Undaria pinnatifida (Laminariales,Phaeophyta) 12 years after its introduction into the Atlantic Ocean

Since its first introduction into the Atlantic for farming purposes in 1983, the brown alga Undaria pinnatifida has been found at several sites along the European coasts. The present study deals with an ecological experiment that was performed at the island of Ushant (Brittany, France) in order to check the ability of Undaria to recruit to a rocky substratum in situ. On the one hand, the authors were unsuccessful in getting Undaria to colonize a previously denuded area of shore, which was subsequently colonized by other native species, a result that may speak for a competition between Undaria and the local kelps. On the other hand, evidence is presented that Undaria pinnatifida has become a member of the Atlantic flora for good. The sporophytes can be found growing from low water mark of neap tide (+1.5 m) down to at least –15 m below lower water mark of spring tide. Its propagation is thought to occur in at least two ways: either step by step from cultivation sites along the coasts, or from the hulls of ships sailing from harbour to harbour. The preference of Undaria for settlement is on artificial structures.     

Fronts,jumps and secondary introductions suggested as different invasion patterns in marine species,with an increase in spread rates over time

Not all introduced (invasive) species in a region will spread from a single point of introduction. Long-distance dispersal or further introductions can obscure the pattern of spread, but the regional importance of such processes is difficult to gauge. These difficulties are further compounded when information on the multiple scale process of invasive species range expansion is reduced to one-dimensional estimates of spread (e.g. km yr⁻¹). We therefore compared the results of two different metrics of range expansion: maximum linear rate of spread and accumulation of occupied grid squares (50 × 50 km) over time. An analysis of records for 54 species of introduced marine macrophytes in the Mediterranean and northeast Atlantic revealed cases where the invasion process was probably missed (e.g. Atlantic Bonnemaisonia hamifera) and suggested cases of secondary introductions or erratic jump dispersal (Dasysiphonia sp. and Womersleyella setacea). A majority of species analysed showed evidence for an accumulation of invaded sites without a clear invasion front. Estimates of spread rate are increasing for more recent introductions. The increase is greater than can be accounted for by temporally varying search effort and implies a historical increase in vector efficiency and/or a decreased resistance of native communities to invasion.     

rbcL sequences reveal multiple cryptic introductions of the Japanese red alga Polysiphonia harveyi

In Europe, the last 20 years have seen a spectacular increase in accidental introductions of marine species, but it has recently been suggested that both the actual number of invaders and their impacts have been seriously underestimated because of the prevalence of sibling species in marine habitats. The red alga Polysiphonia harveyi is regarded as an alien in the British Isles and Atlantic Europe, having appeared in various locations there during the past 170 years. Similar or conspecific populations are known from Atlantic North America and Japan. To choose between three competing hypotheses concerning the origin of P. harveyi in Europe, we employed rbcL sequence analysis in conjunction with karyological and interbreeding data for samples and isolates of P. harveyi and various congeners from the Pacific and North Atlantic Oceans. All cultured isolates of P. harveyi were completely interfertile, and there was no evidence of polyploidy or aneuploidy. Thus, this biological species is both morphologically and genetically variable: intraspecific rbcL divergences of up to 2.1% are high even for red algae. Seven rbcL haplotypes were identified. The four most divergent haplotypes were observed in Japanese samples from Hokkaido and south-central Honshu, which are linked by hypothetical 'missing' haplotypes that may be located in northern Honshu. These data are consistent with Japan being the centre of diversity and origin for P. harveyi. Two non-Japanese lineages were linked to Hokkaido and Honshu, respectively. A single haplotype was found in all North Atlantic and Mediterranean accessions, except for North Carolina, where the haplotype found was the same as that invading in New Zealand and California. The introduction of P. harveyi into New Zealand has gone unnoticed because P. strictissima is a morphologically indistinguishable native sibling species. The sequence divergence between them is 4-5%, greater than between some morphologically distinct red algal species. Two different types of cryptic invasions of P. harveyi have therefore occurred. In addition to its introduction as a cryptic sibling species in New Zealand, P. harveyi has been introduced at least twice into the North Atlantic from presumed different source populations. These two introductions are genetically and probably also physiologically divergent but completely interfertile.     

