Oceanic Species Research Articles

Reallocation of the magnificent catshark Proscyllium magnificum Last & Vongpanich, 2004 to the genus Ctenacis Compagno, 1973 (Carcharhiniformes: Proscylliidae).

The magnificent catshark Proscyllium magnificum was described in 2004 based off five specimens collected in the Andaman Sea off Myanmar. It was originally allocated to the genus Proscyllium, but recent molecular analyses suggested it was more closely related to the harlequin catshark Ctenacis fehlmanni from the western Indian Ocean. This study incorporated meristics and external and internal morphology, together with molecular data to reclassify the magnificent catshark as Ctenacis magnificum and provides revised diagnoses for the genera Ctenacis and Proscyllium. Ctenacis consists of two allopatric Indian Ocean species, while Proscyllium is monotypic genus confined to the northwest Pacific. The revised Ctenacis can be distinguished from Proscyllium in having a broader and longer head (head length 21%-23% vs. 16%-18% of total length), distance between pectoral and pelvic bases shorter than head length (vs. greater than head length), more teeth (upper jaw with 80-86 vs. 46-62 tooth files), and a complex colour pattern of dark reddish-brown blotches and saddles (vs. colour pattern of small black spots). A revised key to the genera of proscylliids and species of Ctenacis is provided.

Mercury Stable Isotopes Reveal the Vertical Distribution and Trophic Ecology of Deep-Pelagic Organisms over the North-East Atlantic Ocean Continental Slope.

Deep-pelagic species are central to marine ecosystems and increasingly vulnerable to global change and human exploitation. To date, our understanding of these communities remains limited mainly due to the difficulty of observations, calling for complementary innovative tools to better characterize their ecology. We used mercury (Δ199Hg, δ202Hg, Δ201Hg, and Δ200Hg), carbon (δ13C), and nitrogen (δ15N) stable isotope compositions to segregate deep-pelagic species caught on the continental slope of the Bay of Biscay (NE Atlantic) according to their foraging depth and trophic ecology. Decreasing fish Δ199Hg values with corresponding depth estimates from the surface to down to 1,800 m confirmed that mercury isotopes are able to segregate deep species over a large vertical gradient according to their foraging depth. Results from isotopic compositions also identified different mercury sources, likely reflecting different trophic assemblages over the continental slope, in particular, the demersal influence for some species, compared to purely oceanic species. Overall, our results demonstrate how mercury stable isotopes can inform the vertical foraging habitat of little-known species and communities feeding in the deep.

A preliminary application and its error estimates of a simple stereo-camera measure system for the far-sea fishery of Pacific saury

Pacific saury is one of the most economically important species in the northwestern Pacific Ocean. The management of this resource relies on precise input of biological data such as body length and is often hindered by a lack of such data on captured fish. This study explores the potential of electronic monitoring (EM) using off-the-shelf stereo cameras to overcome the challenges of collecting and measuring saury body length from the Pacific saury fishery. Using a calibrated WEEVIEW SID WV3000 3D camera, a total of 252 paired images with different shooting angles and distances were obtained for further measurement using Sebastes Stereo Image Analysis Software (SSIAS). The treatments for the measurement distance (MD) were 30 cm, 60 cm, and 100 cm, for the depression angle (DA) were 30°, 60°, and 90°, and for the position angle (PA) were 0°, 45°, and 315° (− 45°). An assistant calibration (AC) in the form of a 16 cm black ruler was also added used. The SSIAS measurement results indicated that the best measurement was obtained with 0° position angle, 90° depression angle, and 30 cm distance from the target fish. The use of AC in the SSIAS + AC measurement was proven to reduce the measurement error from 2.45 to 8.64% to − 1.86 to 0.01%. This study set the baseline for the application of EM on collecting saury body length and the use of AC has been proven to increase the measurement accuracy.

Ocean Species Discoveries 1-12 - A primer for accelerating marine invertebrate taxonomy.

