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To minimize the global transfer of harmful aquatic organisms and pathogens, the International Maritime Organization (IMO) has introduced the standard in Regulation D-2 to limit the number of viable organisms in ballast water discharged by ships. To meet the standard, many ships are installing ballast water management systems. Concurrently, regulators are looking for indicative analysis devices able to assess compliance with Regulation D-2, producing rapid, accurate and reliable results while being easy to operate. The purpose of this research is to compare four indicative analysis devices against detailed microscopy for measuring the size class of organisms ≥10 to <50 µm in minimum dimension (e.g., phytoplankton, including autotrophs, heterotrophs or mixotrophs), using field and laboratory tests. Comparisons were conducted on (treated) ballast water discharge samples collected across Canada during three consecutive years (2017-2019). During seven tests in 2019, paired ballast water uptake samples were also obtained, facilitating measurements before and after treatment was applied. Indicative analysis devices also were challenged with natural environmental samples containing different organism abundance levels, ranging from low (nominally <10 cells mL-1) to high (nominally >150 cells mL-1) during laboratory tests. While the indicative analysis devices examined during this research produced numeric estimates having weak correlations with the standard reference method, categorical outcomes (above/below the D-2 standard) had high agreement (89% or better) when assessing ballast water samples, but lower agreement (67% or poorer) during laboratory tests. There was a relatively high rate of false negative results measured by all devices during laboratory tests. Results provided by indicative analysis devices had higher uncertainty when organism abundances in ballast samples are below and close to the D-2 standards.
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Navios , Água , Organismos Aquáticos , Canadá , Espécies Introduzidas , FitoplânctonRESUMO
The movement of ballast water by commercial shipping is a prominent pathway for aquatic invasions. Ships' ballast water management is now transitioning from open ocean exchange to a ballast water performance standard that will effectively require use of onboard treatment systems. Neither strategy is perfect, therefore, combined use of ballast water exchange plus treatment has been suggested to provide greatest protection of aquatic ecosystems. This study compared the performance of exchange plus treatment against treatment alone by modeling establishment rates of nonindigenous zooplankton introduced by ballast water across different habitat types (fresh, brackish, and marine) in Canada. Treatment was modeled under two efficacy scenarios (100% and 50% of ship trips) to consider the possibility that treatment may not always be successful. The model results indicate that exchange plus treatment will be more effective than treatment alone at reducing establishments when recipient ports are freshwater (58â¯140 vs 11â¯338 trips until ≥1 establishment occurs, respectively). Exchange plus treatment also serves as an important backup strategy if treatment systems are partially effective (50% of trips), primarily for freshwater recipient ecosystems (1442 versus 585 trips until ≥1 establishment occurs, respectively).
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Ecossistema , Água , Animais , Canadá , Água Doce , Espécies Introduzidas , NaviosRESUMO
The International Convention for the Control and Management of Ships' Ballast Water and Sediments (BWM Convention) aims to mitigate the introduction risk of harmful aquatic organisms and pathogens (HAOP) via ships' ballast water and sediments. The BWM Convention has set regulations for ships to utilise exceptions and exemptions from ballast water management under specific circumstances. This study evaluated local and regional case studies to provide clarity for situations, where ships could be excepted or exempted from ballast water management without risking recipient locations to new introductions of HAOP. Ships may be excepted from ballast water management if all ballasting operations are conducted in the same location (Regulation A-3.5 of the BWM Convention). The same location case study determined whether the entire Vuosaari harbour (Helsinki, Finland) should be considered as the same location based on salinity and composition of HAOP between the two harbour terminals. The Vuosaari harbour case study revealed mismatching occurrences of HAOP between the harbour terminals, supporting the recommendation that exceptions based on the same location concept should be limited to the smallest feasible areas within a harbour. The other case studies evaluated whether ballast water exemptions could be granted for ships using two existing risk assessment (RA) methods (Joint Harmonised Procedure [JHP] and Same Risk Area [SRA]), consistent with Regulation A-4 of the BWM Convention. The JHP method compares salinity and presence of target species (TS) between donor and recipient ports to indicate the introduction risk (high or low) attributed to transferring unmanaged ballast water. The SRA method uses a biophysical model to determine whether HAOP could naturally disperse between ports, regardless of their transportation in ballast water. The results of the JHP case study for the Baltic Sea and North-East Atlantic Ocean determined that over 97% of shipping routes within these regions resulted in a high-risk indication. The one route assessed in the Gulf of Maine, North America also resulted in a high-risk outcome. The SRA assessment resulted in an overall weak connectivity between all ports assessed within the Gulf of the St. Lawrence, indicating that a SRA-based exemption would not be appropriate for the entire study area. In summary, exceptions and exemptions should not be considered as common alternatives for ballast water management. The availability of recent and detailed species occurrence data was considered the most important factor to conduct a successful and reliable RA. SRA models should include biological factors that influence larval dispersal and recruitment potential (e.g., pelagic larval duration, settlement period) to provide a more realistic estimation of natural dispersal.
