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Genetic and morphological analyses were used to revise historical controversies in identities of Beroe species in the southern seas of Europe. DNA analyses with ITS sequences were used to explore their phylogenetic relationships. Three species of Beroe were initially listed as occurring in the Mediterranean Sea. Recently, the specimens were often re-identified using genetic analyses, causing confusions of identity. In 1997, a non-native Beroe was introduced into the Black Sea, which established there and spread to the adjacent seas, adding more controversy and confusion to species identification in this group of comb jellies. Our revisions suggest the following systematic changes: (1) re-naming of the species from what was initially labelled as Beroe ovata, and later as B. cucumis (sensuMayer, 1912), to Beroe pseudocucumis sp. nov.. (2) Beroe mitrata but not Beroe ‘anatoliensis’ inhabits the Aegean Sea, as was reported previously. (3) Beroe ovata is the only species of Beroidae that became invasive and should be used with the authority of Bruguière, 1789, but not in the sense of Mayer, 1912. (4) Beroe forskalii appears to be represented by two species in the Mediterranean and in the Antarctic, but further justification is needed. (5) Beroe cucumis is a species of bipolar occurrence, inhabiting both subpolar areas and its recent definition as Beroe ‘norvegica’ is incorrect.

Ctenophores are fragile gelatinous organisms whose diversity and distribution are relatively unknown. For the first time, the occurrence of four planktonic species, namely Ocyropsis crystallina, Eurhamphaea vexilligera, Cestum veneris, and Beroe sp., was reported from Madeira archipelago waters (NE subtropical Atlantic). This report represents the northernmost records in the Eastern Atlantic Ocean for O. crystallina and E. vexilligera.

Genus Beroe sp. (cf cucumis) M��ller, 1776 (Table 1, Fig. 4B) Examined material: South Madeira Island, 32��38���22������N 16��49���50������W. One specimen was observed at around 2 m depth during a recreational free dive. Description: Body cylindrical, flattened, opaque, without tentacles. Ctene rows with anastomosed canals. Remark: The resolution of the photographed specimen is low due to the recording quality (GoPro Hero 7). Only two Beroe species have been recorded in Macaronesia: Beroe cucumis (Azores and Canary Islands) and Beroe ovata (Canary Islands) (Gueroun et al. 2021). As B. ovata aboral part tends to be wider than in B. cucumis, the specimen recorded in Madeira is more likely a B. cucumis., Published as part of Gueroun, Sonia Km, Sch��fer, Susanne, Gizzi, Francesca, ��lvarez, Soledad, Monteiro, Jo��o Gama, Andrade, Carlos & Canning-Clode, Jo��o, 2021, Planktonic Ctenophora of the Madeira Archipelago (Northeastern Atlantic), pp. 433-443 in Zootaxa 5081 (3) on page 438, DOI: 10.11646/zootaxa.5081.3.7, http://zenodo.org/record/5775861, {"references":["Muller, O. F. (1776) Zoologiae Danicae Prodromus, seu Animalium Daniae et Norvegiae indigenarum characteres, nomina, et synonyma imprimis popularium. typis Hallageriis, Havniae [Copenhagen], xxxii + 274 pp. https: // doi. org / 10.5962 / bhl. title. 63795","Gueroun, S. K. M., Javidpour, J., Andrade, C., Nogueira, N., Freitas, M. & Canning-Clode, J. (2021) Pelagic Cnidaria and Ctenophora diversity patterns and trends in Macaronesia insular systems (NE Atlantic). Marine Biodiversity, 51, 32. https: // doi. org / 10.1007 / s 12526 - 021 - 01174 - z"]}

