Intraspecific variability of leaf form and function across habitat types.

Trait-based ecology has already revealed main independent axes of trait variation defining trait spaces that summarize plant adaptive strategies, but often ignoring intraspecific trait variability (ITV). By using empirical ITV-level data for two independent dimensions of leaf form and function and 167 species across five habitat types (coastal dunes, forests, grasslands, heathlands, wetlands) in the Italian peninsula, we found that ITV: (i) rotated the axes of trait variation that define the trait space; (ii) increased the variance explained by these axes and (iii) affected the functional structure of the target trait space. However, the magnitude of these effects was rather small and depended on the trait and habitat type. Our results reinforce the idea that ITV is context-dependent, calling for careful extrapolations of ITV patterns across traits and spatial scales. Importantly, our study provides a framework that can be used to start integrating ITV into trait space analyses.

Climate regulation processes are linked to the functional composition of plant communities in European forests, shrublands, and grasslands.

Terrestrial ecosystems affect climate by reflecting solar irradiation, evaporative cooling, and carbon sequestration. Yet very little is known about how plant traits affect climate regulation processes (CRPs) in different habitat types. Here, we used linear and random forest models to relate the community-weighted mean and variance values of 19 plant traits (summarized into eight trait axes) to the climate-adjusted proportion of reflected solar irradiation, evapotranspiration, and net primary productivity across 36,630 grid cells at the European extent, classified into 10 types of forest, shrubland, and grassland habitats. We found that these trait axes were more tightly linked to log evapotranspiration (with an average of 6.2% explained variation) and the proportion of reflected solar irradiation (6.1%) than to net primary productivity (4.9%). The highest variation in CRPs was explained in forest and temperate shrubland habitats. Yet, the strength and direction of these relationships were strongly habitat-dependent. We conclude that any spatial upscaling of the effects of plant communities on CRPs must consider the relative contribution of different habitat types.

Plant invasion risk inside and outside protected areas: Propagule pressure, abiotic and biotic factors definitively matter.

Invasive alien species are among the main global drivers of biodiversity loss posing major challenges to nature conservation and to managers of protected areas. The present study applied a methodological framework that combined invasive Species Distribution Models, based on propagule pressure, abiotic and biotic factors for 14 invasive alien plants of Union concern in Italy, with the local interpretable model-agnostic explanation analysis aiming to map, evaluate and analyse the risk of plant invasions across the country, inside and outside the network of protected areas. Using a hierarchical invasive Species Distribution Model, we explored the combined effect of propagule pressure, abiotic and biotic factors on shaping invasive alien plant occurrence across three biogeographic regions (Alpine, Continental, and Mediterranean) and realms (terrestrial and aquatic) in Italy. We disentangled the role of propagule pressure, abiotic and biotic factors on invasive alien plant distribution and projected invasion risk maps. We compared the risk posed by invasive alien plants inside and outside protected areas. Invasive alien plant distribution varied across biogeographic regions and realms and unevenly threatens protected areas. As an alien's occurrence and risk on a national scale are linked with abiotic factors followed by propagule pressure, their local distribution in protected areas is shaped by propagule pressure and biotic filters. The proposed modelling framework for the assessment of the risk posed by invasive alien plants across spatial scales and under different protection regimes represents an attempt to fill the gap between theory and practice in conservation planning helping to identify scale, site, and species-specific priorities of management, monitoring and control actions. Based on solid theory and on free geographic information, it has great potential for application to wider networks of protected areas in the world and to any invasive alien plant, aiding improved management strategies claimed by the environmental legislation and national and global strategies.

Climate-trait relationships exhibit strong habitat specificity in plant communities across Europe.

Ecological theory predicts close relationships between macroclimate and functional traits. Yet, global climatic gradients correlate only weakly with the trait composition of local plant communities, suggesting that important factors have been ignored. Here, we investigate the consistency of climate-trait relationships for plant communities in European habitats. Assuming that local factors are better accounted for in more narrowly defined habitats, we assigned > 300,000 vegetation plots to hierarchically classified habitats and modelled the effects of climate on the community-weighted means of four key functional traits using generalized additive models. We found that the predictive power of climate increased from broadly to narrowly defined habitats for specific leaf area and root length, but not for plant height and seed mass. Although macroclimate generally predicted the distribution of all traits, its effects varied, with habitat-specificity increasing toward more narrowly defined habitats. We conclude that macroclimate is an important determinant of terrestrial plant communities, but future predictions of climatic effects must consider how habitats are defined.

Dimensions of invasiveness: Links between local abundance, geographic range size, and habitat breadth in Europe's alien and native floras.