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Espécies Introduzidas , Água , Oceano Atlântico , Finlândia , Maine , América do Norte , Navios , Abastecimento de ÁguaRESUMO
Climate change and increased anthropogenic activities are expected to elevate the potential of introducing nonindigenous species (NIS) into the Arctic. Yet, the knowledge base needed to identify gaps and priorities for NIS research and management is limited. Here, we reviewed primary introduction events to each ecoregion of the marine Arctic realm to identify temporal and spatial patterns, likely source regions of NIS, and the putative introduction pathways. We included 54 introduction events representing 34 unique NIS. The rate of NIS discovery ranged from zero to four species per year between 1960 and 2015. The Iceland Shelf had the greatest number of introduction events (n = 14), followed by the Barents Sea (n = 11), and the Norwegian Sea (n = 11). Sixteen of the 54 introduction records had no known origins. The majority of those with known source regions were attributed to the Northeast Atlantic and the Northwest Pacific, 19 and 14 records, respectively. Some introduction events were attributed to multiple possible pathways. For these introductions, vessels transferred the greatest number of aquatic NIS (39%) to the Arctic, followed by natural spread (30%) and aquaculture activities (25%). Similar trends were found for introductions attributed to a single pathway. The phyla Arthropoda and Ochrophyta had the highest number of recorded introduction events, with 19 and 12 records, respectively. Recommendations including vector management, horizon scanning, early detection, rapid response, and a pan-Arctic biodiversity inventory are considered in this paper. Our study provides a comprehensive record of primary introductions of NIS for marine environments in the circumpolar Arctic and identifies knowledge gaps and opportunities for NIS research and management. Ecosystems worldwide will face dramatic changes in the coming decades due to global change. Our findings contribute to the knowledge base needed to address two aspects of global change-invasive species and climate change.
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Organismos Aquáticos/fisiologia , Mudança Climática , Espécies Introduzidas/tendências , Animais , Regiões Árticas , Biodiversidade , Ecossistema , Espécies Introduzidas/estatística & dados numéricos , RiscoRESUMO
Invasions of freshwater habitats by marine and brackish species have become more frequent in recent years with many of those species originating from the Ponto-Caspian region. Populations of Ponto-Caspian species have successfully established in the North and Baltic Seas and their adjoining rivers, as well as in the Great Lakes-St. Lawrence River region. To determine if Ponto-Caspian taxa more readily acclimatize to and colonize diverse salinity habitats than taxa from other regions, we conducted laboratory experiments on 22 populations of eight gammarid species native to the Ponto-Caspian, Northern European and Great Lakes-St. Lawrence River regions. In addition, we conducted a literature search to survey salinity ranges of these species worldwide. Finally, to explore evolutionary relationships among examined species and their populations, we sequenced the mitochondrial cytochrome c oxidase subunit I gene (COI) from individuals used for our experiments. Our study revealed that all tested populations tolerate wide ranges of salinity, however, different patterns arose among species from different regions. Ponto-Caspian taxa showed lower mortality in fresh water, while Northern European taxa showed lower mortality in fully marine conditions. Genetic analyses showed evolutionary divergence among species from different regions. Due to the geological history of the two regions, as well as high tolerance of Ponto-Caspian species to fresh water, whereas Northern European species are more tolerant of fully marine conditions, we suggest that species originating from the Ponto-Caspian and Northern European regions may be adapted to freshwater and marine environments, respectively. Consequently, the perception that Ponto-Caspian species are more successful colonizers might be biased by the fact that areas with highest introduction frequency of NIS (i.e., shipping ports) are environmentally variable habitats which often include freshwater conditions that cannot be tolerated by euryhaline taxa of marine origin.
