Marta Carboni

Environmental Factors Influencing Coastal Vegetation Pattern: New Insights from the Mediterranean Basin

Coastal dune ecosystems show strongly dynamic interactions between abiotic and biotic factors. The relationship between plant communities and environmental factors has been previously studied in oceanic dune systems, but few studies have been conducted along Mediterranean coasts. In this study we analyze the relative contribution of environmental factors associated with two of the main drivers of vegetation zonation: soil and wind. We chose two representative coastal dune systems in the western Mediterranean Basin subject to low levels of human disturbance. Within 54 plots we recorded floristic and environmental data. Vegetation zonation and relationships with environmental variables were investigated through unconstrained and constrained ordinations, correlation, and variance partitioning. Environmental factors shift along the gradient from coastal to inland dunes, concomitantly with the pattern of community types from annual beach communities to shrub-covered fixed dunes. This general gradient is similar both in the Mediterranean and in the oceanic coastal ecosystems, with the same factors that show similar trends along the dune profile. However, our results highlight some peculiarities of the Mediterranean dune systems in relation to the amount of variation explained by environmental factors. While most studies conducted in oceanic ecosystems find that wind-related parameters may control the vegetation zonation, in our study areas we observed a minor importance of the wind-related variables when compared to soil properties. In particular, organic matter and grain-size variability were found to be closely correlated with the distribution of plant communities along the gradient.

The community ecology of invasive species: where are we and what's next?

Alien species are continually introduced in most regions of the world, but not all survive and coexist with the resident native species. Approaches analyzing the functional (or phylogenetic) similarity between invasive species and native communities are increasingly employed to infer the processes underlying successful invasions and to predict future invaders. The relatively simple conceptual foundations have made these approaches very appealing and therefore widely used, often leading to confusion and hampering generalizations. We undertook a comprehensive review and synthesis of the functional similarity approach in invasion community ecology to clarify its advantages and limitations, to summarize what has been learned thus far, and to suggest avenues for future improvement. We first present the methodological state of the art and provide general guidelines. Second, by organizing the published literature around seven key questions in invasion ecology we found cumulative evidence that: at large spatial scales phylogenetic relatedness of invasive and native species is a good predictor of invasion success, but a poor predictor of invasion impacts; at fine spatial scales, community resistance to invasion tends to increase with native species diversity and with similarity to the invaders, consistent with patterns emerging from biotic interactions. In general, the processes filtering invaders appear to vary across species’ invasion stage and along environmental gradients. Nonetheless, we found conflicting evidence for differences in community assembly processes between invasive and native species, and between the invaders native and adventive ranges. Finally, we propose four important avenues for overcoming some of the identified methodological and conceptual limitations of the approach, in particular by integrating observational and experimental analyses and by explicitly considering species’ demographic rates and complex biotic interactions. Although the functional similarity approach bears intrinsic limitations, it still offers many opportunities in invasion community ecology at large spatial scales. This article is protected by copyright. All rights reserved.

Identifying the signal of environmental filtering and competition in invasion patterns – a contest of approaches from community ecology

Summary In 1859, Darwin had already identified environmental constraints and competition with the native community as major drivers of invasion success. Since then, a toolbox of indices and statistical approaches has been developed and commonly applied to test for the relative importance of these drivers. This toolbox is largely based on community ecology theory with the underlying hypothesis that patterns of trait (or phylogenetic) similarities between invaders and native species permit to disentangle the signatures of competition and environmental filtering. However, so far the performance of the indices and statistical approaches has not been thoroughly evaluated, and there exists no study exploring the sensitivity of the different methods given common biases in field data. This severely hampers intercomparisons of invasion studies and ultimately prevents the elaboration of general conclusions. In this study, we developed a mechanistic community assembly model to simulate invasion patterns across a range of communities and tested the performance of four different indices aiming at disentangling environmental filtering vs. competition from these patterns. Furthermore, we evaluated the sensitivity of the statistical methods to biases in the data (resulting from non-equilibrium dynamics or observation errors). Our results indicated that the best performing index was mean distance to the native species (the average functional distance between the invader and all the species of the community), especially in heterogeneous landscapes. Further, we demonstrated that the detection of competition was more sensitive to the presence of biases in the data than the detection of environmental filtering. In conclusion, studying invasion mechanisms based on community patterns is possible when employing the appropriate statistical method, but it is highly sensitive to the quality of the data set used.

