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Positive plant–plant interactions are thought to drive vegetation patterns in harsh environments, such as semi-arid areas. According to the stress-gradient hypothesis (SGH), the role of positive interactions between species (facilitation) is expected to increase with harshness, predicting associated variation in species composition along environmental gradients. However, the relation between stress and facilitation along environmental gradients is debated. Furthermore, differentiating facilitative interactions from other underlying mechanisms, such as microtopographic heterogeneity, is not trivial. We analysed the spatial co-occurrence relationships of vascular plant species that form patchy vegetation in arid lapilli fields (tephra) from recent volcanic eruptions on La Palma, Canary Islands. Assuming a harshness gradient negatively correlated with elevation because the lower elevations are more arid and water availability is considered the most limiting resource, and that an outcome of facilitation is plants co-occurring in the same patch, from the SGH we expected a greater degree of co-occurrence at lower elevation. We tested this at both the species and the individual plant level. We analysed the species composition of 1277 shrubby vegetation patches at 64 different sampling points, ranging from the coast to around 700 m a.s.l. Patch morphology and microtopographic heterogeneity variables were also measured, to account for their potential effects on the species composition of patches. We used generalized linear models and generalized mixed-effects models to analyse species richness, number of individuals in patches and percentage of patches with positive co-occurrences, and a pairwise co-occurrence analysis combined with a graphical network analysis to reveal positive links between 13 of the species. We found that the percentage of patches with positive co-occurrences increased at higher elevations, in contrast to the predictions of the SGH, but in accordance with a refined stress-gradient hypothesis for arid sites, in which characteristics of the interacting species are incorporated.
Questions: Habitat islands are often characterized by the presence of more or less sharp boundaries to adjacent matrix habitats. However, knowledge on boundaries of natural habitat islands is scarce, especially regarding patterns of beta diversity and its two underlying components: species turnover and nestedness. We therefore aim to quantify the effects of fine-scaled and sharp boundaries of quartz islands (quartz gravel-covered soils) on the different components of plant beta diversity and how they are linked to different soil environmental drivers. Location: Knersvlakte, Western Cape, South Africa. Methods: We sampled plant species richness in 56 fine-scale transects of 6 m × 1 m plots across eight different boundary types (four quartz island to matrix, four between habitats on quartz islands). Soil depth and chemistry (pH, electrical conductivity) were analyzed for each 1 m2 plot. Differences in the two beta diversity components (turnover and nestedness) for each boundary type were tested by t tests. We used linear models to test relationships between species and environmental dissimilarity. Results: All boundary types showed high beta diversity. Species turnover was the prevailing component for six boundary types, the nestedness component was only important for two boundary types. We found a significant linear increase of species dissimilarity with increasing dissimilarity in soil pH and distinct plant communities for the habitat types, but no significant increase for electrical conductivity or soil depth. Conclusions: The spatial distinctiveness of the quartz islands leads to sharp boundaries, which result in high beta diversity, mainly through species turnover. This reflects the high levels of diversification and adaptation of the local plant communities. Nestedness occurred at two boundaries to the matrix, indicating that the latter does not necessarily represent an impermeable boundary for all species of the respective ecosystem. Studying diversity patterns across boundaries contributes to the question of applicability of island biogeography theory to habitat islands.
Questions: Both species turnover and intraspecific trait variation can affect plant assemblage dynamics along environmental gradients. Here, we asked how community assemblage patterns in relation to species turnover and intraspecific variation differ between endemic and non-endemic species. We hypothesized that endemic species show lower intraspecific variation than non-endemic species because they tend to have high rates of in situ speciation, whereas non-endemic species are expected to have a larger gene pool and higher phenotypic plasticity.
Location: La Palma, Canary Islands.
Methods: We established 44 sampling sites along a directional gradient of precipitation, heat load, soil nitrogen, phosphorus and pH. Along this gradient, we estimated species abundances and measured three traits (plant height, leaf area and leaf thickness) on perennial endemic and non-endemic plant species. In total, we recorded traits for 1,223 plant individuals of 43 species. Subsequently, we calculated community-weighted mean traits to measure the relative contribution of species turnover, intraspecific variation and their covariation along the analysed gradient.