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Anfípodes/fisiologia , Evolução Biológica , Espécies Introduzidas , Tolerância ao Sal , Animais , Proteínas de Artrópodes/genética , Canadá , Complexo IV da Cadeia de Transporte de Elétrons/genética , Europa (Continente) , Evolução Molecular , Proteínas Mitocondriais/genética , Estados UnidosRESUMO
Biological invasions are largely considered to be a "numbers game", wherein the larger the introduction effort, the greater the probability that an introduced population will become established. However, conditions during transport - an early stage of the invasion - can be particularly harsh, thereby greatly reducing the size of a population available to establish in a new region. Some successful non-indigenous species are more tolerant of environmental and anthropogenic stressors than related native species, possibly stemming from selection (ie survival of only pre-adapted individuals for particular environmental conditions) during the invasion process. By reviewing current literature concerning population genetics and consequences of selection on population fitness, we propose that selection acting on transported populations can facilitate local adaptation, which may result in a greater likelihood of invasion than predicted by propagule pressure alone. Specifically, we suggest that detailed surveys should be conducted to determine interactions between molecular mechanisms and demographic factors, given that current management strategies may underestimate invasion risk.
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Mixed-species assemblages are often unintentionally introduced into new ecosystems. Analysing how assemblage structure varies during transport may provide insights into how introduction risk changes before propagules are released. Characterization of introduction risk is typically based on assessments of colonization pressure (CP, the number of species transported) and total propagule pressure (total PP, the total abundance of propagules released) associated with an invasion vector. Generally, invasion potential following introduction increases with greater CP or total PP. Here, we extend these assessments using rank-abundance distributions to examine how CP : total PP relationships change temporally in ballast water of ocean-going ships. Rank-abundance distributions and CP : total PP patterns varied widely between trans-Atlantic and trans-Pacific voyages, with the latter appearing to pose a much lower risk than the former. Responses also differed by taxonomic group, with invertebrates experiencing losses mainly in total PP, while diatoms and dinoflagellates sustained losses mainly in CP. In certain cases, open-ocean ballast water exchange appeared to increase introduction risk by uptake of new species or supplementation of existing ones. Our study demonstrates that rank-abundance distributions provide new insights into the utility of CP and PP in characterizing introduction risk.
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Distribuição Animal , Organismos Aquáticos , Espécies Introduzidas , Animais , Ecossistema , Modelos Lineares , Modelos Teóricos , Densidade Demográfica , Risco , Água do Mar , NaviosRESUMO
The most effective way to manage species transfers is to prevent their introduction via vector regulation. Soon, international ships will be required to meet numeric ballast discharge standards using ballast water treatment (BWT) systems, and ballast water exchange (BWE), currently required by several countries, will be phased out. However, there are concerns that BWT systems may not function reliably in fresh and/or turbid water. A land-based evaluation of simulated "BWE plus BWT" versus "BWT alone" demonstrated potential benefits of combining BWE with BWT for protection of freshwater ecosystems. We conducted ship-based testing to compare the efficacy of "BWE plus BWT" versus "BWT alone" on voyages starting with freshwater ballast. We tested the hypotheses that there is an additional effect of "BWE plus BWT" compared to "BWT alone" on the reduction of plankton, and that taxa remaining after "BWE plus BWT" will be marine (low risk for establishment at freshwater recipient ports). Our study found that BWE has significant additional effect on the reduction of plankton, and this effect increases with initial abundance. As per expectations, "BWT alone" tanks contained higher risk freshwater or euryhaline taxa at discharge, while "BWE plus BWT" tanks contained mostly lower risk marine taxa unlikely to survive in recipient freshwater ecosystems.
