Biological Invasions

Interactions between invasive species can have important consequences for the speed and impact of biological invasions. Containers occupied by the invasive mosquito, Aedes albopictus Skuse, may be sensitive to invasive plants whose leaves fall into this larval habitat. To examine the potential for interactions between invasive leaf species and larval A. albopictus, we conducted a field survey of leaf material found with A. albopictus in containers in Palm Beach County, Florida and measured density dependent responses of A. albopictus larvae to two invasive and one native leaf species in laboratory experiments. We found increased diversity of leaf species, particularly invasive species, in areas further from the urbanized coast, and a significant positive association between the presence of Schinus terebinthifolious (Brazilian pepper) and the abundance of A. albopictus. In laboratory experiments, we determined that larval growth and survivorship were significantly affected by both larval density and leaf species which, in turn, resulted in higher population performance on the most abundant invasive species (Brazilian pepper) relative to the most abundant native species, Quercus virginiana (live oak). These results suggest invasive leaf species can alleviate density dependent reductions in population performance in A. albopictus, and may contribute to its invasion success and potential to spread infectious disease.

When a species is introduced into a new location, it may escape, at least temporarily, from its natural enemies. In field surveys, we found that when the exotic, invasive mosquito, Aedes albopictus, invades new sites, it initially experiences reduced infection by its gut parasite, Ascogregarina taiwanensis. To determine the effect of this escape from parasitism on the competitive ability of A. albopictus, we performed a laboratory competition experiment in which infected and uninfected A. albopictus larvae were reared in microcosms alone and in competition with larvae of the native mosquito, Ochlerotatus triseriatus. We analyzed the effect of parasitism by A. taiwanensis on A. albopictus performance when subjected to intra- and interspecific competition across a range of larval densities, as well as the effect of A. albopictus parasitism by A. taiwanensis on the competitive impact of A. albopictus on O. triseriatus. At a density of 30 O. triseriatus larvae, O. triseriatus survivorship was significantly reduced by the addition of 30 unifected A. albopictus, but not by addition of 30 infected A. albopictus, and not by addition of 15 A. albopictus whether infected or uninfected. Although estimated finite rate of population increase (λ') showed similar trends, and was significantly affected by treatments, no pairwise differences in rate of increase were significant. Infection by A. taiwanensis also significantly prolonged A. albopictus female development time and reduced the intraspecific competitive effect of increased density of A. albopictus, but did not affect A. albopictus survivorship, mass, or estimated finite rate of population increase. Thus, when A. albopictus escapes from this parasite as it colonizes new sites, this escape may give it a small, but significant, added competitive advantage over O. triseriatus, which may facilitate range expansion of A. albopictus and enhance A. albopictus's initial impact on resident species.

Understanding the migration patterns of invasive organisms is of paramount importance to predict and prevent their further spread. Previous attempts at reconstructing the entire history of the sudden oak death (SOD) epidemic in California were limited by: (1) incomplete sampling; (2) the inability to include infestations caused by a single genotype of the pathogen; (3) collapsing of non-spatially contiguous yet genetically similar samples into large meta-samples that confounded the coalescent analyses. Here, we employ an intensive sampling coverage of 832 isolates of Phytopthora ramorum (the causative agent of SOD) from 60 California forests, genotyped at nine microsatellite loci, to reconstruct its invasion. By using age of infestation as a constraint on coalescent analyses, by dividing genetically indistinguishable meta-populations into highly-resolved sets of spatially contiguous populations, and by using Bruvo genetic distances for most analyses, we reconstruct the entire history of the epidemic and convincingly show infected nursery plants are the original source for the entire California epidemic. Results indicate that multiple human-mediated introductions occurred in most counties and that further disease sources were represented by large wild infestations. The study also identifies minor introductions, some of them relatively recent, linked to infected ornamental plants. Finally, using archival isolates collected soon after the discovery of the pathogen in California, we corroborate that the epidemic is likely to have resulted form 3 to 4 core founder individuals evolved from a single genotype. This is probably the most complete reconstruction ever completed for an invasion by an exotic forest pathogen, and the approach here described may be useful for the reconstruction of invasions by any clonally reproducing organism with a relatively limited natural dispersal range.

Hosts of soybean rust (Phakopsora pachyrhizi) are sensitive to low temperatures, limiting this obligate parasite in the United States to overwintering sites in a restricted area along the Gulf Coast. This temperature sensitivity of soybean rust hosts allowed us to study spatial spread of epidemic invasions over similar territory for seven sequential years, 2005-2011. The epidemic front expanded slowly from early April through July, with the majority of expansion occurring from August through November. There was a 7.4-fold range of final epidemic extent (0.4 to 3.0 million km(2)) from the year of smallest final disease extent (2011) to that of the largest (2007). The final epidemic area of each year was regressed against epidemic areas recorded at one-week intervals to determine the association of final epidemic extent with current epidemic extent. Coefficients of determination for these regressions varied between 0.44 to 0.62 during April and May. The correlation coefficients varied between 0.70 and 0.96 from early June through October, and then increased monotonically to 1.0 by year's end. Thus, the spatial extent of disease when the epidemics began rapid expansion may have been a crucial contributor to subsequent spread of soybean rust. Our analyses used presence/absence data at the county level to evaluate the spread of the epidemic front only; the subsequent local intensification of disease could be strongly influenced by other factors, including weather.

