Small sea with high traffic - what is the biofouling potential of commercial ships in the Baltic Sea.

Despite the Baltic Sea being one of the most intensive shipping regions in the world the potential magnitude of the biofouled hulls in this region is unknown. This study estimated the biofouling load to Baltic Sea Region (BSR) based on the wetted surface area (WSA) method with regard to country, ship type and donor bioregion. WSA flux reached 656 km2, of which 86% was associated with ships operating inside and 14% was WSA flux brought by ships from outside of the Baltic Sea. Most of the WSA was transported to Swedish, Finnish and Danish ports as well. The highest WSA flux was assigned to roll-on/roll-off, passenger and general cargo ships. The high biofouling potential in BSR indicates a potential high risk to the environment and, therefore there is an urgent need for appropriate guidelines to be introduced into daily use by the commercial shipping community.

Integrating invasive species risk assessment into environmental DNA metabarcoding reference libraries.

Environmental DNA (eDNA) metabarcoding has shown promise as a tool for estimating biodiversity and early detection of invasive species. In aquatic systems, advantages of this method include the ability to concurrently monitor biodiversity and detect incipient invasions simply through the collection and analysis of water samples. However, depending on the molecular markers chosen for a given study, reference libraries containing target sequences from present species may limit the usefulness of eDNA metabarcoding. To explore the extent of this issue and how it may be resolved to aid biodiversity and invasive species early detection goals, we focus on fishes in the well-studied Laurentian Great Lakes region. First, we provide a synthesis of species currently known from the region and of non-indigenous species identified as threats by international, national, regional, and introduction pathway-specific fish risk assessments. With these species lists, we then evaluate 23 primer pairs commonly used in fish eDNA metabarcoding with available databases of sequence coverage and species specificity. Finally, we identify established and potentially invasive non-indigenous fish that should be prioritized for genetic sequencing to ensure robust eDNA metabarcoding for the region. Our results should increase confidence in using eDNA metabarcoding for fisheries conservation and management in the Great Lakes region and help prioritize reference sequencing efforts. The ultimate utility of eDNA metabarcoding approaches will come when conservation management of existing fish communities is integrated with early detection efforts for invasive species surveillance to assess total fish biodiversity.

Global Impacts of a Bilateral Trade Policy on Ballast Water-Mediated Species Spread Risk: A Case Study of Sino-US Trade.

A trade policy could generate both economic and environmental impacts. This work is focused on the impacts of a bilateral trade policy on ballast water-mediated nonindigenous species (NIS) spread risk. Taking the hypothetical Sino-US trade restriction as an example, we integrate a computable general equilibrium model and a higher-order NIS spread risk assessment model to examine the impacts of bilateral trade policy on both the economy and NIS spread risks. We have two important findings. First, the Sino-US trade restriction would cause decreases in NIS spread risks to China and the US, as well as to three quarters of worldwide countries/regions. However, the rest one fourth would experience increased NIS spread risks. Second, the relationship between changes in exports and changes in NIS spread risks might not be directly proportional. This is observed with 46% of countries and regions that would see their exports increase but their NIS spread risks drop, with positive impacts on both their economies and environment under the Sino-US trade restriction. These results reveal both broader global impacts as well as the decoupled economic and ecological impacts of a bilateral trade policy. These broader impacts demonstrate the necessity for national governments, which are parties to bilateral agreements to give due consideration to the economic and environmental impacts on countries and regions outside of the agreement.

Facilitation of non-indigenous ascidian by marine eco-engineering interventions at an urban site.

Marine artificial structures often support lower native species diversity and more non-indigenous species (NIS), but adding complex habitat and using bioreceptive materials have the potential to mitigate these impacts. Here, the interacting effects of structural complexity (flat, complex with pits) and concrete mixture (standard, or with oyster shell or vermiculite aggregate) on recruitment were assessed at two intertidal levels at an urban site. Complex tiles had less green algal cover, oyster shell mixtures had less brown (Ralfsia sp.) algal cover. At a low tidal elevation, the non-indigenous ascidian Styela plicata dominated complex tiles. Additionally, mixtures with oyster shell supported higher total cover of sessile species, and a higher cover of S. plicata. There were no effects of complexity or mixture on biofilm communities and native and NIS richness. Overall, these results suggest that habitat complexity and some bioreceptive materials may facilitate colonisation by a dominant invertebrate invader on artificial structures.