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Navios , Purificação da Água/métodos , Animais , Ecossistema , Água Doce , Espécies Introduzidas , Fitoplâncton , ZooplânctonRESUMO
A recent model demonstrated that the adenosine triphosphate (ATP) content of spherical aquatic organisms with a 10 to 50 µm diameter is between 0.16 and 19.9 pg cell-1. Here, the model is validated by comparing microscopy-based counts with ATP concentrations from a commercial ATP kit. The measured ATP content of both freshwater and marine organisms 10 to 50 µm size range falls in the 0.16 to 19.9 pg cell-1 model range. On average, freshwater organisms contain 0.33 pg ATP cell-1, have a spherical equivalent diameter (SED) of 13 µm, while marine organisms have 0.89 pg ATP cell-1 and a SED of 18 µm. In addition, their 13 to 18 µm size is within the 10 to 50 µm ballast water size range and in agreement with the 15 µm mean SED of a coastal plankton size-distribution model. This study concludes that the ATP-model is reliable, emphasizing the need for caution when converting three-dimensional biomass proxies into linear cell concentrations.
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Trifosfato de Adenosina , Plâncton , Organismos Aquáticos , Água , Água DoceRESUMO
Standardised terminology in science is important for clarity of interpretation and communication. In invasion science - a dynamic and rapidly evolving discipline - the proliferation of technical terminology has lacked a standardised framework for its development. The result is a convoluted and inconsistent usage of terminology, with various discrepancies in descriptions of damage and interventions. A standardised framework is therefore needed for a clear, universally applicable, and consistent terminology to promote more effective communication across researchers, stakeholders, and policymakers. Inconsistencies in terminology stem from the exponential increase in scientific publications on the patterns and processes of biological invasions authored by experts from various disciplines and countries since the 1990s, as well as publications by legislators and policymakers focusing on practical applications, regulations, and management of resources. Aligning and standardising terminology across stakeholders remains a challenge in invasion science. Here, we review and evaluate the multiple terms used in invasion science (e.g. 'non-native', 'alien', 'invasive' or 'invader', 'exotic', 'non-indigenous', 'naturalised', 'pest') to propose a more simplified and standardised terminology. The streamlined framework we propose and translate into 28 other languages is based on the terms (i) 'non-native', denoting species transported beyond their natural biogeographic range, (ii) 'established non-native', i.e. those non-native species that have established self-sustaining populations in their new location(s) in the wild, and (iii) 'invasive non-native' - populations of established non-native species that have recently spread or are spreading rapidly in their invaded range actively or passively with or without human mediation. We also highlight the importance of conceptualising 'spread' for classifying invasiveness and 'impact' for management. Finally, we propose a protocol for classifying populations based on (i) dispersal mechanism, (ii) species origin, (iii) population status, and (iv) impact. Collectively and without introducing new terminology, the framework that we present aims to facilitate effective communication and collaboration in invasion science and management of non-native species.
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Espécies Introduzidas , Terminologia como Assunto , AnimaisRESUMO
Nonindigenous species (NIS) cause global biotic homogenization and extinctions, with commercial shipping being a leading vector for spread of aquatic NIS. To reduce transport of NIS by ships, regulations requiring ballast water exchange (BWE) have been implemented by numerous countries. BWE appears to effectively reduce risk for freshwater ports, but provides only moderate protection of marine ports. In the near future, ships may be required to undertake ballast water treatment (BWT) to meet numeric performance standards, and BWE may be phased out of use. However, there are concerns that BWT systems may not operate reliably in fresh or turbid water, or both. Consequently, it has been proposed that BWE could be used in combination with BWT to maximize the positive benefits of both management strategies for protection of freshwater ports. We compared the biological efficacy of "BWE plus BWT" against "BWT alone" at a ballast water treatment experimental test facility. Our comparative evaluation showed that even though BWT alone significantly reduced abundances of all tested organism groups except total heterotrophic bacteria, the BWE plus BWT strategy significantly reduced abundances for all groups and furthermore resulted in significantly lower abundances of most groups when compared to BWT alone. Our study clearly demonstrates potential benefits of combining BWE with BWT to reduce invasion risk of freshwater organisms transported in ships' ballast water, and it should be of interest to policy makers and environmental managers.