Discoveries of new species often depend on one or a few specimens, leading to delays as researchers wait for additional context, sometimes for decades. There is currently little professional incentive for a single expert to publish a stand-alone species description. Additionally, while many journals accept taxonomic descriptions, even specialist journals expect insights beyond the descriptive work itself. The combination of these factors exacerbates the issue that only a small fraction of marine species are known and new discoveries are described at a slow pace, while they face increasing threats from accelerating global change. To tackle this challenge, this first compilation of Ocean Species Discoveries (OSD) presents a new collaborative framework to accelerate the description and naming of marine invertebrate taxa that can be extended across all phyla. Through a mode of publication that can be speedy, taxonomy-focused and generate higher citation rates, OSD aims to create an attractive home for single species descriptions. This Senckenberg Ocean Species Alliance (SOSA) approach emphasises thorough, but compact species descriptions and diagnoses, with supporting illustrations and with molecular data when available. Even basic species descriptions carry key data for distributions and ecological interactions (e.g., host-parasite relationships) besides universally valid species names; these are essential for downstream uses, such as conservation assessments and communicating biodiversity to the broader public. This paper presents thirteen marine invertebrate taxa, comprising one new genus, eleven new species and one re-description and reinstatement, covering wide taxonomic, geographic, bathymetric and ecological ranges. The taxa addressed herein span three phyla (Mollusca, Arthropoda, Echinodermata), five classes, eight orders and twelve families. Apart from the new genus, an updated generic diagnosis is provided for four other genera. The newly-described species of the phylum Mollusca are Placiphorellamethanophila Vončina, sp. nov. (Polyplacophora, Mopaliidae), Lepetodrilusmarianae Chen, Watanabe & Tsuda, sp. nov. (Gastropoda, Lepetodrilidae), Shinkailepasgigas Chen, Watanabe & Tsuda, sp. nov. (Gastropoda, Phenacolepadidae) and Lyonsiellaillaesa Machado & Sigwart, sp. nov. (Bivalvia, Lyonsiellidae). The new taxa of the phylum Arthropoda are all members of the subphylum Crustacea: Lepechinellanaces Lörz & Engel, sp. nov. (Amphipoda, Lepechinellidae), Cuniculomaeragrata Tandberg & Jażdżewska, gen. et sp. nov. (Amphipoda, Maeridae), Pseudionellapumulaensis Williams & Landschoff, sp. nov. (Isopoda, Bopyridae), Mastigoniscusminimus Wenz, Knauber & Riehl, sp. nov. (Isopoda, Haploniscidae), Macrostylispapandreas Jonannsen, Riehl & Brandt, sp. nov. (Isopoda, Macrostylidae), Austroniscusindobathyasellus Kaiser, Kniesz & Kihara, sp. nov. (Isopoda, Nannoniscidae) and Apseudopsisdaria Esquete & Tato, sp. nov. (Tanaidacea, Apseudidae). In the phylum Echinodermata, the reinstated species is Psychropotesbuglossa E. Perrier, 1886 (Holothuroidea, Psychropotidae).The study areas span the North and Central Atlantic Ocean, the Indian Ocean and the North, East and West Pacific Ocean and depths from 5.2 m to 7081 m. Specimens of eleven free-living and one parasite species were collected from habitats ranging from an estuary to deep-sea trenches. The species were illustrated with photographs, line drawings, micro-computed tomography, confocal laser scanning microscopy and scanning electron microscopy images. Molecular data are included for nine species and four species include a molecular diagnosis in addition to their morphological diagnosis.The five new geographic and bathymetric distribution records comprise Lepechinellanaces Lörz & Engel, sp. nov., Cuniculomaeragrata Tandberg & Jażdżewska, sp. nov., Pseudionellapumulaensis Williams & Landschoff, sp. nov., Austroniscusindobathyasellus Kaiser, Kniesz & Kihara, sp. nov. and Psychropotesbuglossa E. Perrier, 1886, with the novelty spanning from the species to the family level. The new parasite record is Pseudionellapumulaensis Williams & Landschoff, sp. nov., found in association with the hermit crab Pagurusfraserorum Landschoff & Komai, 2018.

Ballast water transport of alien phytoplankton species to the brazilian amazon coast

The aim of this study was to investigate the presence of alien phytoplankton species transported through ballast water of ships that docked on the Amazon coast. Phytoplankton samples were collected from 25 ships between 2012 and 2014, revealing 215 identified species, mostly comprising oceanic planktonic marine species. However, several coastal and freshwater species not yet documented on the Maranhão coast were also observed. The identification of several coastal and freshwater species not yet recorded for Amazonian environments in the ballast water of the Ponta da Madeira Maritime Terminal (TMPM), as well as toxic microalgae genera such as the dinoflagellates Alexandrium and Gymnodinium and of some diatom species from the genus Pseudo-nitzchia, raises concerns regarding the possibility of introducing species. This indicates that ballast water can be responsible for the introduction of alien species in Amazonian aquatic environments, thereby highlighting the TMPM as a critical hotspot in the Amazonian region.

