How do introduction characteristics influence the invasion success of Mediterranean alien plants?

Invasive plant species are becoming increasingly widespread following accelerated anthropogenic activity in the Mediterranean region. Humans have played a central role in the expansion process, and it is important to incorporate such considerations into management plans. Using generalized linear models, our first aim was to describe how the invasion success of 862 prominent alien plant species on Mediterranean islands is related to characteristics of the introduction process: introduction frequency, date and region of origin, range size and purpose of import. The importance of each was measured by the numbers of species present and their average invasiveness. The main findings were: (a) accidental imports and ornamentals accounted for a high proportion of all aliens, although neither group had particularly high average invasiveness; (b) introduction frequency had a comparatively modest influence, with the most commonly-introduced species naturalized only three times more widely than those rarely-introduced; (c) rates of species introduction appear to have increased dramatically in the last century, although aliens which have been present in the region for more than 200 years were most widespread, indicating that it may be centuries before some species fill their potential range; (d) there were small tendencies for successful invaders to originate in the Neotropics or in regions with Mediterranean climate biomes and to have large range sizes. Our second aim was to determine whether the number or average invasiveness of species introduced via a given pathway had the most influence on the overall probability of invasion on a given island. An elasticity analysis suggested that the number of species was substantially the best predictor of the two. This finding arises largely because invasion events are rare and remain unpredictable, and has significant implications for assessing invasion risk. We discuss how substantial sources of error and intrinsic variability in invasiveness within species groups limit the potential for developing accurate risk models.     

The historical role of agriculture and gardening in the introduction of alien plants in the western Mediterranean

The aim of this study is to clarify the role of gardening and agriculture in the introduction of alien vascular plants to the western Mediterranean Basin during the last century. The main country in which this study is based is Spain. However, other countries in the Western Mediterranean Basin also form part of it. These are Portugal, France, Italy and Morocco. Three percentages were calculated. Firstly, the percentage of alien flora was calculated. Then, we calculated the percentages of species introduced as either a consequence of gardening or of agriculture. The figures were based on thirty-four floristic studies carried out in the western Mediterranean Basin between 1904 and 2006. A regression model, in which the explanatory variable is time and the dependent variables are the three percentages mentioned above, was drawn up with the data obtained. The results show the existence of positive correlation between time and the percentages of total exotic flora and the species introduced through gardening (R = 0.64 and 0.56, respectively) and the negative correlation between time and the percentage of alien species introduced through agriculture (R = −0.43). During the last century, the roles of agriculture and gardening as sources of the introduction of exotic flora to the western Mediterranean Basin were reversed. However, in the areas where the socio-economic scene is more predominantly agrarian, the role of gardening is still limited.     

Ecological traits and environmental affinity explain Red Sea fish introduction into the Mediterranean

Alien species are considered one of the prime threats to biodiversity, driving major changes in ecosystem structure and function. Identifying the traits associated with alien introduction has been largely restricted to comparing indigenous and alien species or comparing alien species that differ in abundance or impact. However, a more complete understanding may emerge when the entire pool of potential alien species is used as a control, information that is rarely available. In the eastern Mediterranean, the marine environment is undergoing an unparalleled species composition transformation, as a flood of aliens have entered from the Red Sea following the opening of the Suez Canal in 1869. In this study, we compile data on species traits, geographical distribution, and environmental affinity of the entire pool of reef‐associated fish species in the Red Sea and more generally across the Indo‐Pacific. We use this extensive data to identify the prime characteristics separating Red Sea species that have become alien in the Mediterranean from those that have not. We find that alien species occupy a larger range of environments in their native ranges, explaining their ability to colonize the seasonal Mediterranean. Red Sea species that naturally experience high maximum temperatures in their native range have a high probability of becoming alien. Thus, contrary to predictions of an accelerating number of aliens following increased water temperatures, hotter summers in this region may prevent the establishment of many alien species. We further find that ecological trait diversity of alien species is substantially more evenly spaced and more divergent than random samples from the pool of Red Sea species, pointing at additional processes, such as competition, promoting ecological diversity among alien species. We use these results to provide a first quantitative ranking of the potential of Red Sea species to become established in the eastern Mediterranean.     

