Giuseppe Brundu

Inventory of the non‐native flora of Italy

Abstract In this paper we present a comprehensive inventory of the non‐native vascular flora of Italy, which was produced within the project “A survey of the Italian non‐native flora”, funded by the Italian Ministry for the Environment. Previously published floristic accounts were the main source of information. Historical records were critically revised and integrated with recent literature, data from herbaria and some unpublished information, so as to obtain a complete, up‐to‐date catalogue of the non‐native vascular plant species that occur spontaneously in Italy. The inventory lists 1023 non‐native species and subspecies, which account for 13.4% of all the Italian flora. The Italian non‐native flora was divided, according to its residence time, into 103 archaeophytes and 920 neophytes. According to its current invasion status, it was classified into 437 casual (42.7% of all non‐native) and 524 established taxa, the latter being divided into 361 naturalized non‐invasive (35.3%) and 163 invasive taxa (15.9%). The inventory includes a group of 62 species (6.1%) that lack recent records (i.e. since 1950). By combining local expertise into a unified, nationwide scheme using a standardized method and terminology, the inventory provides the essential scientific basis for the development of plant invasion research and management in the country.

No saturation in the accumulation of alien species worldwide

Although research on human-mediated exchanges of species has substantially intensified during the last centuries, we know surprisingly little about temporal dynamics of alien species accumulations across regions and taxa. Using a novel database of 45,813 first records of 16,926 established alien species, we show that the annual rate of first records worldwide has increased during the last 200 years, with 37% of all first records reported most recently (1970–2014). Inter-continental and inter-taxonomic variation can be largely attributed to the diaspora of European settlers in the nineteenth century and to the acceleration in trade in the twentieth century. For all taxonomic groups, the increase in numbers of alien species does not show any sign of saturation and most taxa even show increases in the rate of first records over time. This highlights that past efforts to mitigate invasions have not been effective enough to keep up with increasing globalization.

Non‐native flora of Italy: Species distribution and threats

Abstract In this paper, we provide an overview of the distribution and invasive status of non‐native species in the Italian flora across its administrative regions, biogeographic regions and main land use types, and a synthesis of current knowledge on the threats they pose within the country. The information on non‐native plant species collected during the project “A survey of the non‐native Italian flora” was used to compile comprehensive regional and national databases. The number of non‐native species within a given administrative region increases in proportion to its size, resident population density and latitude, reaching the highest values in the intensively cultivated, heavily industrialized and urbanized Po Plain in northern Italy. The number of casual species is positively correlated with the number of yearly visitors in each region and negatively correlated with the proportion of mountainous terrain within the region. If compared with the Continental and Mediterranean biogeographic regions, the Alpine region yields the lowest number of non‐native species and lowest proportion of casual species. The number and density of introduced species is highest in artificial land use types, particularly in urban areas. A negative impact is reported to be exerted by 203 species, most of which are agricultural weeds.

Consistent performance of invasive plant species within and among islands of the Mediterranean basin

Since the success of an invasive species depends not only upon its intrinsic traits but also on particular characteristics of the recipient habitat, assessing the performance of an invader across habitats provides a more realistic analysis of risk. Such an analysis will not only provide insights into the traits related to invasiveness, but also the habitat characteristics that underpin vulnerability to invasion that, taken together, will facilitate the selection of management strategies to mitigate the invader’s effect. In the present study, we considered the Mediterranean basin islands as an excellent study region to test how the same invasive species perform in different habitats within a single island, and to scale up differences among islands with similar climate. We tested how the performance of three widespread plant invaders with clonal growth but contrasting life-history traits, a deciduous tree Ailanthus altissima, a succulent subshrub Carpobrotus spp., and an annual geophyte Oxalis pes-caprae, varied depending upon the species identity, habitat, and invaded island. The environmental parameters considered were habitat type, elevation, species diversity in the invaded plot, and several soil traits (% C, % N, C/N, pH, and relative humidity). The study documents that the performance of these three important and widespread plant invaders is dependent mainly on species identity, and less upon the invaded island’s general features. Likewise, differences in performance among habitats were only significant in the case of Ailanthus, whereas Carpobrotus and Oxalis appear to perform equally well in different environments. Ailanthus thus appears to have a broader spectrum of invasiveness, being able to invade a larger number of habitat types. On the contrary, Carpobrotus spp. have not yet invaded habitats different from those where the species have been originally introduced and where they are still commonly spread by humans. Oxalis distribution is mainly related to agricultural activities and disturbed sites, and the total area infested by this geophyte may be more reflection of the extent of suitable habitats than of invasiveness or ecological impact. Our results confirm the potential for these species to significantly alter the functioning of ecosystems in the Mediterranean islands and highlight the risk to other islands not yet invaded.

