Invasive feral pigs impact native tree ferns and woody seedlings in Hawaiian forest

Invasive mammals can fundamentally alter native plant communities, especially on isolated islands where plants evolved without them. The globally invasive feral pig (Sus scrofa) can be particularly destructive to native plant communities. Tree ferns are an important understory component in many forests facilitating the establishment of a variety of species. However, the extent and effects of feral pig damage to tree ferns, and associated impacts on plant community regeneration, are largely unknown. We quantified the effect that feral pig damage has on tree fern growth, survival, and epiphytic woody seedling abundance over 1 year on 438 randomly selected tree ferns of three endemic species (Cibotium chamissoi, Cibotium glaucum, and Cibotium menziesii) in a Hawaiian montane wet forest with high tree fern and feral pig densities. Across all tree fern species, feral pigs damaged 13 % of individuals over 1 year. Compared with undamaged tree ferns, moderately- to heavily-damaged individuals had decreases of 4 to 27 % in trunk length increment and lost tenfold more fronds. Tree fern angle (standing, leaning, prone, or semi-prone) and woody seedling abundance co-varied with feral pig damage. Specifically, damaged tree ferns were more often prone or semi-prone and supported more seedlings, but also had annual mortality up to 34 % higher than undamaged tree ferns. Overall, feral pig damage had substantial negative effects on tree ferns by reducing growth and survival. Given the importance of tree ferns as regeneration sites for a variety of native plants, feral pig damage to tree ferns will likely alter future forest composition and structure. Specifically, feral pig damage to tree ferns reduces potential establishment sites for species that either regenerate preferentially as epiphytes or are currently restricted to epiphytic establishment due to ground rooting by feral pigs.     

Spatiotemporal dynamics of the spread of African tilapias (Pisces: Oreochromis spp.) into rivers of northeastern Mesoamerica

African tilapias (Oreochromis spp.) occur in more than 100 countries outside of their native ranges and research on their invasions is largely lacking. We investigated spatiotemporal patterns of tilapia spread into 29 drainage basins in Belize and parts of Guatemala and Mexico, drawing on field data and interviews with fishermen. Habitat-suitability models for tilapias were created from geospatial and species occurrence data, and fishermen interviews were used to reconstruct the chronology of tilapia spread into predicted suitable habitats. Tilapia (predominantly Nile tilapia, O. niloticus) presence was confirmed at 78 sites in 9 of 29 drainage basins. Our habitat-suitability model predicted that 7,510 linear km of river habitat in the study area were vulnerable to colonization by tilapias, predominately in mid- to low elevation main stem rivers, from sea level to 277 m above sea level. The reconstructed spatial chronology of spread showed that the invasion started in 1990 and progressed slowly (2 km yr^?1) through an establishment phase before rapid expansion (~30 km yr^?1) between 1996 and 2002, after which new detections slowed. Human movement of fish for aquaculture was identified as a primary cause of dispersal that interacted with flooding as an important secondary cause. The shortest paths across low elevation drainage divides between major basins revealed several potential corridors for future tilapia spread during flooding. Research into tilapia spatial metapopulation structure and economic fisheries status, more stringent regulation of aquaculture activities, pro-active fisheries management, and development of policies to screen potentially invasive species before importation are recommended to avoid additional releases of tilapia and further spread in the region.     

Insights into the Establishment of the Manila Clam on a Tidal Flat at the Southern End of an Introduced Range in Southern California,USA

Coastal ecosystem modifications have contributed to the spread of introduced species through alterations of historic disturbance regimes and resource availability, and increased propagule pressure. Frequency of occurrence of the Manila clam (Venerupis phillipinarum, Veneridae) in Southern California estuaries has increased from absent or sparse to common since the mid-1990s. Potential invasion vectors include seafood sales and aquaculture, and spread from established northern populations over decades. The clam’s post-settlement habitat preferences are, however, uncertain in this region. Our project aimed to identify factors associated with established patches of the clam within a bay toward the southern end of this introduced range. During summer 2013, we sampled 10 tidal flat sites in Mission Bay, San Diego; each containing an area with and without hard structure (e.g., riprap, boulders). We measured likely environmental influences (e.g., sediment variables, distance to ocean). Manila clam densities across the bay were most strongly associated with site, where highest densities were located in the northern and/or back halves of the bay; and weakly correlated with lower porewater salinities. Within sites, Manila clam density was enhanced in the presence of hard structure in most sites. Prevailing currents and salinity regimes likely contribute to bay wide distributions, while hard structures may provide suitable microhabitats (refuge from predators and physical stress) and larval entrapment within sites. Results provide insights into decisions about future shoreline management efforts. Finally, we identify directions for future study to better understand and therefore predict patterns of establishment of the Manila clam in the southern portion of its introduced range.     