Alien species in the Mediterranean Sea—which, when, where, why?

A critical evaluation of more than 2,200 publications, some dating back to the late 1800s, established the presence, and traced the spatio-temporal spread, of 558 alien metazoan species in the Mediterranean Sea. The majority of aliens in the eastern Mediterranean entered through the Suez Canal, whereas mariculture and shipping are powerful means of introduction in the northwestern Mediterranean and in the Adriatic Sea. Most aliens are thermophilic species. The possible causes for the epic scale of invasion in the Mediterranean Sea are discussed. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: J. Davenport, G. Burnell, T. Cross, M. Emmerson, R. McAllen, R. Ramsay & E. Rogan Challenges to Marine Ecosystems     

Alien molluscan species established along the Italian shores: an update,with discussions on some?Mediterranean “alien species” categories

The state of knowledge of the alien marine Mollusca in Italy is reviewed and updated. Littorina saxatilis (Olivi, 1792), Polycera hedgpethi Er. Marcus, 1964 and Haminoea japonica Pilsbry, 1895are here considered as established on the basis of published and unpublished data, and recent records of the latter considerably expand its known Mediterranean range to the Tyrrhenian Sea. COI sequences obtained indicate that a comprehensive survey of additional European localities is needed to elucidate the dispersal pathways of Haminoea japonica.Recent records and interpretation of several molluscan taxa as alien are discussed both in light of new Mediterranean (published and unpublished) records and of four categories previously excluded from alien species lists. Within this framework, ten taxa are no longer considered as alien species, or their records from Italy are refuted. Furthermore, Trochocochlea castriotae Bellini, 1903 is considered a new synonym for Gibbula albida (Gmelin, 1791). Data provided here leave unchanged as 35 the number of alien molluscan taxa recorded from Italy as well as the percentage of the most plausible vectors of introduction, but raise to 22 the number of established species along the Italian shores during the 2005–2010 period, and backdate to 1792 the first introduction of an alien molluscan species (Littorina saxatilis) to the Italian shores.     

FIRST RECORD OF GRACILARIA VERMICULOPHYLLA (GRACILARIALES,RHODOPHYTA) IN THE PO DELTA LAGOONS,MEDITERRANEAN SEA (ITALY)1

Molecular outcomes led us to report the first occurrence of the invasive alien species Gracilaria vermiculophylla (Ohmi) Papenf. in the Mediterranean Sea. This species was recorded for the first time in the Po Delta lagoons in May and October 2008, probably introduced by the importation of the Manila clam Tapes philippinarum. At present, G. vermiculophylla is spread only near some clam‐farming areas, but its diffusion is expected to increase with the colonization of other lagoons where aquaculture is spread, as already observed for other alien species such as Agardhiella subulata and Solieria filiformis. The present study supplies further information on the morphology of this species, the ecological characteristics of the colonized areas, and the most probable introduction vector, confirming that the species spreading occurs in eutrophic and turbid coastal systems.     

The rise and fall of an alien: why the successful colonizer Littorina saxatilis failed to invade the Mediterranean Sea

Understanding what determines range expansion or extinction is crucial to predict the success of biological invaders. We tackled this long-standing question from an unparalleled perspective using the failed expansions in Littorina saxatilis and investigated its present and past habitat suitability in Europe through Ecological Niche Modelling. This intertidal snail is a typically successful Atlantic colonizer and the earliest confirmed alien species in the Mediterranean Sea, where, however, it failed to thrive despite its high dispersal ability and adaptability. We explored the environmental constraints affecting its biogeography, identified potential glacial refugia in Europe that fuelled its post-glacial colonisations and tested whether the current gaps in its distribution are linked to local ecological features. Our results suggested that L. saxatilis is unlikely to be a glacial relict in the Mediterranean basin. Multiple Atlantic glacial refugia occurred in the Last Glacial Maximum, and abiotic environmental features such as salinity and water temperature have influenced the past and current distributions of this snail and limited its invasion of the Mediterranean Sea. The snail showed a significant overlap in geographic space and ecological niche with Carcinus maenas, the Atlantic predator, but distinct from Pachygrapsus marmoratus, the Mediterranean predator, further pointing to Atlantic-like habitat requirements for this species. Abiotic constrains during introduction rather than dispersal abilities have shaped the past and current range of L. saxatilis and help explaining why some invasions have not been successful. Our findings contribute to clarifying the processes constraining or facilitating shifts in species’ distributions and biological invasions.