Understanding drivers of success for alien species can inform on potential future invasions. Recent conceptual advances highlight that species may achieve invasiveness via performance along at least three distinct dimensions: 1) local abundance, 2) geographic range size, and 3) habitat breadth in naturalized distributions. Associations among these dimensions and the factors that determine success in each have yet to be assessed at large geographic scales. Here, we combine data from over one million vegetation plots covering the extent of Europe and its habitat diversity with databases on species' distributions, traits, and historical origins to provide a comprehensive assessment of invasiveness dimensions for the European alien seed plant flora. Invasiveness dimensions are linked in alien distributions, leading to a continuum from overall poor invaders to super invaders-abundant, widespread aliens that invade diverse habitats. This pattern echoes relationships among analogous dimensions measured for native European species. Success along invasiveness dimensions was associated with details of alien species' introduction histories: earlier introduction dates were positively associated with all three dimensions, and consistent with theory-based expectations, species originating from other continents, particularly acquisitive growth strategists, were among the most successful invaders in Europe. Despite general correlations among invasiveness dimensions, we identified habitats and traits associated with atypical patterns of success in only one or two dimensions-for example, the role of disturbed habitats in facilitating widespread specialists. We conclude that considering invasiveness within a multidimensional framework can provide insights into invasion processes while also informing general understanding of the dynamics of species distributions.

Functional composition and diversity of leaf traits in subalpine versus alpine vegetation in the Apennines.

Mediterranean high mountain grasslands are shaped by climatic stress and understanding their functional adaptations can contribute to better understanding ecosystems' response to global change. The present work analyses the plant functional traits of high-elevation grasslands growing in Mediterranean limestone mountains to explore, at the community level, the presence of different plant strategies for resource use (conservative vs. acquisitive) and functional diversity syndromes (convergent or divergent). Thus, we compared the functional composition and diversity of the above-ground traits related to resource acquisition strategies of subalpine and alpine calcareous grasslands in the central Apennines, a mountain region characterized by a dry-summer Mediterranean climate. We used georeferenced vegetation plots and field-measured plant functional traits (plant maximum height, specific leaf area and leaf dry matter content) for the dominant species of two characteristic vegetation types: the subalpine Sesleria juncifolia community and the alpine Silene acaulis community. Both communities are of particular conservation concern and are rich in endemic species for which plant functional traits are measured here for the first time. We analysed the functional composition and diversity using the community-weighted mean trait index and the functional diversity using Rao's function, and we assessed how much the observed pattern deviated from a random distribution by calculating the respective standardized effect sizes. The results highlighted that an acquisitive resource use strategy and relatively higher functional diversity of leaf traits prevail in the alpine S. acaulis community, optimizing a rapid carbon gain, which would help overcome the constraints exerted by the short growing season. The divergent functional strategy underlines the co-occurrence of different leaf traits in the alpine grasslands, which shows good adaptation to a microhabitat-rich environment. Conversely, in the subalpine S. juncifolia grassland, a conservative resource use strategy and relatively lower functional diversity of the leaf traits are likely related to a high level resistance to aridity over a longer growing season. Our outcomes indicate the preadaptation strategy of the subalpine S. juncifolia grassland to shift upwards to the alpine zone that will become warmer and drier as a result of anthropogenic climate change.

Not just a sandy beach. The multi-service value of Mediterranean coastal dunes.

Coastal sand dunes are complex transitional systems hosting high levels of biodiversity and providing important benefits to society. In this paper we aimed to evaluate the multi-service nature of ecosystem services (ES) supply in the dunes of the Italian Adriatic coast within Natura 2000 (N2K) sites. We i) identified ES indicators and assessed the supply capacity (Climate regulation, Protection from wind and aerosol, Erosion regulation, Recreation and Tourism and Existence value of biodiversity) of natural dune ecosystems of European conservation concern; ii) upscaled this data to create an inventory of ES supply for all dune N2K sites in the study area; iii) explored the trade-offs among ES; and iv) summarized and spatially compared the overall multi-service value of the N2K sites. The study provides a method for quantifying the role of N2K sites in supplying benefits for our society. We found that the multi-service capacity of coastal dunes is uneven within sites and within administrative regions. This variability is related to both ecological (e.g. distribution, ecological integrity, extent and conservation status of dune habitats) and administrative (e.g. local implementation of the Habitats Directive) characteristics of the analysed area. ES are not coupled as several sites with high values for one ES show very low values for others. The results suggest that conservation actions should favour restoration of the natural dune zonation, since this underpins multi-service ES supply. The approach can distinguish regions with high ES values and regions where the paucity of protected areas represents a gap in ES supply, fact that offers an incentive to enhance the protection strategy but also suggests an urgent need to improve the N2K network by enlarging existent sites and including new ones.

Assessing transhumance corridors on high mountain environments by least cost path analysis: the case of yak herds in Gilgit-Baltistan, Pakistan.