The Zooxanthellate Jellyfish Holobiont Cassiopea andromeda, a Source of Soluble Bioactive Compounds.

Cassiopea andromeda (Forsskål, 1775), commonly found across the Indo-Pacific Ocean, the Red Sea, and now also in the warmest areas of the Mediterranean Sea, is a scyphozoan jellyfish that hosts autotrophic dinoflagellate symbionts (family Symbiodiniaceae). Besides supplying photosynthates to their host, these microalgae are known to produce bioactive compounds as long-chain unsaturated fatty acids, polyphenols, and pigments, including carotenoids, with antioxidant properties and other beneficial biological activities. By the present study, a fractionation method was applied on the hydroalcoholic extract from two main body parts (oral arms and umbrella) of the jellyfish holobiont to obtain an improved biochemical characterization of the obtained fractions from the two body parts. The composition of each fraction (i.e., proteins, phenols, fatty acids, and pigments) as well as the associated antioxidant activity were analyzed. The oral arms proved richer in zooxanthellae and pigments than the umbrella. The applied fractionation method was effective in separating pigments and fatty acids into a lipophilic fraction from proteins and pigment-protein complexes. Therefore, the C. andromeda-dinoflagellate holobiont might be considered as a promising natural source of multiple bioactive compounds produced through mixotrophic metabolism, which are of interest for a wide range of biotechnological applications.

Prioritizing management of high-risk routes and ports by vessel type to improve marine biosecurity efforts.

The shipping industry constitutes the main vector of marine bioinvasions. Over 90,000 vessels world-wide create a highly complex shipping network that requires appropriate management tools. Here we characterized a novel vessel category, Ultra Large Container Vessels (ULCV), in terms of potential contribution to the dispersal of Non-Indigenous Species (NIS) in comparison to smaller vessels traveling similar routes. Such approach is essential for providing precise information-based risk analysis necessary to enforce biosecurity regulations and reduce the adverse global effects of marine NIS. We used Automatic Identification System (AIS) based websites to extract shipping data that will enable us to test for differences in two vessel behaviors linked to NIS dispersal: port visit durations and voyage sailing times. We then examined the geographic spread of ULCVs and small vessels, quantifying the accumulation of new port visits, countries, and ecoregions for each vessel category. Finally, Higher Order Network (HON) analysis revealed emergent patterns within shipping traffic, species flow, and invasion risk networks of these two categories. Compared to the smaller vessels, ULCVs spent significantly longer time in 20% of the ports and were more geographically constrained, with fewer port visits, countries, and regions. HON analysis revealed that the ULCV shipping species flow and invasion risk networks were more similar to each other than to those of the smaller vessels. However, HON port importance shifts were discernible for both vessel categories, with major shipping hubs not necessarily being major invasion hubs. Overall, compared to smaller vessels, ULCVs behave differently in ways that potentially increase biofouling risk, albeit in a smaller set of ports. Future studies using HON analysis of other dispersal vectors appears critical for prioritizing management of high-risk routes and ports.

Observations on Predation in Paranthura japonica Richardson, 1909 (Isopoda: Cymothoida: Paranthuridae).

Although isopods in Anthuroidea are predators, there is little information on their predatory behavior. In this study, we investigated predation by the paranthurid Paranthura japonica, which was originally described from northern Japan but has recently been reported as an invasive alien species in western America and Europe. Six crustacean species (two isopods, two amphipods, and two tanaidaceans, one of which does not co-occur with P. japonica in the wild) and one pycnogonid species were used as prey candidates in our experiments. Paranthura japonica preyed on all candidate species except the pycnogonid, grasping them with its falciform pereopods, inserting its piercing-type mouthparts, and sucking out the internal contents of the prey. Cannibalism or scavenging was observed when several P. japonica individuals were put in a single aquarium. This study showed that P. japonica is an aggressive predator; it consumed various crustaceans, including one it never encounters in the wild. Our results suggest that P. japonica will have a high impact on alien ecosystems it invades as a predator on native crustaceans. Cytochrome c oxidase subunit (COI) nucleotide sequences for putative P. japonica from Oshoro, Japan and a topotypic individual from Muroran confirmed that the population we dealt with was P. japonica.

The journey of hull-fouling mobile invaders: basibionts and boldness mediate dislodgement risk during transit.