The Origin of Clonal Diversity and Structure of Populus alba in Sardinia: Evidence from Nuclear and Plastid Microsatellite Markers

BACKGROUND AND AIMS Populus alba is a thermophilic forest tree present in the Mediterranean basin. Its habitat is highly fragmented and its distribution range has been subject to long-term human interference, resulting in debate surrounding whether certain populations are native or exotic in origin. In particular, populations from the islands of Corsica and Sardinia are of uncertain origin. While populations of P. alba mainly reproduce sexually, clonal reproduction is also common. The aims of this study were to locate and molecularly characterize the poorly studied island populations of P. alba and compare these with samples from various spatial scales, in order to provide information on the genetic structure and phylogeography of this species. This information will provide evidence on whether the species is native to Sardinia, which is important for the development of conservation strategies. METHODS DNA extracts were obtained from the following P. alba trees: 159 from Sardinia, 47 from Ticino regional park (northern Italy), 15 acquired from an Italian Germoplasm Bank (IRC; Italian Reference Collection) and 28 from the Mediterranean basin (MB). Genetic polymorphisms were revealed at nuclear and chloroplast DNA (cpDNA) microsatellite loci, both at the island scale (Sardinia) and at broader scales, for comparative assessment of the genetic and genotypic diversity and phylogeography. KEY RESULTS Based on nuclear microsatellite loci, Sardinian white poplar consists of a small number of genets (26), each of which is represented by several ramets. Despite the uniqueness of the Sardinian haplotypes and the very low value of genetic diversity at the cpDNA level (vK = 0.15), the HT (0.60) and the AR (3.61) values, estimated at the nuclear level for Sardinia, were comparable with those of the other populations and collections. CONCLUSIONS The uniqueness of the cpDNA haplotypes, the prevalence of clonality and the restricted number of genets recorded suggest that Sardinian white poplar could be a floristic relict of the native flora of the island, which has spread through available habitats on the island mainly by means of vegetative propagation and human activities.

Consistency in the habitat degree of invasion for three invasive plant species across Mediterranean islands

Habitats are known to vary in their vulnerability to invasion by alien plants and different species often colonise distinct habitats. To assess the consistency in the degree of invasion of particular habitats, this study examined the frequency of occurrence and local abundance of three invasive plant taxa: Ailanthus altissima, Carpobrotus spp., and Oxalis pes-caprae across different habitat types on four representative Mediterranean islands. We conducted systematic field surveys recording the presence-absence and cover of these taxa on the islands of Mallorca, Corsica, Sardinia and Crete. Drawing on the results of 5,285 sample points, the frequency of occurrence of the three invaders tends to be higher than expected in urban, ruderal and roadside habitats. In contrast, scrub habitats rarely contain any of the three invaders, indicating that they may be more resistant to invasion. The degree of invasion, determined by the local abundance of an invasive plant in any one habitat, varies according to the identity of the invader and the island. However, based on average abundance, Oxalis pes-caprae exhibits the highest degree of invasion, and Carpobrotus spp. the least. There is no indication that any one of the four islands is more prone to either higher frequencies or abundances of the three invaders. These patterns suggest that anthropogenic changes in Mediterranean islands will increase the vulnerability of certain habitats to invasion and increase the distribution of these three invasive taxa at any of the four islands.

Determinants of non-native plant species richness and composition across small Mediterranean islands

The main aim of this paper is to assess the drivers of non-native plant species richness and composition on a set of 37 small Mediterranean islands in Italy. We compiled the list of the non-native flora and distinguished species according to their establishment status. The non-native flora of the islands studied consists of 154 species, 105 of which are established on at least one island. Regression tree analysis revealed that the main determinants of both total non-native and established non-native species richness are the variables linked to tourist development and the sprawl of artificial surfaces. The variation partitioning approach shows that the variation in total non-native flora composition is driven above all by environmental variables, whereas when established taxa are considered, human-mediated factors account for most of the explained variation. Mediterranean islands have recently experienced changes in the type and intensity of human activities, leading to a decline in rural landscape and the development of tourist facilities. Our results indicate that tourism and the increase in the extension of artificial surfaces are also the main drivers of non-native plant species occurrence and establishment. These factors outweigh the effects of abiotic variables such as climate and lithology in determining the number of non-native species, whether it be total or established, as well as variations in the composition of established species. High human pressure and ornamental introductions associated with tourist activities appear to be the key elements that need to be considered when planning strategies aimed at managing plant invasions in Mediterranean islands.