The once immense diversity of native Pacific island land snail species, with high single island or archipelago endemism, is declining dramatically. The native/endemic species are being replaced by a much smaller number of widespread tropical tramps, that is, those species that are most readily transported by humans. The 82 introduced (including 14 cryptogenic) land snail species recorded include some that were distributed accidentally by Pacific islanders before European exploration of the Pacific and that are now widespread. However, the majority are modern introductions, with many recent accidental introductions often associated with the horticultural trade. Native freshwater faunas were less diverse than the terrestrial faunas and exhibited much lower endemism. Among the 59 alien freshwater species recorded (including 38 cryptogenic species), the most diverse and widespread are the thiarids. Predation by and competition with these aliens (as well as habitat loss) are probably important mechanisms underlying the loss of native taxa, but almost no quantitative or experimental work has been done to demonstrate such ecological interactions. Prevention of further spread and of new introductions should be the main approach. Increased public education and development of public trust is essential to the success of these efforts.

The rusty crayfish, Orconectes rusticus, is one of America’s best-known non-indigenous crayfishes, having been identified as extirpating native crayfishes and disrupting local aquatic ecosystems. Over the past 40–50years, rusty crayfish have spread from its historical range in the Ohio River drainage (U.S.A.), to waters throughout much of Illinois, Michigan, Wisconsin, and Minnesota and parts of 11 other states, Ontario (Canada) and the Laurentian Great Lakes. Using a comprehensive dataset based on all known historical records and extensive present-day surveys (n = 2775) this study reports on the invasion history of rusty crayfish, with observations on concomitant declines of native crayfishes in Wisconsin over the past 130years (1870–2004). We found that rusty crayfish occurrences have increased from 7% of all crayfish records collected during the first 20years of their invasion (1965–1984) to 36% of all records during the last 20years, and that rusty crayfish have replaced the northern clearwater crayfish (O. propinquus) and virile crayfish (O. virilis) as the most dominant member of the contemporary crayfish fauna. In light of our results we discuss the introduction, establishment and integration phases of the rusty crayfish invasion and provide preliminary predictions of the potential distribution of rusty crayfish in Wisconsin lakes based on critical environmental requirements.

The many qualifying terms attributed to invasive species reveal the lack of precision surrounding the notion of biological invasion itself. In spite of several proposed definitions, some basic disagreements persist concerning characterization of the phenomenon. These primarily arise from the lack of pertinence of both of the main current criteria—the geographic (or biogeographic) criterion and the impact criterion—to what is really intended by “invasion.” Faced with this situation, it seems preferable to adopt an ontological approach allowing a return to the basic principles of the elaboration of a definition. Starting with the nature of the phenomenon itself (i.e., its essence), we try to elucidate the notion of biological invasion and we suggest a general definition compatible with most of the ideas already expressed.

China’s current invasive alien plant species were analyzed with regard to their floristic status, biological attributes and invasion status elsewhere. Most of the 270 species identified were annuals, followed by perennial herbs. Woody perennials made only about 10% of the species. The invasives were comprised of 59 families, the largest being Asteraceae, Poaceae, and Brassicaceae. The genera with most invasive species were Amaranthus, Ipomoea, and Solanum. Most of the species originated from the New World, notably from South America. About one-third of the species were serious invaders of natural habitats in countries other than China. The proportion of invasive alien plants in province floras ranged from 0.5 to 3.8%, absolute numbers from nine to 117 species per province. Density of invasive species was correlated positively with native species density at provincial scale. The results demonstrate that in China invasive plants are present throughout the country, with a particularly high species richness in the Southeast. The ecological diversity of invasive plants suggests wide ranging impacts which need to be assessed.

Small seed mass is regarded as a robust trait related to invasion success, especially in pines. However, few studies have explored whether invasiveness related to small seed mass is also associated to low levels of seed predation in the recipient community. We conducted field cafeteria seed removal experiments comparing 16 Pinus species that differ in seed mass to test if seed removal might impose biotic resistance to Pinus spp. and if there are differences between species related to seed mass. Seeds were removed rapidly and in high proportion. In the Mediterranean shrublands, where the experiments were conducted, rodents and ants were the main seed removers. Mean seed survival time was significantly different between species. However, smaller seeds were not the most predated. Our study suggests that, in pine species with high invasiveness, the potential higher seed removal of small seeds can be counterbalanced by larger seed crops. KeywordsConifers-Biotic resistance-Invasiveness- Pinus spp.-Seed mass-Seed removal

Despite the substantial impacts of nonindigenous plant pests and weeds, relatively little is known about the pathways by which these organisms arrive in the U.S. One source of such information is the Port Information Network (PIN) database, maintained by the U.S. Department of Agriculture, Animal and Plant Health Inspection Service (APHIS) since 1984. The PIN database is comprised of records of pests intercepted by APHIS personnel during inspections of travelers’ baggage, cargo, conveyances and related items arriving at U.S. ports of entry and border crossings. Each record typically includes the taxonomic identify of the pest, its country of origin, and information related to the commodity and interception site. We summarized more than 725,000 pest interceptions recorded in PIN from 1984 to 2000 to examine origins, interception sites and modes of transport for nonindigenous insects, mites, mollusks, nematodes, plant pathogens and weeds. Roughly 62% of intercepted pests were associated with baggage, 30% were associated with cargo and 7% were associated with plant propagative material. Pest interceptions occurred most commonly at airports (73%), U.S.-Mexico land border crossings (13%) and marine ports (9%). Insects dominated the database, comprising 73 to 84% of the records annually, with the orders Homoptera, Lepidoptera and Diptera collectively accounting for over 75% of the insect records. Plant pathogens, weeds and mollusks accounted for 13, 7 and 1.5% of all pest records, respectively, while mites and nematodes comprised less than 1% of the records. Pests were intercepted from at least 259 different locations. Common origins included Mexico, Central and South American countries, the Caribbean and Asia. Within specific commodity pathways, richness of the pest taxa generally increased linearly with the number of interceptions. Application of PIN data for statistically robust predictions is limited by nonrandom sampling protocols, but the data provide a valuable historical record of the array of nonindigenous organisms transported to the U.S. through international trade and travel.