Introduced or glacial relict? Phylogeography of the cryptogenic tunicate Molgula manhattensis (Ascidiacea,Pleurogona)

Aim The tunicate Molgula manhattensis has a disjunct amphi‐Atlantic distribution and a recent history of world‐wide introductions. Its distribution could be the result of regional extinctions followed by post‐glacial recolonization, or anthropogenic dispersal. To determine whether the North Atlantic distribution of M. manhattensis is natural or human‐mediated, we analysed mtDNA cytochrome c oxidase subunit I (COI) sequence variation in individuals from cryptogenic and introduced ranges. Location North Atlantic Europe and America; Black Sea; San Francisco Bay; Osaka Bay. Methods Nuclear 18S rDNA sequences were used to resolve phylogenetic relationships and mtDNA COI sequences for phylogeographic analyses. Results Phylogenetic analyses confirmed that M. manhattensis and M. socialis, which are frequently confused, are distinct species. MtDNA haplotype diversity was nearly three times higher with deeper relationships among haplotypes on the North‐east American coast than in Europe. Diversity declined from south to north in America but not in Europe. In areas of known introductions (Black Sea, Japan, San Francisco Bay), M. manhattensis showed variable levels of haplotype diversity. Medium‐to‐high‐frequency haplotypes originating from the North‐west Atlantic were present in two locations of known introductions, but not in Europe. Private haplotypes were found on both sides of the Atlantic and in introduced populations. The mismatch distribution for the North‐east Atlantic coast indicates a recent expansion. Main conclusions Molgula manhattensis is native in North‐east America. However, whether it was introduced or is native to Europe remains equivocal. Additional sampling might or might not reveal the presence of putative private European haplotypes in America. The low European diversity may be explained by low effective population size and a recent expansion, or by low propagule pressure of anthropogenic introduction. Absence of medium‐to‐high‐frequency American haplotypes in Europe may be the result of exclusive transport from southern ports, or long‐term residence. These arguments are ambiguous, and M. manhattensis remains cryptogenic in Europe.     

Introduced marine species of the North Sea coasts

About 80 non-indigenous species are assumed to have been introduced into the North Sea by transoceanic shipping and aquaculture. The number is certainly underestimated as most small organisms received insufficient attention at the species level. Also, the seafaring tradition of the North Sea countries is much longer than our biological surveys are. Most exotic invertebrates originate from the western Atlantic and were introduced by shipping, while most algae stem from the Pacific and came with the introduced oysters. A peak of newcomers was observed in the 1970s. Most of the arrivals became established in brackish environments, at harbor sites and in the vicinity of oyster farms, fouling on hard substrates or living as epibionts. A few live in sediments, are holoplanktonic or are parasites. At the open coast, approximately 6% of the macrobenthic species are exotics, while in estuaries their share is up to 20%. Most exotics have been encountered in the southern North Sea first, and many did not spread further north. About 25% of the established non-natives are widespread and attain locally high abundances. As a consequence, some inshore habitats are entirely dominated by exotics. The overall effect on the ecosystem seems to be more additive than one of displacement. This suggests that the coastal biota of the North Sea are quite capable of accommodating newcomers. However, this is no guarantee that the next introduced species may not cause severe ecological change or economic harm. There is a need to minimize the risk of unintentional introductions by ballast water treatment and by adhering to quarantine procedures in aquaculture. Current research on exotics in the North Sea is regarded as inadequate for proper evaluation and management requirements.     