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Conservação dos Recursos Naturais/métodos , Espécies Introduzidas , Enterococcus/crescimento & desenvolvimento , Escherichia coli/crescimento & desenvolvimento , Great Lakes Region , Processos Heterotróficos , Navios , Purificação da ÁguaRESUMO
The environmental similarity scores between source and recipient locations are essential in ballast water risk assessment (BWRA) models used to estimate the potential for non-indigenous species (NIS) introduction, survival, and establishment, and to guide management strategies aiming to minimize biodiversity loss and economic impacts. Previous BWRA models incorporate annual-scale environmental data, which may overlook seasonal variability. In this study, temporal variation in sea surface temperature and salinity data were examined at global ports, and the influence of this variation on environmental distance calculations (and corresponding risk of NIS) was examined for ballast water discharges in Canada by comparing outputs from monthly and annual scale assessments in a BWRA model. Except for some outliers in the Pacific region, the environmental distances based on monthly scale data generally become smaller in all regions, demonstrating that the model using annual decadal average environmental data to inform environmental matching can underestimate risk of NIS survival and establishment in comparison to monthly data. The results of this study suggest future evaluations incorporating the date of ballast water uptake and discharge can provide a more sensitive assessment of risk reflecting seasonal variability compared to an annual average risk model.
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A number of ballast water compliance monitoring devices (CMDs) have been made commercially available to verify the efficacy of ballast water management systems by quantifying the living organisms for both plankton size classes (≥50 µm and ≥10-<50 µm). This study aimed to examine whether new CMDs can provide a reliable indication of compliance regarding Regulation D-2 and to evaluate their performance for indicative analysis of organisms by assessing their accuracy (comparison to microscopy) and precision (comparison within measurement). Challenge fresh water samples were collected in four locations of Lake Ontario, Canada, whereas marine challenge water samples were collected around the Bay of Fundy, New Brunswick, Canada. Ballast water samples were collected from ships visiting several ports across Canada. Overall, accuracy was higher (>80%) in estimating organisms from prepared-challenge water (Ballast Eye and BallastWISE) than from ballast water samples (>70%) (B-QUA only). The sensitivity ranged from 50 to 100% for the ≥50 µm organism size class, whereas for the ≥10-<50 µm organism size class, it was higher for freshwater samples (>75%) than for marine samples (>50%). The performance of CMDs should be assessed under real-world conditions for a better understanding and to improve their use.
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Increasing empirical evidence indicates the number of released individuals (i.e. propagule pressure) and number of released species (i.e. colonization pressure) are key determinants of the number of species that successfully invade new habitats. In view of these relationships, and the possibility that ships transport whole communities of organisms, we collected 333 ballast water and sediment samples to investigate the relationship between propagule and colonization pressure for a variety of diverse taxonomic groups (diatoms, dinoflagellates and invertebrates). We also reviewed the scientific literature to compare the number of species transported by ships to those reported in nature. Here, we show that even though ships transport nearly entire local communities, a strong relationship between propagule and colonization pressure exists only for dinoflagellates. Our study provides evidence that colonization pressure of invertebrates and diatoms may fluctuate widely irrespective of propagule pressure. We suggest that the lack of correspondence is explained by reduced uptake of invertebrates into the transport vector and the sensitivity of invertebrates and diatoms to selective pressures during transportation. Selection during transportation is initially evident through decreases in propagule pressure, followed by decreased colonization pressure in the most sensitive taxa.