Return of the native facilitated by the invasive? Population composition,substrate preferences and epibenthic species richness of a recently discovered shellfish reef with native European flat oysters (Ostrea edulis) in the North Sea

After being ecologically extinct for almost a century, the discovery of a shellfish reef with native European flat oysters (Ostrea edulis) in the Dutch coastal area of the North Sea by the authors of this study called for an extensive survey to better understand some of the key requirements for the return of the native oyster in coastal waters. We assessed habitat conditions, its potential for increasing biodiversity, and the role of substrate provision by other bivalves such as the invasive alien Pacific oyster (Crassostrea gigas). Using underwater visual census, O. edulis size-frequency distributions and attachment substrate was investigated, as well as the composition of the epibenthic community and substrata types inside quadrats that were distributed across the reef. This reef was found to be composed of native European flat oysters, invasive alien Pacific oysters and blue mussels (Mytilus edulis), alternated with sandy patches. The O. edulis population (6.8?±?0.6 oysters m^?2) consisted of individuals of different size classes. In quadrats with native and non-native oysters the number of epibenthic species was 60% higher compared to adjacent sand patches within the reef. Notably, our results showed that the native oyster predominantly used shell (fragments) of the invasive Pacific oyster as settlement substrate (81% of individuals). Our results optimistically show that conditions for native oyster restoration can be suitable at a local scale in the coastal North Sea area and suggest that the return of native oysters may be facilitated by novel substrate provided by invasive oysters at sites where their distribution overlap.     

Inferring the invasion history of coral berry Ardisia crenata from China to the USA using molecular markers

Genetic comparisons between native and invasive populations of a species can provide insights into its invasion history information, which is useful for guiding management and control strategies. The coral berry Ardisia crenata was introduced to Florida last century as a cultivated ornament plant, and has since spread widely throughout the southern regions of the USA. Previously, the genetic variation among 20 natural populations of A. crenata across its distribution center in southern China was quantified using seven microsatellite markers. Here we expand on that work by additionally sampling individuals from four other native populations in Taiwan and Japan, and from five invasive populations in the USA. We also examined the results from one chloroplast intergenic spacer region (trnF-trnL) in all 29 populations. Our aim is to identify the invasion source and subsequent history of the species?? spread throughout the southern USA. We observed lower genetic diversity in the invasive populations based on both microsatellite and chloroplast markers. Our data show that the invasive populations can be clustered with native populations in southeastern China, inferring this region as the geographic origin of A. crenata cultivars invading the USA. We further classified invasive individuals into invasive I and invasive II clusters. Nantou in Taiwan and Xihu in mainland China are the most closely related populations to those, which identify the former as potential sources for host-specific control agents. Our results, combined with the known introduction records, suggest that A. crenata was first multiply introduced into Florida and then secondarily colonized Louisiana and Texas from Florida.     

Occurrence of Transgenic Feral Alfalfa (Medicago sativa subsp. sativa L.) in Alfalfa Seed Production Areas in the United States

The potential environmental risks of transgene exposure are not clear for alfalfa (Medicago sativa subsp. sativa), a perennial crop that is cross-pollinated by insects. We gathered data on feral alfalfa in major alfalfa seed-production areas in the western United States to (1) evaluate evidence that feral transgenic plants spread transgenes and (2) determine environmental and agricultural production factors influencing the location of feral alfalfa, especially transgenic plants. Road verges in Fresno, California; Canyon, Idaho; and Walla Walla, Washington were surveyed in 2011 and 2012 for feral plants, and samples were tested for the CP4 EPSPS protein that conveys resistance to glyphosate. Of 4580 sites surveyed, feral plants were observed at 404 sites. Twenty-seven percent of these sites had transgenic plants. The frequency of sites having transgenic feral plants varied among our study areas. Transgenic plants were found in 32.7%, 21.4.7% and 8.3% of feral plant sites in Fresno, Canyon and Walla Walla, respectively. Spatial analysis suggested that feral populations started independently and tended to cluster in seed and hay production areas, places where seed tended to drop. Significant but low spatial auto correlation suggested that in some instances, plants colonized nearby locations. Neighboring feral plants were frequently within pollinator foraging range; however, further research is needed to confirm transgene flow. Locations of feral plant clusters were not well predicted by environmental and production variables. However, the likelihood of seed spillage during production and transport had predictive value in explaining the occurrence of transgenic feral populations. Our study confirms that genetically engineered alfalfa has dispersed into the environment, and suggests that minimizing seed spillage and eradicating feral alfalfa along road sides would be effective strategies to minimize transgene dispersal.     