Otolith radiocarbon signatures provide distinct migration history of walleye pollock around Hokkaido, Japan in the North-Western Pacific.

Trace elements and stable isotope ratios in otoliths have been used as proxies for the migration history of teleosts; however, their application in oceanic fishes remains limited. This study reports the first use of radiocarbons in otoliths to evaluate the horizontal migration histories of an oceanic fish species, the walleye pollock Gadus chalcogrammus. We conducted radiocarbon analyses of three stocks sourced from Hokkaido, Japan. The radiocarbon concentrations from the outermost portion of the otoliths from the Japanese Pacific, Northern Japan Sea (JS), and Southern Okhotsk Sea (OS) stocks were in general agreement with the seawater radiocarbon concentration of the sampling region, suggesting that pollock of all three stocks generally inhabited the within the sea region where each pollocks were sampled throughout their life cycle. However, the radiocarbon signals also provided some indications that some JS and OS stocks may be migrating between different sea regions. The proposed novel approach of reconstructing the individual migration history of marine fish using radiocarbon in otoliths may help examine fish migration with a higher temporal and spatial resolution that could not be achieved by trace elements and stable isotope ratios.

First record of the deep-sea anglerfish Ceratias uranoscopus Murray, 1877 (Lophiiformes, Ceratiidae) from the Tropical Eastern Pacific Ocean

n 1973, a specimen of the deep‑sea anglerfish Ceratias uranoscopus Murray, 1877 was collected at a depth of 1000 m off the Pacific coast of Panama. This specimen represents (1) the first verified occurrence record of this species and genus in the Tropical Eastern Pacific Ocean and (2) an extension of about 7000 km east on this species’ previously known distribution. In this contribution, we provide and discuss unpublished and comparative morphometric, meristic, and occurrence data, corroborating the identity of the specimen and species and validating the new geographic range.

Investigation of the Link between Per- and Polyfluoroalkyl Substances and Stress Biomarkers in Bottlenose Dolphins (Tursiops truncatus).

Bottlenose dolphins (Tursiops truncatus) are keystone and sentinel species in the world's oceans. We studied correlations between per- and polyfluoroalkyl substances (PFAS) and their stress axis. We investigated associations between plasma biomarkers of 12 different PFAS variants and three cortisol pools (total, bound, and free) in wild T. truncatus from estuarine waters of Charleston, South Carolina (n = 115) and Indian River Lagoon, Florida (n = 178) from 2003 to 2006, 2010-2013, and 2015. All PFAS and total cortisol levels for these dolphins were previously reported; bound cortisol levels and free cortisol calculations have not been previously reported. We tested null hypotheses that levels of each PFAS were not correlated with those of each cortisol pool. Free cortisol levels were lower when PFOS, PFOA, and PFHxS biomarker levels were higher, but free cortisol levels were higher when PFTriA was higher. Bound cortisol levels were higher when there were higher PFDA, PFDoDA, PFDS, PFTeA, and PFUnDA biomarkers. Total cortisol was higher when PFOA was lower, but total cortisol was higher when PFDA, PFDoDA, PFTeA, and PFTriA were higher. Additional analyses indicated sex and age trends, as well as heterogeneity of effects from the covariates carbon chain length and PFAS class. Although this is a cross-sectional observational study and, therefore, could reflect cortisol impacts on PFAS toxicokinetics, these correlations are suggestive that PFAS impacts the stress axis in T. truncatus. However, if PFAS do impact the stress axis of dolphins, it is specific to the chemical structure, and could affect the individual pools of cortisol differently. It is critical to conduct long-term studies on these dolphins and to compare them to populations that have no or little expose to PFAS.

Combining mesocosms with models reveals effects of global warming and ocean acidification on a temperate marine ecosystem.

