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  • 1
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    Ascidians as models for studying invasion success (2015)
    Springer
    In:  Marine Biology, 162 (12). pp. 2449-2470.
    Details
    Publication Date: 2018-06-20
    Description: During the past three decades, coastal marine waters have become among the most invaded habitats globally. Ascidians are among the most notorious invaders in these ecosystems. Owing to their rapid spread, frequent population outbreaks, and associated negative ecological and economic impacts, invasive ascidians have become a global problem. Thus, the study of ascidian invasions has become a prominent area of invasion biology. Here, we review current knowledge and conclude that ascidians are good models for studying invasion success in the marine realm. Firstly, we summarize the reconstruction of invasion pathways or colonization histories and associated negative impacts of invasive ascidians, and address the urgent need to clarify ambiguous taxonomy of ascidians. Secondly, we discuss factors that underlie or facilitate invasion success of ascidians, including vectors of introduction and spread, environmental changes, biological traits, and possible genetic issues. Finally, we summarize current science-based policies and management solutions that are in place to prevent and control spread of invasive ascidians. We conclude by highlighting key research questions that remain to be answered, and propose future research to investigate mechanisms of invasion success in the marine realm using ascidians as model systems.
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  • 2
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    Population attenuation in zooplankton communities during transoceanic transfer in ballast water (2016)
    Wiley
    In:  Ecology and Evolution, 6 (17). pp. 6170-6177.
    Details
    Publication Date: 2019-02-01
    Description: Successful biological invasion requires introduction of a viable population of a nonindigenous species (NIS). Rarely have ecologists assessed changes in populations while entrained in invasion pathways. Here, we investigate how zooplankton communities resident in ballast water change during transoceanic voyages. We used next-generation sequencing technology to sequence a nuclear small subunit ribosomal DNA fragment of zooplankton from ballast water during initial, middle, and final segments as a vessel transited between Canada and Brazil. Operational taxonomic unit (OTU) diversity decreased as voyage duration increased, indicating loss of community-based genetic diversity and development of bottlenecks for zooplankton taxa prior to discharge of ballast water. On average, we observed 47, 26, and 24 OTUs in initial, middle, and final samples, respectively. Moreover, a comparison of genetic diversity within taxa indicated likely attenuation of OTUs in final relative to initial samples. Abundance of the most common taxa (copepods) declined in all final relative to initial samples. Some taxa (e.g., Copepoda) were represented by a high number of OTUs throughout the voyage, and thus had a high level of intraspecific genetic variation. It is not clear whether genotypes that were most successful in surviving transit in ballast water will be the most successful upon introduction to novel environments. This study highlights that population bottlenecks may be common prior to introduction of NIS to new ecosystems
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  • 3
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    Efficacy of ‘saltwater flushing’ in protecting the Great Lakes from biological invasions by invertebrate eggs in ships’ ballast sediment (2010)
    Wiley
    In:  Freshwater Biology, 55 (11). pp. 2414-2424.
    Details
    Publication Date: 2014-08-08
    Description: 1. Mid-ocean exchange and saltwater flushing were implemented as management practices to reduce the likelihood of new biological invasions in the Laurentian Great Lakes associated with ships’ ballast water and sediments. Despite this, there has been no formal assessment of the efficacy of these procedures. Here, we conduct a comparative analysis of community composition of dormant taxa transported by ballast sediment before and after regulations came into effect in 2006. 2. Ballast sediment samples were collected from 17 ships during the post-regulation interval of 2007 and 2008. Invertebrate eggs were counted, hatched and species identified in the laboratory. Results were compared to similar samples collected from 39 ships between 2000 and 2002, prior to implementation of saltwater flushing regulations. 3. The estimated amount of residual ballast sediment transported by vessels was significantly lower during the post-regulation period, ranging from 〈1 to 45 tonnes per ship, with an average of 5 tonnes. Mean density and number of dormant viable eggs per ship declined 91 and 81%, respectively. 4. Community composition also changed through time, with Rotifera accounting for 78% of taxa transported prior to regulation, whereas Cladocera and Copepoda each accounted for 38% of abundance post-regulation. Although the number of non-indigenous species (NIS) declined 73% per ship after 2006, the reduction was not statistically significant; however, the number of freshwater NIS – which pose the greatest risk of invasion for the Great Lakes – was significantly lowered. 5. Our comparative analysis suggests that ballast management regulations enacted in 2006 markedly reduced the probability of introduction of NIS via dormant eggs carried in ballast sediments.