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Diatomáceas/fisiologia , Dinoflagellida/fisiologia , Ecossistema , Invertebrados/fisiologia , Água do Mar , Navios , Animais , Incrustação Biológica , Diatomáceas/classificação , Dinoflagellida/classificação , Sedimentos Geológicos/microbiologia , Sedimentos Geológicos/parasitologia , Espécies Introduzidas , Invertebrados/classificação , Água do Mar/microbiologia , Água do Mar/parasitologiaRESUMO
Ballast water is a leading pathway for the global introduction of aquatic nonindigenous species. Most international ships are expected to install ballast water management systems (BWMS) by 2024 to treat ballast water before release. This study examines if ballast water discharges managed by BWMS are meeting standards for organisms ≥50 µm in minimum dimension (i.e., <10 organisms per m3; typically zooplankton). Representative samples of ballast water were collected from 29 ships (using 14 different BWMS) arriving to Canada during 2017-2018. Fourteen samples (48 %) had zooplankton concentrations clearly exceeding the standard (ranging from 18 to 3822 organisms per m3). Nonetheless, compared to earlier management strategies, BWMS appear to reduce the frequency of high-risk introduction events. BWMS filter mesh size was an important predictor of zooplankton concentration following treatment. Greater rates of compliance may be achieved as ship crews gain experience with operation and maintenance of BWMS.
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Navios , Zooplâncton , Animais , Espécies Introduzidas , Água , Abastecimento de ÁguaRESUMO
Enactment of any environmental policy should be followed by an evaluation of its efficacy to ensure optimal utilization of limited resources, yet measuring the success of these policies can be a challenging task owing to a dearth of data and confounding factors. We examine the efficacy of ballast water policies enacted to prevent biological invasions in the Laurentian Great Lakes. We utilize four criteria to assess the efficacy of this environmental regulation: (1) Is the prescribed management action demonstrably effective? (2) Is the management action effective under operational conditions? (3) Can compliance be achieved on a broad scale? (4) Are desired changes observed in the environment? The four lines of evidence resulting from this analysis indicate that the Great Lakes ballast water management program provides robust, but not complete, protection against ship-mediated biological invasions. Our analysis also indicates that corresponding inspection and enforcement efforts should be undertaken to ensure that environmental policies translate into increased environmental protection. Similar programs could be implemented immediately around the world to protect the biodiversity of the many freshwater ecosystems which receive ballast water discharges by international vessels. This general framework can be extended to evaluate efficacy of other environmental policies.
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Organismos Aquáticos , Política Ambiental , Espécies Introduzidas/legislação & jurisprudência , Navios/legislação & jurisprudência , Conservação dos Recursos Naturais , Regulamentação Governamental , Espécies Introduzidas/estatística & dados numéricos , Navios/métodosRESUMO
To verify ships' compliance with ballast water regulations, samples may be collected and tested for viable organisms. This task is completed using a sample probe, which is placed in the ballast discharge pipe through a sample port (a flanged opening). To collect representative samples, the placement of the sample port and the size of the sample probe must be appropriate for the shipboard piping arrangement and ballast water flows. The placement of sample ports was evaluated on 72 ships to assess the current condition of ballast water sampling installations against available guidance. Few ships (15%) had sample ports fully aligned with International Organization for Standardization (ISO) standard 11711-1. While current configurations may present challenges in collecting representative samples, these installations likely occurred before the ISO standard was available. Future installations should be in accordance with the standard to facilitate representative sampling.