Ocean warming and species exploitation have already caused large-scale reorganization of biological communities across the world. Accurate projections of future biodiversity change require a comprehensive understanding of how entire communities respond to global change. We combined a time-dynamic integrated food web modeling approach (Ecosim) with previous data from community-level mesocosm experiments to determine the independent and combined effects of ocean warming, ocean acidification and fisheries exploitation on a well-managed temperate coastal ecosystem. The mesocosm parameters enabled important physiological and behavioral responses to climate stressors to be projected for trophic levels ranging from primary producers to top predators, including sharks. Through model simulations, we show that under sustainable rates of fisheries exploitation, near-future warming or ocean acidification in isolation could benefit species biomass at higher trophic levels (e.g., mammals, birds, and demersal finfish) in their current climate ranges, with the exception of small pelagic fishes. However, under warming and acidification combined, biomass increases at higher trophic levels will be lower or absent, while in the longer term reduced productivity of prey species is unlikely to support the increased biomass at the top of the food web. We also show that increases in exploitation will suppress any positive effects of human-driven climate change, causing individual species biomass to decrease at higher trophic levels. Nevertheless, total future potential biomass of some fisheries species in temperate areas might remain high, particularly under acidification, because unharvested opportunistic species will likely benefit from decreased competition and show an increase in biomass. Ecological indicators of species composition such as the Shannon diversity index decline under all climate change scenarios, suggesting a trade-off between biomass gain and functional diversity. By coupling parameters from multilevel mesocosm food web experiments with dynamic food web models, we were able to simulate the generative mechanisms that drive complex responses of temperate marine ecosystems to global change. This approach, which blends theory with experimental data, provides new prospects for forecasting climate-driven biodiversity change and its effects on ecosystem processes.

Southernmost record of Coryphaena hippurus Linnaeus, 1758 (Coryphaenidae) in the southwest Atlantic Ocean, off Mar del Plata, Argentina

This paper reports the presence of the common dolphinfish, Coryphaena hippurus, in Argentinian waters for the first time, with a specimen collected by experienced sport fishermen near Mar del Plata (38°S). This finding also represents the first record of a species from the Coryphaenidae family in Argentinian waters. The collection of these specimens coincided with sightings of other rarely-seen pelagic and oceanic fish species during the same period, providing further evidence supporting the influx of subtropical fish species from warmer Brazilian waters. These observations highlight the need for continued monitoring of fish populations and their distribution and the ecological dynamics within the region.

Correction: Dietary and niche analyses of four sympatric batoid species of the subtropical South Atlantic Ocean

Correction: Dietary and niche analyses of four sympatric batoid species of the subtropical South Atlantic Ocean

The climate-induced Spatio-temporal Distributions of tuna species in the tropical Atlantic Ocean

The climate-induced Spatio-temporal Distributions of tuna species in the tropical Atlantic Ocean

Dietary and niche analyses of four sympatric batoid species of the subtropical South Atlantic Ocean

Dietary and niche analyses of four sympatric batoid species of the subtropical South Atlantic Ocean

The dataset of the CLU lichen herbarium (Calabria, Italy).

Calabria, the southernmost tip of the Italian Peninsula, is a biogeographically very interesting region for lichenologists, characterised by the abundance of oceanic and suboceanic species with subtropical affinities, but also by the presence of the southernmost outposts of several boreal species on the highest peaks. The lichen biota of Calabria, which began to be intensively studied only from the 1980s, hosts more than 1000 infrageneric taxa. The lichen herbarium of the Botanical Garden of the University of Calabria (CLU) is the most relevant lichen collection from this region. It was established in 1985 and it presently includes 16926 specimens, most of which were collected in Calabria, although there are also several specimens from other parts of Italy and from abroad. This dataset contains 16926 records of lichens for a total of 1316 species. Of the 15219 georeferenced specimens, 10254 were collected in Calabria, while 4965 in other administrative regions of Italy. The dataset is available through GBIF, as well as in ITALIC, the Information System of Italian Lichens.