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  • 4
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    Invasion risk posed by macroinvertebrates transported in ships’ ballast tanks (2012)
    Springer
    In:  Biological Invasions, 14 (9). pp. 1843-1850.
    Details
    Publication Date: 2018-01-25
    Description: Invasions by non-indigenous macroinvertebrates often cause ecological and economic problems, and commercial ships have been implicated as a principal mechanism for their dispersal. We explored the presence and species diversity of adult macroinvertebrates transported by transoceanic and coastal vessels arriving to ports on the Atlantic coast of Canada. We sampled 67 ballast tanks from 62 ships operating along discrete geographic pathways and tested whether mid-ocean exchange or voyage length affects the probability for translocation of macroinvertebrates. Additionally, we assessed the relationship between macroinvertebrate presence and the amount of sediment in ballast tanks. We document the presence of highly invasive European green crab (Carcinus maenas), mud crab (Rhithropanopeus harrisii), common periwinkle (Littorina littorea), soft shell clam (Mya arenaria) and blue mussel (Mytilus galloprovincialis) in ballast tanks of surveyed ships. Mid-ocean exchange did not affect macroinvertebrate occurrence, suggesting that current ballast water management regulations are ineffective for this taxonomic group. Viable individuals were recorded in vessels undertaking shorter voyages (average and maximum of 4.5 and 15 days, respectively) and presence was not related to the amount of sediment in tanks. While presence and densities of macroinvertebrates were low, invasion risk may nonetheless be significant during reproductive seasons owing to high fecundity of some taxa. The highest risk may be posed by decapods since gravid females may carry thousands to several million eggs per clutch, and after several weeks of brooding, two or more subsequent clutches may be fertilized by retained sperm from an earlier mating.
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  • 5
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    Genetic Diversity in Introduced Golden Mussel Populations Corresponds to Vector Activity (2013)
    Public Library of Science
    In:  PLoS ONE, 8 (3). e59328.
    Details
    Publication Date: 2014-11-06
    Description: We explored possible links between vector activity and genetic diversity in introduced populations of Limnoperna fortunei by characterizing the genetic structure in native and introduced ranges in Asia and South America. We surveyed 24 populations: ten in Asia and 14 in South America using the mitochondrial cytochrome c oxidase subunit I (COI) gene, as well as eight polymorphic microsatellite markers. We performed population genetics and phylogenetic analyses to investigate population genetic structure across native and introduced regions. Introduced populations in Asia exhibit higher genetic diversity (HE = 0.667–0.746) than those in South America (HE = 0.519–0.575), suggesting higher introduction effort for the former populations. We observed pronounced geographical structuring in introduced regions, as indicated by both mitochondrial and nuclear markers based on multiple genetic analyses including pairwise ФST, FST, Bayesian clustering method, and three-dimensional factorial correspondence analyses. Pairwise FST values within both Asia (FST = 0.017–0.126, P = 0.000–0.009) and South America (FST = 0.004–0.107, P = 0.000–0.721) were lower than those between continents (FST = 0.180–0.319, P = 0.000). Fine-scale genetic structuring was also apparent among introduced populations in both Asia and South America, suggesting either multiple introductions of distinct propagules or strong post-introduction selection and demographic stochasticity. Higher genetic diversity in Asia as compared to South America is likely due to more frequent propagule transfers associated with higher shipping activities between source and donor regions within Asia. This study suggests that the intensity of human-mediated introduction vectors influences patterns of genetic diversity in non-indigenous species.
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  • 6
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    A conceptual model of community dynamics during the transport stage of the invasion process: a case study of ships’ ballast (2014)
    Wiley
    In:  Diversity and Distributions, 20 (2). pp. 236-244.
