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Non-indigenous species (NIS) are of concern for biodiversity conservation and ecosystem functioning. We present an updated list of NIS, including cryptogenic species, from Danish marine waters containing 123 species. Benthic invertebrates (36%) and phytoplankton (28%) dominate the list, but fish (15%) and macroalgae (13%) are also important. The Limfjord in Northern Jutland emerges as a hotspot for the introduction of NIS. Data from multiple sources were included, i.e., the National Monitoring Program (NOVANA), the National Fish Atlas project, the citizen science project Arter.dk, research articles, and annual national reports of the ICES working group ITMO. Forty-six NIS species were subject to expert judging using a modified Harmonia protocol; 19 were found to fulfil the four selected criteria identifying a species as being ‘invasive’. Additionally, 38 species, not yet recorded in Danish waters, were evaluated using the same method, and 31 were found to fulfil the ‘invasive’ criteria. For nine selected species, introduction history, distribution maps, and time-series diagrams are presented. Our data document that the national monitoring efforts should be expanded to record macrozooplankton, coastal fish, and mobile epibenthic species. Furthermore, the national data repository, Arter.dk, should be expanded to enable more detailed documentation of new NIS records.

Larvaceans are gelatinous zooplankton abundant throughout the ocean. Larvaceans have been overlooked in research because they are difficult to collect and are perceived as being unimportant in biogeochemical cycles and food-webs. We synthesise evidence that their unique biology enables larvaceans to transfer more carbon to higher trophic levels and deeper into the ocean than is commonly appreciated. Larvaceans could become even more important in the Anthropocene because they eat small phytoplankton that are predicted to become more prevalent under climate change, thus moderating projected future declines in ocean productivity and fisheries. We identify critical knowledge gaps and argue that larvaceans should be incorporated into ecosystem assessments and biogeochemical models to improve predictions of the future ocean.

Highlights: • Sightings of non-native of range expanding marine species in the SW Baltic Sea • Novel approach for classification of hydrographic conditions in the SW Baltic Sea • Linking sightings of species to hydrographic environmental anomalies Abstract: The Baltic Sea, located in northern Europe, is one of the largest brackish water bodies in the world. Salinity levels range from fresh water conditions in the northeast to full strength saline waters at its transition zone to the North Sea in the west. Most of the water exchange happens in the SW Baltic Sea, the Belt Sea, where fresh water exits the Baltic Sea at the surface, while high saline, oxygen rich water is entering the Baltic at depth. Due to the extended salinity gradient of the Baltic Sea, a variety of species occur at the limit of their physiological tolerance and preference, i.e. in areas and habitats not representing their marine or fresh water origin. Additionally, the Baltic Sea is known for its high share of non-indigenous species, which have established. In this study, we compiled extraordinary sightings of transient, non-native or potentially range expanding species in the SW Baltic Sea for a period from 2001 to 2018. We focused on jellyfish, squid, fishes and marine mammals and linked their occurrences to the local hydrography. Hydrographic conditions, such as water temperature and salinity, were obtained from a high spatio-temporally resolved hydrodynamic Baltic Sea model, covering a daily resolved 40-year time series. We investigated that changes in the occurrence of exceptional species reflect the dynamics of water mass exchange between the Kattegat/Skagerrak and the SW Baltic Sea. Our analyses show that these changes could be related to the presence of anomalously high saline water masses. However, only a minor part of the sightings was caused by major Baltic inflow events, which are important to sustain oxygen rich deep water in the central Baltic Sea. This documents that the hydrographic

The goal of the ICES Working Group on the Introduction and Transfers of Marine Organisms (WGITMO) is to contribute to the ICES vision and mission by addressing specific science objectives related to the introduction, spread and impacts of non-indigenous marine species (NIS). The expert working group provides information and advice on the impacts of human activities (e.g. pressures caused by invasive species, litter, aquaculture, biofouling) on marine ecosystems including the Arctic. The Terms of Reference include annual national reporting on NIS detections and research, climate change impact on the spread of NIS, the risk and impacts of NIS, vectors of spread of NIS including biofouling, marine debris and aquaculture. This report summarizes the key findings and outcomes from the 2020–2022 term. A joint meeting in (2020) with WGBOSV [Ballast Water and Other Ship Vectors] and WGHABD [Harmful Algal Bloom Dynamics] was held to discuss scientific issues of joint interest including climate change in the Arctic environment, and early detection of and response to species through the use of innovative molecular tools. The discussion on molecular tools was used by the groups to develop the framework for a targeted Molecular Tools workshop in conjunction with at the International Conference for Marine Bioinvasions (ICMB) in May 2023. National activities were presented and discussed from over 20 countries each year. Common themes included, new species introduction and monitoring, recent activities on traditional and innovative methods for detection and surveillance (including molecular tools). Science highlights include the submission of the ICES biofouling viewpoint to the International Maritime Organization (IMO) regarding biofouling guidelines, development and publication of a global scale NIS screening tool, and publication of a global review of NIS trends over 50 years. The Information system on aquatic non-indigenous and cryptogenic species AquaNIS continu

