Induction of settlement of larvae of the sea urchin Holopneustes purpurascens by histamine from a host alga

Larvae of the Australian sea urchin Holopneustes purpurascens are induced to settle and metamorphose (termed settlement herein) by a water-soluble compound produced by the red alga Delisea pulchra, the main host plant of new recruits. The settlement cue for H. purpurascens had previously been identified as a floridoside-isethionic acid complex, and this paper presents new evidence correcting that finding. The actual settlement cue produced by D. pulchra was isolated from the polar extract by cation-exchange chromatography and identified as histamine, using one- and two-dimensional nuclear magnetic resonance spectrometry. The chemical identity of the cue was confirmed by gas chromatography-mass spectrometry (GC-MS) and matrix-assisted laser desorption/ionization-time-of-flight mass spectrometry. Synthetic histamine and histamine at 4.5 microM isolated from D. pulchra both induced rapid settlement in 80%-100% of the larvae of H. purpurascens. Lower concentrations of histamine (0.9-2.3 micro M) induced larval settlement, but this response varied from 0%-90%. The histamine content of two host plants of H. purpurascens--D. pulchra and Ecklonia radiata--and of four other common species was quantified using GC-MS. D. pulchra had the highest histamine content, which is consistent with H. purpurascens recruiting to this species. Histamine was also detected in the seawater surrounding these host algae. This is the first time that a settlement cue has been quantified in the habitat of a marine organism.     

Mitochondrial cytochrome b sequence yields new insight into the speciation of social voles in south-west Asia

We established a cytochrome b (cyt b ) phylogeny for six species of social voles. A Bayesian approach to phylogenetic reconstruction (BI) and a maximum likelihood (ML) tree revealed a dichotomy into two major clusters, namely a Microtus guentheri cluster and a M. socialis cluster. The three main lineages that emerged within each of these two clusters were separated by the K2P divergences which are above the intraspecific variation in Microtus . All six species were also retrieved in the minimum spanning network. Within its present taxonomic scope, M. guentheri is paraphyletic and consists of two allopatric sibling species: M. guentheri (Syria, Israel) and M. hartingi (Anatolia and the Balkans). The closest relative to these two species is M. dogramacii , which is possibly a sister species to M. hartingi . The two geographic samples were identified as M. irani , one from Shiraz (Iran) and the other from Balkusan (Turkey). The cyt b sequence confirmed the specific status of M. anatolicus within the M. socialis cluster. Although five species of social voles occur within a radius of < 500 km in the north-eastern corner of the Mediterranean, small-scale sympatry is exceptional. Species richness in this region possibly originates from past fragmentation with subsequent allopatric speciation in refugial areas.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 121–128.     

Induction of metamorphosis in the sea urchin Holopneustes purpurascens by a metabolite complex from the algal host Delisea pulchra

Most benthic invertebrates have complex life cycles with planktonic larvae that return to the substratum to settle and metamorphose into a benthic stage. Although naturally produced chemical cues have long been thought to be important for the settlement or metamorphosis of invertebrate larvae, few ecologically relevant chemical cues have been clearly identified. The marine echinoid Holopneustes purpurascens has a complex life cycle, with a planktonic, nonfeeding dispersive larva that metamorphoses into a benthic stage that lives in the canopy of subtidal benthic algae such as the red alga Delisea pulchra and the kelp Ecklonia radiata. Recently recruited juveniles are found primarily on D. pulchra, and we hypothesized that this was in response to a chemical cue produced by this alga. Competent larvae metamorphosed in the presence of D. pulchra, or seawater surrounding this alga, but not in response to the presence of E. radiata or its extracts. A cue for metamorphosis was isolated and characterized from D. pulchra and found to be a water-soluble complex of the sugar floridoside and isethionic acid in a 1:1 molar ratio. The floridoside-isethionic acid complex also triggered settlement in H. purpurascens; however, this response was less specific than metamorphosis and was reversible. Larvae of H. purpurascens also metamorphosed in the presence of several other species of red, but not brown or green, algae from their habitat. Floridoside is found only in red algae, suggesting that the floridoside-isethionic acid complex may be acting as a cue for metamorphosis in other red algae as well as in D. pulchra.     