     

Altitudinal distribution of alien plant species in the Swiss Alps

In summer 2003 we recorded the presence and abundance of alien plant species at 232 sites (107 railway stations and 125 road sites) along mountain passes in the Swiss Alps. The altitudinal distribution of species was related to the current abundance of the species in Switzerland and time since introduction. A total of 155 alien taxa were recorded. Numbers of species per site declined exponentially with altitude, and only a few species were found in the alpine zone (>2000 m). In contrast, species richness among comparable native taxa appeared to be nearly independent of altitude over the range investigated. Maximum altitude reached by alien species was related positively to both total area occupied in Switzerland and to time since introduction. A comparison of the results with earlier records suggests that many species, particularly those previously restricted to low or intermediate altitudes, have advanced their altitudinal limits over the past few decades. Various hypotheses are presented to explain the declining abundance of alien species with altitude: low-altitude filter effects, low propagule pressure, and genetic swamping of peripheral populations at higher altitudes. However, at present we do not have sufficient evidence to determine the relative importance of these effects. We conclude that invasion into mountain areas such as the Swiss Alps tends to proceed rather slowly, though the process may be accelerated by climatic warming. For this reason, further research to investigate the processes determining how plants invade mountain areas is urgently needed. And more generally, investigations into the distribution of alien species along strong altitudinal gradients may provide valuable insights into the mechanisms driving the spread of alien organisms.     

Four arguments why so many alien species settle into estuaries, with special reference to the German river Elbe

In one of the largest European rivers, the Elbe, from its source in the Czech Republic to the German North Sea, 31 alien macrozoobenthic species have been recorded in total. Most of these species have been introduced by shipping activities. With a total number of 21 species, many of the established aliens occur—partly exclusively—in the brackish area of the Elbe estuary. In order to explain this observed settlement characteristic, four main arguments come into consideration: (1) estuaries with intensive international shipping have a higher potential infection rate than other aquatic zones; (2) brackish water species have, due to specific physiological characteristics, a better chance of being transported alive than euhaline or freshwater species and they also probably have a higher perennation and establishment potential after release; (3) brackish waters have the greatest natural ‘indigenous species minimum’, so that more alien species can potentially establish; and (4) salt-tolerant limnetic alien species introduced into inland water reached the coast at first in the estuaries. It seems that the combination of brackish water with its unsaturated ecological niches and intensive international ship traffic has the highest potential infection rate for aquatic systems with alien macrozoobenthic species. And, estuaries are subjected to a two-sided invasion pressure by alien species, via the ocean (mainly shipping) and via inland waters (mainly shipping canal construction). The identification of such patterns is an important prerequisite for the development of a forward-looking alien monitoring and management strategy.     

Bark beetles and pinhole borers (Curculionidae, Scolytinae, Platypodinae) alien to Europe