High throughput low-density SNP arrays provide a cost-effective solution for population genetic studies and monitoring of genetic diversity as well as population structure commonly implemented in real time stock assessment of commercially important fish species. However, the application of high throughput SNP arrays for monitoring of invasive species has so far not been implemented. We developed a species-specific SNP array for the invasive comb jelly Mnemiopsis leidyi based on whole genome re-sequencing data. Initially, a total of 1,395 high quality SNPs were identified using stringent filtering criteria. From those, 192 assays were designed and validated, resulting in the final panel of 116 SNPs. Markers were diagnostic between the northern and southern M. leidyi lineages and highly polymorphic to distinguish populations. Despite using a reduced representation of the genome, our SNP panel yielded comparable results to using a whole genome re-sequencing approach (832,323 SNPs), recovering similar values of genetic differentiation between samples and detecting the same clustering groups when performing Structure analyses. The resource presented here provides a cost-effective, high throughput solution for population genetic studies, allowing to routinely genotype large number of individuals. Monitoring of genetic diversity and effective population size estimations in this highly invasive species will allow for the early detection of new introductions from distant source regions or hybridization events. Thereby, this SNP chip represents an important management tool in order to understand invasion dynamics and opens the door for implementing such methods for a wider range of non-indigenous invasive species.

This data article includes a qualitative and quantitative description of the gelatinous macrozooplankton community of the North Sea during January-February 2021. Sampling was conducted during the 1st quarter International Bottom Trawl Survey (IBTS) on board the Danish R/V DANA (DTU Aqua Denmark) and the Swedish R/V Svea (SLU Sweden), as part of the ichthyoplankton investigation during night-time. A total of 147 stations were investigated in the western, central and eastern North Sea as well as the Skagerrak and Kattegat. Sampling was conducted with a 13 m long Midwater Ring Net (MIK net, Ø 2 m, mesh size 1.6 mm, cod end with smaller mesh size of 500 µm), equipped with a flow meter. The MIK net was deployed in double oblique hauls from the surface to c. 5 m above the sea floor [1,2]. Samples were visually analysed unpreserved on a light table and/or with a stereomicroscope or magnifying lamp within 2 hours after catch. A total of 13,510 individuals were counted/sized. Twelve gelatinous macrozooplankton species or genera were encountered, namely the hydrozoan Aequorea vitrina, Aglantha digitale, Clytia spp., Leuckartiara octona, Tima bairdii, Muggiaea atlantica; the scyphozoans Cyanea capillata and Cyanea lamarckii and the ctenophores Beroe spp., Bolinopsis infundibulum, Mnemiopsis leidyi, Pleurobrachia pileus. Abundance data are presented on a volume specific (m−3) and area specific (m−2) basis. Size data have been used to estimate wet weights based on published length-weight regressions (see Table 1). For the groups i) hydrozoan jellyfish, ii) scyphozoan jellyfish, iii) ctenophores, as well as iv) grouped gelatinous macrozooplankton, spatial weight specific distribution patterns are presented. This unpublished dataset contributes baseline information about the gelatinous macrozooplankton diversity and its specific distribution patterns in the extended North Sea area during winter (January-February) 2021. These data can be an important contribut

Global change has led to manifold changes of marine ecosystems and biodiversity world-wide. While it has been shown that certain jellyfish and comb jelly species have increased regionally, it remains to be investigated if this is a general trend or localized phenomenon. Especially for the economically important Baltic Sea, which is characterized by an extreme physical environmental gradient, this question has not been addressed to date. Here we present a detailed account of the gelatinous macro-zooplankton community including their physiological tolerance towards abiotic conditions and resulting distribution ranges in the Baltic. We show that the arrival and establishment of non-indigenous species has led to a rising importance of jellyfish and comb jellies in the Baltic. This accounts for the comb jelly Mnemiopsis leidyi, which was first observed in Northern Europe in 2005, as well as for the hydromedusae Blackfordia virginica, first sighted in 2014. Both species have been shown to attain high population densities with pronounced grazing impact in other invasive regions. Given the current and anticipated changes of the physical environment of the Baltic Sea, especially ongoing warming, amplification of their impact can be expected.

The associated microbiota of marine invertebrates plays an important role to the host in relation to fitness, health, and homeostasis. Cooperative and competitive interactions between bacteria, due to release of, for example, antibacterial substances and quorum sensing (QS)/quorum quenching (QQ) molecules, ultimately affect the establishment and dynamics of the associated microbial community. Aiming to address interspecies competition of cultivable microbes associated with emerging model species of the basal animal phyla Cnidaria (Aurelia aurita) and Ctenophora (Mnemiopsis leidyi), we performed a classical isolation approach. Overall, 84 bacteria were isolated from A. aurita medusae and polyps, 64 bacteria from M. leidyi, and 83 bacteria from ambient seawater, followed by taxonomically classification by 16S rRNA gene analysis. The results show that A. aurita and M. leidyi harbor a cultivable core microbiome consisting of typical marine ubiquitous bacteria also found in the ambient seawater. However, several bacteria were restricted to one host suggesting host-specific microbial community patterns. Interbacterial interactions were assessed by (a) a growth inhibition assay and (b) QS interference screening assay. Out of 231 isolates, 4 bacterial isolates inhibited growth of 17 isolates on agar plates. Moreover, 121 of the 231 isolates showed QS-interfering activities. They interfered with the acyl-homoserine lactone (AHL)-based communication, of which 21 showed simultaneous interference with autoinducer 2. Overall, this study provides insights into the cultivable part of the microbiota associated with two environmentally important marine non-model organisms and into interbacterial interactions, which are most likely considerably involved in shaping a healthy and resilient microbiota.