Showing papers in "Biological Invasions in 2009"

Ecological effects of invasive alien insects

Abstract: A literature survey identified 403 primary research publications that investigated the ecological effects of invasive alien insects and/or the mechanisms underlying these effects. The majority of these studies were published in the last 8 years and nearly two-thirds were carried out in North America. These publications concerned 72 invasive insect species, of which two ant species, Solenopsis invicta and Linepithema humile, accounted for 18% and 14% of the studies, respectively. Most publications investigated effects on native biodiversity at population or community level. Genetic effects and, to a lesser extent, effects on ecosystem services and processes were rarely explored. We review the effects caused by different insect invaders according to: their ecosystem roles, i.e. herbivores, predators, parasites, parasitoids and pollinators; the level of biological organisation at which they occur; and the direct and indirect mechanisms underlying these effects. The best documented effects occur in invasive ants, Eurasian forest herbivores invasive in North America, and honeybees. Impacts may occur through simple trophic interactions such as herbivory, predation or parasitism. Alien species may also affect native species and communities through more complex mechanisms such as competition for resources, disease transmission, apparent competition, or pollination disruption, among others. Finally, some invasive insects, particularly forest herbivores and ants, are known to affect ecosystem processes through cascading effects. We identify biases and gaps in our knowledge of ecological effects of invasive insects and suggest further opportunities for research.

Non-indigenous invasive bivalves as ecosystem engineers

Abstract: Several non-indigenous bivalve species have been colonising aquatic ecosystems worldwide, in some cases with great ecological and economic impacts. In this paper, we focus on the ecosystem engineering attributes of non-indigenous invasive bivalves (i.e., the capacities of these organisms to directly or indirectly affect the availability of resources to other species by physically modifying the environment). By reviewing the ecology of several invasive bivalves we identify a variety of mechanisms via which they modify, maintain and/or create habitats. Given the usually high densities and broad spatial distributions of such bivalves, their engineering activities can significantly alter ecosystem structure and functioning (e.g., changes in sediment chemistry, grain size, and organic matter content via bioturbation, increased light penetration into the water column due to filter feeding, changes in near bed flows and shear stress due to the presence of shells, provision of colonisable substrate and refuges by shells). In addition, changes in ecosystem structure and functioning due to engineering by invasive bivalves often have very large economic impacts. Given the worldwide spread of non-indigenous bivalves and the varied ways in which they physically modify habitats, their engineering effects should receive more serious consideration in restoration and management initiatives.

We can eliminate invasions or live with them. Successful management projects

Abstract: Pessimism about the prospects of eradicating invasive species or managing them at acceptably low densities is fostered by publicity about failures, but it is often unwarranted. Many species, including insects, plants, and aquatic invaders of various taxa, have been eradicated, and a variety of management techniques have maintained others at low densities for long periods. Many of these projects entailed low-tech, scorched-earth approaches, often using mechanical, physical, or chemical means. Others required sophisticated scientific research. There are at best rough guidelines rather than general rules about what approach to undertake. It is therefore important in each case to consider the full range of possible methods for eradication or long-term control, including the possibility of marshalling a massive amount of physical labor.

Hybridization and the evolution of invasiveness in plants and other organisms

Abstract: Less than a decade ago, we proposed that hybridization could serve as a stimulus for the evolution of invasiveness in plants (Ellstrand and Schierenbeck Proc Nat Acad Sci USA 97:7043–7050, 2000). A substantial amount of research has taken place on that topic since the publication of that paper, stimulating the symposium that makes up this special issue. Here we present an update of this emergent field, based both on the papers in this volume and on the relevant literature. We reevaluate the lists that we presented in our earlier paper of reports in which hybridization has preceded the evolution of invasiveness. We discard a few cases that were found to be in error, published only as abstracts, or based on personal communication. Then we augment the list from examples in this volume and a supplementary literature search. Despite the omissions, the total number of cases has increased. Many have been strengthened. We add a list of cases in which there has been evidence that intra-taxon hybridization has preceded the evolution of invasiveness. We also provide a number of examples from organisms other than plants. We consider how our examples suggest mechanisms whereby hybridization may act to stimulate the evolution of invasiveness. Hybridization does not represent the only evolutionary pathway to invasiveness, but it is one that can explain why the appearance of invasiveness often involves a long lag time and/or multiple introductions of exotics.