Invasive bark beetles are posing a major threat to forest resources around the world. DAISIE's web-based and printed databases of invasive species in Europe provide an incomplete and misleading picture of the alien scolytines and platypodines. We present a review of the alien bark beetle fauna of Europe based on primary literature through 2009. We find that there are 18 Scolytinae and one Platypodinae species apparently established in Europe, from 14 different genera. Seventeen species are naturalized. We argue that Trypodendron laeve, commonly considered alien in Europe, is a native species; conversely, we hypothesize that Xyleborus pfeilii, which has always been treated as indigenous, is an alien species from Asia. We also point out the possibility that the Asian larch bark beetle Ips subelongatus is established in European Russia. We show that there has been a marked acceleration in the rate of new introductions to Europe, as is also happening in North America: seven alien species were first recorded in the last decade.We present information on the biology, origins, and distributions of the alien species. All but four are polyphagous, and 11 are inbreeders: two traits which increase invasiveness. Eleven species are native to Asia, six to the Americas, and one is from the Canary Islands. The Mediterranean is especially favorable for invasives, hosting a large proportion of the aliens (9/19). Italy, France and Spain have the largest numbers of alien species (14, 10 and 7, respectively). We point out that the low numbers for at least some countries is likely due to under-reporting.Finally, we discuss the difficulties associated with identifying newly invasive species. Lack of good illustrations and keys hinder identification, particularly for species coming from Asia and Oceania.     

On the occurrence of the fireworm Eurythoe?complanata complex (Annelida,Amphinomidae) in the Mediterranean Sea with an updated revision of the alien Mediterranean amphinomids

The presence of two species within the Eurythoe complanata complex in the Mediterranean Sea is reported, as well as their geographical distributions. One species, Eurythoe laevisetis, occurs in the eastern and central Mediterranean, likely constituting the first historical introduction to the Mediterranean Sea and the other, Eurythoe complanata, in both eastern and Levantine basins. Brief notes on their taxonomy are also provided and their potential pathways for introduction to the Mediterranean are discussed. A simplified key to the Mediterranean amphinomid genera and species of Eurythoe and Linopherus is presented plus an updated revision of the alien amphinomid species reported previously from the Mediterranean Sea. A total of five exotic species have been included; information on their location, habitat, date of introduction and other relevant features is also provided.     

江西省外来植物现状

[目的] 完善江西省外来植物本底调查数据,为江西省外来入侵植物的预防和预警机制构建提供依据。[方法] 通过文献、标本研究以及野外实地调查,对江西外来植物种类、原产地、生活型、引入途径、引入时间、在各地级市的分布情况以及外来入侵植物的种类进行分析、探讨。[结果] 江西省外来植物有106科407属599种：菊科最多（85种,14.19%）,其次是豆科（52种,8.68%）,10种以下的科有91个,占总科的85.85%;主要源于美洲（49.25%）,其次是亚洲（20.80%）,欧洲第三（13.81%）,地中海和大洋洲较少;外来植物中草本植物种类最多,乔木次之;大部分被用于观赏、饲料、药物等有意引入。江西省的外来入侵植物有53科168属267种;九江、赣州和南昌由于地理位置优越及经济地位突出被入侵较严重。[结论] 江西省外来植物种类丰富,引入途径多样,入侵物种较多。建议构建风险评价系统,加强外来植物监管监测,科学引种;制订和更新外来植物防控的具体政策法规;同时,加大入侵危害宣传,及时防控和阻止外来植物逃逸入侵对本省生态环境造成侵害。     

A fast snail’s pace: colonization of Central Europe by Mediterranean gastropods

Central Europe faces an influx of terrestrial invertebrates from more southerly regions, in particular from the Mediterranean. This process is particularly noticeable among terrestrial gastropods. In the last 30 years, the number of non-native land snail species found outdoors in the Czech Republic increased from 5 to 15 (8% of all species); more than half of these have presumed Mediterranean origin. The trend has accelerated recently; seven new alien species (six Mediterranean) have been recorded since 2000. As the most recent example, we report the finding of a large helicid species from the South-East Mediterranean, Helix lucorum. A well-established overwintering population was found in Prague, over 400 km from the nearest known localities of the species. This trend of newly established snails corresponds with increases in average temperatures as well as the intensity of foreign trade in the past six decades, suggesting a synergistic effect of both climatic conditions and socioeconomic factors.     