Escapees of potentially invasive fishes from an ornamental aquaculture facility: the case of topmouth gudgeon Pseudorasbora parva

Accidental escapees from aquacultural facilities are an ongoing problem facing scientists, conservationists, policy makers and naturalists throughout the world. The topmouth gudgeon Pseudorasbora parva , a small cyprinid native to Southeast Asia, was introduced to Romanian waters in 1960 via the aquaculture trade and has since spread throughout Europe. It first appeared in England in an ornamental fish farm near Romsey in Hampshire. The aim of the present study was to quantify the dispersal of this potentially highly invasive species from an ornamental fish farm into the wild. In order to measure the dispersal of escapees, intensive electrofishing surveys were carried out between June and September 2003 in the catchment downstream of the fish farm. The distribution of topmouth gudgeon in the catchment was found to be limited and patchy. The presence of small individuals suggests that the species has successfully established itself in the wild, but this remains to be confirmed. The results were examined in light of invasion theory principles of biological resistance and ecological fitness.     

The ecological impact of invasive cane toads (Bufo marinus) in Australia

Although invasive species are viewed as major threats to ecosystems worldwide, few such species have been studied in enough detail to identify the pathways, magnitudes, and timescales of their impact on native fauna. One of the most intensively studied invasive taxa in this respect is the cane toad (Bufo marinus), which was introduced to Australia in 1935. A review of these studies suggests that a single pathway-lethal toxic ingestion of toads by frog-eating predators-is the major mechanism of impact, but that the magnitude of impact varies dramatically among predator taxa, as well as through space and time. Populations of large predators (e.g., varanid and scincid lizards, elapid snakes, freshwater crocodiles, and dasyurid marsupials) may be imperilled by toad invasion, but impacts vary spatially even within the same predator species. Some of the taxa severely impacted by toad invasion recover within a few decades, via aversion learning and longer-term adaptive changes. No native species have gone extinct as a result of toad invasion, and many native taxa widely imagined to be at risk are not affected, largely as a result of their physiological ability to tolerate toad toxins (e.g., as found in many birds and rodents), as well as the reluctance of many native anuran-eating predators to consume toads, either innately or as a learned response. Indirect effects of cane toads as mediated through trophic webs are likely as important as direct effects, but they are more difficult to study. Overall, some Australian native species (mostly large predators) have declined due to cane toads; others, especially species formerly consumed by those predators, have benefited. For yet others, effects have been minor or have been mediated indirectly rather than through direct interactions with the invasive toads. Factors that increase a predator's vulnerability to toad invasion include habitat overlap with toads, anurophagy, large body size, inability to develop rapid behavioral aversion to toads as prey items, and physiological vulnerability to bufotoxins as a result of a lack of coevolutionary history of exposure to other bufonid taxa.     

The stock of invasive insect species and its economic determinants

Invasions of nonindigenous organisms have long been linked to trade, but the contribution of individual trade pathways remains poorly understood, because species are not observed immediately upon arrival and the number of species arriving annually is unknown. Species interception records may count both new arrivals and species long introduced. Furthermore, the stock of invasive insect species already present is unknown. In this study, a state-space model is used to infer the stock of detected as well as undetected invasive insect species established in the United States. A system of equations is estimated jointly to distinguish the patterns of introduction, identification, and eradication. Introductions of invasive species are modeled as dependent on the volume of trade and arrival of people. Identifications depend on the public efforts at invasive species research, as well as on the established stock of invasive species that remain undetected. Eradications of both detected and undetected invasive species depend on containment and quarantine efforts, as well as on the stock of all established invasive species. These patterns are estimated by fitting the predicted number of invasive species detections to the observed record in the North American Non-Indigenous Arthropod Database. The results indicate that agricultural imports are the most important pathway of introduction, followed by immigration of people. Expenditures by the U.S. Department of Agriculture and the Agricultural Research Service are found to explain the species identification record well. Between three and 38 invasive insect species are estimated to be established in the United States undetected.     