Genetic diversity and relatedness in aquaculture and marina populations of the invasive tunicate Didemnum vexillum in the British Isles

Introductions of invasive, non-native species in the marine environment are increasing as human activity within coastal areas rises. Genetic datasets are useful tools to identify source populations, track routes of invasions, and illuminate the role of genetic variation in the establishment and subsequent spread of novel introductions. Here, a microsatellite dataset is used to estimate the genetic diversity and population structure of 7 introduced Didemnum vexillum populations in Britain and Ireland, 4 of which are associated with aquaculture and 3 with marinas. Genetic differentiation observed between these populations indicates human-mediated transport as the main mechanism underlying the population structure of D. vexillum in Britain and Ireland. In addition to elucidating patterns of population structure we found that aquaculture sites showed significantly higher genetic diversity (measured as allelic richness) in comparison to the marina sites. We discuss these findings in relation to the history of each invasion, the complex life history of D. vexillum, and available evidence of the relative invasiveness of these populations. Our results show numerous interesting patterns which highlight further research avenues to elucidate the complex factors underlying the global spread of this successful invader.

     

Abiotic factors controlling the establishment and abundance of the invasive golden mussel Limnoperna fortunei

Limnoperna fortunei (Bivalvia, Mytilidae), a freshwater bivalve native to Southern Asia, has been an invasive species in South America since 1991. It spread upstream in the La Plata basin reaching the Paraguay River in the vicinity of the Pantanal wetland, Brazil, around 1998. The role of abiotic factors in controlling establishment and abundance of this species is not well known, making projections of its risk of further spread difficult. This study evaluates the importance of abiotic factors to L. fortunei populations established in rivers of the Pantanal, focusing on larval and juvenile densities and taking advantage of a wide range of seasonal variability in water temperature, flow, dissolved oxygen, and suspended material. Temperature, river stage (influencing several water characteristics) and water velocity are the main variables related to the larval and juvenile densities. In the Pantanal, environmental variables vary over a broader range compared with other South American locations, subjecting L. fortunei to oxygen depletion, low calcium, low pH, and high water velocity and suspended solids, associated with low chlorophyll a concentrations. The combined effect of several of these conditions may explain the relatively low densities in some Pantanal sites. However, they probably will not prevent the persistence of populations in the Pantanal and the eventual establishment of viable populations in upriver systems connected to the Pantanal. These results are pertinent not only to this species but also to other aquatic invasive invertebrates whose expansion may be limited by thermal extremes, episodic oxygen depletion, and waters that are too dilute or acidic for optimal biocalcification.     

Multiple dispersal vectors drive range expansion in an invasive marine species

The establishment and subsequent spread of invasive species is widely recognized as one of the most threatening processes contributing to global biodiversity loss. This is especially true for marine and estuarine ecosystems, which have experienced significant increases in the number of invasive species with the increase in global maritime trade. Understanding the rate and mechanisms of range expansion is therefore of significant interest to ecologists and conservation managers alike. Using a combination of population genetic surveys, environmental DNA (eDNA) plankton sampling and hydrodynamic modelling, we examined the patterns of introduction of the predatory Northern Pacific seastar (Asterias amurensis) and pathways of secondary spread within southeast Australia. Genetic surveys across the invasive range reveal some genetic divergence between the two main invasive regions and no evidence of ongoing gene flow, a pattern that is consistent with the establishment of the second invasive region via a human‐mediated translocation event. In contrast, hydrodynamic modelling combined with eDNA plankton sampling demonstrated that the establishment of range expansion populations within a region is consistent with natural larval dispersal and recruitment. Our results suggest that both anthropogenic and natural dispersal vectors have played an important role in the range expansion of this species in Australia. The multiple modes of spread combined with high levels of fecundity and a long larval duration in A. amurensis suggests it is likely to continue its range expansion and significantly impact Australian marine ecosystems.     