Influence of past land use and current human disturbance on non-native plant species on small Italian islands

Biological invasions are regarded as one of the main drivers of habitat degradation in island ecosystems. Mediterranean islands have been subjected to a high degree of land conversion over the past 60 years, resulting in a massive reduction in the amount of rural land and the sprawl of tourist activities. The aims of this paper are to evaluate the current level of invasion of alien plant species in semi-natural vegetation types that have developed after the abandonment of agriculture and to analyze the relationships between non-native species, native flora, and environmental characteristics. Two Italian islands (Ponza and Ventotene) were surveyed using a random-stratified sampling. The occurrence and relative cover of alien plant species were compared and separate analyses were performed for the native flora. Abundance patterns of both native and alien species were then studied in the light of the environmental and anthropogenic features. Although we found that some non-native species are extremely widespread, their relative cover at the plot level is low. Permutational Multivariate Analysis of Variance and Indicator Species Analysis revealed dissimilarities in the native species composition, while Mann–Whitney and Kruskal–Wallis tests showed differences in the ecological requirements (moisture, soil reaction, and nitrogen) of the native species pool. Canonical Correspondence Analysis pointed to the importance of the proximity to agricultural areas, human disturbance, and past land management, particularly residual terraces, in determining the difference between plant communities on the two islands. The results of our study suggest that traditional forms of agriculture may represent a key element for countering the establishment and spread of non-native plants in Mediterranean areas.     

How strongly do interactions with closely‐related native species influence plant invasions? Darwin's naturalization hypothesis assessed on Mediterranean islands

Aim Recent works have found the presence of native congeners to have a small effect on the naturalization rates of introduced plants, some suggesting a negative interaction (as proposed by Charles Darwin in The Origin of Species), and others a positive association. We assessed this question for a new biogeographic region, and discuss some of the problems associated with data base analyses of this type. Location Islands of the Mediterranean basin. Methods Presence or absence of congeners was assessed for all naturalized alien plants species at regional, local and habitat scales. Using general linear models, we attempted to explain the abundance of the species (as measured by the number of islands where recorded) from their congeneric status, and assessed whether the patterns could be alternatively accounted for by a range of biological, geographical and anthropogenic factors. A simulation model was also used to investigate the impact of a simple bias on a comparable but hypothetical data set. Results Data base analyses addressing Darwin's hypothesis are prone to bias from a number of sources. Interaction between invaders and congenerics may be overestimated, as they often do not co‐occur in the same habitats. Furthermore, intercorrelations between naturalization success and associated factors such as introduction frequency, which are also not independent from relatedness with the native flora, may generate an apparent influence of congenerics without implying a biological interaction. We detected no true influence from related natives on the successful establishment of alien species of the Mediterranean. Rarely‐introduced species tended to fare better in the presence of congeners, but it appears that this effect was generated because species introduced accidentally into highly invasible agricultural and ruderal habitats have many relatives in the region, due to common evolutionary origins. Main conclusions Relatedness to the native flora has no more than a marginal influence on the invasion success of alien plants in the Mediterranean, although apparent trends can easily be generated through artefacts of the data base.     

The new threat to Italian inland waters from the alien crayfish “gang”: the Australian Cherax destructor Clark, 1936

Biological invasions inflict damage to the ecology, economy, and human health, and pose serious threats to the native communities. Among the many invasive taxa, crayfish have attracted much attention by scientists and policy makers. Recently, an established population of an alien species of crayfish, new for Italy, was found in the Natural Preserve of “Laghi di Ninfa” (central Italy). Based on morphological and genetic evidence (molecular barcoding of COI and 16S rDNA), we classified it at subspecies level as the Parastacidae Cherax destructor destructor, native to Australia. Its introduction possibly occurred at the end of the 1980s but the species seems to be still confined in the preserve. The low temperature of the adjacent waters may be a barrier against its natural spreading but not against its intentional translocation into natural waters by man. Because of the invasive history of C. destructor, eradication of this population is urgent and still economically profitable. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: Pierluigi Viaroli     

The tropical caprellid amphipod Paracaprella pusilla: a new alien crustacean in the Mediterranean Sea