Taking stock: inventory of alien species in the Mediterranean sea

Abstract: 573 alien marine metazoan species have been recorded in the Mediterranean Sea. The present checklist is the first to present the species’ native range, presumed mode of introduction, spatial extent, and the date of the first record in each country. The majority of aliens are thermophilic species originating from the Indo-Pacific or Indian Oceans, which have entered the Mediterranean through the Suez Canal. However, the means of introduction differ greatly among the phyla, and the basins of the Mediterranean. The temporal records of the alien species reflect political crises, economic development and scientific interest in studying the phenomenon—in the past two decades on average about 10 alien species new to the Mediterranean are recorded annually. Many have established durable populations and extended their range: 125 alien species have been recorded from four or more countries. The possible impacts of regulatory instruments and environmental management options are examined.

Ecological impacts of non-indigenous invasive fungi as forest pathogens

Abstract: Non-indigenous pathogenic fungi increasingly threaten North American tree species. Ecosystems may be fundamentally changed when abundant tree species are functionally eliminated, as occurred with American chestnut (Castanea dentata (Marsh.) Borkh.). Conversely, changes may be more subtle but still significant and long lasting when populations are lost, or all trees in the larger size classes are killed. Proposed approaches for characterizing the magnitude of ecological impacts use characteristics of both the non-indigenous pathogen and the host species. Impacts are most significant when highly successful invading pathogens attack foundation species, setting in motion a long-lasting cascade of effects on the host and associated species. Such impacts have generally not been well documented at the ecosystem level. Several North American forest tree species have been functionally eliminated or severely reduced by non-indigenous pathogens. Historical invasions, such as that of chestnut blight (Cryphonectria parasitica (Murr.) Bar), caused very significant ecological impacts that will never be completely understood because of lack of quantitative data. Beech bark disease, caused by a combination of an introduced scale insect (Cryptococcus fagisuga Lindinger) and a fungus (Neonectria faginata (Lohman et al.) Castl. & Rossman), is still advancing and provides opportunities for studying ecosystem-level impacts when a major tree species is removed or markedly reduced in abundance from the overstorey. Butternut canker, caused by the fungus, Sirococcus clavigignenti-juglandacearum N.B. Nair, Kostichka & Kuntz, has spread throughout the host range, endangering species’ survival. Other non-indigenous invasive fungi such as Cronartium ribicola J.C. Fisch and Phytophthora lateralis Tucker & Milbrath continue to move into new populations, causing high mortality and associated restructuring of these ecosystems. Global trade and environmental change trends will ensure new challenges by non-indigenous fungal pathogens, presenting an urgent need for improved understanding of long-term impacts across ecological systems.

Diet overlap among two Asian carp and three native fishes in backwater lakes on the Illinois and Mississippi rivers

Abstract: Bighead and silver carp are well established in the Mississippi River basin following their accidental introduction in the 1980s. Referred to collectively as Asian carp, these species are filter feeders consuming phytoplankton and zooplankton. We examined diet overlap and electivity of Asian carp and three native filter feeding fishes, bigmouth buffalo, gizzard shad, and paddlefish, in backwater lakes of the Illinois and Mississippi rivers. Rotifers, Keratella spp., Brachionus spp., and Trichocerca spp., were the most common prey items consumed by Asian carp and gizzard shad, whereas crustacean zooplankton were the preferred prey of paddlefish. Bigmouth buffalo diet was broad, including both rotifers and crustacean zooplankton. Dietary overlap with Asian carp was greatest for gizzard shad followed by bigmouth buffalo, but we found little diet overlap for paddlefish. Diet similarity based on taxonomy correlated strongly with diet similarity based on size suggesting filtration efficiency influenced the overlap patterns we observed. Although rotifers were the most common prey item consumed by both bighead and silver carp, we found a negative relation between silver carp CPUE and cladoceran density. The competitive effect of Asian carp on native fishes may be forestalled because of the high productivity of Illinois and Mississippi river habitats, yet the potential for negative consequences of Asian carp in less productive ecosystems, including Lake Michigan, should not be underestimated.