A Faunistic Revision of Longnose Skates of the Genus Dipturus (Rajiformes: Rajidae) from the Southern Southwestern Atlantic Ocean, Based on Morphological and Molecular Evidence

The identity of longnose skates (Dipturus-Zearaja-like skates) in the southern cone of the Americas has been a topic of extensive debate. This study employs a comprehensive analysis encompassing morphometrics, claspers, and the examination of COI and NADH2 sequence data to conclusively demonstrate the existence of only two longnose skate species in the southwestern Atlantic Ocean, extending south of 35 °S. Notably, Dipturus argentinensis Díaz de Astarloa, Mabragaña, Hanner and Figueroa, 2008 is revealed as a junior synonym of D. trachyderma (Krefft and Stehmann, 1975). Dipturus leptocaudus (Krefft and Stehmann, 1975) remains a northern valid species, but the specimen from the Falkland Islands (Malvinas) is recognized as a misidentification of D. trachyderma. Zearaja flavirostris (Philippi, 1892) and Dipturus lamillai Concha, Caira, Ebert and Pompert, 2019 are confirmed as junior synonyms of Zearaja brevicaudata (Marini, 1933). These findings contradict the previous report of six species within the same region over the last decade and underscore the presence of D. trachyderma and Z. brevicaudata south of 35 °S in the southwestern Atlantic. Additionally, this study notes the occurrence of only one specimen of Z. chilensis (Guichenot, 1848) in the Falkland Islands (Malvinas), suggesting an unusual frequency of this eastern Pacific skate in the southern Southwest Atlantic. Given that clasper morphology serves as the key distinguishing trait between Dipturus and Zearaja species, we provided a detailed analysis of the clasper characteristics of the Atlantic D. trachyderma, unequivocally situating it within Dipturus. The diagnostic characteristics include: (i) the presence of cartilage with the distal portion referred to as the sentinel, a feature absent in Zearaja; (ii) longer ventral terminal cartilage with the distal end referred to as the funnel, compared to Zearaja; and (iii) a non-spatulate distal lobe, a distinctive trait specific to Dipturus.

Species range shifts, biological invasions and ocean warming

Marine species are moving beyond their native geographical ranges in response to climate change and shifts in other abiotic conditions, establishing populations at higher latitudes. At the same time, species are introduced into new marine regions as a consequence of an increasingly connected world. The success of such range expansions, whether from shifts associated with climate change or human-mediated introductions, depends on the biotic interactions and environmental conditions at the recipient site and/or the capacity of species to adapt to these new conditions. Originating from a special session held at the Association for the Sciences of Limnology and Oceanography (ASLO) Aquatic Sciences Meeting 2021, the present Theme Section (TS) brings together the current research on 2 interrelated marine ecological topics: (1) invasive species in a warming ocean, and (2) climate-mediated species range shifts. Our investigation of the temporal trends in publications highlights that only one-tenth of studies on marine invasive species examine the species’ interaction with ocean warming. The topic of range shifting species receives slightly more attention than climate driven invasions, but both exhibited a comparable, similarly fast-growing publication rate. Overall, the papers in this TS provide evidence that global warming, and occasionally environmental degradation, are crucial factors altering the distribution of both marine invasive species and range-shifting species. Even though the published literature and even the contributions to this TS only infrequently focus on the interaction between these 2 topics, we propose that future collaboration between scientists from both fields would advance our understanding of the impact of global warming on the shifting distributions of native and invasive species and would allow the development of conservation and management plans in a warming ocean.

Targeting ocean conservation outcomes through threat reduction

Nations have committed to reductions in the global rate of species extinctions through the Sustainable Development Goals 14 and 15, for ocean and terrestrial species, respectively. Biodiversity loss is worsening despite rapid growth in the number and extent of protected areas, both at sea and on land. Resolving this requires targeting the locations and actions that will deliver positive conservation outcomes for biodiversity. The Species Threat Abatement and Restoration (STAR) metric, developed by a consortium of experts, quantifies the contributions that abating threats and restoring habitats in specific places offer towards reducing extinction risk based on the IUCN Red List of Threatened SpeciesTM. STAR is now recommended as an appropriate metric by recent disclosure frameworks for companies to report their impacts on nature and STAR has seen widespread uptake within the private sector. However, it is currently only available for the terrestrial realm. We extend the coverage of the threat abatement component of the STAR metric (START), used to identify locations where positive interventions could make a large contribution to reducing global species extinction risk and where developments that increase threats to species should be mitigated, to the marine realm for 1646 marine species. Reducing unsustainable fishing provides the greatest opportunity to lower species extinction risk, comprising 43% of the marine START score. Three-quarters (75%) of the global marine START score falls entirely outside the boundaries of protected areas and only 2.7% falls within no-take protected areas. The STAR metric can be used both to guide protected area expansion and to target other actions, such as establishment and enforcement of fishing limits, to recover biodiversity.