Paracaprella pusilla Mayer (Fauna und Flora des Golfes von Neapel 17:1–55, 1890), originally described from Brazil, is one of the most abundant caprellid amphipod species in tropical and subtropical seas around the world. During a survey of caprellid amphipods from marinas along the Balearic Island (Western Mediterranean Sea) carried out between November 2011 and August 2012, we found two established populations of P. pusilla in Mallorca and Ibiza, respectively. So far, its occurrence in European waters was reported only from southwestern Spain in 2010. This record represents a northward range expansion of the species’ distribution, which is found for the first time in the Mediterranean. This is also the first record of the genus Paracaprella in the Mediterranean Sea. The most probable introduction vector was ship fouling. We also found the invasive caprellid Caprella scaura Templeton (Trans Entomol Soc Lond 1:185–198, 1836) in Mallorca and Menorca, which is recorded for the first time in the Balearic Islands, confirming its rapid expansion along the Mediterranean. When comparing reproductive traits between both alien species, we found that P. pusilla has a higher fecundity than C. scaura for the same female size. Taking into account this evidence, the species may be expected to appear in other Mediterranean and adjacent areas.     

A global analysis of the determinants of alien geographical range size in birds

Aim Determining the causes of range size variation in the distributions of alien species is important for understanding the spread of invasive species. Factors influencing alien range size have been explored for some species at a regional level, but to date there has been no global analysis of an entire class. Here, we present such an analysis for birds, testing for the effects of introduction event, location and species‐level variables on alien range sizes. Location Global. Methods We used a novel dataset on the global distributions of alien bird species to test for relationships between alien range size and colonization pressure, residence time, extent of the global climatic niche, native range size, body mass and specialization, using a statistical approach based on phylogenetic generalized least squares models. We performed this analysis globally, and for separate biogeographical realms. Results Approximately half of the variation in alien bird range size is explained by colonization pressure in univariate analysis. We identified consistent effects of higher colonization pressure at global and realm levels, as well as support for effects of native range size and residence time. We found less support for effects of body mass, specialization or extent of the global climatic niche on alien range size. Main conclusions Alien bird range sizes are generally small relative to their native range sizes, and many are continuing to expand. Nevertheless, current variation is predictable, most strongly by the event‐level factor of colonization pressure. Whether a species is widespread is a better predictor of alien range size than whether a species could be widespread (estimated by global climatic niche extent), while we also find effects of residence time on alien range size. These relationships may help to identify those alien species that are more likely to spread and hence have greater environmental and economic impacts where they have been introduced.     

Little,but increasing evidence of impacts by alien bryophytes

Based on data of bryophyte invasions into 82 regions on five continents of both hemispheres, we aim here at a first comprehensive overview of the impacts that bryophytes may have on biodiversity and socio-economy. Of the 139 bryophytes species which are alien in the study regions seven cause negative impacts on biodiversity in 26 regions, whereas three species cause negative impacts on socio-economic sectors in five regions. The vast majority of impacts stem from anecdotal observations, whereas only 14 field or experimental studies (mostly on Campylopus introflexus in Europe) have quantitatively assessed the impacts of an alien bryophyte. The main documented type of impact on biodiversity is competition (8 alien bryophytes), with native cryptogams being most affected. In particular, C. introflexus (9 regions) and Pseudoscleropodium purum (7 regions) affect resident species composition. The few socio-economic impacts are caused by alien bryophytes which form dense mats in lawns and are then considered a nuisance. Most negative impacts on biodiversity have been recorded in natural grasslands, forests, and wetlands. Impacts of alien bryophytes on biodiversity and socio-economy are a recent phenomenon, with >85 % of impacts on biodiversity, and 80 % of impacts on socio-economy recorded since 1990. On average, 40 years (impacts on biodiversity) and 25 years (impacts on socio-economy) elapsed between the year a bryophyte species has been first recorded as alien in a region and the year impacts have been recorded first. Taking into account the substantial time lag between first record and first recorded impact in a region, it seems to be likely that the currently moderate impacts of alien bryophytes will continue to increase. As quantitative studies on impacts of alien bryophytes are rare and restricted to few environments and biogeographic regions, there is a need for addressing potential impacts of alien bryophytes in yet understudied settings.