Vessel hull-fouling is responsible for most bioinvasion events in the marine environment, yet it lacks regulation in most countries. Although experts advocate a preventative approach, research efforts on pre-arrival processes are limited. The performance of mobile epifauna during vessel transport was evaluated via laboratory simulations, using the well-known invasive Japanese skeleton shrimp (Caprella mutica), and its native congener C. laeviuscula as case study. The invader did not possess any advantage in terms of inherent resistance to drag. Instead, its performance was conditioned by the complexity of secondary substrate. Dislodgement risk was significantly reduced when sessile fouling basibionts were added, which provided refugia and boosted the probability of C. mutica remaining attached from 7 to 65% in flow exposure trials. Interestingly, the invader exhibited significantly higher exploratory tendency and motility than its native congener at zero-flow conditions. Implications in terms of en-route survivorship, invasion success and macrofouling management are discussed.

Bacterial communities in ballast tanks of cargo vessels - Shaped by salinity, treatment and the point of origin of the water but "hatch" its typical microbiome.

Ballast water is a main vector of introduction of potentially harmful or pathogenic aquatic organisms. The development of genetic tools for ballast water monitoring has been underway and highlighted as a source for accurate and reliable data for decision making. We used 16S rRNA gene amplicon sequencing to analyze the microbial communities found in the ballast water of fifteen commercial ships routed through two Estonian ports. In parallel, samples from the port area were collected at the same time each ship visited. Fluorescence microscopy was utilized to assess the effectiveness of the treatment applied to ballast water. In addition, supplemental samples were collected from Hamburg Port (Germany) and a ballast tank decontamination system used at this port. The composition and diversity of bacterial communities varied greatly between obtained samples. The application of UV treatment did not demonstrate significant reduction in species richness estimates. The composition of microbial communities was significantly influenced by salinity, treatment (mainly untreated or UV treated) and the point of origin of the ballast water. Over a hundred potentially pathogenic bacterial taxa were found in relatively high abundance, including in ballast water that had received UV treatment. These shortcomings of stand-alone UV treatment of ballast water, especially when weak treatment is applied insufficiently, highlight the danger of possible harmful effects arising over time and the need for genetic tools for ballast water monitoring and management.

Enhancing the resilience of Zostera noltei seagrass meadows against Arenicola spp. bio-invasion: A decision-making approach.

Seagrass meadows provide important and valuable ecosystem services. They are affected by several natural and human-induced stressors, but a combination of natural recovery and management actions have recently inverted the worldwide reduction. The main objectives of this study were to provide science-based knowledge on ecology and restoration, framed on environmental-related policies. By coupling the general guidelines with practical experience, obtained from sequential in situ experiments carried out for several months in a show-case study area, this study provides guidelines useful for restoration practitioners. A decision-making approach is proposed to answer the following questions: 1) What is the best Zostera noltei transplanting method? 2) What is the best technique to reduce the bioturbation activity of Arenicola spp.?, 3) Do bioturbation reduction techniques affect the survival rate of Z. noltei transplants?, and finally, 4) What are the key steps to maximize the success of a Z. noltei transplant and increase the species' resilience? Having a Portuguese coastal lagoon as show-case (Mira Channel, Ria de Aveiro), different transplant and restoration methodologies were tested (i.e. metal frames, nails, bamboo sticks, shoots inserted unanchored into the sediment, and intact units of sediment with seagrasses, named as SODs) to assure low environmental impact on donor meadows, high survival rate of transplanted shoots and the recovery of fragmented or lost meadows. Moreover, to potentially reverse a degraded Arenicola spp. colonized seagrass habitat, different types of natural membranes were tested. Results showed that the best transplanting method is the use of SODs as the self-facilitation process of Z. noltei is enhanced, while being the least invasive for the donor population. The use of a natural membrane can significantly decrease the bioturbation stress caused by Arenicola spp., with jute membrane being the best option, given its cost-handling-benefit trade-offs. Enhancing the success of seagrass restoration requires the implementation of effective measures by environmental restoration practitioners. We defined a three-step process to improve the resilience of Z. noltei. This stepwise approach consists on 1) Characterization of the donor population, 2) Identification of the constraints and implementation of measures to prevent them, and 3) Scale-up the restoration plan. The application of this stepwise approach in intertidal coastal and estuarine systems management will, therefore, facilitate the success of Z. noltei restoration plans.

Parental whole life cycle exposure modulates progeny responses to ocean acidification in slipper limpets.