Hybridization, polyploidy and invasion: lessons from Spartina (Poaceae)

Abstract: In this paper, we examine how the Spartina system has helped our understanding of the genomic aspects of allopolyploid speciation in the context of biological invasion. More specifically the respective roles of hybridization and genome duplication in the success of newly formed allopolyploid species are explored. Hybridization appears to have triggered genetic and epigenetic changes in the two recently formed European homoploid hybrids S. × towsendii and S. × neyrautii. Deviation from parental structural additivity is observed in both hybrids, with different patterns when considering transposable element insertions or AFLP and methylation alteration. No important changes are observed in the invasive allopolyploid Spartina anglica that inherited the identical genome to S. × townsendii. The repeated rRNA genes are not homogenized in the allopolyploid, and both parental repeats are expressed in the populations examined. Transcriptomic changes suggest possible gene silencing in both hybrids and allopolyploid. In the long-term of evolutionary time, older hexaploid Spartina species (Spartina alterniflora, Spartina maritima and Spartina foliosa) appear to have selectively retained differential homeologous copies of nuclear genes. Waxy gene genealogies suggest a hybrid (allopolyploid) origin of this hexaploid lineage of Spartina. Finally, nuclear and chloroplast DNA data indicate a reticulate origin (alloheptaploid) of the invasive Spartina densiflora. All together these studies stress hybridization as a primary stimulus in the invasive success of polyploid Spartina species.

Predicting establishment success for alien reptiles and amphibians: a role for climate matching

Abstract: We examined data comprising 1,028 successful and 967 failed introduction records for 596 species of alien reptiles and amphibians around the world to test for factors influencing establishment success. We found significant variations between families and between genera. The number of jurisdictions where a species was introduced was a significant predictor of the probability the species had established in at least one jurisdiction. All species that had been introduced to more than 10 jurisdictions (34 species) had established at least one alien population. We also conducted more detailed quantitative comparisons for successful (69 species) and failed (116 species) introductions to three jurisdictions (Great Britain, California and Florida) to test for associations with climate match, geographic range size, and history of establishment success elsewhere. Relative to failed species, successful species had better climate matches between the jurisdiction where they were introduced and their geographic range elsewhere in the world. Successful species were also more likely to have high establishment success rates elsewhere in the world. Cross-validations indicated our full model correctly categorized establishment success with 78–80% accuracy. Our findings may guide risk assessments for the import of live alien reptiles and amphibians to reduce the rate new species establish in the wild.

Fish fauna destruction after the introduction of a non-native predator ( Cichla kelberi ) in a Neotropical reservoir

Abstract: In South America, the introduction of peacock-bass (Cichla), a voracious predator fish, has been an underestimated threat for native fish communities. Although this predator is widespread in many reservoirs, few studies have explored its impact on biodiversity. To investigate the relationship between invasion and fish diversity, the present study followed a natural experiment in the Rosana Reservoir (Parana River basin), where Cichla kelberi were introduced in 2004. We monitored fish assemblages associated with submerged macrophytes between 2003 and 2007, using a 1 m2 throw trap. In the years following the introduction, fish diversity dramatically changed. For example, in March 2007, mean fish density and richness were reduced by ca. 95 and 80%, respectively, and many small-sized species had vanished. One aspect was the gradual change of biodiversity, which unfolded at two times during each year: (1) impacts during summer/autumn periods, which coincided with large shoals of young C. kelberi in the patches; and (2) assemblage recovery during the spring. The sequence of extinction-colonization events, however, might not be able to maintain fish assemblages due to the decrease in recovery intensity each spring; assuming a constant decline rate in the coming years, we predict complete assemblage extinction by the summer of 2010. Results from this natural experiment provided evidence supporting the collapse of fish assemblages soon after the introduction of C. kelberi. Such rapid destruction (2 years) reveals an important homogenizing force behind this predator and stresses the need for control measures that prevent new transferences among South American basins.

The habitat and conduit functions of roads in the spread of three invasive plant species

Abstract: Nonnative plant species commonly occur along roadsides, and populations are often assumed to invade by spread along the road axis. To distinguish between the function of roadsides as movement corridors and as habitat, nonnative plant species were surveyed along roads in deciduous forest sites in southeastern Ohio, USA. The importance of road proximity was tested by comparing nonnative species abundance in 100 m transects along roads with transects in undisturbed forest. Nonnative species were most abundant and most frequently observed in roadside sites in valleys. Three common species were chosen for closer scrutiny. In a seed sowing experiment roads and open sites proved to be better locations for the germination and growth of Microstegium vimineum than non-roadside and closed-canopy sites. Tussilago farfara and Rosa multiflora occurred in a small number of disjunct patches suggesting infrequent arrival in the sampled transects. Both species were strongly clustered at scales consistent with diffusive spread by vegetative growth and short-range seed dispersal. Comparisons of distributions parallel and perpendicular to roads showed no evidence for enhanced dispersal along the road axis. Microstegium distributions were correlated with local light availability implying site saturation. Microstegium micro-distributions suggested that spread along the road axis was facilitated by movement of dormant seeds in road maintenance. Thus, roadsides appear to function as both habitat and a conduit for population expansion, with the rate of spread dependent on the life history of the individual species. These results suggest a hierarchical process of regional invasion, with different dispersal mechanisms functioning at different spatial scales.