Cleft,Crevice, or the Inner Thigh: ‘Another Place’ for the Establishment of the Invasive Barnacle Austrominius modestus (Darwin, 1854)

The proliferation of anthropogenic infrastructure in the marine environment has aided the establishment and spread of invasive species. These structures can create novel habitats in areas normally characterised as void of suitable settlement sites. The habitat requirements of the invasive acorn barnacle Austrominius modestus (Darwin, 1854) were assessed using a novel sampling site at Crosby Beach, Liverpool. Austrominius modestus has spread rapidly around the UK since its initial introduction, becoming locally dominant in many estuarine areas including the Antony Gormley art installation, ‘Another Place’, at Crosby Beach. The installation consists of 100 replicate solid cast-iron life-size human figures, located at a range of heights on the shore. We recorded the distribution and abundance of A. modestus present on all of the statues at various positions during the summer of 2006. The positions varied in location, exposure, direction, and rugosity. Although parameters such as rugosity and exposure did influence patterns of recruitment, they were less important than interactions between shore height and direction, and specific location on the beach. The addition of a suitable substrate to a sheltered and estuarine region of Liverpool Bay has facilitated the establishment of A. modestus. Understanding the habitat requirements of invasive species is important if we are to make predictions about their spread and the likelihood of invasion success. Austrominius modestus has already become locally dominant in some regions of the UK and, with projections of favourable warming conditions and the global expansion of artificial structures, the continued spread of this species can be expected. The implications of this on the balance between native and invasive species dominance should be considered.     

High mitochondrial DNA diversity of an introduced alien carnivore: comparison of feral and ranch American mink Neovison vison in Poland

Aim Invasive alien species usually exhibit very high adaptation and rapid evolution in a new environment, but they often have low levels of genetic diversity (invasive species paradox). Genetic variation and population genetic structure of feral American mink, Neovison vison, in Poland was investigated to explain the invasion paradox and to assess current gene flow. Furthermore, the influence of mink farming on adaptation of the feral population was evaluated by comparing the genetic structure of feral and ranch mink. Location Samples from feral mink were collected in 11 study areas in northern and central Poland and from ranch mink at 10 farms distributed throughout the country. Methods A 373‐bp‐long mtDNA control region fragment was amplified from 276 feral and 166 ranch mink. Results Overall, 31 haplotypes, belonging to two groups from genetically diverse sources, were detected: 11 only in feral mink, 12 only in ranch mink and eight in both. The genetic differentiation of feral mink from the trapping sites was high, while that among ranch mink from various farms was moderate. There was no significant relationship between genetic and geographic distance. The number of trapping sites where given haplotypes occurred correlated with the number of farms with these haplotypes. The mink from two sites were the most divergent, both from all other feral mink and from ranch mink. Comparison of mtDNA and microsatellite differentiation suggests male‐biased dispersal in this species. Main conclusions American mink in Poland exhibit high genetic diversity and originate from different source populations of their native range. The process of colonization was triggered by numerous escapees from various farms and by immigrants from Belarus. The genetic structure of local feral mink populations was shaped by the founder effect and multiple introductions. The genomic admixture that occurred during mixing of different populations might have increased the fitness of individuals and accelerated the invasiveness of this species.     

Ecological impacts of non-native Pacific oysters (<Emphasis Type="Italic">Crassostrea gigas</Emphasis>) and management measures for protected areas in Europe

Pacific oysters are now one of the most ‘globalised’ marine invertebrates. They dominate bivalve aquaculture production in many regions and wild populations are increasingly becoming established, with potential to displace native species and modify habitats and ecosystems. While some fishing communities may benefit from wild populations, there is now a tension between the continued production of Pacific oysters and risk to biodiversity, which is of particular concern within protected sites. The issue of the Pacific oyster therefore locates at the intersection between two policy areas: one concerning the conservation of protected habitats, the other relating to livelihoods and the socio-economics of coastal aquaculture and fishing communities. To help provide an informed basis for management decisions, we first summarise evidence for ecological impacts of wild Pacific oysters in representative coastal habitats. At local scales, it is clear that establishment of Pacific oysters can significantly alter diversity, community structure and ecosystem processes, with effects varying among habitats and locations and with the density of oysters. Less evidence is available to evaluate regional-scale impacts. A range of management measures have been applied to mitigate negative impacts of wild Pacific oysters and we develop recommendations which are consistent with the scientific evidence and believe compatible with multiple interests. We conclude that all stakeholders must engage in regional decision making to help minimise negative environmental impacts, and promote sustainable industry development.     