Mountain environments in the world host highly specialized flora and fauna which are vulnerable because of land use and climate change. Transhumance and other land use traditional practices are present in most of the mountains of the world, and management tools able to cope with new socioeconomic settings and environmental changes are urgently needed. During past centuries, yak (Bos grunniens) herding in Northern Pakistan involved the migration of herds to high mountain international rangelands, but the recent establishment of international borders breaks the traditional transhumance paths, promoting several ecological problems. In this paper, we propose the use of least cost path (LCP) algorithm to identify the most efficient corridors of transhumance for yak herds on northern high altitudes of Pakistan. Specifically, LCP was implemented to identify the critical grazing areas and the connecting zones to be accounted in a new management plan for the yaks in the region. The LCP analysis showed that some grazing areas are connected with several paths, whereas other areas are connected to a lesser extent. The analyses identified a set of best minimum cost paths able to guarantee local connectivity. We also delineated several medium and low efficient paths that could play a crucial role for maintaining regional connectivity which is essential for reducing the isolation of herds and the consequent inbreeding problems. The analytical framework implemented in this study allowed to (1) provide valuable information concerning the movement of yak herds in Gilgit-Baltistan, (2) identify potential corridors that are able to promote herd movement between villages and high mountain rangelands, and (3) identify critical areas for the connectivity of yaks by ranking of the potential corridors according to their length and permeability. The analysis would be extended to other transhumant herds and high mountain areas that are facing sociopolitical transformations and environmental changes.

A systematic conservation planning approach to fire risk management in Natura 2000 sites.

A primary challenge in conservation biology is to preserve the most representative biodiversity while simultaneously optimizing the efforts associated with conservation. In Europe, the implementation of the Natura 2000 network requires protocols to recognize and map threats to biodiversity and to identify specific mitigation actions. We propose a systematic conservation planning approach to optimize management actions against specific threats based on two fundamental parameters: biodiversity values and threat pressure. We used the conservation planning software Marxan to optimize a fire management plan in a Natura 2000 coastal network in southern Italy. We address three primary questions: i) Which areas are at high fire risk? ii) Which areas are the most valuable for threatened biodiversity? iii) Which areas should receive priority risk-mitigation actions for the optimal effect?, iv) which fire-prevention actions are feasible in the management areas?. The biodiversity values for the Natura 2000 spatial units were derived from the distribution maps of 18 habitats and 89 vertebrate species of concern in Europe (Habitat Directive 92/43/EEC). The threat pressure map, defined as fire probability, was obtained from digital layers of fire risk and of fire frequency. Marxan settings were defined as follows: a) planning units of 40 × 40 m, b) conservation features defined as all habitats and vertebrate species of European concern occurring in the study area, c) conservation targets defined according with fire sensitivity and extinction risk of conservation features, and d) costs determined as the complement of fire probabilities. We identified 23 management areas in which to concentrate efforts for the optimal reduction of fire-induced effects. Because traditional fire prevention is not feasible for most of policy habitats included in the management areas, alternative prevention practices were identified that allows the conservation of the vegetation structure. The proposed approach has potential applications for multiple landscapes, threats and spatial scales and could be extended to other valuable natural areas, including protected areas.

Changes in composition, ecology and structure of high-mountain vegetation: a re-visitation study over 42 years.

High-mountain ecosystems are increasingly threatened by climate change, causing biodiversity loss, habitat degradation and landscape modifications. However, very few detailed studies have focussed on plant biodiversity in the high mountains of the Mediterranean. In this study, we investigated the long-term changes that have occurred in the composition, structure and ecology of high-mountain vegetation in the central Apennines (Majella) over the last 42 years. We performed a re-visitation study, using historical and newly collected vegetation data to explore which ecological and structural features have been the most successful in coping with climatic changes. Vegetation changes were analysed by comparing geo-referenced phytosociological relevés collected in high-mountain habitats (dolines, gentle slopes and ridges) on the Majella massif in 1972 and in 2014. Composition analysis was performed by detrended correspondence analysis, followed by an analysis of similarities for statistical significance assessment and by similarity percentage procedure (SIMPER) for identifying which species indicate temporal changes. Changes in ecological and structural indicators were analysed by a permutational multivariate analysis of variance, followed by a post hoc comparison. Over the last 42 years, clear floristic changes and significant ecological and structural variations occurred. We observed a significant increase in the thermophilic and mesonitrophilic plant species and an increment in the frequencies of hemicryptophytes. This re-visitation study in the Apennines agrees with observations in other alpine ecosystems, providing new insights for a better understanding of the effects of global change on Mediterranean high-mountain biodiversity. The observed changes in floristic composition, the thermophilization process and the shift towards a more nutrient-demanding vegetation are likely attributable to the combined effect of higher temperatures and the increase in soil nutrients triggered by global change. The re-visitation approach adopted herein represents a powerful tool for studying climate-related changes in sensitive high-mountain habitats.

