Impact of invasive alien plants on native plant communities and Natura 2000 habitats: State of the art, gap analysis and perspectives in Italy.

Invasive alien plants are a major threat to biodiversity and they contribute to the unfavourable conservation status of habitats of interest to the European Community. In order to favour implementation of European Union Regulation no. 1143/2014 on invasive alien species, the Italian Society of Vegetation Science carried out a large survey led by a task force of 49 contributors with expertise in vegetation across all the Italian administrative regions. The survey summed up the knowledge on impact mechanisms of invasive alien plants in Italy and their outcomes on plant communities and the EU habitats of Community Interest, in accordance with Directive no. 92/43/EEC. The survey covered 241 alien plant species reported as having deleterious ecological impacts. The data collected illustrate the current state of the art, highlight the main gaps in knowledge, and suggest topics to be further investigated. In particular, the survey underlined competition as being the main mechanism of ecological impact on plant communities and Natura 2000 habitats. Of the 241 species, only Ailanthus altissima was found to exert an ecological impact on plant communities and Natura 2000 habitats in all Italian regions; while a further 20 species impact up to ten out of the 20 Italian administrative regions. Our data indicate that 84 out of 132 Natura 2000 Habitats (64%) are subjected to some degree of impact by invasive alien plants. Freshwater habitats and natural and semi-natural grassland formations were impacted by the highest number of alien species, followed by coastal sand dunes and inland dunes, and forests. Although not exhaustive, this research is the first example of nationwide evaluation of the ecological impacts of invasive alien plants on plant communities and Natura 2000 Habitats.

Persistence of the Strictly Endemic Plants of Forest Margins: The Case of Cirsium alpis-lunae in the Northern Apennines (Italy).

Narrow endemic plants constitute a pivotal group for conservation, being often reduced to a small contingent of individuals and frequently threatened. However, effective conservation actions require reliable basic information about distribution range, ecological requirements, and population traits. Nevertheless, such knowledge results are incomplete or even completely missing for some neglected or recently described plants, such as Cirsium alpis-lunae, a thistle exclusive to the N-Apennines (Italy). To fill this gap, all sites where C. alpis-lunae grow were monitored, and data on the site and population traits were collected. Our results indicated that this plant is restricted to 16 scattered sites, varied in surface area and number of individuals. Reproductive and juvenile plants showed to be affected by roughly the same variables, in particular the surface of the site, the slope aspect, and the canopy cover. The narrow ecological niche of C. alpis-lunae was mainly determined by the canopy cover, and where coverage increases, the number of individuals decreases. The individuals only grow at forest edges, where the peculiar ecological conditions are limiting factors for the development of forestry cover; some other factors (i.e., high inclination and instability of the substrate) contribute to limiting the development of forestry vegetation and guarantee the persistence of these ecotones. Despite the great difficulties in accessing the sites where this species grows, this study presents, for the first time, a complete picture of the C. alpis-lunae population and yielded important data to identify effective conservation measures.

Impacts of the Invasive Alien Carpobrotus spp. on Coastal Habitats on a Mediterranean Island (Giglio Island, Central Italy).

Carpobrotus acinaciformis and C. edulis are well-known invasive alien plants native to South Africa, whose detrimental effects on native communities are widely documented in the Mediterranean basin and thus largely managed in coastal ecosystems. Most of the literature on these species focuses on their impacts on habitats of sandy coastal dunes, while the effects of Carpobrotus spp. invasion on other habitats such as rocky cliffs and coastal scrubs and garrigues are almost neglected. We present a study case conducted on a small Mediterranean island where Carpobrotus spp. invaded three different natural habitats listed within the Habitat Directive 92/43/CEE (Natura 2000 codes 1240, 1430, and 5320). We surveyed the presence and abundance of native species and Carpobrotus spp. on 44 permanent square plots of 4 m2 in invaded and uninvaded areas in each of the three habitats. We found impacts on plant alpha diversity (intended as the species diversity within each sampled plot) in all the habitats investigated in terms of a decrease in species richness, Shannon index, and abundance. Invaded communities also showed a severe change in species composition with a strong homogenization of the floras of the three habitats. Finally, the negative effect of invasion emerged even through the analyses of beta diversity (expressing the species diversity among sampled plots of the same habitat type), with Carpobrotus spp. replacing a large set of native species.

Effects of a Dominant Species on the Functional Diversity of Coexisting Species in Temperate Deciduous Understorey.

The herb layer plays a significant role in maintaining forest functions, and its community composition is determined by various abiotic factors and biotic interactions. This study attempted to investigate the interspecific plant-plant biotic interactions using a functional traits approach. Specifically, the effects of a dominant species coverage on the functional diversity of coexisting species in the temperate forest understory were studied. Species coverage and soil moisture data were collected using a 1 m2 quadrat couplet (2 × 1 m2) from six sites alongside a 20 m linear transect encompassing a cover gradient of Allium ursinum in southwest Hungary. Major plant functional dimensions i.e., aboveground, and clonal functional traits were considered. Linear and nonlinear mixed models to quantify the effects of biotic interaction on the functional diversity of every single trait and multiple traits were employed. Both aboveground traits and clonal traits of persistent clonal growth organs responded positively to the A. ursinum L., cover gradient. The coexistence of understory species in the presence of a monodominant species seems to be mainly influenced by aboveground traits as compared to the clonal traits suggesting, a role of niche differentiation. The consistent impact of A. ursinum coverage on coexisting species dynamics highlights a need for similar in-depth studies in various forest settings.

Small-scale drivers on plant and ant diversity in a grassland habitat through a multifaceted approach.