Review of impacts of the introduced house mouse on islands in the Southern Ocean: are mice equivalent to rats?

Abstract: Research on the impacts of house mice Mus musculus introduced to islands is patchy across most of the species’ global range, except on islands of the Southern Ocean. Here we review mouse impacts on Southern Ocean islands’ plants, invertebrates, land birds and seabirds, and describe the kinds of effects that can be expected elsewhere. A key finding is that where mice occur as part of a complex of invasive mammals, especially other rodents, their densities appear to be suppressed and rat-like impacts have not been reported. Where mice are the only introduced mammal, a greater range of native biota is impacted and the impacts are most severe, and include the only examples of predation on seabird eggs and chicks. Thus mice can have devastating, irreversible and ecosystem-changing effects on islands, impacts typically associated with introduced rats Rattus spp. Island restoration projects should routinely include mouse eradication or manage mouse impacts.

Consensual predictions of potential distributional areas for invasive species: a case study of Argentine ants in the Iberian Peninsula

Abstract: Invasive species are known to influence the structure and function of invaded ecological communities, and preventive measures appear to be the most efficient means of controlling these effects. However, management of biological invasions requires use of adequate tools to understand and predict invasion patterns in recently introduced areas. The present study: (1) estimates the potential geographic distribution and ecological requirements of the Argentine ant (Linepithema humile Mayr), one of the most conspicuous invasive species throughout the world, in the Iberian Peninsula using ecological niche modeling, and (2) provides new insights into the process of selection of consensual areas among predictions from several modeling methodologies. Ecological niche models were developed using 5 modeling techniques: generalized linear models (GLM), generalized additive models (GAM), generalized boosted models (GBM), Genetic Algorithm for Rule-Set Prediction (GARP), and Maximum Entropy (Maxent). Models for the eastern and western portions of the Iberian Peninsula were built using subsets of occurrence and environmental data to investigate the potential for ecological niche differences between the invading populations. Our results indicate geographic differences between predictions of different approaches, and the utility of ensemble predictions in identifying areas of uncertainty regarding the species’ invasive potential. More generally, our models predict coastal areas and major river corridors as highly suitable for Argentine ants, and indicate that western and eastern Iberian Peninsula populations occupy similar environmental conditions.

Invaders are not a random selection of species

Abstract: We assembled information on 119 species of freshwater macroinvertebrate invaders in North America and Europe, and compared them to all native freshwater species in North America and Europe. We tested whether the invaders were a random or selected group among taxa (phylum or class), water quality requirements, and feeding habit. We found that freshwater macroinvertebrate invaders are not a random selection of species, and are over-represented by molluscs and crustaceans, while taxa richness of native communities are dominated by insects. Over 35% of native species of aquatic invertebrates in North America are only able to live in areas with excellent or very good water quality, and are intolerant of organic pollution. In contrast, all invaders are tolerant of at least moderate amounts of organic pollution. There was a significant difference in the distribution of feeding habits between native species and invaders: collector-filterers (including suspension feeders) were 2.5-3 times more abundant, and predators were 3-4 times less abundant among invaders than among native invertebrates. The ongoing spread of exotic species affects the biodiversity of selected taxa, shifts communities toward greater tolerance of organic pollution and increases the numbers of suspension feeders, thereby enhancing benthic pelagic coupling in waterbodies with high densities of invaders. Because these processes are very similar in Europe and North America, we suggest that the observed patterns may have a common global effect.

The perfect invader: a parthenogenic crayfish poses a new threat to Madagascar’s freshwater biodiversity

Abstract: In 2007 an unusual crayfish found in food markets in the capital of Madagascar was preliminarily identified as Procambarus ‘Marmorkrebs’: a new world taxa and the only decapod known to reproduce by parthenogenesis. We present information on the identity, distribution and ecology of this recent invader and attempt to evaluate the threat it poses to Madagascar’s biodiversity and to livelihoods. The species appears to be currently limited to the area close to Antananarivo, but is being sold alive on major transport routes. We present molecular evidence of its taxonomic relationships and confirm that the Procambarus present in Madagascar is indeed the parthenogenic taxa. We investigate its reproductive ecology and find Procambarus ‘Marmorkrebs’ to have an extremely high fecundity; more than six times that of the native crayfish Astacoides. The limited evidence we have suggests that this species poses a serious threat to freshwater biodiversity and that it is likely to damage human livelihoods (through its impact on fishing and possibly rice agriculture). More research is urgently needed but in the meantime action is needed to reduce the rate of spread before it is too late.

