New species and new records of bryozoan species from fouling communities in the Madeira Archipelago (NE Atlantic).

Hull fouling is considered to be the most significant vector of introduction of marine non-indigenous species (NIS) in the Madeira Archipelago (NE Atlantic) because these islands provide a vital passage route for many ships. The transfer of species between boat hulls and artificial substrates in marinas is known to be high. Bryozoans are among the most common groups of marine invertebrates growing on this type of substrate. In recent years, significant advances have been made in our knowledge about the biodiversity of bryozoans in the Madeira Archipelago. Nonetheless, the currently recognized numbers remain far from reflecting the actual bryozoan species richness. In this context, we examine bryozoan samples stemming from NIS monitoring surveys on artificial substrates along the southern coast of the Madeira Archipelago, in four recreational marinas and in two offshore aquaculture farms. This has yielded new information about ten bryozoan species. Two of them, Crisia noronhai sp. nov. and Amathia maderensis sp. nov., are described for the first time, although at least the first one was previously recorded from Madeira but misidentified. Bugula ingens, Cradoscrupocellaria insularis, Scruparia ambigua, and Celleporaria brunnea are recorded for the first time in Madeira. Moreover, the material of C. brunnea was compared with the type, and a biometric analysis was performed with material from the Atlantic and Mediterranean. All samples identified as C. brunnea in both regions are the same species, and the variations described in the literature apparently reflect high intracolonial variability. Finally, we provide new information for the descriptions of 4 additional bryozoans, namely, Crisia sp. aff. elongata, Cradoscrupocellaria bertholletii, Scrupocaberea maderensis, and Tricellaria inopinata.

Testing differences of marine non-indigenous species diversity across Macaronesia using a standardised approach.

The introduction of non-indigenous species (NIS) induces severe impacts on marine biodiversity and ecosystems. Macaronesia is an ecologically relevant region where several NIS were detected recently. For the first time, a standard experimental approach was designed to examine biofouling assemblages and investigate NIS across the region. In this context, sessile biofouling assemblages were examined in four recreational marinas in all the Macaronesian archipelagos from 2018 to 2020: the Azores, Madeira, Canary Islands, and Cabo Verde. We hypothesised that NIS numbers, abundance, and recruitment differed in each location due to abiotic and biotic features. From the Azores (higher latitudes) to Cabo Verde (lower latitudes), NIS recruitment and percentage cover decreased following a partial latitude gradient. The present study unveiled 25 NIS, with new records for the Azores (two cryptogenic species), Canary Islands (one NIS and two cryptogenic species), and Cabo Verde (three NIS and three cryptogenic species). The present research represents a pioneer and relevant step in advancing our current understanding of marine biological invasions in Macaronesia, employing a standard and low-cost approach.

Location and building material determine fouling assemblages within marinas: A case study in Madeira Island (NE Atlantic, Portugal).

Marinas are hubs for non-indigenous species (NIS) and constitute the nodes of a network of highly modified water bodies (HMWB) connected by recreational maritime traffic. Floating structures, such as pontoons, are often the surfaces with higher NIS abundance inside marinas and lead the risk for NIS introduction, establishment and spread. However, there is still little information on how the location within the marina and the substratum type can influence the recruitment of fouling assemblages depending on water parameters and substratum chemical composition. In this study, fouling recruitment was studied using an experimental approach with three materials (basalt, concrete and HDPE plastic) in two sites (close and far to the entrance) in two marinas of Madeira Island (NE Atlantic, Portugal). The structure of benthic assemblages after 6- and 12-months colonization, as well as biotic abundance, NIS abundance, richness, diversity, assemblages' volume, biomass and assemblages' morphology were explored. Differences between marinas were the main source of variation for both 6- and 12-month assemblages, with both marinas having different species composition and biomass. The inner and outer sites of both marinas varied in terms of structure and heterogeneity of assemblages and heterogeneity of morphological traits, but assemblages did not differ among substrata. However, basalt had a higher species richness and diversity while concrete showed a higher bioreceptivity in terms of total biotic coverage than the rest of materials. Overall, differences between and within marinas could be related to their structural morphology. This study can be valuable for management of urban ecosystems, towards an increase in the environmental and ecological status of existing marinas and their HMWB and mitigation coastal ecosystems degradation.

Anthropogenic pressure leads to more introductions: Marine traffic and artificial structures in offshore islands increases non-indigenous species.

