. Mean relative turnover was 2.06 (species per islet) and 3.26 (islets per species). Turnover was not correlated with area, elevation or distance from the nearest large island. Nestedness and co-occurrence levels were very similar. Tables of islet by islet floral similarity (Jaccard's index) did not differ between the 1974 and 1994 data sets.Main conclusions The turnover rates found are among the highest recorded for plants; at the same time the islet communities exhibit notable stability in overall properties. Our results provide evidence for rapid shifts in species number that may nonetheless be considered as equilibrial dynamics, as these islets are able to respond rapidly to environmental change and disturbance. Human activities, notably the application of grazing, have a significant complicating effect on community dynamics, enhancing observed turnover rates.
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