Belowground mutualists and the invasive ability of Acacia longifolia in coastal dunes of Portugal

Abstract: The ability to form symbiotic associations with soil microorganisms and the consequences for plant growth were studied for three woody legumes grown in five different soils of a Portuguese coastal dune system. Seedlings of the invasive Acacia longifolia and the natives Ulex europaeus and Cytisus grandiflorus were planted in the five soil types in which at least one of these species appear in the studied coastal dune system. We found significant differences between the three woody legumes in the number of nodules produced, final plant biomass and shoot 15N content. The number of nodules produced by A. longifolia was more than five times higher than the number of nodules produced by the native legumes. The obtained 15N values suggest that both A. longifolia and U. europaeus incorporated more biologically-fixed nitrogen than C. grandiflorus which is also the species with the smallest distribution. Finally, differences were also found between the three species in the allocation of biomass in the different studied soils. Acacia longifolia displayed a lower phenotypic plasticity than the two native legumes which resulted in a greater allocation to aboveground biomass in the soils with lower nutrient content. We conclude that the invasive success of A. longifolia in the studied coastal sand dune system is correlated to its capacity to nodulate profusely and to use the biologically-fixed nitrogen to enhance aboveground growth in soils with low N content.

Asexual spread versus sexual reproduction and evolution in Japanese Knotweed s.l. sets the stage for the “Battle of the Clones”

Abstract: The invasion of NW Europe by a single clone of male-sterile Fallopia japonica var. japonica by purely vegetative reproduction has demonstrated that reproduction by seed and the genetic diversity associated with it may not be an essential feature of plant invasions, at least in the short term. What is less well known is that a significant proportion of the Japanese Knotweed s.l. involved is not F. japonica var. japonica, but the hybrid between it and F. sachalinensis–F. × bohemica. This hybrid is able to backcross to either parent with the potential to replace the missing male F. japonica; by the same process, the hybrid is generating the genetic diversity so conspicuously lacking in F. japonica. In terms of understanding the population structure in a particular country, it is important to be able to identify hybrids and putative back-crosses. We bring together a mixture of published and unpublished information to provide a comprehensive section on morphological and anatomical aids to identification. Regeneration of plants from stem and rhizome fragments varies by taxon, as do responses to control techniques, underlining the importance of correct identification of these plants. In this paper we look closely at this group of taxa, with special emphasis on the role played by hybridization, with an examination of the genetic make up of seedlings produced in the wild or by artificial hybridizations, and the implications that this will have on the future directions of the invasion process.

Increasing southern invasion enhances congruence between endemic and exotic Mediterranean fish fauna.

Abstract: Species movements in relation with global warming may increase the spatial overlap between exotic and endemic species, which is a critical issue for the conservation of biodiversity. The Mediterranean Sea, which is a receptacle for exotic species while being a hotspot for endemism, provides exceptional material for a case study. The aim of our study was to quantify (i) the increasing invasion from southern fish exotic species (Red Sea and Atlantic Ocean) that the Mediterranean biota is experiencing and (ii) the spatial overlap between exotic and endemic Mediterranean fish fauna following the northward movement of exotic species within the Mediterranean Sea in the context of global warming. The historical invasion dynamic of exotic fish species and the sea surface temperature series were reconstructed from 1810 to 2006 in order to estimate the correlation between invasion rate and climate. The geographical distributions of exotic and endemic fish richness before and after the period of global warming were used to assess the dynamic of spatial congruence. The results revealed (i) an acceleration of successful introductions from the Red Sea and (ii) the introduction of Atlantic species from lower latitudes in correlation with the increasing temperature of the Mediterranean Sea. We also showed an increasing overlap between the spatial distributions of endemic and exotic species richness. Taken together, our results suggest that endemic fish species are facing a growing number of exotic species because the Mediterranean Sea is acting as a catchment basin for southern species.