Results: The contribution of intraspecific variation to total variation was similar in endemic and non-endemic assemblages. For plant height, intraspecific variation explained roughly as much variation as species turnover. For leaf area and leaf thickness, intraspecific variation explained almost no variation. Species turnover effects mainly drove trait responses along the environmental gradient, but intraspecific variation was important for responses in leaf area to precipitation.
Conclusions: Despite their distinct evolutionary history, endemic and non-endemic plant assemblages show similar patterns in species turnover and intraspecific variation. Our results indicate that species turnover is the main component of trait variation in the underlying study system. However, intraspecific variation can increase individual species’ fitness in response to precipitation. Overall, our study challenges the theory that intraspecific trait variation is more important for the establishment of non-endemic species compared with endemic species.
Climatic and biogeographical drivers of functional diversity in the flora of the Canary Islands
(2022)
Aim: Functional traits can help us to elucidate biogeographical and ecological processes driving assemblage structure. We analysed the functional diversity of plant species of different evolutionary origins across an island archipelago, along environmental gradients and across geological age, to assess functional aspects of island biogeographical theory.
Location: Canary Islands, Spain.
Major taxa studied: Spermatophytes.
Time period: Present day.
Methods: We collected data for four traits (plant height, leaf length, flower length and fruit length) associated with resource acquisition, competitive ability, reproduction and dispersal ability of 893 endemic, non-endemic native and alien plant species (c. 43% of the Canary Island flora) from the literature. Linking these traits to species occurrences and composition across a 500 m × 500 m grid, we calculated functional diversity for endemic, non-endemic native and alien assemblages using multidimensional functional hypervolumes and related the resulting patterns to climatic (humidity) and island biogeographical (geographical isolation, topographic complexity and geological age) gradients.
Results: Trait space of endemic and non-endemic native species overlapped considerably, and alien species added novel trait combinations, expanding the overall functional space of the Canary Islands. We found that functional diversity of endemic plant assemblages was highest in geographically isolated and humid grid cells. Functional diversity of non-endemic native assemblages was highest in less isolated and humid grid cells. In contrast, functional diversity of alien assemblages was highest in arid ecosystems. Topographic complexity and geological age had only a subordinate effect on functional diversity across floristic groups.
Main conclusions: We found that endemic and non-endemic native island species possess similar traits, whereas alien species tend to expand functional space in ecosystems where they have been introduced. The spatial distribution of the functional diversity of floristic groups is very distinct across environmental gradients, indicating that species assemblages of different evolutionary origins thrive functionally in dissimilar habitats.
Aim: Biological invasions are likely determined by species dispersal strategies as well as environmental characteristics of a recipient region, especially climate and human impact. However, the contribution of climatic factors, human impact, and dispersal strategies in driving invasion processes is still controversial and not well embedded in the existing theoretical considerations. Here, we study how climate, species dispersal strategies, and human impact determine plant invasion processes on islands distributed in all major oceans in the context of directional ecological filtering.
Location: Six mountainous, tropical, and subtropical islands in three major oceans: Island of Hawai'i and Maui (Pacific), Tenerife and La Palma (Atlantic), and La Réunion and Socotra (Indian Ocean).
Taxon: Vascular Plants.
Methods: We recorded 360 non-native species in 218 plots along roadside elevational transects covering the major temperature, precipitation and human impact (i.e., road density) gradients of the islands. We collected dispersal strategies for a majority of the recorded species and calculated the environmental niche per species using a hypervolume approach.
Results: Non-native species’ generalism (i.e., mean community niche width) increased with precipitation, elevation and human impact but showed no relationship with temperature. Increasing precipitation led to environmental filtering of non-native species resulting in more generalist species under high precipitation conditions. We found no directional filtering for temperature but an optimum range of most species between 10 and 20°C. Niche widths of non-native species increased with the prevalence of certain dispersal strategies, particularly anemochory and anthropochory.
Main conclusions: Plant invasion on tropical and subtropical islands seems to be mainly driven by precipitation and human impact, while temperature seems to be of little importance. Furthermore, anemochory and anthropochory are dispersal strategies associated with large niche widths of non-native species. Our study allows a more detailed look at the mechanisms behind directional ecological filtering of non-native plant species in non-temperature-limited ecosystems.