Introductions and developments of oysters in the North Sea area: a review

To replenish the exploited native stocks ofOstrea edulis, imports from almost all European coasts have arrived in the North Sea, particularly in the Dutch Oosterschelde estuary. The American oysterCrassostrea virginica and the Portuguese oysterC. angulata have also been imported into the North Sea several times. However, only the introductions of various genetic strains of the Pacific oysterC. gigas have been of lasting success. Spat from British Columbia (Canada) was first imported to the Oosterschelde in 1964. First spatfalls in this area took place in the warm summers of 1975 and 1976. Further larval outbursts occurred in 1982 and 1989, and good settlements took place on culture plots as well as along the dikes of the Oosterschelde. Since 1977 no more cupped oysters have been imported from overseas. The population maintained itself and was able to spread in a northern direction along the Dutch North Sea coast. In Britain, hatchery-producedC. gigas were transferred to several sites, in the 1970s including the British North Sea coast. Here, occurrence in the wild seems to be rather limited up to now. Into the German Wadden Sea,C. gigas larvae and spat from a Scottish hatchery have been introduced since 1971, as were medium-sized oysters from a variety of European sources in the 1980s. Strong spatfalls on intertidal mussel beds in the northern German Wadden Sea occured in 1991 and 1994. For the introductions ofC. gigas along the west European coasts, precautionary measures to minimize unintentional transfers of associated organisms, as recommended by the ICES Code of Practice in 1994, came too late.     

Evidence for multiple introductions of Centaurea stoebe micranthos (spotted knapweed,Asteraceae) to North America

Invasive species’ success may depend strongly on the genetic resources they maintain through the invasion process. We ask how many introductions have occurred in the North American weed Centaurea stoebe micranthos (Asteraceae), and explore whether genetic diversity and population structure have changed as a result of introduction. We surveyed individuals from 15 European native range sites and 11 North American introduced range sites at six polymorphic microsatellite loci. No significant difference existed in the total number of alleles or in the number of private alleles found in each range. Shannon–Weaver diversity of phenotype frequencies was also not significantly different between the ranges, while expected heterozygosity was significantly higher in the invasive range. Population structure was similar between the native range and the invasive range, and isolation by distance was not significant in either range. Traditional assignment methods did not allocate any North American individuals to the sampled European populations, while Bayesian assignment methods grouped individuals into nine genetic clusters, with three of them shared between North America and Europe. Invasive individuals tended to have genetically admixed profiles, while natives tended to assign more strongly to a single cluster. Many North American individuals share assignment with Romania and Bulgaria, suggesting two separate invasions that have undergone gene flow in North America. Samples from three other invasive range sites were genetically distinct, possibly representing three other unique introductions. Multiple introductions and the maintenance of high genetic diversity through the introduction process may be partially responsible for the invasive success of C. stoebe micranthos.     

A risk assessment of aquarium trade introductions of seaweed in European waters

Aquaculture and maritime traffic have been identified as the main vectors for introductions of alien marine species. Except for one notorious case of Caulerpa taxifolia, the role of aquarium trade towards the introduction of alien seaweeds has been largely unassessed. Here, we address the risk of accidental release of seaweed species from the aquarium trade market in European waters. We assessed the importance and diversity of seaweed species in the European online aquarium retail circuit. Our web survey revealed more than 30 genera available for online sale into Europe, including known introduced and invasive species. A second aspect of the study consisted in sampling algal diversity found in aquaria. While allowing direct and accurate identification of the specimens, this approach was targeting not only ornamental species, but also seaweeds that may be accidentally present in the aquarium circuit. By DNA-barcoding we identified no less than 134 taxa, 7 of which are flagged as introduced in Europe and 5 reported as invasive. Climate envelope models show that at least 23 aquarium species have the potential to thrive in European waters. As expected by the tropical conditions in most aquaria, southern Atlantic regions of Europe and the Mediterranean are the most vulnerable towards new introductions. Further predictions show that this risk will increase and shift northwards as global warming proceeds. Overall our data indicate that aquarium trade poses a potential risk of new seaweed introductions, and calls for a cautious approach.     

