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Aim Patterns that relate species richness with fragment area (the species-area relationship, SAR) and with isolation (the species-isolation relationship, SIR) are well documented. However, those that relate species density - the number of species within a standardized area - with fragment area (D-SAR) or isolation (D-SIR) have not been sufficiently explored, despite the potential for such an analysis to disentangle the underlying mechanisms of SARs and SIRs. Previous spatial theory predicts that a significant D-SAR or D-SIR is unlikely to emerge in taxa with high dispersal limitation, such as plants. Furthermore, a recent model predicts that the detection and the significance of D-SARs or D-SIRs may decrease with grain size. We combined a literature review with grain size-dependent sampling in a fragmented landscape to evaluate the prevalence and grain size-dependent nature of D-SARs and D-SIRs in plants. Location Worldwide (review) and a semi-arid agro-ecosystem in Israel (case study). Methods We combined an extensive literature review of 31 D-SAR studies of plants in fragmented landscapes with an empirical study in which we analysed grain size-dependent D-SARs and D-SIRs using a grain size-dependent hierarchical sampling of species density and species richness in a fragmented, semi-arid agro-ecosystem. Results We found that significantly increasing D-SARs are rare in plant studies. Furthermore, we found that the detection of a significant D-SAR is often possible only after the data have been stratified by species, habitat or landscape characteristics. The results from our case study indicated that the significance and the slopes of both D-SARs and D-SIRs increase as grain size decreases. Main conclusions These results call for a careful consideration of scale while analysing and interpreting the responses of species richness and species density to fragmentation. Our results suggest that grain size-dependent analyses of D-SARs and D-SIRs may help to disentangle the mechanisms that generate SARs and SIRs and may enable early detection of the effects of fragmentation on plant biodiversity.
1. Worldwide, the floristic composition of temperate forests bears the imprint of past land use for decades to centuries as forests regrow on agricultural land. Many species, however, display significant interregional variation in their ability to (re)colonize post-agricultural forests. This variation in colonization across regions and the underlying factors remain largely unexplored.
2. We compiled data on 90 species and 812 species x study combinations from 18 studies across Europe that determined species' distribution patterns in ancient (i.e. continuously forested since the first available land use maps) and post-agricultural forests. The recovery rate (RR) of species in each landscape was quantified as the log-response ratio of the percentage occurrence in post-agricultural over ancient forest and related to the species-specific life-history traits and local (soil characteristics and light availability) and regional factors (landscape properties as habitat availability, time available for colonization, and climate).
3. For the herb species, we demonstrate a strong (interactive) effect of species' life-history traits and forest habitat availability on the RR of post-agricultural forest. In graminoids, however, none of the investigated variables were significantly related to the RR.
4. The better colonizing species that mainly belonged to the short-lived herbs group showed the largest interregional variability. Their recovery significantly increased with the amount of forest habitat within the landscape, whereas, surprisingly, the time available for colonization, climate, soil characteristics and light availability had no effect.
5. Synthesis. By analysing 18 independent studies across Europe, we clearly showed for the first time on a continental scale that the recovery of short-lived forest herbs increased with the forest habitat availability in the landscape. Small perennial forest herbs, however, were generally unsuccessful in colonizing post-agricultural forest even in relatively densely forested landscapes. Hence, our results stress the need to avoid ancient forest clearance to preserve the typical woodland flora.