The prehistory and biogeography of terrestrial vertebrates on Guam, Mariana Islands

Aim By establishing a prehistoric fossil record for the vertebrate fauna of Guam, we can document the composition of the island’s fauna prior to Western contact. It will also complement the extensive prehistoric fossil data already known from the nearby islands of Rota, Tinian and Aguiguan, and improve our understanding of natural distributional patterns through a chronologically deeper view of the biogeography of the Mariana Islands. Location Northern Guam, Mariana Islands, Micronesia. Methods Fossil‐bearing sediment was removed from karst features with vertically and horizontally controlled excavation methods. Age determinations of the fossils were estimated by radiocarbon dating associated wood charcoal samples. Results The 3314 bones (fossils) of terrestrial vertebrates date to the past two millennia and represent 34 species. Among 12 species of squamate reptiles, one is extinct (a gekkonid lizard) and another, the monitor lizard, Varanus indicus, was introduced to Guam no later than about 1600 years ago. The 17 species of birds feature five that are extinct. Two others not recorded previously on Guam are extirpated. Eight others were lost on Guam in historic times. We also recovered bones of two species of indigenous bats, (Pteropus sp., extirpated; Emballonura semicaudata, probably extirpated), and non‐native rodents, (Rattus spp.). Main conclusions Guam’s contemporary faunal losses have been well documented since invasion of the Brown Treesnake (Boiga irregularis) during or shortly after WWII. Other exotics, as well as habitat deterioration, have abetted this snake’s deleterious presence. Our data further show that human impact led to declines or losses of vertebrate populations by late prehistoric times. Numerous anthropogenic extirpations would not be evident without studying fossils. Our chrono‐stratigraphic evidence argues for the arrival of non‐native rats (Rattus sp.) only about 1000 years ago, which helps to explain why a flightless rail survived on Guam into modern times, unlike on most other islands in Oceania.     

Description of Perkinsus andrewsi n. sp. isolated from the Baltic clam (Macoma balthica) by characterization of the ribosomal RNA locus, and development of a species-specific PCR-based diagnostic assay

A Perkinsus species was isolated from the baltic clam Macoma balthica and an in vitro culture established under conditions described for P. marinus. As reported previously, morphological features remarkable enough to clearly indicate that this isolate is a distinct Perkinsus species were lacking. In this study, regions of the rRNA locus (NTS, 18S, ITS1, 5.8S, and ITS2) of this isolate were cloned, sequenced, and compared by alignment with those available for other Perkinsus species and isolates. Sequence data from the rRNA locus and species-specific PCR assays indicated not only that Perkinsus sp. from M. balthica was not P. marinus, but it was different from P. atlanticus and P. olseni. The degree of difference was comparable to or greater than differences between accepted Perkinsus species. In particular, NTS sequence and length were dramatically different from that of P. marinus and P. atlanticus. Therefore, we formally propose to designate the Perkinsus sp. from M. balthica as a separate species, P. andrewsi n. sp. Primers based on P. andrewsi NTS sequence were used to develop a PCR-based diagnostic assay that was validated for species-specificity and sensitivity. PCR-based assays specific for either P. andrewsi or P. marinus were used to test for their presence in bivalve species sympatric to M. balthica. Although isolated from M. balthica, P. andrewsi was also detected in the oyster Crassostrea virginica and clams Macoma mitchelli and Mercenaria mercenaria, and could coexist with P. marinus in all four bivalve species tested.     

The bioinvasion of Guam: inferring geographic origin,pace, pattern and process of an invasive lizard (<Emphasis Type="Italic">Carlia</Emphasis>) in the Pacific using multi-locus genomic data

Invasive species often have dramatic negative effects that lead to the deterioration and loss of biodiversity frequently coupled with the burden of expensive biocontrol programs and subversion of socioeconomic stability. The fauna and flora of oceanic islands are particularly susceptible to invasive species and the increase of global movements of humans and their products since WW II has caused numerous anthropogenic translocations and increased the ills of human-mediated invasions. We use a multi-locus genomic dataset to identify geographic origin, pace, pattern and historical process of an invasive scincid lizard (Carlia) that has been inadvertently introduced to Guam, the Northern Marianas, and Palau. This lizard is of major importance as its introduction is thought to have assisted in the establishment of the invasive brown treesnake (Boiga irregularis) on Guam by providing a food resource. Our findings demonstrate multiple waves of introductions that appear to be concordant with movements of Allied and Imperial Japanese forces in the Pacific during World War II.     