    Details
    Publication Date: 2014-12-09
    Description: Aim: After J. L. Lockwood, P. Cassey and T. Blackburn (2009, Diversity and Distributions, 15, 904–910) first described a theoretical relationship between propagule pressure and colonization pressure, two empirical studies demonstrated that the transport stage of the invasion process can profoundly influence the strength of the relationship among multiple events, as well as predictions of introduction risk. However, studies exploring dynamics of transported communities are rare, as repeated-measures sampling during transportation by any vector is logistically difficult. We constructed a conceptual model of community dynamics during transportation and supported it by empirical data for propagule pressure and colonization pressure of plankton. Location: Global. Methods: A conceptual model of community dynamics was developed based on lognormal species abundance distribution and the simulation model of J. L. Lockwood, P. Cassey and T. Blackburn (2009, Diversity and Distributions, 15, 904–910). We considered four cases: case ‘A’ – no reduction in propagule nor colonization pressure; case ‘B’ – strong reduction in propagule and mild reduction in colonization pressure; case ‘C’ – mild reduction in propagule and strong reduction in colonization pressure; and case ‘D’ – strong reduction in both propagule and colonization pressures. Results: The cases ‘B’, ‘C’ and ‘D’ were supported by empirical data for invertebrates, dinoflagellates and diatoms from ships’ ballast tanks, respectively. Propagule pressure of invertebrates, dinoflagellates and diatoms decreased 99.95%, 80% and 94% in 25 days, respectively, while colonization pressure decreased 34%, 57% and 64%. Main conclusions: Transport affects both propagule pressure and colonization pressure of taxa, with the magnitude of change dependent on length of transport and taxon-specific survival and reproduction. Our model demonstrates that introduction risk varies substantially across and within taxa depending on the occurrence and severity of selection pressures during transportation which serve to change species abundance distributions.
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  • 7
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    Assessing invasion risk across taxa and habitats: life stage as a determinant of invasion success (2011)
    Wiley
    In:  Diversity and Distributions, 17 (4). pp. 593-602.
    Details
    Publication Date: 2014-08-08
    Description: Aim  Many aquatic invertebrates produce dormant life-history stages as a means to endure inhospitable environments and to facilitate natural long-distance dispersal, yet we have little understanding of the role of dormant stages as a mechanism for human-mediated introductions of non-indigenous species. We explore the survival of invertebrate dormant eggs in collected ships’ ballast sediment over a 1-year period to determine relative invasion potential across taxa (i.e. rotifers, copepods, cladocerans and bryozoans) and different habitats (freshwater, marine). Location  Canadian Atlantic and Pacific coasts and Laurentian Great Lakes. Methods  During 2007 and 2008, 19 ballast samples were collected as a part of a larger study. The degradation rate of dormant eggs was assessed by enumerating dormant eggs and by conducting viability hatching experiments. Results  Taxa examined included rotifers, copepods, anomopods, onychopods and bryozoans. Dormant eggs of rotifers degraded at the highest rate of all taxa examined, with no viable eggs remaining within 10 months. Copepods showed a less rapid degradation rate than rotifers. The degradation rate of anomopod dormant eggs was significantly slower than that of both rotifers and copepods. Onychopods and bryozoans did not visibly degrade at all over 12 months. Viability hatching experiments were successful for rotifers, copepods, and anomopods. Onychopods and bryozoans did not hatch during any of the three hatching trials. Main conclusions  Dormancy is not equally beneficial to all invertebrate taxa. Our results indicate that dormant eggs of rotifers and copepods degrade at a rapid rate and may not pose high invasion risk. In contrast, the slow degradation rate of anomopod dormant eggs and the lack of degradation of onychopod and bryozoan dormant eggs could result in high invasion risk because of their accumulation in ballast tanks. Species having resistant dormant eggs mostly belong to freshwater taxa making freshwater habitats at higher invasion risk by dormant invertebrates than marine habitats.
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  • 8
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    Use of DNA barcoding to detect invertebrate invasive species from diapausing eggs (2011)
    Springer
    In:  Biological Invasions, 13 (6). pp. 1325-1340.
    Details
    Publication Date: 2018-01-25
    Description: The global transhipment of ballast water and associated flora and fauna by cargo vessels has increased dramatically in recent decades. Invertebrate species are frequently carried in ballast water and sediment, although identification of diapausing eggs can be extremely problematic. Here we test the application of DNA barcoding using mitochondrial cytochrome c oxidase subunit I and 16S rDNA to identify species from diapausing eggs collected in ballast sediment of ships. The accuracy of DNA barcoding identification was tested by comparing results from the molecular markers against each other, and by comparing barcoding results to traditional morphological identification of individuals hatched from diapausing eggs. Further, we explored two public genetic databases to determine the broader applicability of DNA barcodes. Of 289 diapausing eggs surveyed, sufficient DNA for barcoding was obtained from 96 individuals (33%). Unsuccessful DNA extractions from 67% of eggs in our study were most likely due to degraded condition of eggs. Of 96 eggs with successful DNA extraction, 61 (64%) were identified to species level, while 36% were identified to possible family/order level. Species level identifications were always consistent between methodologies. DNA barcoding was suitable for a wide range of taxa, including Branchiopoda, Copepoda, Rotifera, Bryozoa and Ascidia. Branchiopoda and Copepoda were respectively the best and worst represented groups in genetic databases. Though genetic databases remain incomplete, DNA barcoding resolved nearly double the number of species identified by traditional taxonomy (19 vs. 10). Notorious invaders are well represented in existing databases, rendering these NIS detectable using molecular methods. DNA barcoding provides a rapid and accurate approach to identification of invertebrate diapausing eggs that otherwise would be very difficult to identify.