Multigenerational exposure is needed to assess the evolutionary potential of organisms in the rapidly changing seascape. Here, we investigate if there is a transgenerational effect of ocean acidification exposure on a calyptraeid gastropod such that long-term exposure elevates offspring resilience. Larvae from wild type Crepidula onyx adults were reared from hatching until sexual maturity for over 36 months under three pH conditions (pH 7.3, 7.7, and 8.0). While the survivorship, growth, and respiration rate of F1 larvae were unaffected by acute ocean acidification (OA), long-term and whole life cycle exposure significantly compromised adult survivorship, growth, and reproductive output of the slipper limpets. When kept under low pH throughout their life cycle, only 6% of the F1 slipper limpets survived pH 7.3 conditions after ~2.5 years and the number of larvae they released was ~10% of those released by the control. However, the F2 progeny from adults kept under the long-term low pH condition hatched at a comparable size to those in medium and control pH conditions. More importantly, these F2 progeny from low pH adults outperformed F2 slipper limpets from control conditions; they had higher larval survivorship and growth, and reduced respiration rate across pH conditions, even at the extreme low pH of 7.0. The intragenerational negative consequences of OA during long-term acclimation highlights potential carryover effects and ontogenetic shifts in stress vulnerability, especially prior to and during reproduction. Yet, the presence of a transgenerational effect implies that this slipper limpet, which has been widely introduced along the West Pacific coasts, has the potential to adapt to rapid acidification.

Exceptions and exemptions under the ballast water management convention - Sustainable alternatives for ballast water management?

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.

The Role of Biofilms Developed under Different Anthropogenic Pressure on Recruitment of Macro-Invertebrates.

Microbial biofilms can be key mediators for settlement of macrofoulers. The present study examines the coupled effects of microbial biofilms and local environmental conditions on the composition, structure and functioning of macrofouling assemblages. Settlement of invertebrates over a gradient of human-impacted sites was investigated on local biofilms and on biofilms developed in marine protected areas (MPAs). Special attention was given to the presence of non-indigenous species (NIS), a global problem that can cause important impacts on local assemblages. In general, the formation of macrofouling assemblages was influenced by the identity of the biofilm. However, these relationships varied across levels of anthropogenic pressure, possibly influenced by environmental conditions and the propagule pressure locally available. While the NIS Watersipora subatra seemed to be inhibited by the biofilm developed in the MPA, Diplosoma cf. listerianum seemed to be attracted by biofilm developed in the MPA only under mid anthropogenic pressure. The obtained information is critical for marine environmental management, urgently needed for the establishment of prevention and control mechanisms to minimize the settlement of NIS and mitigate their threats.

Citizen science provides added value in the monitoring for coastal non-indigenous species.

Continuous and comprehensive monitoring is one of the most important practices to trace changes in the state of the environment and target management efforts. Yet, governmental resources are often insufficient for monitoring all required environmental parameters, and therefore authorities have started to utilize citizen observations to supplement and increase the scale of monitoring. The aims of the present study were to show the potential of citizen science in environmental monitoring by utilising citizen observations of the non-indigenous Harris mud crab Rhithropanopeus harrisii in Finnish waters, where coastal monitoring is insufficient to estimate the distribution and spread of non-indigenous species. Harris mud crab has shown measurable impact locally and is considered invasive. For reporting the status of invasions to national and European authorities and planning for potential eradication efforts, up to date knowledge on NIS ranges are needed. Citizen observations on the species were collected from the first observation onwards between 2009 and 2018, at first via email and later through an active citizen observation web portal (Invasive Alien Species Portal). The outcomes of the study indicate that species-specific citizen observations can be a beneficial addition to supplement national monitoring programs to fulfil legislative reporting requirements and to target potential management. Recognizable species and geographical areas with low biodiversity provide a good opportunity to utilize citizen observations. Moreover, citizen observations may enable distribution assessments for certain species that would otherwise require excessive resources and sampling efforts.

Considerations for incorporating real-time PCR assays into routine marine biosecurity surveillance programmes: a case study targeting the Mediterranean fanworm (Sabella spallanzanii) and club tunicate (Styela clava) 1.