Review of negative effects of introduced rodents on small mammals on islands

Abstract: In this first comprehensive review of negative effects of introduced rodents on insular small mammals, the focal species Rattus rattus, R. norvegicus, R. exulans and Mus musculus are implicated in at least 11 extinctions. Furthermore, removal experiments, eradication campaigns and control programmes provide evidence for negative effects on extant populations. While data are currently insufficient for meaningful generalisation with regard to the most threatening rodents, the most threatened small mammals, and the true extent of the problem, it is interesting that R. rattus is implicated in the majority of impacts. This may be explained by its extensive distribution and ecological plasticity. I conclude with methodological recommendations to guide data collection for impact quantification and the study of impact mechanism. This information should facilitate the prioritisation and justification of eradication campaigns, control programmes and biosecurity measures while ensuring that much-needed attention is paid to the conservation of insular small mammals.

Modelling non-equilibrium distributions of invasive species: a tale of two modelling paradigms

Abstract: Invasive species, biological control and climate change are driving demand for tools to estimate species’ potential ranges in new environments. Flawed results from some tools are being used to inform policy and management in these fields. Independent validation of models is urgently needed so we compare the performance of the ubiquitous, logistic regression and the CLIMEX model in predicting recent range extensions of the livestock tick, Rhipicephalus (Boophilus) microplus, in Africa. Both models have been applied to the tick so new, independent data can be used to test their ability to model non-equilibrium distributions. Logistical regression described the spatial data well but failed to predict the range extensions. CLIMEX correctly predicted the extensions without fitting the non-equilibrium data accurately. Our results question the validity of using descriptive, statistical models to predict changes in species ranges with translocation and climate change. More test cases that include independent validation are needed.

Soil recovery after removal of the N2-fixing invasive Acacia longifolia: consequences for ecosystem restoration

Abstract: Invasion by Acacia longifolia alters soil characteristics and processes. The present study was conducted to determine if the changes in soil C and N pools and processes induced by A. longifolia persist after its removal, at the Sao Jacinto Dunes Nature Reserve (Portugal). Some areas had been invaded for a long time (>20 years) and others more recently ( 30%, β-glucosaminidase activity (N mineralization index) >60% and potential nitrification >95%. Removal of plants and litter resulted in a >35% decrease in C and N content after four and half years. In recently invaded areas, β-glucosaminidase activity and potential nitrification showed a marked decrease (>54% and >95%, respectively) after removal of both A. longifolia and litter. Our results suggest that after removal of an N2-fixing invasive tree that changes ecosystem-level processes, it takes several years before soil nutrients and processes return to pre-invasion levels, but this legacy slowly diminish, suggesting that the susceptibility of native areas to (re)invasion is a function of the time elapsed since removal. Removal of the N-rich litter layer facilitates ecosystem recovery.

Distribution and impacts of invasive earthworms in Canadian forest ecosystems

Abstract: In Canada it is generally accepted that most indigenous earthworms did not survive glaciation, and that the majority of the earthworms now inhabiting Canadian soils are relatively recent introductions of European origin. Although these exotic earthworms are generally considered to be beneficial in agricultural soils, their effects can be less benign in forested ecosystems. Studies have shown that invading earthworms can significantly alter the forest floor, affecting the distribution of carbon, nitrogen and other chemicals, roots, microbes and other elements of the soil fauna, and even understory vegetation. This paper summarizes the current distribution of exotic and native earthworm species in Canadian forests and draws on the results of studies of invasion patterns and environmental impacts in northern forests in North America and Europe to discuss potential outcomes for forests in Canada. The potential for variables such as temperature, pH, litter palatability and dispersal, to limit or promote the invasion of exotic earthworms in Canadian forests is discussed, and areas for future research are proposed. The same earthworm species that are invading forests in northern Europe and the US are also invading Canadian forests. Several species of exotic earthworms are already established in a wide range of deciduous and coniferous forest types, including the boreal. Evidence is presented to suggest that further expansion into Canadian forests is likely.