Alien algae <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> causes habitat loss for rocky reef fishes in north Patagonia

Since the introduction of Undaria into Nuevo Gulf, Argentina, around 1992, this alien seaweed has now colonized different sites over 700 km of coast, forming dense seasonal forests in waters from 0 to 15 meters in depth. In the spring it is common for plants of Undaria to break away from the substrate and be transported by sea currents. As Undaria gets stuck onto reefs it has the potential to reduce habitat quality for reef fish by physically obstructing refuges. This study aims to assess the impact of Undaria on the abundance of four species of rocky-reef fishes by an observational experiment. Fish abundance on reefs with and without Undaria was estimated by underwater visual census methods. Sites were classified according to their topographical relief, as this was expected to influence the effect of Undaria on the abundance of reef fishes. Fish abundance decreased markedly in low-relief reefs that had been covered by Undaria. In contrast, the drifting Undaria had no effect on the abundance of any of the fish species considered in high-relief reefs, where it tends to cover only the lowest-lying areas, leaving much of the refuges for fish unaffected. In conclusion, the presence of Undaria off the coast of Argentina results in transitory habitat loss for reef fishes inhabiting low-relief reefs during late spring and early summer. Although we do not know how much of a threat this habitat loss represents for the conservation of reef fish populations of northern Patagonia, the documented local impact of Undaria within the gulfs is strong and may affect a number of recreational and commercial activities which are centered on the reefs and their fish assemblages.     

Red algal exotics on North Sea coasts

A total of ten red seaweed species are recognized as introduced into the North Sea from other parts of the world. These areAsparagopsis armata andBonnemaisonia hamifera (Bonnemaisoniales),Grateloupia doryphora (Halymeniales),Antithamnionella spirographidis, Antithamnionella ternifolia, Anotrichium furcellatum, Dasya baillouviana, ?Dasysiphonia sp.,Polysiphonia harveyi andPolysiphonia senticulosa (Ceramiales). The oldest of these isB. hamifera, introduced prior to 1890, while the most recent,?Dasysiphonia sp., was first found in 1994 and still requires taxonomic investigation. A variety of distribution patterns is seen, with geographical ranges varying from general within the North Sea to very restricted. The diversity of introduced red algae on eastern coasts of the North Sea is much greater than in the west. The most likely explanation for this pattern is that French coasts were the initial site of introduction for many of the seaweeds, which were then distributed northwards by the residual surface currents. Their increasing success in the Netherlands has probably been promoted by the drastically changed local hydrodynamic conditions which have also permitted the recent introduction of many native European species. Of the biological features of species that may favour their success as introductions, clonal vegetative propagation, often with specialized propagules or fragmentation mechanisms, is almost ubiquitous. Low-temperature tolerances can be inferred, but data are sparse. Many of the alien red algae in the North Sea contain anti-grazing compounds such as bromophenols, which may contribute to their invasive potential by deterring grazing sufficiently to permit establishment of an inoculum.     

A global assessment of freshwater fish introductions in mediterranean-climate regions

Mediterranean-climate regions (med-regions) are global hotspots of endemism facing mounting environmental threats associated with human-related activities, including the ecological impacts associated with non-native species introductions. We review freshwater fish introductions across med-regions to evaluate the influences of non-native fishes on the biogeography of taxonomic and functional diversity. Our synthesis revealed that 136 freshwater fish species (26 families, 13 orders) have been introduced into med-regions globally. These introductions, and local extirpations, have increased taxonomic and functional faunal similarity among regions by an average of 7.5% (4.6–11.4%; Jaccard) and 7.2% (1.4–14.0%; Bray–Curtis), respectively. Faunal homogenisation was highest in Chile and the western Med Basin, whereas sw Cape and the Aegean Sea drainages showed slight differentiation (decrease in faunal similarity) over time. At present, fish faunas of different med-regions have widespread species in common (e.g. Gambusia holbrooki, Cyprinus carpio, Oncorhynchus mykiss, Carassius auratus, and Micropterus salmoides) which are typically large-bodied, non-migratory, have higher physiological tolerance, and display fast population growth rates. Our findings suggest that intentional and accidental introductions of freshwater fish have dissolved dispersal barriers and significantly changed the present-day biogeography of med-regions across the globe. Conservation challenges in med-regions include understanding the ecosystem consequences of non-native species introductions at macro-ecological scales.     