Molecular techniques may provide effective tools to enhance marine biosecurity surveillance. Prior to routine implementation, evidence-based consideration of their benefits and limitations is needed. In this study, we assessed the efficiency and practicality of visual diver surveys and real-time PCR assays (targeting DNA and RNA) for detecting two marine invasive species whose infestation levels varied between species and location: Sabella spallanzanii and Styela clava. Filtered water samples (n = 171) were collected in parallel with dive surveys at two locations as part of the New Zealand Marine High Risk Site Surveillance programme: Nelson Harbour (27 sites) and Waitemata Harbour (30 sites). Diver surveys resulted in a greater number of detections compared to real-time PCR: S. clava - 21 versus 5 sites in Nelson, 6 versus 1 in Auckland; S. spallanzanii - 18 versus 10 in Auckland, no detections in Nelson. Occupancy modelling derived detection probabilities for the real-time PCR for S. clava were low (14%), compared to S. spallanzanii (66%). This could be related to abundances, or species-specific differences in DNA shedding. Only one RNA sample was positive, suggesting that most detections were from extracellular DNA or non-viable fragments. While molecular methods cannot yet replace visual observations, this study shows they provide useful complementary information.

The Asian cyclopoid copepod Mesocyclops pehpeiensis reported from the western basin of Lake Erie.

The Asian cyclopoid copepod Mesocyclops pehpeiensis Hu, 1943 has been reported as an introduced species at several locations in the western hemisphere. In the United States, reports of this exotic species are restricted to localities in Louisiana, Mississippi, and Washington D.C. This report documents a new record of occurrence for M. pehpeiensis from the western basin of Lake Erie. The detection of M. pehpeiensis in Lake Erie constitutes the first record of this species from the Laurentian Great Lakes, and the northernmost record in the western hemisphere. The species was found in 2016, 2017 and 2018, including females with egg sacks, and can therefore be considered established in the area. The occurrence of M. pehpeiensis in Lake Erie suggests that this Asian copepod may be more widely distributed in North America than is currently understood.

Non-indigenous species contribute equally to biofouling communities in international vs local ports in the Biobío region, Chile.

Growing coastal urbanization together with the intensification of maritime traffic are major processes explaining the increasing rate of biological introductions in marine environments. To investigate the link between international maritime traffic and the establishment of non-indigenous species (NIS) in coastal areas, biofouling communities in three international and three nearby local ports along 100 km of coastline in south-central Chile were compared using settlement panels and rapid assessment surveys. A larger number of NIS was observed in international ports, as expected in these 'invasion hubs'. However, despite a few environmental differences between international and local ports, the two port categories did not display significant differences regarding NIS establishment and contribution to community structure over the studied period (1.5 years). In international ports, the free space could be a limiting factor for NIS establishment. The results also suggest that local ports should be considered in NIS surveillance programs in Chile.

Human introduction or natural dispersion? Atlantic Ocean occurrence of the Indo-Pacific whitetip reef shark Triaenodon obesus.

Occurrence of multiple whitetip reef sharks Triaenodon obesus in the Atlantic Ocean is reported for the first time from near a sunken ferry off the Paraná coast in south-eastern Brazil. This occurrence is hypothesized to have been caused by either a human introduction or a remarkably long oceanic displacement.

First record of Pomacanthus maculosus (Perciformes, Pomacanthidae) in the south-western Atlantic Ocean.

A single Pomacanthus maculosus was filmed during a scuba diving survey on a rocky reef from southern Brazil. The body shape and coloration pattern confirmed the species identification. The biological and ecological characteristics of P. maculosus and the long distance of Brazil from its original distribution strongly suggest of an anthropogenic pathway of dispersion.

Benthic assemblages, biodiversity and invasiveness in marinas and commercial harbours: an investigation using a bioindicator group.

Fouling communities on artificial marine structures are generally different from benthic communities in natural rocky habitats. However, they may also differ among different types of artificial structures. Two artificial structures in direct contact with arriving vessels were compared: floating pontoons within recreational marinas, and sea-walls within commercial harbours. Natural rocky habitats were used as a reference, and the genus Eudendrium (Cnidaria, Hydrozoa) was chosen as a bioindicator. The assemblages were different among the three types of habitat studied, with different species characterising each habitat. The probability of finding an invasive Eudendrium species was significantly higher on pontoons. Diversity was the lowest on pontoons, but it was not significantly different between sea-walls and natural rocks. In general, a barrier to the spread of exotic species exists between harbours and natural rocky habitats. Floating pontoons seem to be a less suitable habitat for native fauna and a key element in marine biological invasions.