Genetic structure and introgression in riparian populations of Populus alba L.

Abstract White poplar (Populus alba) is a widespread species of the northern hemisphere. Introgressed populations or hybrid zones with the related species of the European aspen (Populus tremula) have been suggested as potential venues for the identification of functionally important variation for germplasm conservation, restoration efforts and tree breeding. Data on the genetic diversity and structure of introgressed P. alba are available only for sympatric populations from central Europe. Here, clonality, introgression and spatial genetic patterns were evaluated in three riparian populations of P. alba along the Ticino, Paglia‐Tevere and Cesano river drainages in Italy. Samples of all three populations were typed for five nuclear microsatellite markers and 137 polymorphic amplified fragment length polymorphisms. Microsatellite‐based inbreeding co‐efficients (F IS) were significantly positive in all three populations. Genetic diversity was consistently highest in Ticino, the population with the highest level of introgression from P. tremula. Population differentiation (F ST) was low between the Ticino valley in northern Italy and the Cesano valley in central Italy and between the central Italian populations of Cesano and Paglia‐Tevere, consistent with a role of the Appenine mountain range as a barrier to gene flow between adjacent drainage areas. Introgression was not the primary determinant of within‐population spatial genetic structure (SGS) in the studied populations.

Plant invaders in European and Mediterranean inland waters: profiles, distribution, and threats

Abstract The present brief and selective review focuses on a set of invasive alien aquatic plant species (notably the 21 listed by the European and Mediterranean Plant Protection Organisation, i.e., five A2 species, 11 invasive, 2 in the Observation List, 1 in the Alert List, 2 noteworthy), on definitions in use, on introduction pathways, and ecological effects, although the social and economic effects of freshwater invaders can be diverse and substantial. Legislation, literature, and scientific research dealing with plant invasions in inland waters often merge different types of invasive plant species and invaded habitats, as there are some overlappings between the concepts of macrophyte, aquatic plants, inland waters, wetlands, and riparian habitats. Freshwater ecosystems may well be the most endangered ecosystems in the world. Countless aquatic plants have been introduced in Europe and in the Mediterranean region, either deliberately because they were thought to be ornamental or otherwise desirable, or accidentally as releases from aquaria, dumping from water gardens, or contaminants. The escape of alien plants from managed environments is also frequent. Aquatic invaders have strong negative impacts on native biodiversity and many different impacts are reported in the literature for Europe and worldwide. Plant invasions in inland waters may be considered as one of a syndrome of factors that characterize degraded aquatic ecosystems. There are a number of risk analysis, impact assessment, ranking systems, decision trees, and prioritization methods available throughout Europe and the EPPO region, but the lack of a common framework for assessing the risks posed by invasive alien aquatic species is seen as a key gap. Successful management should include prevention, habitat restoration, dedicated risk assessment, prioritization, awareness raising, dedicated legislation, collaboration between different stakeholders, development of codes of conduct with the horticultural sector, and prohibiting the sale, purchase, and intentional release of some species in the wild. Actions to protect the European and Mediterranean inland waters from plants invasions are urgently needed and could provide overarching benefits.

Impacts of alien plants and man‐made disturbance on soil‐growing bryophyte and lichen diversity in coastal areas of Sardinia (Italy)

Abstract Seventy phytosociological relevés were performed in 1 m × 1 m plots at 14 study sites spread along sandy shores in northern and southern Sardinia (Italy). The plots were selected in different habitat types (open dunes, native Juniperus woodlands, maquis, and plantations with Acacia, Eucalyptus and Pinus) according to a stratified sampling method in order to investigate impacts deriving from different levels of Carpobrotus spp. cover, dry litter from exotic trees, and other disturbance types. The quantile regression and logistic regression analyses revealed that the reduction in the amount of bryophyte and lichen cover on sand dunes of the study area is caused either by a high cover of Carpobrotus spp. mats or by a high cover of dry exotic litter in dense, unmanaged or poorly managed forest plantations. Additional detrimental effects are often driven by other kinds of man‐made disturbances. Forest management in the coastal areas of Sardinia should be gradually modified to take into account the conservation of bryophytes and lichens. Some of the biological indicators used are quite widespread in the Mediterranean coastal habitats or are exclusively associated with sand dunes; therefore, they can also be conveniently used as indicators of biological impacts in other countries or islands of the same biogeographical region.