Introgression between invasive saltcedars (Tamarix chinensis and T. ramosissima) in the USA

Abstract: Saltcedars (Tamarix ramosissima and T chinensis) are native to Asia, but since introduction into the USA have become common and invasive in many western riparian habitats Recent molecular analysis of a single locus nuclear DNA sequence marker has shown that in their native range the two species are genetically distinct, but within the USA populations many of the plants (23%) are novel hybrids Here, we used multilocus DNA markers (amplified fragment length polymorphisms) to determine the levels of introgression in USA plants Species-specific diagnostic markers, principal coordinates analysis, and a Bayesian model-based clustering analysis all indicate a much higher incidence of hybridization (83–87%) than was revealed by the single locus marker, with USA plants forming a genetic continuum between the two parental types Additionally, the level of introgression toward Tamarix ramosissima or T chinensis was strongly correlated with latitude Concordance of level of introgression was highest between principal coordinates analysis and the Bayesian analysis The high percentage of novel hybrids may have implications for classical biological control efforts

First evidence of hybridization between endangered sterlets ( Acipenser ruthenus ) and exotic Siberian sturgeons ( Acipenser baerii ) in the Danube River

Abstract: Most natural populations of Eurasian sturgeons have declined dramatically during recent decades, reaching historic low levels today. During the same period, sturgeon has become very popular in European aquaculture. Because many hatcheries are located near rivers, their unintentional escape is often documented, especially during floods. Until now, no cases of successful hybridization of these escaped fish with wild stocks have been reported. In this study, the genetic structure of a highly threatened population of sterlets (Acipenser ruthenus) from the Upper Danube was analysed as a requirement for their conservation. Surprisingly, we observed genotypes and morphotypes of Siberian sturgeon (Acipenser baerii), as well as hybrids between this species and native sterlets. This hybridization poses a serious threat for the survival of this isolated sterlet population in the upper part of the Danube. For the first time, natural reproduction is documented for Siberian sturgeon outside their natural range in Europe. This finding demonstrates the risk of extinction by hybridisation of endangered populations. We would like to stress that taking into consideration the risks for native sturgeon populations, farmed sturgeon should not be released into the wild, and all measures should be taken to prevent their accidental escape.

Direct and indirect effects of rats: does rat eradication restore ecosystem functioning of New Zealand seabird islands?

Abstract: Introduced rats (Rattus spp.) can affect island vegetation structure and ecosystem functioning, both directly and indirectly (through the reduction of seabird populations). The extent to which structure and function of islands where rats have been eradicated will converge on uninvaded islands remains unclear. We compared three groups of islands in New Zealand: islands never invaded by rats, islands with rats, and islands on which rats have been controlled. Differences between island groups in soil and leaf chemistry and leaf production were largely explained by burrow densities. Community structure of woody seedlings differed by rat history and burrow density. Plots on islands with high seabird densities had the most non-native plant species. Since most impacts of rats were mediated through seabird density, the removal of rats without seabird recolonization is unlikely to result in a reversal of these processes. Even if seabirds return, a novel plant community may emerge.

A hierarchical framework for integrating invasibility experiments incorporating different factors and spatial scales

Abstract: Results from experiments studying different factors determining invasibility (e.g. land use, disturbance, biotic interactions) at different spatial scales are mainly used in isolation, probably because a methodology for integration is lacking. Recent studies show that factors affecting invasibility most likely do so in a hierarchical manner, with different factors acting more strongly at different spatial scales. Climate can be considered the dominant factor at the continental scale, while at regional and landscape scale topography, land cover and land use become increasingly important. At smaller spatial scales, soil type, disturbance, biotic interactions, resources, and microclimate may become significant. In the current paper, we propose a hierarchical framework for combining results from different types of studies. In this hierarchical system, factors operating at a smaller scale are subordinate to factors operating at a larger scale, but if conditions at higher levels are satisfied, the small-scale factors may become indispensable for making accurate predictions. Depending on the aim of the study, the accuracy of prediction can be selected by the researcher, which in its turn determines which data are required. We discuss several applications of the framework and indicate some options for future research. Although the complexity of natural systems presents fundamental limits to predictions, we think this framework can provide a useful tool for the identification of areas of risk for biological invasions, for improving our understanding of invasibility, and for identifying gaps in our current knowledge.

Assessing the residual effects of Carpobrotus edulis invasion, implications for restoration

Abstract: We examined whether the residual effects on soil caused by the invasion of Carpobrotus edulis, common iceplant, would inhibit the reestablishment of a native plant species. Carpobrotus edulis interacts both directly by suppressing the growth and establishment of other plants and indirectly by altering soil chemistry. We tested whether the residual effects of C. edulis resulted in lowered germination, survival, growth, and reproduction of Gilia millefoliata, a rare dune annual. We compared G. millefoliata planted in plots previously occupied by C. edulis to G. millefoliata planted in plots that previously had native vegetation. Each plot received three treatments: seed, transplant, and unplanted, and were censused every three weeks until senescence. Carpobrotus edulis had strong negative effects on the germination, survival, growth, and reproduction of G. millefoliata. C. edulis lowers soil pH and increases organic content due to the recalcitrance of tissue to decomposition, which may have evolved as a mechanism to facilitate recolonization and invasion.