First Report of the Asian kelp Undaria pinnatifida in the Northeastern Pacific Ocean

The Asian kelp Undaria pinnatifida was discovered in southern California in the spring of 2000, and by the summer of 2001 had been collected at several California sites from Los Angeles to Long Beach Harbors and Catalina Island as far north as Monterey Harbor. From its native range in the northwestern Pacific, Undaria has now been introduced to the northeastern and southwestern Atlantic and the southwestern and northeastern Pacific, through a combination of intentional transport for cultivation, accidental transport with oysters, as fouling on vessel hulls, and possibly other means. In the northeastern Pacific, water temperatures are suitable for its establishment from at least Baja California to British Columbia, where it is likely to grow well in sheltered and partially sheltered waters.     

Environmental and ecological factors influencing the spillover of the non-native kelp, <Emphasis Type="Italic">Undaria pinnatifida</Emphasis>, from marinas into natural rocky reef communities

The non-native kelp, Undaria pinnatifida, is considered one of the world’s worst invasive species. The northeast Atlantic is a hotspot of Undaria invasion, yet there is limited knowledge on its invasion dynamics. In the UK its distribution is strongly associated with artificial structures, primarily marina and harbour pontoons, with relatively few records of Undaria on natural substrates. Here, the southwest UK is used as a case region, to explicitly link Undaria distribution-abundance patterns in artificial marina habitats with those in natural rocky reef habitats. Using a mixture of in situ recording and video survey techniques, Undaria was found at all thirteen marina sites surveyed; but in only 17 of 35 rocky reef sites, all of which were in 2 of the 5 larger systems surveyed (Plymouth Sound and Torbay). The distribution-abundance patterns of Undaria at reef sites were analysed using zero-inflated models. The probability of finding Undaria on rocky reef increased with increasing proximity to marinas with high abundances of Undaria. Total propagule pressure from marinas also increased the probability of occurrence, and was positively related to Undaria abundance and cover at reef sites. Increases in the cover of native kelps, Laminaria spp., and wave exposure at reef sites were linked to a reduced probability of Undaria occurrence, and lower abundance and cover. Identifying high risk areas, natural boundaries and factors affecting the spread and abundance of non-native species in natural habitats is key to future management prioritisation. Where Undaria is confined to artificial substrates management may be deemed a low priority. However, the results of this study suggest that controlling the abundance and propagule pressure in artificial habitats may limit, to some extent, the spillover of Undaria into natural rocky reef habitats, where it has the potential to interact with and influence native communities.     

Global population genetic structure of the starlet anemone <Emphasis Type="Italic">Nematostella vectensis</Emphasis>: multiple introductions and implications for conservation policy

Distinguishing natural versus anthropogenic dispersal of organisms is essential for determining the native range of a species and implementing an effective conservation strategy. For cryptogenic species with limited historical records, molecular data can help to identify introductions. Nematostella vectensis is a small, burrowing estuarine sea anemone found in tidally restricted salt marsh pools. This species’ current distribution extends over three coast lines: (i) the Atlantic coast of North America from Nova Scotia to Georgia, (ii) the Pacific coast of North America from Washington to central California, and (iii) the southeast coast of England. The 1996 IUCN Red List designates N. vectensis as “vulnerable” in England. Amplified fragment length polymorphism (AFLP) fingerprinting of 516 individuals from 24 N. vectensis populations throughout its range and mtDNA sequencing of a subsample of these individuals strongly suggest that anthropogenic dispersal has played a significant role in its current distribution. Certain western Atlantic populations of N. vectensis exhibit greater genetic similarity to Pacific populations or English populations than to other western Atlantic populations. At the same time, F-statistics showing high degrees of genetic differentiation between geographically proximate populations support a low likelihood for natural dispersal between salt marshes. Furthermore, the western Atlantic harbors greater genetic diversity than either England or the eastern Pacific. Collectively, these data clearly imply that N. vectensis is native to the Atlantic coast of North America and that populations along the Pacific coast and in England are cases of successful introduction.     