The presence and distribution in Argentina of the invasive snail species Theba pisana is presented on the basis of a survey carried out in 10 beach resorts along the Atlantic coast of Buenos Aires Province. Additionally, complementary information and complete revision of material of other exotic terrestrial gastropods housed in museum collections was carried out, reporting herein seven new species for Argentina (Otala punctata, Hawaiia minuscula, Paralaoma servilis, Opeas goodalli, Vallonia pulchella, Vertigo ovata and Pupisoma dioscoricola), one for Colombia (Subulina octona), and two for Peru (S. octona and P. dioscoricola). At present, 42 introduced species of terrestrial gastropods have been recorded in nine countries of South America (Argentina, Bolivia, Brazil, Chile, Colombia, Paraguay, Peru, Uruguay and Venezuela). The most likely pathways for introduction of terrestrial gastropods, at least in Argentina, are horticultural development and urban and suburban transformation of original natural habitats. KeywordsAlien species-Argentina-Land molluscs- Theba pisana

In this paper we provide the first report of the presence of exotic Chinook salmon (Oncorhynchus tshawytscha) in two rivers off the Beagle Channel, Lapataia and Ovando, in southern Tierra del Fuego. We also confirm that successful reproduction occurred in the fall of 2007, as we captured yearlings in freshwater. Scale pattern analyses of adult fish caught were all of the “stream” ecotype, with ages ranging between 3 and 5 (average 4.2year). Stable isotope analysis of Ovando-Lapataia Chinook population indicates general patterns consistent with those of other populations in the region, but characteristically enriched levels of C indicates a distinct ocean feeding location as compared to Atlantic populations in the Santa Cruz River. Two different haplotypes, one identical to the unique haplotype of the Caterina River population, were found in the Ovando-Lapataia rivers, providing partial evidence for some level of contemporary segregation between these two populations. As an exotic species, Chinook salmon have been able to use the ocean as a waterway to rapidly colonize new habitats both in New Zealand and in several Pacific and Atlantic river basins of continental Patagonia. This record expands the known distribution of this species in Patagonia further south and into the Island of Tierra del Fuego. Its presence in the Beagle Channel creates the conditions for its expansion to a significant collection of new rivers, as well as to adjacent marine areas in and around the Southern Fuegian Channels. Our results provide support to the idea that, in practice, no district of Patagonia is sheltered from the colonization by invasive anadromous Salmonids. KeywordsChinook salmon-Beagle Channel-Patagonia-Exotic anadromous salmon

Ecological niche modeling based on surface seawater (SST) and atmospheric (AT) temperature records was used to predict the potential range of distribution of Crassostrea gigas, focusing on the South American coast as a study case. In its native range, the species maintains self-sustaining populations at thermal regimes with mean SST ranging from 14.0° to 28.9°C for the warmest month and from −1.9° to 19.8°C for the coldest month of the year. Settlement is also constrained to mean AT varying between 15° and 31°C for the warmest month and between −23° and 14°C for the coldest month of the year. Latitudinal limits for the species’ distribution in South America predicted by the analysis of AT regimes were Valdivia-Chiloe Island (39.8º–42.0°S, Chile) and Pisco (13.4°S, Peru) on the Pacific coast, and San Julian port (49.3°S) (Argentina) and Garopaba-Rio Grande (28.0°–32.0°S, Brazil) on the Atlantic coast. Geographical limits of distribution predicted by analysis of SST regimes were Chiloe Island (42.0°S, Chile) and Mancora (4.1°S, Peru) on the Pacific coast, and Puerto Deseado ria (47.7°S, Argentina) and Paranaguá (25.7°S, Brazil) on the Atlantic coast. Therefore, SST regimes would expand the potential range on the Pacific coast equatorward relative to AT. KeywordsEcological niche modeling-Non-indigenous species-Temperature range-Ecological risk-Oyster introduction

The African clawed frog Xenopus laevis has been introduced at several locations, including Mediterranean climate-type regions. In 2006, several individuals of this species were found inhabiting a stream at Oeiras, about 20kmW of Lisbon, Portugal. Although the place and time of introduction are not clearly identified, there are reasons to believe that this population may be the result of an almost 30-year-old introduction proceeding from research laboratories located nearby. Field surveys were conducted in 2007 and 2008 on the distribution and abundance of this species in the region. The species was found in two streams, about 5km from each other. While being locally abundant, the adults of X. laevis are smaller than those from South Africa and California. In spite of an abundant production of eggs and tadpoles at one of the streams, no tadpoles were found in advanced developmental stages. Until now, most individuals were found in heavily urbanized areas that should constrain their ability to cross overland to other water bodies. An eradication program may be feasible, but the presence of adults on two streams indicates that the species may be expanding, in spite of the urban landscape. KeywordsAmphibian-Iberian Peninsula-Urban-Distribution-Reproduction