Integrating invasive species policies across ornamental horticulture supply chains to prevent plant invasions

Research was supported by COST Action TD1209 “Alien Challenge”. The authors are grateful to John David and Franziska Humair for valuable discussions on this topic. PP and JP were supported by project no. 14-36079G Centre of Excellence PLADIAS (Czech Science Foundation) and RVO 67985939 (The Czech Academy of Sciences). FE, SD, MC and MvK were supported by the ERA-Net BiodivERsA through the Austrian Science Fund, German Research Foundation and French National Research Agency. AN was supported by the Working for Water (WfW) Programme and the DST-NRF Centre of Excellence for Invasion Biology. HS acknowledges support by the DFG (grant SE 1891/2-1).

Multifaceted Analysis of Patch-Level Plant Diversity in Response to Landscape Spatial Pattern and History on Mediterranean Dunes

Landscape structure is known to critically affect biodiversity. However, although the multi-facetted character of biodiversity is widely recognized, few studies have linked landscape spatial pattern and history simultaneously to multiple facets (taxonomic, functional, and phylogenetic) and spatial components (α, β, and γ) of plant diversity. We set out to reveal whether landscape parameters have specific effects on the separate diversity facets and components of plant diversity at a patch scale on coastal dune landscapes of Central Italy. For each landscape patch, we computed a set of patch-based metrics relying on multi-temporal land-cover maps. Based on a database of plant community plots, on functional traits from field measurements and on a dated phylogenetic tree, we calculated taxonomic (TD), functional (FD), and phylogenetic diversity (PD) within each patch at α, β, and γ level. Diversity measures were then related to the landscape metrics via linear mixed-effect models. Landscape pattern and transformations affected TD only moderately in coastal dune ecosystems. We found much stronger and contrasted effects on FD and PD. FD increased in patches surrounded by human-dominated habitats; PD was higher in fragmented patches, particularly in the Mediterranean macchia. Moreover, landscape pattern affected differently the single communities, the turnover among communities and the pool of species within the patch (α, β, and γ components). Our results call for the combined inclusion of FD and PD and their partitions into ecological analyses, being TD too crude to capture the comprehensive and contrasted response of plant diversity to landscape spatial pattern.

Extreme climate events counteract the effects of climate and land‐use changes in Alpine tree lines

1. Climate change and extreme events, such as drought, threaten ecosystems worldwide and in particular mountain ecosystems, where species often live at their environmental tolerance limits. In the European Alps, plant communities are also influenced by land-use abandonment leading to woody encroachment of subalpine and alpine grasslands. 2. In this study, we explored how the forest-grassland ecotone of Alpine treelines will respond to gradual climate warming, drought events and land-use change in terms of forest expansion rates, taxonomic diversity and functional composition. We used a previously validated dynamic vegetation model, FATE-HD, parameterised for plant communities in the Ecrins National Park in the French Alps. 3. Our results showed that intense drought counteracted the forest expansion at higher elevations driven by land-use abandonment and climate change, especially when combined with high drought frequency (occurring every 2 or less than 2 years). 4. Furthermore, intense and frequent drought accelerated the rates of taxonomic change and resulted in overall higher taxonomic spatial heterogeneity of the ecotone than would be expected under gradual climate and land-use changes only. 5. Synthesis and applications. The results from our model show that intense and frequent drought counteracts forest expansion driven by climate and land-use changes in the forest-grassland ecotone of Alpine treelines. We argue that land-use planning must consider the effects of extreme events, such as drought, as well as climate and land-use changes, since extreme events might interfere with trends predicted under gradual climate warming and agricultural abandonment.

The changing role of ornamental horticulture in alien plant invasions

The number of alien plants escaping from cultivation into native ecosystems is increasing steadily. We provide an overview of the historical, contemporary and potential future roles of ornamental horticulture in plant invasions. We show that currently at least 75% and 93% of the global naturalised alien flora is grown in domestic and botanical gardens, respectively. Species grown in gardens also have a larger naturalised range than those that are not. After the Middle Ages, particularly in the 18th and 19th centuries, a global trade network in plants emerged. Since then, cultivated alien species also started to appear in the wild more frequently than non‐cultivated aliens globally, particularly during the 19th century. Horticulture still plays a prominent role in current plant introduction, and the monetary value of live‐plant imports in different parts of the world is steadily increasing. Historically, botanical gardens – an important component of horticulture – played a major role in displaying, cultivating and distributing new plant discoveries. While the role of botanical gardens in the horticultural supply chain has declined, they are still a significant link, with one‐third of institutions involved in retail‐plant sales and horticultural research. However, botanical gardens have also become more dependent on commercial nurseries as plant sources, particularly in North America. Plants selected for ornamental purposes are not a random selection of the global flora, and some of the plant characteristics promoted through horticulture, such as fast growth, also promote invasion. Efforts to breed non‐invasive plant cultivars are still rare. Socio‐economical, technological, and environmental changes will lead to novel patterns of plant introductions and invasion opportunities for the species that are already cultivated. We describe the role that horticulture could play in mediating these changes. We identify current research challenges, and call for more research efforts on the past and current role of horticulture in plant invasions. This is required to develop science‐based regulatory frameworks to prevent further plant invasions.