Widespread primary,but geographically restricted secondary,human introductions of wall lizards,Podarcis muralis

Establishing the introduction pathways of alien species is a fundamental task in invasion biology. The common wall lizard, Podarcis muralis, has been widely introduced outside of its native range in both Europe and North America, primarily through escaped pets or deliberate release of animals from captive or wild populations. Here, we use Bayesian clustering, approximate Bayesian computation (ABC) methods and network analyses to reconstruct the origin and colonization history of 23 non‐native populations of wall lizards in England. Our analyses show that established populations in southern England originate from at least nine separate sources of animals from native populations in France and Italy. Secondary introductions from previously established non‐native populations were supported for eleven (47%) populations. In contrast to the primary introductions, secondary introductions were highly restricted geographically and appear to have occurred within a limited time frame rather than being increasingly common. Together, these data suggest that extant wall lizard populations in England are the result of isolated accidental and deliberate releases of imported animals since the 1970s, with only local translocation of animals from established non‐native populations. Given that populations introduced as recently as 25 years ago show evidence of having adapted to cool climate, discouraging further translocations may be important to prevent more extensive establishment on the south coast of England.     

Pathways of spread of the introduced ascidian Styela clava (Tunicata) in Northern Europe, as revealed by microsatellite markers

Styela clava, a solitary ascidian native to the NW Pacific, has become a conspicuous member of fouling communities in NW European waters. As its natural dispersal appears to be limited, the wide distribution of S. clava along coasts within its introduced range may be attributed to secondary spread assisted by human activities. Here, we used six microsatellite loci to examine the genetic diversity and extent of gene flow among S. clava populations in its European introduced range. Samples were collected from 21 populations within Europe (N = 808), 4 populations within the USA and two populations within the native range (Japan). Large variation in genetic diversity was observed among the European populations but were not explained either by the geographic distance from the first introduction area (i.e. Plymouth, UK) nor by the time elapsed since the introduction. No founder effect was observed in the introduced populations, except possibly in Puget Sound (USA). At least two different introductions occurred in Europe, identified as distinct genetic clusters: northern Danish populations (resembling one Japanese population), and the rest of Europe; a sample from Shoreham (England) possibly represents a third introduction. In North America, the population from the Atlantic was genetically similar to the majority of European populations, suggesting a European origin for populations on this seaboard, while populations from the Pacific coast were genetically similar to the same Japanese population as the Danish populations.     

A population model for predicting the successful establishment of introduced bird species

One of the strongest generalities in invasion biology is the positive relationship between probability of establishment and the numbers of individuals introduced. Nevertheless, a number of significant questions remain regarding: (1) the relative importance of different processes during introduction (e.g., demographic, environmental, and genetic stochasticity, and Allee effects); (2) the relative effects of propagule pressure (e.g., number of introductions, size of introductions, and lag between introductions); and (3) different life history characteristics of the species themselves. Here, we adopt an individual-based simulation modeling approach to explore a range of such details in the relationship between establishment success and numbers of individuals introduced. Our models are developed for typical exotic bird introductions, for which the relationship between probability of establishment and the numbers of individuals introduced has been particularly well documented. For both short-lived and long-lived species, probability of establishment decreased across multiple introductions (compared with a single introduction of the same total size), and this decrease was greater when inbreeding depression was included. Sensitivity analyses revealed four predictors that together accounted for >95 % of model performance. Of these, R ₀ (the average number of daughters produced per female over her lifetime) and propagule pressure were of primary importance, while random environmental effects and inbreeding depression exerted lesser influence. Initial founder size is undoubtedly going to be important for ensuring the persistence of introduced populations. However, we found the demographic traits, which influence how introduced individuals behave, to have the greatest effect on establishment success.     