The American bullfrog (Lithobates catesbeianus) has been introduced throughout the world over the past two centuries. This taxa is a potentially devastating invader because of its large size, prolific reproductive output, and broad ecological niche. Consequently, the extent of this ongoing biological invasion is an increasing conservation concern. In Argentina, several introduced populations have been reported. In most cases, these introductions have been intentional or incidental releases from breeding facilities, yet the consequences and effective controls for captive-breeding programs have not been assessed by government environmental agencies. Further studies are needed regarding the trophic ecology, reproductive biology, ecological niche, and chytrid fungus infection prevalence to predict the ultimate impacts of this species on native ecosystems. The aim of this work is to report a new alien population of L. catesbeianus at La Candelaria, Salta province, Argentina. This record represents the first population of American bullfrogs detected in northwestern Argentina. KeywordsAmerican bullfrog- Lithobates catesbeianus -Invasive introduced species-Salta province-Argentina

The screening of 2,735 eels from European waters and aquaculture farms was conducted using mitochondrial Cytochrome b and 16S rRNA gene fragments amplified by polymerase chain reaction. Reaction products were either sequenced directly or subjected to analysis using restriction fragment length polymorphism which resulted in species-specific restriction patterns. Beside the expected European eel, Anguilla anguilla (Linnaeus, 1758), the American eel, Anguilla rostrata (Le Sueur, 1817), was also identified in samples from both aquaculture (N=40 out of 1,025) and from natural waters (N=44 out of 1,710). The life stages of American eels identified from several German waters draining to either the Baltic Sea and the North Sea ranged from elver to silver eels. This indicates that stocking with glass eels or elvers must have occurred several times most likely in the period from 1998 to 2002. The application of a fast and precise method for species identification and genetic monitoring of eels delivered for stocking is therefore essential for maintaining the autochthonous species composition in future.

The North American beaver (Castor Canadensis) was introduced into Isla Grande de Tierra del Fuego, Argentina in 1946 as a potential source of wild fur. The species showed high growth potential, reaching close to 100,000 individuals from an original founding stock of 25 females and 25 males. Beavers adapted rapidly to their new environment and became invasive, providing an excellent model of successful adaptation of introduced populations to a new habitat. In this study, we used polymorphic mitochondrial (mt) DNA to evaluate genetic variation in the introduced beaver population from Tierra del Fuego. Nucleotide variation in partial sequences of Cytochrome b (500bp) and 12S rRNA (421bp) genes and the main non-coding D-loop region (521bp) were analyzed. Our study allowed to identify 10 different mtDNA lineages in the invasive population, none of them shared among the source populations. The pattern observed is a consequence of cessation of gene flow following expansion of the founding beaver population since the time of its introduction. This approach contributes to the understanding of effects of genetic changes on survival ability and reproductive success of invasive species. It also has important management implications to invasive species.

We record here the invasion of the African sharp-tooth catfish Clarias gariepinus in the South China biodiversity hotspot, an area rich in endemic and endangered fish fauna. C. gariepinus was introduced into the natural systems as escapees from aquaculture ponds. These catfishes are very large, top predators, and thus have the potential to cause serious threats to the native fish fauna. The impact of C. gariepinus needs more investigation with emphasis on developing techniques for controlling its dispersal. KeywordsAlien species–African catfish–Invasion–Control measures–South China

We record here the introduction of the African catfish Clarias gariepinus into the Guaraguaçu River basin in Paraná State, Brazil, an area with an extremely rich endemic fish fauna, including many catfishes. C. gariepinus was introduced as escapees from ponds built for recreational angling. These catfishes are very large and hard predators, thus posing serious potential for impact on the native fish fauna. The impact of C. gariepinus needs study with emphasis on finding means for controlling its spread.

In their native ranges, muricid gastropods feed on similar prey, often bivalves and barnacles, which they usually drill. Throughout its wide southern Australian distribution, the intertidal Lepsiella vinosa feeds on a range of prey from barnacles and littorinid gastropods in the southeast to mussels in the southwest. A number of muricids have been introduced throughout the world, either with oysters or in ship ballast water. In their new environments, they switch to native prey but their feeding responses to them have never been studied in the laboratory. The object of this study was to study the feeding of L.vinosa on a suite of non-native species. Australian Lepsiella vinosa was taken to Hong Kong, offered five different possible prey species and allowed to feed to satiation for many weeks. Replacement of consumed prey items by similar-sized conspecifics was undertaken until trends emerged. Lepsiella vinosa readily attacked thick- and thin-shelled bivalves from Hong Kong’s sandy beaches, Anomalocardia squamosa and Caecella chinensis, respectively, and from rocky shores, Septifer virgatus and Hormomya mutabilis, again respectively. It attacked them all, as it does its major prey item, Xenostrobus inconstans, in its native southwestern Australia, by either drilling or marginal probing with its proboscis. It also preferred intermediate sized prey (10–15mm shell length), as with its natural prey. It quickly attacked the sandy shore species, and Hormomya mutabilis and Mytilopsis sallei, the latter two being closely similar in shell form and size to its natural prey X.inconstans. Hormomya mutabilis was the most favored prey, and was most similar in shell form and thickness to X.inconstans. This study therefore suggests that if introduced elsewhere, L.vinosa could radically affect intertidal community structure.