Anthropogenic pressures such as the introduction of non-indigenous species (NIS) have impacted global biodiversity and ecosystems. Most marine species spreading outside their natural biogeographical limits are promoted and facilitated by maritime traffic through ballast water and hull biofouling. Propagule pressure plays a primary role in invasion success mixed with environmental conditions of the arrival port. Moreover, with the current ocean sprawl, new substrates are offered for potential NIS recruits. Here, differences in the fouling assemblages thriving inside three different ports/marinas facilities in Madeira Island were assessed for comparison. The locations showed significant differences concerning assemblage structure. Most NIS were detected in plastic floating pontoons. Funchal harbour receives most of the marine traffic in Madeira, acting as the main hub for primary NIS introductions, being recreational boating involved in NIS secondary transfers. Our results highlight the need for future management actions in island ecosystems, particularly monitoring and sampling of recreational boating.

Plasticrusts derive from maritime ropes scouring across raspy rocks.

Plasticrusts are a novel form of plastic debris which has only recently been discovered in Madeira Island, NE Atlantic Ocean. Plasticrusts consist of plastic encrusting wave-exposed rocky intertidal habitats and are presumably generated by waves smashing plastic debris against intertidal rocks. However, direct observations of this process are lacking and it is unknown which type of plastic debris the plasticrusts derive from. Therefore, we examined the Madeira rocky intertidal for signs of plasticrust formation and collected plasticrust and co-occurring plastic debris pieces of matching colors. We examined all collected materials using digital microscopy and Fourier-transform infrared spectroscopy. We found that plasticrusts can result from maritime ropes being scoured across raspy intertidal rocks and that the plasticrusts and the corresponding ropes consisted of polypropylene (PP) and high-density polyethylene (HDPE). Furthermore, we show that high temperatures contribute to plasticrust formation. Thereby, our study provides first insights into the complex plasticrust formation process.

A global-scale screening of non-native aquatic organisms to identify potentially invasive species under current and future climate conditions.

The threat posed by invasive non-native species worldwide requires a global approach to identify which introduced species are likely to pose an elevated risk of impact to native species and ecosystems. To inform policy, stakeholders and management decisions on global threats to aquatic ecosystems, 195 assessors representing 120 risk assessment areas across all six inhabited continents screened 819 non-native species from 15 groups of aquatic organisms (freshwater, brackish, marine plants and animals) using the Aquatic Species Invasiveness Screening Kit. This multi-lingual decision-support tool for the risk screening of aquatic organisms provides assessors with risk scores for a species under current and future climate change conditions that, following a statistically based calibration, permits the accurate classification of species into high-, medium- and low-risk categories under current and predicted climate conditions. The 1730 screenings undertaken encompassed wide geographical areas (regions, political entities, parts thereof, water bodies, river basins, lake drainage basins, and marine regions), which permitted thresholds to be identified for almost all aquatic organismal groups screened as well as for tropical, temperate and continental climate classes, and for tropical and temperate marine ecoregions. In total, 33 species were identified as posing a 'very high risk' of being or becoming invasive, and the scores of several of these species under current climate increased under future climate conditions, primarily due to their wide thermal tolerances. The risk thresholds determined for taxonomic groups and climate zones provide a basis against which area-specific or climate-based calibrated thresholds may be interpreted. In turn, the risk rankings help decision-makers identify which species require an immediate 'rapid' management action (e.g. eradication, control) to avoid or mitigate adverse impacts, which require a full risk assessment, and which are to be restricted or banned with regard to importation and/or sale as ornamental or aquarium/fishery enhancement.

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.

Removal of an established invader can change gross primary production of native macroalgae and alter carbon flow in intertidal rock pools.

The impact of invasive species on recipient communities can vary with environmental context and across levels of biological complexity. We investigated how an established invasive seaweed species affected the biomass, eco-physiology, carbon and nitrogen storage capacity of native seaweeds at sites with a different environmental setting due to a persistent upwelling in northern Spain. We removed the invasive Japanese wireweed Sargassum muticum from intertidal rock pools once every month during a one-year period and used an in-situ stable isotope pulse-chase labeling to estimate gross primary production (GPP), nitrogen uptake rate, 13C-carbon and 15N-nitrogen storage capacities. Following the addition of 13C-enriched bicarbonate and 15N-enriched nitrate to the seawater in the rock pools during the period of the low tide, we sampled macroalgal thalli at incoming tide to determine label uptake rate. After four days, we sampled macroalgal assemblages to determine both label storage capacity and biomass. After one year of removal there was no change in the macroalgal assemblage. However, both the GPP and 13C-carbon storage capacity were higher in the turf-forming Corallina spp. and, sometimes, in the canopy-forming Bifurcaria bifurcata. Nitrogen uptake rate followed similar, but more variable results. Although S. muticum inhibited carbon storage capacity of native species, the assemblage-level 13C-carbon storage was similar in the S. muticum-removed and control rock pools because the presence of the invasive species compensated for the functional loss of native species, particularly at sites where it was most abundant. No obvious effects were observed in relation to the environmental setting. Overall, the effect of the invasive S. muticum on carbon flow appeared to be mediated both by the effects on resource-use efficiency of native species and by its own biomass. Integrating physiological and assemblage-level responses can provide a broad understanding of how invasive species affect recipient communities and ecosystem functioning.