Invaders interfere with native parasite–host interactions

Abstract: The introduction of species is of increasing concern as invaders often reduce the abundance of native species due to a variety of interactions like habitat engineering, predation and competition. A more subtle and not recognized effect of invaders on their recipient biota is their potential interference with native parasite–host interactions. Here, we experimentally demonstrate that two invasive molluscan filter-feeders of European coastal waters interfere with the transmission of free-living infective trematode larval stages and hereby mitigate the parasite burden of native mussels (Mytilus edulis). In laboratory mesocosm experiments, the presence of Pacific oysters (Crassostrea gigas) and American slipper limpets (Crepidula fornicata) reduced the parasite load in mussels by 65–77% and 89% in single and mixed species treatments, respectively. Both introduced species acted as decoys for the trematodes thus reducing the risk of hosts to become infected. This dilution effect was density-dependent with higher reductions at higher invader densities. Similar effects in a field experiment with artificial oyster beds suggest the observed dilution effect to be relevant in the field. As parasite infections have detrimental effects on the mussel hosts, the presence of the two invaders may elicit a beneficial effect on mussels. Our experiments indicate that introduced species alter native parasite–hosts systems thus extending the potential impacts of invaders beyond the usually perceived mechanisms.

The rat and the octopus: initial human colonization and the prehistoric introduction of domestic animals to Remote Oceania

Abstract: Remote Oceania was colonized initially in three migratory phases: the western archipelagos of Micronesia plus eastern Melanesia out to west Polynesia in the period 3500–2800 cal BP (all dates hereafter are cal BP), central and eastern Micronesia 2200–2000 BP and east and south Polynesia 1100–700 BP. The early and late migration phases are best-known archaeologically. During these phases a number of plants and animals were introduced. Of the latter, the pig (Sus scrofa), dog (Canis familiaris), fowl or chicken (Gallus gallus) and rats (Rattus spp., especially R. exulans) were most deliberately associated with human settlement. The pattern of introductions appears to be only partly in agreement with an implication of widespread early distribution derived from the orthodox colonisation model of ‘transported landscape’ coupled with sophisticated seafaring. Within the two main migrations the pattern of introductions is similar. Excepting in the movement to West Micronesia, all four taxa were transported into the islands nearest their proximate sources at, or soon after, the beginning of migration, but their introduction to more remote islands was partial and patchy. Evaluation of invasibility, invasiveness and transportability characteristics amongst the four taxa suggest that island size and complexity, propagule pressure and seafaring capability were important factors in differential distribution and survival. Seafaring capability was especially important because it determined the extent of accessibility to islands near and far and the degree of propagule or introduction pressure that was exerted. Framing the archaeological data within a model of invasion biology offers a richer and more systematic approach to the complexities of introduction than adopting a culture-historical perspective.

Invasion of alien macroalgae in different Mediterranean habitats

Abstract: The study evaluated different macroalgal invasions in the main Mediterranean coastal habitats on hard bottom. Biodiversity, species composition and structure of macroalgal assemblages were compared among non-invaded areas and areas invaded by the Chlorophyta Caulerpa racemosa var. cylindracea and by the turf-forming Rhodophyta Womersleyella setacea in three different habitats: shallow rocky bottom, deep rocky bottom and dead matte of the seagrass Posidonia oceanica. Results showed that alien macroalgae constituted a relevant component of benthic assemblages in invaded areas of the Mediterranean Sea. Assemblages invaded by Womersleyella setacea and Caulerpa racemosa showed lower values of diversity and large differences in the structure and species composition related to non-nvaded assemblages. The species that mostly suffered from invasion were erect species reproducing sexually; moreover, the dominance of W. setacea led to low abundance of native filamentous algae, while C. racemosa colonization seemed particularly threatening for encrusting algae. All the studied habitats appeared highly invasible by alien macroalgae, even if W. setacea appeared more invasive in deeper habitats, while colonization of C. racemosa seemed more serious in shallower habitats; the dead matte of P. oceanica represented a suitable substrate for the spread of both species. Differences among assemblages in different habitats were reduced in invaded areas.