This paper records the new occurrence of spaghetti bryozoan Zoobotryon verticillatum at the Port of Natal, Rio Grande do Norte, Brazil. The study, carried out between 2006 and 2007, also monitored its proliferation. Six observation stations were selected from the estuarine area, as well as samples of benthic invertebrates. The species was initially detected on the pilings of the port and on the hull of a fishing boat. 9months later it was also found in four stations and on another fishing boat. The luxuriant colonial growth in the lower intertidal zone by the end of the study indicates that this species is well established in the estuarine area. This observation is consistent with the species’ biological characteristics; thus, it can be defined as an invasive organism due to its aggressive behavior when occupying the substrate. Its presence in port installations and vessels provide evidence that biofouling on ships’ hulls has been the most likely vector of introduction. Keywords Zoobotryon verticillatum -Introduced species-Harbor area-Vessels’ hull-Brazil

In 2006 an unusual find of Procambarus clarkii, today considered one of the most dangerous exotic crayfish, was made within the preserved brackish Mediterranean biotope “Palude di Torre Flavia” (Rome, Italy). Here the investigation on the population structure and dynamics of P. clarkii inhabiting this brackish wetland is presented in an attempt to describe the life history pattern in this type of habitat. The present study showed that this species has a lower growth rate and higher longevity than other studied European P. clarkii populations, and that it can damage brackish habitats and human livelihoods. The main results of this study chiefly concern the differences found between several life-history parameters of the studied population and those obtained in previous studies. These results showed the high ecological plasticity of P. clarkii, indicate the need to intervene before the species spreads further in brackish water, for the conservation of Mediterranean coastal habitats. Keywords Procambarus clarkii -Red swamp crayfish-Cambaridae-Life history pattern-Coastal habitat

The red crayfish Procambarus clarkii, which is native to southcentral USA and northeastern Mexico, has been successfully introduced into several countries around the world. This study documents the geographic expansion of the exotic red crayfish in Mexico and discusses the consequence of a greater propagation of this species in Mexican inland waters. New state records of this crayfish in the Baja California peninsula and in the states of Durango and Sinaloa indicate its progressive dispersion. The propagation of P.clarkii in Mexico has been caused mainly by human introduction, but it is also facilitated because of the species’ tolerance to an ample range of environmental conditions. Because of the invasive capability of P.clarkii, we suspect that this exotic species is competing for habitat and food with native freshwater shrimp of the genus Macrobrachium in many sites of northern Mexico.

Eriocheir sinensis H. Milne Edwards, 1853 is on the list of top 100 invaders compiled by the International Union for Conservation of Nature and Natural Resources. The recent establishment of a large Chinese mitten crab population in San Francisco Bay and the potential for introductions from California, Asia and Europe pose a significant invasion potential for estuaries and rivers from California to Alaska. This alien species would place at risk the catchment areas of the Pacific Northwest including the economic and social activities that depend upon intact aquatic systems. An analysis of ecological conditions that define the mitten crab’s native and introduced range suggests that large stable estuaries with long flushing times are necessary to sustain significant populations. Most Pacific Northwest estuaries have limited salinity intrusion, estuarine habitat and short flushing times and face a reduced risk of population establishment. Large, stable estuaries, such as the Puget Sound, may support significant populations. River-dominated estuaries, such as the Columbia River, have flushing times less than the duration of larval development and wouldn’t support populations. An application of a temperature based larval development rate to near-shore and estuary sea surface temperatures suggests that estuaries in Oregon and Washington have sufficient thermal regimes to support larval development. Most estuary systems in Alaska have limited periods where water temperatures are above the mortality threshold for the larval stages and are at a low risk for the establishment of populations. A potential sea temperature rise of two degrees Celsius would permit larval development in Alaskan estuaries, where sufficient estuarine and freshwater habitats exist.

Balanus glandula, an east Pacific acorn barnacle from rocky shores, was introduced to Mar del Plata, Argentina more than 40years ago and has spread over 17 latitudinal degrees southward. Here we report the first record of this species living in a soft-bottom environment colonizing the salt marsh plant species Limonium brasiliense, Spartina densiflora, S.alterniflora and Sarcocornia perennis. In addition, we describe the size frequency distribution, density and spatial distribution of the barnacles colonizing the different plant species. The size frequency distribution of Balanus showed a bimodal pattern in all plants. Barnacles were mostly large in S.densiflora, but small in S.alterniflora, with more balanced distributions of small and large barnacles on S.perennis and L.brasiliense. The highest density of barnacles was observed on S.perennis (x=35.8ind/cm2, SD=40.5) and S.alterniflora (x=33.8ind/cm2, SD=23), while the lowest on L.brasiliense (x=1.5ind/cm2, SD=1.18) and S.densiflora (x=0.17ind/cm2, SD=0.09). More than 90% of the barnacles on any given plant were found living. While barnacles colonized only the first few centimeters above the soil surface level in S.alterniflora and L.brasiliense, they reached their highest point on S.perennis. The finding of a rocky shore species successfully colonizing soft-bottom marshes within an invaded region brings new perspectives to discussions in biological invasion ecology, and raises additional considerations for coastal environmental management.