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  • 9
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    Assemblage structure: an overlooked component of human-mediated species movements among freshwater ecosystems (2014)
    CNR
    In:  Journal of Limnology, 73 (s1). pp. 112-119.
    Details
    Publication Date: 2014-10-29
    Description: The spread and impact of alien species among freshwater ecosystems has increased with global trade and human movement; therefore, quantifying the role of anthropogenic and ecological factors that increase the risk of invasion is an important conservation goal. Two factors considered as null models when assessing the potential for invasion are colonization pressure (i.e., the number of species introduced) and propagule pressure [i.e., the number (propagule size), and frequency (propagule number), of individuals of each species introduced]. We translate the terminology of species abundance distributions to the invasion terminology of propagule size and colonization size (PS and CS, respectively). We conduct hypothesis testing to determine the underlying statistical species abundance distribution for zooplankton assemblages transported between freshwater ecosystems; and, on the basis of a lognormal distribution, construct four hypothetical assemblages spanning assemblage structure, rank-abundance gradient (e.g., even vs uneven), total abundance (of all species combined), and relative contribution of PS vs CS. For a given CS, many combinations of PS and total abundance can occur when transported assemblages conform to a lognormal species abundance distribution; therefore, for a given transportation event, many combinations of CS and PS are possible with potentially different ecological outcomes. An assemblage exhibiting high PS but low CS (species poor, but highly abundant) may overcome demographic barriers to establishment, but with lower certainty of amenable environmental conditions in the recipient region; whereas, the opposite extreme, high CS and low PS (species rich, but low abundance per species) may provide multiple opportunities for one of n arriving species to circumvent environmental barriers, albeit with lower potential to overcome demographic constraints. Species abundance distributions and the corresponding influence of CS and PS are some of many influential factors (e.g., demographic and genetic stochasticity, environmental variability, composition of recipient ecosystems) that will help refine an understanding of establishment risk following the human-mediated movement of species.
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  • 10
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    Invasion Pathway of the Ctenophore Mnemiopsis leidyi in the Mediterranean Sea (2013)
    Public Library of Science
    In:  PLoS ONE, 8 (11). e81067.
    Details
    Publication Date: 2014-11-06
    Description: Gelatinous zooplankton outbreaks have increased globally owing to a number of human-mediated factors, including food web alterations and species introductions. The invasive ctenophore Mnemiopsis leidyi entered the Black Sea in the early 1980s. The invasion was followed by the Azov, Caspian, Baltic and North Seas, and, most recently, the Mediterranean Sea. Previous studies identified two distinct invasion pathways of M. leidyi from its native range in the western Atlantic Ocean to Eurasia. However, the source of newly established populations in the Mediterranean Sea remains unclear. Here we build upon our previous study and investigate sequence variation in both mitochondrial (Cytochrome c Oxidase subunit I) and nuclear (Internal Transcribed Spacer) markers in M. leidyi, encompassing five native and 11 introduced populations, including four from the Mediterranean Sea. Extant genetic diversity in Mediterranean populations (n = 8, Na = 10) preclude the occurrence of a severe genetic bottleneck or founder effects in the initial colonizing population. Our mitochondrial and nuclear marker surveys revealed two possible pathways of introduction into Mediterranean Sea. In total, 17 haplotypes and 18 alleles were recovered from all surveyed populations. Haplotype and allelic diversity of Mediterranean populations were comparable to populations from which they were likely drawn. The distribution of genetic diversity and pattern of genetic differentiation suggest initial colonization of the Mediterranean from the Black-Azov Seas (pairwise FST = 0.001–0.028). However, some haplotypes and alleles from the Mediterranean Sea were not detected from the well-sampled Black Sea, although they were found in Gulf of Mexico populations that were also genetically similar to those in the Mediterranean Sea (pairwise FST = 0.010–0.032), raising the possibility of multiple invasion sources. Multiple introductions from a combination of Black Sea and native region sources could be facilitated by intense local and transcontinental shipping activity, respectively.
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