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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.
Understanding the regional dynamics of plant communities is crucial for predicting the response of plant diversity to habitat fragmentation. However, for fragmented landscapes the importance of regional processes, such as seed dispersal among isolated habitat patches, has been controversially debated. Due to the stochasticity and rarity of among-patch dispersal and colonization events, we still lack a quantitative understanding of the consequences of these processes at the landscape-scale. In this study, we used extensive field data from a fragmented, semi-arid landscape in Israel to parameterize a multi-species incidence-function model. This model simulates species occupancy pattern based on patch areas and habitat configuration and explicitly considers the locations and the shapes of habitat patches for the derivation of patch connectivity. We implemented an approximate Bayesian computation approach for parameter inference and uncertainty assessment. We tested which of the three types of regional dynamics - the metacommunity, the mainland-island, or the island communities type - best represents the community dynamics in the study area and applied the simulation model to estimate the extinction debt in the investigated landscape. We found that the regional dynamics in the patch-matrix study landscape is best represented as a system of highly isolated island' communities with low rates of propagule exchange among habitat patches and consequently low colonization rates in local communities. Accordingly, the extinction rates in the local communities are the main drivers of community dynamics. Our findings indicate that the landscape carries a significant extinction debt and in model projections 33-60% of all species went extinct within 1000 yr. Our study demonstrates that the combination of dynamic simulation models with field data provides a promising approach for understanding regional community dynamics and for projecting community responses to habitat fragmentation. The approach bears the potential for efficient tests of conservation activities aimed at mitigating future losses of biodiversity.
Quantifying the association of plant functional traits to environmental gradients is a promising approach for understanding and projecting community responses to land use and climatic changes. Although habitat fragmentation and climate are expected to affect plant communities interactively, there is a lack of empirical studies addressing trait associations to fragmentation in different climatic regimes.
In this study, we analyse data on the key functional traits: specific leaf area (SLA), plant height, seed mass and seed number. First, we assess the evidence for the community assembly mechanisms habitat filtering and competition at different spatial scales, using several null-models and a comprehensive set of community-level trait convergence and divergence indices. Second, we analyse the association of community-mean traits with patch area and connectivity along a south-north productivity gradient.
We found clear evidence for trait convergence due to habitat filtering. In contrast, the evidence for trait divergence due to competition fundamentally depended on the null-model used. When the null-model controlled for habitat filtering, there was only evidence for trait divergence at the smallest sampling scale (0.25 m x 0.25 m). All traits varied significantly along the S-N productivity gradient. While plant height and SLA were consistently associated with fragmentation, the association of seed mass and seed number with fragmentation changed along the S-N gradient.
Our findings indicate trait convergence due to drought stress in the arid sites and due to higher productivity in the mesic sites. The association of plant traits to fragmentation is likely driven by increased colonization ability in small and/or isolated patches (plant height, seed number) or increased persistence ability in isolated patches (seed mass).
Our study provides the first empirical test of trait associations with fragmentation along a productivity gradient. We conclude that it is crucial to study the interactive effects of different ecological drivers on plant functional traits.
Intraspecific trait variability plays an important role in species adaptation to climate change. However, it still remains unclear how plants in semi-arid environments respond to increasing aridity. We investigated the intraspecific trait variability of two common Mediterranean annuals (Geropogon hybridus and Crupina crupinastrum) with similar habitat preferences. They were studied along a steep precipitation gradient in Israel similar to the maximum predicted precipitation changes in the eastern Mediterranean basin (i.e. -30% until 2100). We expected a shift from competitive ability to stress tolerance with decreasing precipitation and tested this expectation by measuring key functional traits (canopy and seed release height, specific leaf area, N-and P-leaf content, seed mass). Further, we evaluated generative bet-hedging strategies by different seed traits. Both species showed different responses along the precipitation gradient. C. crupinastrum exhibited only decreased plant height toward saridity, while G. hybridus showed strong trends of generative adaptation to aridity. Different seed trait indices suggest increased bet-hedging of G. hybridus in arid environments. However, no clear trends along the precipitation gradient were observed in leaf traits (specific leaf area and leaf N-/P-content) in both species. Moreover, variance decomposition revealed that most of the observed trait variation (>> 50%) is found within populations. The findings of our study suggest that responses to increased aridity are highly species-specific and local environmental factors may have a stronger effect on intraspecific trait variation than shifts in annual precipitation. We therefore argue that trait-based analyses should focus on precipitation gradients that are comparable to predicted precipitation changes and compare precipitation effects to effects of local environmental factors. (C) 2017 Gesellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.
Spatial environmental heterogeneity is considered a fundamental factor for the maintenance of plant species richness. However, it still remains unclear whether heterogeneity may also facilitate coexistence at fine grain sizes or whether other processes, like mass effects and source sink dynamics due to dispersal, control species composition and diversity at these scales. In this study, we used two complimentary analyses to identify the role of heterogeneity within 15 m x 15 m plots for the coexistence of species-rich annual communities in a semi-arid environment along a steep precipitation gradient. Specifically, we: (a) analyzed the effect of environmental heterogeneity on species, functional and phylogenetic diversity within microsites (alpha diversity, 0.06 m(2) and 1 m(2)), across microsites (beta diversity), and diversity at the entire plot (gamma diversity); (b) further we used two null models to detect non-random trait and phylogenetic patterns in order to infer assembly processes, i.e. whether co-occurring species tend to share similar traits (trait convergence) or dissimilar traits (trait divergence). In general, our results showed that heterogeneity had a positive effect on community diversity. Specifically, for alpha diversity, the effect was significant for functional diversity, and not significant for either species or phylogenetic diversities. For beta diversity, all three measures of community diversity (species, functional, and phylogenetic) increased significantly, as they also did for gamma diversity, where functional measures were again stronger than for species or phylogenetic measures. In addition, the null model approach consistently detected trait convergence, indicating that species with similar traits tended to co-occur and had high abundances in a given microsite. While null model analysis across the phylogeny partly supported these trait findings, showing phylogenetic underdispersion at the 1m(2) grain size, surprisingly when species abundances in microsites were analyzed they were more evenly distributed across the phylogenetic tress than expected (phylogenetic overdispersion). In conclusion, our results provide compelling support that environmental heterogeneity at a relatively fine scale is an important factor for species co-existence as it positively affects diversity as well as influences species assembly. Our study underlines the need for trait-based approaches conducted at fine grain sizes in order to better understand species coexistence and community assembly. (C) 2017 Elsevier GmbH. All rights reserved.