Assessment of the northern distribution range of selected Perkinsus species in eastern oysters (Crassostrea virginica) and hard clams (Mercenaria mercenaria) with the use of PCR-based detection assays

Perkinsus species are protistan parasites of molluscs. In Chesapeake Bay, Perkinsus marinus, Perkinsus chesapeaki, and Perkinsus andrewsi are sympatric, infecting oysters and clams. Although P. marinus is a pathogen for Crassostrea virginica, it remains unknown whether P. andrewsi and P. chesapeaki are equally pathogenic. Perkinsus species have been reported in C. virginica as far north as Maine, sometimes associated with high prevalence, but low mortality. Thus, we hypothesized that, in addition to P. marinus, Perkinsus species with little or no pathogenicity for C. virginica may be present. Accordingly, we investigated the distribution of Perkinsus species in C. virginica and Mercenaria mercenaria, collected from Maine to Virginia, by applying PCR-based assays specific for P. marinus, P. andrewsi, and a Perkinsus sp. isolated from M. mercenaria. DNA samples of M. mercenaria possessed potent PCR inhibitory activity, which was overcome by the addition of 1 mg/ml BSA and 5% (v/v) DMSO to the PCR reaction mixture. All 3 Perkinsus species were found in both host species throughout the study area. Interestingly, the prevalence of P. marinus in M. mercenaria was significantly lower than in C. virginica, suggesting that M. mercenaria is not an optimal host for P. marinus.     

How many and which ant species are being accidentally moved around the world?

Human transportation facilitates the dispersal of exotic ants, but few studies have quantified the magnitude and geography of these movements. We used several non-parametric indexes to estimate the number of species successfully introduced to or established in new regions. We also compared their source biogeographic realms to assess the importance of the geographical origin in determining the likelihood of establishment after introduction. Data on exotic ants derive from studies of three temperate regions. Our results suggest that the numbers of introduced or established ants may be much larger than the numbers so far documented. Ants introduced or established in new regions tend to arrive from the same or neighbouring realms, as would be expected if exotic species tend to match climates and if arrival/establishment is dependent upon higher trade rates from neighbouring countries.     

Callosciurus squirrels: worldwide introductions,ecological impacts and recommendations to prevent the establishment of new invasive populations

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Genetic structure of invasive earthworms Dendrobaena octaedra in the boreal forest of Alberta: insights into introduction mechanisms

Population genetic studies can help to determine whether invasive species are established via single vs. multiple introduction events and also to distinguish among various colonization scenarios. We used this approach to investigate the introduction of Dendrobaena octaedra , a non-native earthworm species, to the boreal forest of northern Alberta. The spread of non-native earthworms in forested systems is not well understood, although bait abandonment and vehicular transport are believed to be important. Mitochondrial DNA sequencing revealed that multiple introductions of this species have occurred in northern Alberta, although individual populations may have been established by either single or multiple invaders introduced on one or more occasions. There was no relationship between genetic distances and either geographical distances or distances along road networks, suggesting that human-mediated jump dispersal is more common than diffusive spread via road networks or via active dispersal. As well, genetic diversity was significantly greater at boat launches than roads, indicating that multiple introductions may be more likely to occur at those locations. Focusing management efforts on areas where multiple introductions are likely to occur may help to reduce invasive species' potential for adaptive evolution and subsequent rapid spread.     

Direct and indirect effects of the introduced toad Bufo marinus (Anura: Bufonidae) on populations of native anuran larvae in Australia

Introduced species are frequently believed to have adverse effects on native biota and ecosystems. However, much of our knowledge of the ecological impacts of introduced species is anecdotal, and the mechanisms controlling these effects are often poorly understood. I used replicated artificial pond experiments to investigate the impact of eggs and hatchlings of the introduced toad Bufo marinus on populations of native anuran larvae (Limnodynastes ornatus and Litoria rubella) in Australia. Bufo marinus eggs and hatchlings are highly toxic to predatory native tadpoles. Under naturalistic conditions, populations of predatory L. ornatus tadpoles experienced significantly reduced survival when exposed to Bufo eggs and hatchlings. The impact of Bufo on L. ornatus survival was positively correlated with Bufo density. However, the toxic effects of Bufo on L. ornatus indirectly facilitated the survival of later-breeding L. rubella by altering predator-prey interactions between L. ornatus and L. rubella. Limnodynastes ornatus tadpoles are voracious predators of L. rubella eggs and hatchlings. Consequently, the negative impact of Bufo on populations of L. ornatus tadpoles reduced the intensity of predation by L. ornatus tadpoles on L. rubella eggs and hatchlings, thereby increasing L. rubella survival. The results demonstrate that B. marinus plays an important role in structuring native larval anuran communities via direct and indirect mechanisms, and that Bufo may have both negative and positive effects on populations of native anuran larvae. As far as I am aware, these are the first quantitative data to demonstrate that introduced fauna may affect populations of native biota via toxic effects.     