Massive settlements of the Pacific oyster, Crassostrea gigas, in Scandinavia

The Pacific oyster (Crassostrea gigas) is an important aquaculture species world-wide. Due to its wide environmental tolerance and high growth rate, it has also become a successful invader in many areas, leading to major ecosystem changes. Low water temperatures were previously believed to restrict the establishment of Pacific oysters in Scandinavia. However, recent surveys reveal that the Pacific oyster is now established in many areas in Scandinavia. The biomass of oysters in the Danish Wadden Sea has increased dramatically between 2005 and 2007, large numbers were observed along the Swedish west coast from settlement in 2006, and in Norway, populations are established along the southwest coast to 60°N.     

Vegetation response to removal of non-native feral pigs from Hawaiian tropical montane wet forest

Globally, non-native ungulates threaten native biodiversity, alter biotic and abiotic factors regulating ecological processes, and incur significant economic costs via herbivory, rooting, and trampling. Removal of non-native ungulates is an increasingly common and crucial first step in conserving and restoring native forests. However, removal is often controversial and there is currently little information on plant community responses to this management action. Here, we examine the response of native and non-native understory vegetation in paired sites inside and outside of exclosures across a 6.5–18.5 year chronosequence of feral pig (Sus scrofa) removal from canopy-intact Hawaiian tropical montane wet forest. Stem density and cover of native plants, species richness of ground-rooted native woody plants, and abundance of native plants of conservation interest were all significantly higher where feral pigs had been removed. Similarly, the area of exposed soil was substantially lower and cover of litter and bryophytes was greater with feral pig removal. Spatial patterns of recruitment were also strongly affected. Whereas epiphytic establishment was similar between treatments, the density of ground-rooted woody plants was four times higher with feral pig removal. Abundance of invasive non-native plants also increased at sites where they had established prior to feral pig removal. We found no patterns in any of the measured variables with time, suggesting that commonly occurring species recover within 6.5 years of feral pig removal. Recovery of species of conservation interest, however, was highly site specific and limited to areas that possessed remnant populations at the time of removal, indicating that some species take much longer (>18.5 years) to recover. Feral pig removal is the first and most crucial step for conservation of native forests in this area, but subsequent management should also include control of non-native invasive plants and outplanting native species of conservation interest that fail to recruit naturally.     

Utilization of Sugarcane Habitat by Feral Pig (Sus scrofa) in Northern Tropical Queensland: Evidence from the Stable Isotope Composition of Hair

Feral pigs (Sus scrofa) are an invasive species that disrupt ecosystem functioning throughout their introduced range. In tropical environments, feral pigs are associated with predation and displacement of endangered species, modification of habitat, and act as a vector for the spread of exotic vegetation and disease. Across many parts of their introduced range, the diet of feral pigs is poorly known. Although the remote location and difficult terrain of far north Queensland makes observing feral pig behavior difficult, feral pigs are perceived to seek refuge in World Heritage tropical rainforests and seasonally 'crop raid' into lowland sugarcane crops. Thus, identifying how feral pigs are using different components of the landscape is important to the design of management strategies. We used the stable isotope composition of captured feral pigs to determine the extent of rainforest and sugarcane habitat usage. Recently grown hair (basal hair) from feral pigs captured in remote rainforest indicated pigs met their dietary needs solely within this habitat. Stable carbon and nitrogen isotope values of basal hair from feral pigs captured near sugarcane plantations were more variable, with some individuals estimated to consume over 85% of their diet within a sugarcane habitat, while a few consumed as much as 90% of their diet from adjacent forested environments. We estimated whether feral pigs switch habitats by sequentially sampling δ(13)C and δ(15)N values of long tail hair from a subset of seven captured animals, and demonstrate that four of these individuals moved between habitats. Our results indicate that feral pigs utilize both sugarcane and forest habitats, and can switch between these resources.     

Aquaculture Enclosures Relate to the Establishment of Feral Populations of Introduced Species

Many species introduced by humans for social and economic benefits have invaded new ranges by escaping from captivity. Such invasive species can negatively affect biodiversity and economies. Understanding the factors that relate to the establishment of feral populations of introduced species is therefore of great importance for managing introduced species. The American Bullfrog (Lithobates catesbeianus) is one species that has escaped from farms, and it is now found in the wild in China. In this study, we examined influences of two types of bullfrog farm (termed simple and elaborate farm enclosures) on the establishment of feral populations of this species in 137 water bodies in 66 plots in four provinces of China. The likelihood of establishment of bullfrog populations in water bodies in plots with simple enclosures (49/89 = 55.1%) was higher than those with elaborate enclosures (3/48 = 6.3%). Based on the Akaike Information Criterion, the minimum adequate model of generalized linear mixed models with a binomial error structure and a logit link function showed that the establishment or failure of bullfrog populations in water bodies was positively correlated with the presence of a simple enclosure, the number of bullfrogs raised and the presence of permanent water in a plot, but negatively correlated with distance from a bullfrog farm and the occurrence of frequent hunting. Results therefore suggest that a simple farm enclosure can increase the establishment of feral bullfrog populations compared with an elaborate enclosure. Our findings are the first to quantify the importance of improving farming enclosures to control and minimize the risk from introduced species.     