Plasticrusts: A new potential threat in the Anthropocene's rocky shores.

Plastic debris is one of the most extensive pollution problems our planet is facing today and a particular concern for marine environment conservation. The dimension of the problem is so large that it is possible our current era will generate an anthropogenic marker horizon of plastic in earth's sedimentary record. Here we present a new type of plastic pollution, the 'plasticrusts', plastic debris encrusting the rocky surface, recently discovered in the intertidal rocky shores of a volcanic Atlantic island. The potential impact that these new 'plasticrusts' may have needs to be further explored, as e.g. potential ingestion by intertidal organisms could suppose a new pathway for entrance of plastics into marine food webs. Consequently, its inclusion as a potential new marine debris category in management and monitoring actions should be pondered.

Metal pollution affects both native and non-indigenous biofouling recruitment in a subtropical island system.

Hull fouling has been a driving force behind the development of most modern marine antifouling coatings that mainly contain copper based biocides to inhibit growth of fouling organisms. Despite these efforts, several non-indigenous species continue to be transferred via hull-fouling worldwide. In this study we designed a disturbance gradient with three commercial antifouling paints applied to PVC settling plates with different concentrations of copper oxide and allowed colonization of fouling communities in four marinas located at the Madeira Archipelago (NE Atlantic). Overall, the antifouling treatments were effective in decreasing the diversity of fouling communities and spatial variability across marinas was observed. Increasing exposure to metal pollutants decreases both species cover and total diversity, independently of their native or NIS condition. However, evidences found suggest that long-term effects of copper based antifouling coatings can be modulated by metal-resistant species allowing a secondary substrate for the epibiosis of other species to establish.

Response of Two Mytilids to a Heatwave: The Complex Interplay of Physiology, Behaviour and Ecological Interactions.

Different combinations of behavioural and physiological responses may play a crucial role in the ecological success of species, notably in the context of biological invasions. The invasive mussel Xenostrobus securis has successfully colonised the inner part of the Galician Rias Baixas (NW Spain), where it co-occurs with the commercially-important mussel Mytilus galloprovincialis. This study investigated the effect of a heatwave on the physiological and behavioural responses in monospecific or mixed aggregations of these species. In a mesocosm experiment, mussels were exposed to simulated tidal cycles and similar temperature conditions to those experienced in the field during a heat-wave that occurred in the summer of 2013, when field robo-mussels registered temperatures up to 44.5°C at low tide. The overall responses to stress differed markedly between the two species. In monospecific aggregations M. galloprovincialis was more vulnerable than X. securis to heat exposure during emersion. However, in mixed aggregations, the presence of the invader was associated with lower mortality in M. galloprovincialis. The greater sensitivity of M. galloprovincialis to heat exposure was reflected in a higher mortality level, greater induction of Hsp70 protein and higher rates of respiration and gaping activity, which were accompanied by a lower heart rate (bradycardia). The findings show that the invader enhanced the physiological performance of M. galloprovincialis, highlighting the importance of species interactions in regulating responses to environmental stress. Understanding the complex interactions between ecological factors and physiological and behavioural responses of closely-related species is essential for predicting the impacts of invasions in the context of future climate change.

Shifts from native to non-indigenous mussels: enhanced habitat complexity and its effects on faunal assemblages.

Ecosystem engineers such as mussels may affect strongly both the structure of benthic assemblages and the ecosystem functioning. The black-pygmy mussel Limnoperna securis is an invasive species that is spreading along the Galician coast (NW Spain). Its current distribution overlaps with the distribution of the commercial native mussel species Mytilus galloprovincialis, but only in the inner part of two southern Galician rias. Here, we analysed the assemblages associated with clumps of the two mussel species and evaluated if the invasive species increased complexity of habitat. To measure complexity of clumps we used a new method modified from the "chain and tape" method. Results showed that the identity of the mussel influenced macrofaunal assemblages, but not meiofauna. L. securis increased the complexity of clumps, and such complexity explained a high percentage of variability of macrofauna. The shift in dominance from M. galloprovincialis to L. securis may alter habitat structure and complexity, affecting the macrofaunal assemblages with unpredictable consequences on trophic web relations.