Temporal Changes in Forest Contexts at Multiple Extents: Three Decades of Fragmentation in the Gran Chaco (1979-2010), Central Argentina.

The context in which a forest exists strongly influences its function and sustainability. Unveiling the multi-scale nature of forest fragmentation context is crucial to understand how human activities affect the spatial patterns of forests across a range of scales. However, this issue remains almost unexplored in subtropical ecosystems. In this study, we analyzed temporal changes (1979-2010) in forest contexts in the Argentinean dry Chaco at multiple extents. We classified forests over the last three decades based on forest context amount (Pf) and structural connectivity (Pff), which were measured using a moving window approach fixed at eight different extents (from local, ~ 6 ha, to regional, ~ 8300 ha). Specific multi-scale forest context profiles (for the years 1979 and 2010) were defined by projecting Pf vs. Pff mean values and were compared across spatial extents. The distributions of Pf across scales were described by scalograms and their shapes over time were compared. The amount of agricultural land and rangelands across the scales were also analyzed. The dry Chaco has undergone an intensive process of fragmentation, resulting in a shift from landscapes dominated by forests with gaps of rangelands to landscapes where small forest patches are embedded in agricultural lands. Multi-scale fragmentation analysis depicted landscapes in which local exploitation, which perforates forest cover, occurs alongside extensive forest clearings, reducing forests to small and isolated patches surrounded by agricultural lands. In addition, the temporal diminution of Pf's variability along with the increment of the mean slope of the Pf 's scalograms, indicate a simplification of the spatial pattern of forest over time. The observed changes have most likely been the result of the interplay between human activities and environmental constraints, which have shaped the spatial patterns of forests across scales. Based on our results, strategies for the conservation and sustainable management of the dry Chaco should take into account both the context of each habitat location and the scales over which a forest pattern might be preserved, altered or restored.

Seasonal habitat preference by the flagship species Testudo hermanni: Implications for the conservation of coastal dunes.

In this study, we explored if, how, and when the European Union habitats (EU sensu Habitats Directive 92/43/CEE) are used by the flagship species Testudo hermanni in a well-preserved coastal dune system of the Italian peninsula. Radio telemetry data and fine-scale vegetation habitat mapping were used to address the following questions: (a) is each EU habitat used differentially by Hermann's tortoises? (b) is there any seasonal variation in this utilization pattern? (c) how does each habitat contribute to the ecological requirements of the tortoises? Nine tortoises were fitted with transmitters and monitored for the entire season of activity. The eight EU habitats present in the study area were surveyed and mapped using GIS. The seasonal preferential use or avoidance of each habitat was tested by comparing, through bootstrap tests, the proportion of habitat occupied (piTh) with the proportion of available habitat in the entire landscape (piL). The analysis of 340 spatial locations showed a marked preference for the Cisto-Lavanduletalia dune sclerophyllous scrubs (EU code 2260) and a seasonal selection of Juniperus macrocarpa bushes (EU code 2250(*)), wooded dunes with Pinus (EU code 2270) and mosaic of dune grasslands and sclerophyllous scrubs (EU codes 2230, 2240, 2260). Seasonal variation of habitat preference was interpreted in light of the different feeding, thermoregulation and reproductive needs of the tortoises. Our results stress the ecological value of EU coastal dune habitats and suggest prioritization of conservation efforts in these ecosystems.

Ecosystem classification for EU habitat distribution assessment in sandy coastal environments: an application in central Italy.

Many recent developments in coastal science have gone against the demands of European Union legislation. Coastal dune systems which cover small areas of the earth can host a high level of biodiversity. However, human pressure on coastal zones around the world has increased dramatically in the last 50 years. In addition to direct habitat loss, the rapid extinction of many species that are unique to these systems can be attributed to landscape deterioration through the lack of appropriate management. In this paper, we propose to use of an ecosystem classification technique that integrates potential natural vegetation distribution as a reference framework for coastal dune EU Habitats (92/43) distribution analysis and assessment. As an example, the present study analyses the EU Habitats distribution within a hierarchical ecosystem classification of the coastal dune systems of central Italy. In total, 24 land elements belonging to 8 land units, 5 land facets, 2 land systems and 2 land regions were identified for the coastal dunes of central Italy, based on diagnostic land attributes. In central Italy, coastal dune environments including all the beach area, mobile dunes and all the fixed-dune land elements contain or could potentially hold at least one EU habitat of interest. Almost all dune slack transitions present the potentiality for the spontaneous development of EU woodlands of interest. The precise information concerning these ecosystems distribution and ecological relationships that this method produces, makes it very effective in Natura 2000 European network assessment. This hierarchical ecosystem classification method facilitates the identification of areas to be surveyed and eventually bound, under the implementation of EU Habitat directive (92/43) including areas with highly disturbed coastal dune ecosystems.