Massive settlements of the Pacific oyster, <Emphasis Type="Italic">Crassostrea gigas</Emphasis>, in Scandinavia

The Pacific oyster (Crassostrea gigas) is an important aquaculture species world-wide. Due to its wide environmental tolerance and high growth rate, it has also become a successful invader in many areas, leading to major ecosystem changes. Low water temperatures were previously believed to restrict the establishment of Pacific oysters in Scandinavia. However, recent surveys reveal that the Pacific oyster is now established in many areas in Scandinavia. We present data on the current distribution, abundance and age-structure in Denmark, Sweden and Norway. The biomass of oysters in the Danish Wadden Sea increased from 1,056 to 6,264 tonnes between 2005 and 2007. Massive settlements were observed along the Swedish west coast in 2007, with densities >400 oysters per m⁻². In Norway, populations are established on the southern coast, and specimens have been found as far north as 60°N. The potential impacts and probable causes of this recent large-scale establishment are discussed.     

Tolerance of the invasive tunicate Styela clava to air exposure

Styela clava is a subtidal invasive marine species in Northern Europe, Atlantic Canada, Australia and New Zealand. It grows attached to solid substrata, including boat hulls, ropes, moorings, piers and aquaculture equipment, all of which can aid its spread to new locations. It interferes with feeding of mussels and oysters, and increases their harvesting costs. Being subtidal, it could be assumed that tunicates would rapidly die in air and thus exposure to air would be a practical method to prevent their spread on boats and equipment. This study tested their survival when exposed to air for up to (1) 120?h at a constant temperature of 10?°C, (2) shade ambient 15–27?°C, and (3) full sun ambient 15–29?°C. Humidity was consistently high (78–100%). The results indicated that survival was longer when the air temperature was cooler. Larger individuals of S. clava generally survived for longer out of seawater than smaller individuals. The results predict that two weeks of exposure to air for two weeks could be an effective management method to eradicate S. clava from marine equipment when the air temperature is 10?°C. However, drying time would be less under conditions of low humidity and under direct sunlight.     

Development of a real-time PCR assay for detection of Mytilus species specific alleles: Application to a sampling survey in Scotland

Shellfish aquaculture is a growing industry in Scotland, dominated by the production of the mussel Mytilus edulis, the native species. Recently the discovery of Mytilus galloprovincialis and Mytilus trossulus together with M. edulis and all 3 hybrids in cultivation in some Scottish sea lochs led to questions regarding the distribution of mussel species in Scotland. The establishment of an extensive sampling survey, involving the collection of mussels at 34 intertidal sites and 10 marinas around Scotland, motivated the development of a high-throughput method for identification of Mytilus alleles from samples. Three Taqman®-MGB probes and one set of primers were designed, based on the previously described Me 15/16 primers targeting the adhesive protein gene sequence, and samples were screened for the presence of M. edulis, M. galloprovincialis and M. trossulus alleles using real-time PCR. Mytilus edulis alleles were identified in samples from all 44 sites. Mytilus galloprovincialis alleles were found together with M. edulis alleles extensively in northern parts of the west and east coasts. Mytilus trossulus alleles were identified in samples from 6 sites in the west and south-west of Scotland. Because M. trossulus is generally undesirable in cultivation and therefore preventing the geographical spread of this species across Scotland is considered beneficial by the shellfish aquaculture industry, these 6 samples were further analysed for genotype frequencies using conventional PCR. Although distribution of the non-native species M. galloprovincialis and M. trossulus have proven to be more widespread than previously thought, there is no evidence from our study of either M. trossulus or M. galloprovincialis acting as an invasive species in Scotland. The real-time PCR method developed in this study has proven to be a rapid and effective tool for the identification of M. edulis, M. galloprovincialis and M. trossulus alleles from samples and should prove useful in future surveys, ecological or aquaculture management related studies in both unispecific and mixed species areas of these species.     

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.