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Navios , Água , Espécies Introduzidas , Padrões de ReferênciaRESUMO
Aim: The introduction of aquatic non-indigenous species (ANS) has become a major driver for global changes in species biogeography. We examined spatial patterns and temporal trends of ANS detections since 1965 to inform conservation policy and management. Location: Global. Methods: We assembled an extensive dataset of first records of detection of ANS (1965-2015) across 49 aquatic ecosystems, including the (a) year of first collection, (b) population status and (c) potential pathway(s) of introduction. Data were analysed at global and regional levels to assess patterns of detection rate, richness and transport pathways. Results: An annual mean of 43 (±16 SD) primary detections of ANS occurred-one new detection every 8.4 days for 50 years. The global rate of detections was relatively stable during 1965-1995, but increased rapidly after this time, peaking at roughly 66 primary detections per year during 2005-2010 and then declining marginally. Detection rates were variable within and across regions through time. Arthropods, molluscs and fishes were the most frequently reported ANS. Most ANS were likely introduced as stowaways in ships' ballast water or biofouling, although direct evidence is typically absent. Main conclusions: This synthesis highlights the magnitude of recent ANS detections, yet almost certainly represents an underestimate as many ANS go unreported due to limited search effort and diminishing taxonomic expertise. Temporal rates of detection are also confounded by reporting lags, likely contributing to the lower detection rate observed in recent years. There is a critical need to implement standardized, repeated methods across regions and taxa to improve the quality of global-scale comparisons and sustain core measures over longer time-scales. It will be fundamental to fill in knowledge gaps given that invasion data representing broad regions of the world's oceans are not yet readily available and to maintain knowledge pipelines for adaptive management.
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Zooplankton populations are spatially heterogeneous in nature and inside ship ballast tanks. Sampling methods should take heterogeneity into account, particularly when estimating quantitative variables such as abundance or concentration. It is particularly important to generate unbiased estimates of zooplankton concentration in ballast water when assessing compliance with new international ballast water discharge standards. We measured spatial heterogeneity of zooplankton within ballast water using three sampling methodologies. In-tank pump samples were collected at fixed depths within the vertical part of the ballast tank (side tank). Vertical net-haul samples were collected from the upper portion of the tank as a depth-integrated and historically relevant method. In-line, time-integrated samples were collected during ballast discharge by an isokinetic sample probe, likely representing the double bottom part of the ballast tank. The bias and precision associated with each sampling method were evaluated in reference to the estimated average abundance of the entire ballast tank, which was modeled from the data collected by all methods. In-tank pump samples provided robust evidence for vertical stratification of zooplankton concentration in the side tank. A consistent trend was also observed for in-line discharge samples, with zooplankton concentration decreasing through time as the ballast tank is being discharged. Sample representativeness, as compared to the tank average, varied depending on the depth or tank volume discharged. In-line discharge samples provided the least biased and most precise estimate of average tank abundance (having lowest mean squared error) when collected during the time frame of 20%-60% of the tank volume being discharged. Results were consistent across five trips despite differences in ballast water source, season, and age.
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Human-mediated vectors often inadvertently translocate species assemblages to new environments. Examining the dynamics of entrained species assemblages during transport can provide insights into the introduction risk associated with these vectors. Ship biofouling is a major transport vector of nonindigenous species in coastal ecosystems globally, yet its magnitude in the Arctic is poorly understood. To determine whether biofouling organisms on ships can survive passages in Arctic waters, we examined how biofouling assemblage structure changed before, during, and after eight round-trip military voyages from temperate to Arctic ports in Canada. Species richness first decreased (~70% loss) and then recovered (~27% loss compared to the original assemblages), as ships travelled to and from the Arctic, respectively, whereas total abundance typically declined over time (~55% total loss). Biofouling community structure differed significantly before and during Arctic transits as well as between those sampled during and after voyages. Assemblage structure varied across different parts of the hull; however, temporal changes were independent of hull location, suggesting that niche areas did not provide protection for biofouling organisms against adverse conditions in the Arctic. Biofouling algae appear to be more tolerant of transport conditions during Arctic voyages than are mobile, sessile, and sedentary invertebrates. Our results suggest that biofouling assemblages on ships generally have poor survivorship during Arctic voyages. Nonetheless, some potential for transporting nonindigenous species to the Arctic via ship biofouling remains, as at least six taxa new to the Canadian Arctic, including a nonindigenous cirripede, appeared to have survived transits from temperate to Arctic ports.