Semi-natural grasslands are characterized by high biodiversity and require multifaceted approaches to monitor their biodiversity. Moreover, grasslands comprise a multitude of microhabitats, making the scale of investigation of fundamental importance. Despite their wide distribution, grasslands are highly threatened and are considered of high conservation priority by Directive no. 92/43/EEC. Here, we investigate the effects of small-scale ecological differences between two ecosites present within the EU habitat of Community Interest of semi-natural dry grasslands on calcareous substrates (6210 according to Dir. 92/43/EEC) occurring on a Mediterranean mountain. We measured taxonomic and functional diversity of plant and ant communities, evaluating the differences among the two ecosites, how these differences are influenced by the environment and whether vegetation affects composition of the ant community. Our results show that taxonomic and functional diversity of plant and ant communities are influenced by the environment. While vegetation has no effect on ant communities, we found plant and ant community composition differed across the two ecosites, filtering ant and plant species according to their functional traits, even at a small spatial scale. Our findings imply that small-scale monitoring is needed to effectively conserve priority habitats, especially for those that comprise multiple microhabitats.

Patterns of change in α and ß taxonomic and phylogenetic diversity in the secondary succession of semi-natural grasslands in the Northern Apennines.

We studied the secondary succession in semi-natural grasslands (dry grasslands and hay meadows) located in the eastern side of the Tuscan Apennines (Tuscany, Central Italy). We compared these habitats, investigating: (i) the changes in species richness, composition and phylogenetic diversity during the succession; (ii) whether the trends in species loss and species turnover in taxonomic diversity matched those in phylogenetic diversity. We performed a stratified random sampling, in a full factorial design between habitat type and succession stage (60 sampled plots, 10 × 2 types of habitat × 3 stages of succession). We constructed a phylogenetic tree of the plant communities and compared the differences in taxonomic/phylogenetic α- and ß-diversity between these two habitats and during their succession. We identified indicator species for each succession stage and habitat. Looking at α-diversity, both habitats displayed a decrease in species richness, with a random process of species selection in the earlier succession stages from the species regional pool. Nevertheless, in the latter stage of dry grasslands we recorded a shift towards phylogenetic overdispersion at the higher-level groups in the phylogenetic tree. In both habitats, while the richness decreased with succession stage, most species were replaced during the succession. However, the hay meadows were characterized by a higher rate of new species' ingression whereas the dry grasslands became dominated with Juniperus communis. Accordingly, the two habitats showed similar features in phylogenetic ß-diversity. The main component was true phylogenetic turnover, due to replacement of unique lineages along the succession. Nevertheless, in dry grasslands this trend is slightly higher than expected considering the major importance of difference in species richness of dry grasslands sites and this is due to the presence of a phylogenetically very distant species (J. communis).

Applying predictive models to decipher rhizobacterial modifications in common reed die-back affected populations.

The microbiota inhabiting the soil, as well as the rhizosphere, represents a key determinant of several plant functions. Like for humans, dysbiosis of the plant-associated microbiota may be a co-causal agent in disease with still obscure eziology. In the last decades, the common reed Phragmites australis has been deeply studied for its disappearance from natural stands, but no clear causative agents have been identified and no laboratory models of such "reed die-back syndrome" (RDBS) have been developed. In this study, we try to shed light on the RDBS, by comparing the rhizosphere microbiota of five Italian P. australis populations with different degrees of decline. Results obtained showed a biogeographical meaningful pattern of rhizosphere microbiota, coupled with an impact of RDBS. Obtained data allowed to construct a two-steps predictive model which enabled the prediction of the plant health status from the microbiota taxonomic composition, independently from their geographic location. In conclusion, this study represents one of the first overviews that statistically links RDBS to alteration of rhizosphere microbiota and suggests a model for the analysis of plant-bacteria relationships in nature.

AFLP Approach Reveals Variability in Phragmites australis: Implications for Its Die-Back and Evidence for Genotoxic Effects.

Phragmites australis is a subcosmopolitan species typical of wetlands being studied in Europe for its disappearance from natural stands, a phenomenon called reed die-back syndrome (RDBS). Although it is conjectured that low genetic variability contributes to RDBS, this aspect remains neglected to this day. Using a molecular fingerprinting approach and a sequence analysis of the trnT-trnL/rbcL-psaI regions of cpDNA, this study aimed to compare the genetic structure of stable vs. RDBS-affected P. australis stands from five wetlands of central Italy. Beforehand, in order to characterize the health condition of reed populations, the occurrence of the main macromorphological descriptors for RDBS was considered on 40 reed stands. Soil samples were also collected to examine the total content of heavy metals. The current study analyzed cpDNA in 19 samples and AFLP profiles in 381 samples to investigate the genetic structure of Phragmites populations. Based on the multinomial-Dirichlet model, an analysis of candidate loci under selective pressure was also performed. The relationships among AFLP data, RDBS descriptors and chemicals were evaluated with the use of Linear Mixed Models. The analysis of the cpDNA shows the occurrence of the haplotypes M (the most widespread), and K here recorded for the first time in Italy. Three new haplotypes were also described. The DNA fingerprinting analysis has produced a total of 322 loci (98% polymorphic) and shows the medium-to-high amount of genetic diversity. The significant genetic differentiation among wetlands (Fst = 0.337) suggests either low gene flow or small effective population size. Moreover, the low amount of outlier loci (only 5; l.5% of the total), seems to indicate the scarce occurrence of selective pressure upon the reed's genome. Genetic diversity increased in relationship to the decrease in diameter and of flowering buds of the reed, two of the trends associated with the die-back. The current study rejects the hypothesis that genetic diversity massively contributed to RDBS. Moreover, significant relationships between genetic diversity and the total concentration of some heavy metals (Cr, Cu, and Zn) were highlighted, indicating possible genotoxic effects on P. australis. The current study represents a fact-finding background useful for the conservation of common reed.