How many and which ant species are being accidentally moved around the world?

Human transportation facilitates the dispersal of exotic ants, but few studies have quantified the magnitude and geography of these movements. We used several non-parametric indices to estimate the number of species successfully introduced to or established in new regions. We also compared their source biogeographic realms to assess the importance of geographical origin in determining the likelihood of establishment after introduction. Occurrence data on exotic ants derive from studies of three temperate regions. Our results suggest that the numbers of introduced or established ants may be much larger than the numbers so far documented. Ants introduced or established in new regions tend to arrive from the same or neighbouring realms, as would be expected if exotic species tend to match climates and if arrival/establishment is dependent upon higher trade rates from neighbouring countries.     

Impact of the invasive cane toad (Bufo marinus) on an Australian frog (Opisthodon ornatus) depends on minor variation in reproductive timing

Invasive species are widely viewed as unmitigated ecological catastrophes, but the reality is more complex. Theoretically, invasive species could have negligible or even positive effects if they sufficiently reduce the intensity of processes regulating native populations. Understanding such mechanisms is crucial to predicting ultimate ecological impacts. We used a mesocosm experiment to quantify the impact of eggs and larvae of the introduced cane toad (Bufo marinus) on fitness-related traits (number, size and time of emergence of metamorphs) of a native Australian frog species (Opisthodon ornatus). The results depended upon the timing of oviposition of the two taxa, and hence the life-history stages that came into contact. Growth and survival of O. ornatus tadpoles were enhanced when they preceded B. marinus tadpoles into ponds, and reduced when they followed B. marinus tadpoles into ponds, relative to when tadpoles of both species were added to ponds simultaneously. The dominant tadpole-tadpole interaction is competition, and the results are consistent with competitive priority effects. However, these priority effects were reduced or reversed when O. ornatus tadpoles encountered B. marinus eggs. Predation on toxic toad eggs reduced the survival of O. ornatus and B. marinus. The consequent reduction in tadpole densities allowed the remaining O. ornatus tadpoles to grow more rapidly and to metamorphose at larger body sizes (>60% disparity in mean mass). Thus, exposure to B. marinus eggs reduced the number of O. ornatus metamorphs, but increased their body sizes. If the increased size at metamorphosis more than compensates for the reduced survival, the effective reproductive output of native anurans may be increased rather than decreased by the invasive toad. Minor interspecific differences in the seasonal timing of oviposition thus have the potential to massively alter the impact of invasive cane toads on native anurans.     

Identification of Hotspots in the European Union for the Introduction of Four Zoonotic Arboviroses by Live Animal Trade

Live animal trade is considered a major mode of introduction of viruses from enzootic foci into disease-free areas. Due to societal and behavioural changes, some wild animal species may nowadays be considered as pet species. The species diversity of animals involved in international trade is thus increasing. This could benefit pathogens that have a broad host range such as arboviruses. The objective of this study was to analyze the risk posed by live animal imports for the introduction, in the European Union (EU), of four arboviruses that affect human and horses: Eastern and Western equine encephalomyelitis, Venezuelan equine encephalitis and Japanese encephalitis. Importation data for a five-years period (2005-2009, extracted from the EU TRACES database), environmental data (used as a proxy for the presence of vectors) and horses and human population density data (impacting the occurrence of clinical cases) were combined to derive spatially explicit risk indicators for virus introduction and for the potential consequences of such introductions. Results showed the existence of hotspots where the introduction risk was the highest in Belgium, in the Netherlands and in the north of Italy. This risk was higher for Eastern equine encephalomyelitis (EEE) than for the three other diseases. It was mainly attributed to exotic pet species such as rodents, reptiles or cage birds, imported in small-sized containments from a wide variety of geographic origins. The increasing species and origin diversity of these animals may have in the future a strong impact on the risk of introduction of arboviruses in the EU.     