Limnoperna fortunei (Bivalvia, Mitylidae) was introduced into South America in 1991 in the La Plata River (Argentina). It arrived in the ballast water of ships coming from Asia, where this species is native. It was first observed in 1998 in the Paraguay River. Limnoperna was introduced into the Pantanal region as hull fouling of vessels using the Paraguay–Parana waterway. This study describes how L. fortunei came to the Pantanal region, and provides details of its occurrence, density, and impacts. From 1999 to 2002, observations and sampling on natural and artificial substrates in the Paraguay River were made. Some aspects of the spread and impacts, based on local community information, were also analyzed. On artificial substrate the density reached 523.8individualsm−2 and on natural substrate (rocks), up to 10,000individualsm−2 were found. The densities observed were quite low compared to those found in Southern Brazil, where values up to 100,000individualsm−2 have been recorded in the last 3years. In the Paraguay River, the population density of L. fortunei can be negatively impacted by periodic low levels of dissolved oxygen and decreases in pH to between 5 and 6. Such conditions are frequently present during the periodic flooding or inundation of this area. Under these conditions, a high mortality of L. fortunei was recorded in March of 2002, on both natural and artificial substrates. Despite low densities, L. fortunei can colonize water cooling systems of boats, obstructing water circulation and causing motor overheating. Accumulation in water supply equipment, such as pumps and pipes has also been observed.

The serpulid tubeworm Hydroides sanctaecrucis is recorded from tropical northern Australia for the first time. This species is native to the Caribbean, where it is common in coastal lagoons, and has not previously been reliably recorded from the western Pacific. The species was identified after heavy and unusual serpulid fouling was noted on vessel hulls in Cairns, north Queensland, and hull fouling is considered the most probable vector for its translocation to the western Pacific. In northern Australia, H. sanctaecrucis displays similar traits to other fouling and now widely dispersed fouling serpulids such as Hydroides elegans, H. ezoensis and Ficopomatus enigmaticus.

The virile crayfish (Orconectes virilis) represents a cryptic species complex with several lineages known in the USA, and a wide introduced range. In Europe, O. virilis is an emerging invader, established during the last decade in at least two areas—one in the United Kingdom and another in the Netherlands. We assessed the position of both known European populations within the species complex by sequencing part of the mitochondrial gene for cytochrome c oxidase subunit I. Tested UK and Dutch individuals did not belong to any mitochondrial lineage recorded in North America so far but formed a separate clade, the original distribution area of which is unknown. Additionally sequenced virile crayfish from Iowa (USA) also represented a new clade, suggesting that undiscovered lineage variation within O. virilis remains high. This exemplifies that genetic analyses of invading populations may provide new insights into diversity of a taxon in its original range. KeywordsInvasive crayfish-Cryptic diversity-North America-Europe-COI

Populations of the Australian redback spider, Latrodectus hasseltii Thorell 1870, were first recorded in New Zealand in the early 1980s and in Osaka, Japan in 1995. Reliable records suggest that naturalised populations of L. hasseltii in New Zealand are present only in Central Otago and New Plymouth. In Central Otago, L. hasseltii feeds on endangered invertebrates, such as Prodontria modesta (Broun 1909). Latrodectus hasseltii is also a hazard to the New Zealand endemic L. katipo through interbreeding and competitive displacement. CLIMEXTM was used to model the potential global distribution of L. hasseltii based on current climate, and using ArcGIS® 9.2, areas of suitable climate in New Zealand were overlaid with favourable habitats to identify areas most suitable for L. hasseltii establishment. In addition, shelter that urban areas offer L. hasseltii were modelled in CLIMEX and incorporated into ArcGIS to produce maps indicating cities and built up areas where the species could establish. The presence of L. hasseltii in New Zealand and Japan, and its possible spread to other areas, is of human health significance, and the species may also impact on native biodiversity. KeywordsCLIMEX–GIS–Human health–Invasive species–Biodiversity threat

We report the occurrence of the orange-striped green anemone Diadumene lineata (Verrill 1871) (=Haliplanella lineata) in salt marshes at the Bahía Blanca Estuary for the first time in August 2005. We also found this species attached to roots and stems of Spartina alterniflora, an association that has never been registered before. After their determination, sampling was performed during a year to evaluate seasonal abundance of this sea anemone. Results showed that D. lineata was present through the whole year, indicating the existence of a stable population. All individuals sampled were found attached to roots or stems of S. alterniflora, with the higher abundances detected in summer. Further studies are necessary to precise the potential effects of this exotic sea anemone on salt marsh communities.

This paper represents the first record of the freshwater jellyfish Craspedacusta sowerbyi (Lankester 1880) in México; the medusa was collected in Adolfo Lopez Mateos (ALM) reservoir, one of the main reservoirs in Sinaloa State NW in Mexico. This report of the population of C.sowerbyi represents an exceptional case of freshwater jellyfish on this latitude in American Continent. This study provides additional information on physico chemical characteristics of the reservoir and qualitative and quantitative analysis of the trophic spectrum of C.sowerbyi.