Simulating plant invasion dynamics in mountain ecosystems under global change scenarios

Across the globe, invasive alien species cause severe environmental changes, altering species composition and ecosystem functions. So far, mountain areas have mostly been spared from large-scale invasions. However, climate change, land-use abandonment, the development of tourism and the increasing ornamental trade will weaken the barriers to invasions in these systems. Understanding how alien species will react and how native communities will influence their success is thus of prime importance in a management perspective. Here, we used a spatially and temporally explicit simulation model to forecast invasion risks in a protected mountain area in the French Alps under future conditions. We combined scenarios of climate change, land-use abandonment and tourism-linked increases in propagule pressure to test if the spread of alien species in the region will increase in the future. We modelled already naturalized alien species and new ornamental plants, accounting for interactions among global change components, and also competition with the native vegetation. Our results show that propagule pressure and climate change will interact to increase overall species richness of both naturalized aliens and new ornamentals, as well as their upper elevational limits and regional range-sizes. Under climate change, woody aliens are predicted to more than double in range-size and herbaceous species to occupy up to 20% of the park area. In contrast, land-use abandonment will open new invasion opportunities for woody aliens, but decrease invasion probability for naturalized and ornamental alien herbs as a consequence of colonization by native trees. This emphasizes the importance of interactions with the native vegetation either for facilitating or potentially for curbing invasions. Overall, our work highlights an additional and previously underestimated threat for the fragile mountain flora of the Alps already facing climate changes, land-use transformations and overexploitation by tourism.

Will climate change increase hybridization risk between potential plant invaders and their congeners in Europe

Abstract Aim Interspecific hybridization can promote invasiveness of alien species. In many regions of the world, public and domestic gardens contain a huge pool of non‐native plants. Climate change may relax constraints on their naturalization and hence facilitate hybridization with related species in the resident flora. Here, we evaluate this possible increase in hybridization risk by predicting changes in the overlap of climatically suitable ranges between a set of garden plants and their congeners in the resident flora. Location Europe. Methods From the pool of alien garden plants, we selected those which (1) are not naturalized in Europe, but established outside their native range elsewhere in the world; (2) belong to a genus where interspecific hybridization has been previously reported; and (3) have congeners in the native and naturalized flora of Europe. For the resulting set of 34 alien ornamentals as well as for 173 of their European congeners, we fitted species distribution models and projected suitable ranges under the current climate and three future climate scenarios. Changes in range overlap between garden plants and congeners were then assessed by means of the true skill statistic. Results Projections suggest that under a warming climate, suitable ranges of garden plants will increase, on average, while those of their congeners will remain constant or shrink, at least under the more severe climate scenarios. The mean overlap in ranges among congeners of the two groups will decrease. Variation among genera is pronounced; however, and for some congeners, range overlap is predicted to increase significantly. Main conclusions Averaged across all modelled species, our results do not indicate that hybrids between potential future invaders and resident species will emerge more frequently in Europe when climate warms. These average trends do not preclude, however, that hybridization risk may considerably increase in particular genera.

Functional trait differences and trait plasticity mediate biotic resistance to potential plant invaders

1Dipartimento di Scienze, Roma Tre University, Rome, Italy; 2Plant Evolutionary Ecology, University of Tübingen, Tübingen, Germany; 3Laboratoire d’Écologie Alpine (LECA), UMR-CNRS, University of Grenoble Alpes, Grenoble, France; 4Ecology Group, Department of Biology, University of Konstanz, Konstanz, Germany; 5Division of Conservation Biology, Vegetation and Landscape Ecology, Department of Botany and Biodiversity Research, University of Vienna, Vienna, Austria and 6Institute of Social Ecology, Faculty for Interdisciplinary Studies, Alps Adria University, Vienna, Austria