Human-mediated introduction of geoengineering earthworms in the Fennoscandian arctic

It is now well established that European earthworms are re-shaping formerly glaciated forests in North America with dramatic ecological consequences. However, few have considered the potential invasiveness of this species assemblage in the European arctic. Here we argue that some earthworm species (Lumbricus rubellus, Lumbricus terrestris and Aporrectodea sp.) with great geomorphological impact (geoengineering species) are non-native and invasive in the Fennoscandian arctic birch forests, where they have been introduced by agrarian settlers and most recently through recreational fishing and gardening. Our exploratory surveys indicate no obvious historical dispersal mechanism that can explain early arrival of these earthworms into the Fennoscandian arctic: that is, these species do not appear to establish naturally along coastlines mimicking conditions following deglaciation in Fennoscandia, nor were they spread by early native (Sami) cultures. The importance of anthropogenic sources and the invasive characteristics of L. rubellus and Aporrectodea sp. in the arctic is evident from their radiation outwards from abandoned farms and modern cabin lawns into adjacent arctic birch forests. They appear to outcompete previously established litter-dwelling earthworm species (i.e. Dendrobaena octaedra) that likely colonized the Fennoscandian landscape rapidly following deglaciation via hydrochory and/or dispersal by early Sami settlements. The high geoengineering earthworm biomasses, their recognized ecological impact in other formerly glaciated environments, and their persistence once established leads us to suggest that geoengineering earthworms may pose a potent threat to some of the most remote and protected arctic environments in northern Europe.     

Aquarium hitchhikers: attached commensals imported with freshwater shrimps via the pet trade

The keeping of home aquaria is one of the most popular hobbies globally. In contrast to the ornamental fish trade, decapod crustaceans such as shrimps, crayfish and crabs are relatively new to the pet trade. Nevertheless, the popularity of ornamental shrimp in freshwater aquaria has rapidly increased in recent years. Indonesia is one of the leading producers and exporters of ornamental aquatic animals globally and the Czech Republic is a gateway for these animals into the European Union territory. The pathway for introductions of organisms unintentionally moved in association with ornamental shrimps via the international trade has to date not been evaluated. We examined a small number of shrimps imported from Indonesia into the Czech Republic in May 2015 and found large numbers of the protozoan Vorticella sp., one species of scutariellid temnocephalidan (Caridinicola sp.), and one species of bdelloid rotifer, associated with two species of atyid shrimps, indicating an invasion risk from fauna carried unintentionally by this vector. Although our observations were limited in scale, we estimate the total number of commensal fauna imported into the Czech Republic with ornamental shrimps via the pet trade to be in the order of hundreds of thousands per month. As attached organisms can directly or indirectly cause diseases in certain species of decapod crustaceans, we recommend five steps to reduce risks of introduction of “hitchhikers” to aquaria and wildlife.     

Establishment risk and potential invasiveness of the selected exotic amphibians from pet trade in the European Union

The pet trade has grown in recent years and become the most important pathway for the introduction of non-indigenous species of amphibians and reptiles worldwide. Amphibians traded on the pet market have been widely overlooked in systematic invasion studies, so their establishment potential and invasion dynamics remain poorly understood, despite the fact that the impact of invaders from this taxonomic group on native biota may be considerable. The determination of the most common species of amphibians traded as pets was based on a survey of the market in the Czech Republic, which is an export hub for ornamental aquatic animals into the European Union (EU). Subsequently, the establishment potential of the most common species for EU was determined using a proven risk assessment model. Amphibians that scored higher than the established species Xenopus laevis were additionally evaluated for their invasion potential using the Amphibian Invasiveness Screening Kit (AmphISK).Sixteen species of amphibians in pet market (14 Anura, 2Caudata) have an establishment potential in the EU. However, none of these species have reached the invasion potential score of already spreading species X. laevis and Lithobates catesbeianus. The species closest to the threshold value were Lithobates pipiens and Pelophylax saharicus. Amphibians represent a rather small group within all traded animals but this fact should not lead us to an underestimation of their potential invasiveness.