During an experimental trawl survey carried out by the R/V “Hannibal” in June 2006 several specimens of the Western-Atlantic penaeid shrimp Rimapenaeus similis (Smith, 1885) were caught in the Gulf of Gabes (southern Tunisia, Central Mediterranean). This represents the first recording of Rimapenaeus similis in Tunisian waters and for the Mediterranean Sea. Biological information on the sampled population is also included. Keywords Rimapenaeus similis -Penaeidae-Non-indigenous species-Tunisia-Mediterranean Sea

Live individuals of Dreissena rostriformis bugensis were found in a trawlnet sample taken in April 2006 near Willemstad in the Hollands Diep, part of the main distributary in the Rhine delta in The Netherlands. This represented the first record of this species in Western Europe and a major westward range extension for this Ponto-Caspian species. Density was very low, with the ratio of Dreissena polymorpha to D. rostriformis bugensis in the order of 100:1. Mean length (n=5) was 22.5mm (range 20.4–24.6mm), suggesting that they were at least two years old and that the original introduction into the Hollands Diep had occurred in 2004 or earlier. Species identity was confirmed by morphological and molecular methods. We speculate that the invasion occurred through the Main-Danube Canal – a pathway that is becoming increasingly recognized as a southern corridor for the potential movement of non-indigenous aquatic species between Eastern and Western Europe.

Adults of the palaemonid prawn Palaemon macrodactylus Rathbun, 1902 were reported for the first time from the southwestern Atlantic waters. Males and ovigerous females were collected in Mar del Plata harbor (38°03′S; 57°31′W), Argentina; larvae obtained from one of the collected females were reared in the laboratory until the juvenile stage. The geographical range of P. macrodactylus was previously restricted to the Pacific Ocean: it was originally described in Japan and has been reported as an invasive species in the western coast of USA in the 1950’s and in Australia in the 1970’s. Recently, this prawn has been also reported from European waters.

In various studies encompassing large spatiotemporal scales, the densities of non-indigenous species and indigenous species in the same habitat increased and decreased, respectively, prior to reaching an equilibrium. We examined whether this is true for a non-indigenous mussel Mytilus galloprovincialis, introduced into Japan before 1934, and the sympatric indigenous bivalves: Septifer virgatus, Septifer bilocularis, Hormomya mutabilis, Saccostrea kegaki, Saccostrea mordax and Barbatia (Savignyarca) virescens. Seven quadrat surveys during 1978–2006 on 19 rocky shores (26.6–41.4°N) showed unexpected results. Density of M. galloprovincialis decreased on 89% of the shores where this mussel had previously been found. In contrast, densities of the indigenous bivalves decreased on lower percentages (17–71%) of shores where the bivalves had been found. Shore-to-shore mean density of M. galloprovincialis decreased from the 1970s (27.0–88.7m−2) to the 2000s (0.0–1.1m−2), more sharply than any of the indigenous bivalves (from 4.6–9.1m−2 to 0.7–1.5m−2 for S. bilocularis, which showed the most marked reduction). The conspicuous decrease for M. galloprovincialis suggests that some factors on the studied shores have become disadvantageous especially for this non-indigenous mussel from the earlier to the later surveys. KeywordsInvasive mussel-Decline-Indigenous bivalves-Hard-bottom habitat-Global warming-Quadrat survey

Populations of Partulina redfieldi, an achatinelline tree snail studied in four isolated trees, grew 100–900% between 1983 and 1995. Beginning in 1995, populations declined by 85%, and shells of rat-killed snails accumulated beneath the trees. While rat-marked shells were always present in the study area, numbers increased significantly. Despite a rat-abatement program begun in 1995, the snails continued to disappear, which we conclude was due to continued rat migration into the study area, despite baiting, and a switch in rat-food preference toward the snails. In neighboring forest where tree canopies are more continuous, snail density is lower and rat predation is not apparent. Captive-bred snails were successfully introduced to a small unoccupied tree in the same area in 1989, and this population suffered the same fate as the natural snail populations. Since 2000, P. redfieldi populations have remained low and rat predation continues.

In the Mediterranean, the survival of endemic long-lived seabirds despite the long-standing introduction of one of the most damaging alien predator, the ship rat (Rattus rattus), on most islands constitutes an amazing conservation paradox. A database gathering information on approximately 300 Western Mediterranean islands was analyzed through generalized linear models to identify the factors likely to influence ship rat presence and to account for how ship rat presence and island characteristics may have driven the presence and abundance of seabirds. Our review showed that few Mediterranean islands remain rat-free. At the regional scale, rat presence was only a limiting factor in the abundance of the smallest seabird, the storm petrel (Hydrobates pelagicus), while the distribution and abundance of the three shearwaters were more influenced by island characteristics. We hypothesized that the long-term persistence of these seabirds may have been facilitated by the various biogeographical contexts of Mediterranean islands, likely to provide intra-island refuges.

Without Abstract

The invasion process comprises not only the characteristics of nonindigenous species but also the attributes of the invaded environment which make it susceptible to the establishment of nonindigenous species. Habitat attributes operate like filters in determining the establishment of introduced species and the invasibility of a region. In the Upper Paraná River Basin, Brazil, the practice of introducing species was quite common and frequently carried out by hydroelectric companies. The target species of the present study, the peacock-bass Cichla kelberi, is native to the Amazon Basin. This species was introduced into several reservoirs of the Upper Paraná River Basin and is dispersing rapidly throughout the system. This study shows which characteristics of the reservoirs facilitate their invasibility after testing for the effect of propagule pressure. We conclude that a set of abiotic factors favors the invasibility of these reservoirs. To be more precise, the largest, deepest, most transparent and warmest reservoirs are the most likely to be colonized by Cichla kelberi. It is possible that other environments with similar characteristics to these reservoirs, such as the lagoons from the Upper Paraná River Basin floodplain, can also be colonized by Cichla kelberi. KeywordsAbiotic factors- Cichla kelberi -Invasibility-Paraná River Basin-Reservoirs