Modeling the importance of subsurface environmental variables in driving swordfish (Xiphias gladius) catchability in the Western Indian Ocean

AbstractMany oceanic species in pelagic habitats move vertically through the water column, highlighting the ecological importance of that spatial environment for modeling habitats of marine species. The role and importance of multiple oceanic subsurface environmental variables in modeling the habitat suitability of swordfish (Xiphias gladius), a highly migratory large pelagic fish, is poorly understood. In this study, we analyzed adult swordfish data from the 2017–2019 Chinese Indian Ocean tuna longline fishery observer. We used the maximum entropy model (MaxEnt) and random forest model (RF) to compare modeling schemes that included multiple subsurface environmental datasets. The area under receiver operating characteristic curve (AUC) from training and test sets was evaluated to investigate whether the inclusion of subsurface variables could enhance model performance and affect the simulation results. This analysis showed that model performance was significantly enhanced after addition of subsurface environmental variables, and the best model fit was achieved at 200–300 m depth. Sea water temperature, dissolved oxygen, net primary production, and ocean mixed layer depth were the critical environmental factors constituting catchability for swordfish in the Western Indian Ocean. As the depth increased, dissolved oxygen became the most important environmental factor, replacing surface temperature. Compared with the surface model, the location and extent of areas of high catchability in certain months changed significantly after the addition of subsurface variables. The results of this study provide evidence for a better understanding of the selection of critical environmental variables and improvement of model performance in 3D habitat modeling of pelagic fish.

Feeding behavior of yellowfin tuna around two insular regions of the western Atlantic Ocean.

The yellowfin tuna is a very abundant tropical tuna species in the western equatorial Atlantic Ocean and an important fishery resource for the Brazilian tuna fleet. In this study we performed stable isotope analysis to better understand the spatial trophodynamics and dietary changes in yellowfin tuna around two insular marine protected areas in Brazil. A total of 65 yellowfin tuna specimens measuring between 47 and 138 cm LT (total length) were sampled around the archipelagos of Fernando de Noronha (FNA; n = 34) and Saint Peter and Saint Paul (SPSPA; n = 31) between July 2018 and September 2019. Bayesian mixing models and generalized additive models were used to investigate the contributions of four different prey items (zooplankton, cephalopods, fish larvae, and flying fish) to yellowfin tuna diet in each area and their potential changes in relation to predator growth. The four prey items were found to have different overall contributions between the two studied areas, with zooplankton being the most important prey in FNA, whereas flying fish was the most relevant prey to the species' diet in SPSPA. Significant changes in the species diet by size were also found, with fish smaller than 90 cm (TL) having a more generalist diet and larger animals relying more on consuming larger and more nutritious prey (i.e., flying fish). Our results suggest that these two marine protected areas play an important role in ocean dynamics, providing important and different foraging grounds for the development of this predator species.

Reproductive biology, elemental composition and diel vertical migration of the cosmopolitan warm-temperate pelagic tunicate Soestia zonaria

Abstract Pelagic tunicates (appendicularians, doliolids, pyrosomes, salps) are cosmopolitan members of open-ocean food webs that serve as a link to the microbial loop and play a disproportional role in vertical carbon flux. Soestia zonaria is an oceanic salp species studied for more than a century, but little information exists on its ecology. Specimens of Soestia collected between 2008 and 2021 during four research expeditions (three to the Chatham Rise, New Zealand, and one to the Northeast Atlantic) using MOCNESS-1 net and large midwater trawl were analyzed for reproductive biology, stoichiometry and vertical distribution. Populations at the Chatham Rise sampled in early winter were more developed than in spring/early summer and dominated by small and medium-sized sexually reproducing blastozooids. Whole Soestia specimens had high organic content (mean ± SD = 32.8 ± 7.5%) and carbon-to-nitrogen values (6.8 ± 0.9) compared with other salp species, indicating a stronger dependency on carbon to meet its nutritional needs. Depth-stratified sampling showed that Soestia is primarily a low-amplitude diel vertical migrator occurring in the top 150 m of the water column, but also found at depths exceeding 500 m. Soestia is primarily an epipelagic salp species adapted to living in warm-temperate nutrient-depleted and more productive ocean regions.