An investigation of dispersal of the introduced brown alga Undaria pinnatifida (Harvey) Suringar and its competition with some species on the man-made structures of Torquay Marina (Devon, UK)

As part of a study into the ecology of the recently introduced macroalga Undaria pinnatifida in the British Isles, the competitive ability of this adventive kelp with native species was investigated in Torquay Marina (Devon, UK). Sites of introduction were surveyed for the presence of native species filling a similar ecological niche and long-term competition experiments set up over 6 years. Experiments involving clearance, removal and monitoring showed that Undaria has a low competitive ability in the infralittoral and sublittoral of a rocky breakwater. Undaria did not manage to colonise the nearby (200 m) shoreline for at least 7 years from the date of its original introduction to the floating pontoons of Torquay Marina. Undaria did out-compete native species in a marina environment: two kelp species, Laminaria digitata and Laminaria saccharina, and a sea squirt, Styela clava.     

Tracing the introduction history of the invasive swimming crab <Emphasis Type="Italic">Charybdis hellerii</Emphasis> (A. Milne-Edwards, 1867) in the Western Atlantic: evidences of high genetic diversity and multiple introductions

The swimming crab Charybdis hellerii is an invader with global distribution in warm waters. Native to the Indo-Pacific, this species invaded the eastern Mediterranean Sea after the Suez Canal opening. In 1987, it was first reported in the Western Atlantic, probably transported via ballast water of ships. Since then, it has been registered from many localities along the American coast from the USA to southern Brazil where it has rapidly established reproducing populations. Our main aim was to investigate the introduction history of this species along the American coast, using a phylogeographic approach. Additionally, we attempted to clarify the identity of this invasive species by molecular analyses and morphological assessment in order to provide a basis for our main investigation. C. hellerii was confirmed as a single species, but both cytochrome c oxidase subunit I and 16S rDNA revealed a genetic structure, splitting the potential source populations of American introductions into two groups: “western Indian Ocean and Mediterranean Sea” (WIO + MS) versus “eastern Indian + western Pacific oceans” (EIO + WPO). Most specimens from America clustered with the former group, supporting the hypothesis that the Mediterranean Sea represented the main source of Western Atlantic populations. However, the clustering of animals from southern Brazil with the latter group indicates that introductions from the eastern Indian or Pacific oceans must also have occurred. The existence of a third group, found exclusively within the American range and genetically related to EIO + WPO, also indicates an independent introduction from an unsampled locality from the native range. The haplotype and nucleotide diversities of American localities were comparable to those of source populations, contradicting a founder effect prediction. This finding might be related to the high propagule pressure associated with introductions via ballast water and the occurrence of multiple introductions from genetic distinct sources. The direct comparison of the haplotype numbers suggested no genetic bottleneck during introduction from the Mediterranean Sea, but a bottleneck might have occurred during introductions from the eastern Indian or Pacific oceans.     

Colonization History,Host Distribution,Anthropogenic Influence and Landscape Features Shape Populations of White Pine Blister Rust,an Invasive Alien Tree Pathogen

White pine blister rust is caused by the fungal pathogen Cronartium ribicola J.C. Fisch (Basidiomycota, Pucciniales). This invasive alien pathogen was introduced into North America at the beginning of the 20th century on pine seedlings imported from Europe and has caused serious economic and ecological impacts. In this study, we applied a population and landscape genetics approach to understand the patterns of introduction and colonization as well as population structure and migration of C. ribicola. We characterized 1,292 samples of C. ribicola from 66 geographic locations in North America using single nucleotide polymorphisms (SNPs) and evaluated the effect of landscape features, host distribution, and colonization history on the structure of these pathogen populations. We identified eastern and western genetic populations in North America that are strongly differentiated. Genetic diversity is two to five times higher in eastern populations than in western ones, which can be explained by the repeated accidental introductions of the pathogen into northeastern North America compared with a single documented introduction into western North America. These distinct genetic populations are maintained by a barrier to gene flow that corresponds to a region where host connectivity is interrupted. Furthermore, additional cryptic spatial differentiation was identified in western populations. This differentiation corresponds to landscape features, such as mountain ranges, and also to host connectivity. We also detected genetic differentiation between the pathogen populations in natural stands and plantations, an indication that anthropogenic movement of this pathogen still takes place. These results highlight the importance of monitoring this invasive alien tree pathogen to prevent admixture of eastern and western populations where different pathogen races occur.