The number of marine alien species reported in Galil (Biol Invasions, 2009) is incomplete. Recent literature suggests that the number of aliens is nearly 1,000 and that the introduction rate is way off the 10 species per year suggested by Galil (Biol Invasions, 2009). KeywordsAlien species-Mediterranean-Atlantic origin

Four species of exotic cordgrass (Spartina sp.) occur in the San Francisco estuary in addition to the California native Spartina foliosa. Our goal was to map the location and extent of all non-native Spartina in the estuary. Hybrids of S. alterniflora and S. foliosa are by far the most numerous exotic and are spreading rapidly. Radiating from sites of deliberate introduction, S. alterniflora and hybrids now cover ca. 190 ha, mainly in the South and Central Bay. Estimates of rate of aerial increase range from a constant value to an accelerating rate of increase. This could be due to the proliferation of hybrid clones capable of rapid expansion and having superior seed set and siring abilities. The total coverage of 195 ha by hybrids and other exotic cordgrass species is slightly less than 1% of the Bay's tidal mudflats and marshes. Spartina anglica has not spread beyond its original 1970s introduction site. Spartina densiflora has spread to cover over 5 ha at 3 sites in the Central Bay. Spartina patens has expanded from 2 plants in 1970 to 42 plants at one site in Suisun Bay. Spartina seed floats on the tide, giving it the potential to export this invasion throughout the San Francisco estuary, and to estuaries outside of the Golden Gate. We found isolated plants of S. alterniflora and S. densiflora in outer coast estuaries north of the Bay suggesting the likelihood for the San Francisco Bay populations to found others on the Pacific coast.

In this paper, we examine the hypothesis that reduced intraspecific aggression underlies the competitive prowess of Argentine ants in their introduced range. Specifically, we test three predictions of this hypothesis by comparing the genetic diversity, behavior, and ecology of Argentine ants in their native range to introduced populations. Differences between native and introduced populations of Argentine ants were consistent with our predictions. Introduced populations of the Argentine ant appear to have experienced a population bottleneck at the time of introduction, as evidenced by much reduced variation in polymorphic microsatellite DNA markers. Intraspecific aggression was rare in introduced populations but was common in native populations. Finally, in contrast to the Argentine ant's ecological dominance throughout its introduced range, it did not appear dominant in the native ant assemblages studied in Argentina. Together these results identify a possible mechanism for the widespread success of the Argentine ant in its introduced range.

Propagule pressure and disturbance have both been found to facilitate invasion. Therefore, knowledge on the history of introduction and disturbance is vital for understanding an invasion process, and research should focus on areas in which the invasive species has not been deliberately introduced or managed to study unconfounded colonization patterns. Comparing the outcome of such spontaneous colonization processes for different ecosystems might provide a useful framework for setting management priorities for invasive species that enter new, uninvaded areas. We focused on the 70-year spontaneous spread of the invasive tree species Prunus serotina in a pine forest in the Netherlands. To reconstruct the invasion pattern, we combined historical maps, tree ring analysis, spatially explicit tree inventory data, seed density data, and regeneration data for both native and non-native species. Prunus serotina was the only species that showed successful regeneration: the species was present throughout the forest in the tree, shrub, and herb layer. Native species were not able to outgrow the seedling stage. Our data demonstrate that P. serotina is a gap-dependent species with high seed production that builds up a seedling bank. We also compared the results of this study with a similar study on P. serotina colonization in a deciduous forest in Belgium, where P. serotina invasion was not successful. The sharp contrast between the outcomes of the two invasion processes shows the importance of studying an invasive species and the recipient ecosystem jointly and made us raise the hypothesis that herbivore pressure may facilitate P. serotina invasion.

Since separating from its super-continental origin 80million years ago, New Zealand has effectively been isolated from the impacts of terrestrial mammals. The arrival of Polynesians in 13th C heralded the end of this era, with the introduction of kiore, (Rattus exulans, or Pacific rat), which had far-reaching effects on plant regeneration, survival of small ground vertebrates, larger invertebrates, and seabird breeding colonies. This paper reviews the evidence available from raptor nest sites and Quaternary beetle fossils to summarise extinctions thought to be caused by kiore in New Zealand. It also utilises invertebrate comparisons between islands with and without rats, or where rats have been eradicated, in order to document the impacts of rats (R. exulans, R. norvegicus) on invertebrate abundance, body mass, and the behavioural responses of some large New Zealand insects to the presence of rats. The role of a ‘mammal-free’ evolutionary history is discussed.

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. KeywordsAquatic invasive species–Freshwater mollusks–Wetlands–South America–Oxygen–Calcium

The Argentine ant is a common pest of mediterranean-type ecosystems worldwide, causing widespread extirpation of many native ant species. This study examines spatial and temporal patterns of invasion at a local scale, investigates the effects of this invader on native ant communities and assesses causal mechanisms for these spatial, temporal and numerical patterns, particularly in relation to local climate. Argentine ant workers were more active in the drier months of summer and fall, and abundance was correlated with cumulative precipitation the previous winter. Argentine ant worker abundance greatly increased in conjunction with EI Niño events. No similar relationship existed for native ants. This invasive ant affected native ant abundances differently by species: some species were able to tolerate its presence temporarily. Overall, this study provides a framework of factors to consider when devising control strategies for the invasive Argentine ant.