BEROE MITRATA (MOSER, 1907) Synonym: Pandora mitrata Moser, 1907. Adult individuals of Beroe mitrata are not large compared to other Beroe species. Their average length is about 60 mm (Wrobel & Mills, 1998). The body is mitre-shaped, flattened in cross-section. Its length to width ratio (l/w) is 1.33���1.78. Inside the large mouth, a macrocilia carpet is located, covering about 50% of the total area of the stomodaeum cavity. Macrocilia of B. mitrata are similar to that of B. forskaIii and have blunt, serrated tips, as viewed at the end of the effective stroke. The aboral end is oval or slightly tapered (Fig. 11A, B). The aboral pole is fringed with a row of short branched papillae (Fig. 11B). Like all Beroida, B. mitrata has eight meridional canals with eight comb rows, which are located beginning from the aboral organ and elongated maximally to the middle of the body. Comb rows might be of various lengths, extending from about half or two-thirds to threequarters of the body length. Meridional canals have many diverticulae that all bend toward the mouth. The numbers of branches of meridional canals increase with the growth of the individual (Moser, 1907; Komai, 1915). They are simple, showing only a few bifurcations and no anastomoses appear among them. Meridional canals are extremely flexible and can even turn inside-out as it was noted by Moser (1907) and Wrobel & Mills (1998). The overall colour is white or pale pink, with diffuse orange or red spots or pair of stripes on either side of the body (Fig. 11A, B). Geographical distribution: Beroe mitrata was first described from Japanese waters (Moser, 1907; Komai, 1915). Other findings in this area and elsewhere in the Indo-Pacific are somewhat doubtful (Moss, 2009; Minemizu et al., 2015; Lungren, 2020). In addition, Beroe mitrata was found in the southern Pacific, northwestern Pacific, off South Africa (Wrobel & Mills 1998; Mills & Haddock, 2007) and in the Gulf of Mexico (Moss, 2009). In the Atlantic Ocean, B. mitrata has been recorded from the western Atlantic (Report of RV Ronald H. Brown Cruise 06-03 to the western subtropical and tropical north Atlantic 10 April to 30 April, 2006). It was also recorded from the Mediterranean, in the Ligurian Sea (Madin, 1991; Tamm & Tamm, 1993). Mills (2005) has mentioned B. mitrata as an infrequent visitor to the Mediterranean. Observations of Beroe mitrata were performed in the southern Aegean Sea in Turkish Bays: G��kova, Marmaris, Bodrum and Fethiye Bays (Killi et al., 2019). In G��kova Bay, the invader M. leidyi and the native B. mitrata first arrived from the northern Aegean Sea with currents from the north (Killi & Tarkan, 2013). Now both ctenophores are observed regularly in G��kova Bay and in its several smaller bays and coves, where B. mitrata controls the M. leidyi population by preying on it (Killi & Tarkan, 2013). Habitat: Occurs in the warm surface waters, on the outer continental shelf and on the slopes (Wrobel & Mills, 1998; Moss, 2009)., Published as part of Shiganova, Tamara A. & Abyzova, Galina A., 2022, Revision of Beroidae (Ctenophora) in the southern seas of Europe: systematics and distribution based on genetics and morphology, pp. 297-322 in Zoological Journal of the Linnean Society 194 on pages 311-312, DOI: 10.1093/zoolinnean/zlab021, http://zenodo.org/record/5799206, {"references":["Moser F. 1907. Neues uber Ctenophoren. Mitteilung II. Zoologischer Anzeiger 32: 449 - 454.","Wrobel D, Mills C. 1998. Pacific coast pelagic invertebrates. A guide to the common gelatinous animals. Monterey: Sea Challengers & Monterey Bay Aquarium.","Komai T. 1915. On ctenophores of the neighbourhood of Misaki. Annotationes Zoologicae Japonenses / Nihon Dobutsugaku Iho 9: 451 - 474.","Moss AG. 2009. Ctenophora of the Gulf of Mexico. In: Felder DL, Camp DK, eds. Gulf of Mexico - origins, waters, and biota. Biodiversity. College Station, Texas: Texas A & M University Press, 403 - 411.","Minemizu R, Kubota S, Hirano Y, Lindsay D. 2015. A photographic guide to the jellyfishes of Japan. Tokyo: Heibonsha, 358 [in Japanese].","Lungren M. 2020. Comb jelly (Beroe mitrata) Balayan Bay, Luzon Island, Philippines. In: Nature picture library. Available at: https: // www. naturepl. com / stock-photo / comb-jelly- (beroemitrata) - balayan-bay-luzon-island-philippines-minimum / search / detail- 0 _ 01643972. html (accessed 18 March 2021).","Mills CE, Haddock SHD. 2007. Ctenophora. In: Carlton JT, ed. The Light and Smith manual: intertidal invertebrates from central California to Oregon. Berkeley and Los Angeles: University of California Press, 189 - 199.","Madin LP. 1991. Distribution and taxonomy of zooplankton in the Alboran Sea and adjacent western Mediterranean: a literature survey and field guide (No. WHOI- 91 - 26). Woods Hole: Woods Hole Oceanographic Institution.","Tamm SL, Tamm S. 1993. Diversity of macrociliary size, tooth patterns, and distribution in Beroe (Ctenophora). Zoomorphology 113: 79 - 89.","Mills CCE. 2005. Ctenophora. In Rellini La Checklist della fauna Italiana (Rellini et al. ed.), Fasc. 3. Available at: http: // www. sibm. it / CHECKLIST / menusxcheck. htm. Published by the author, web page established March 1998, last updated January 2017.","Killi N, Abyzova G, Shiganova T. 2019. Occurrence of Beroe mitrata (Beroidae) associated with its prey aggressive invader ctenophore Mnemiopsis leidyi A. Agassiz from in Gokova Bay. Istanbul: BioEco 2019 - International Biodiversity & Ecology Sciences Symposium.","Killi, N, Tarkan AN. 2013. Seasonal distribution of Scyphozoa (Cnidaria) and Ctenophora species in Gokova Bay, Mugla, Turkey. Rapports et Proces-verbaux des Reunions Commission Internationale pour l'Exploration Scientifique de la Mer Mediterranee 40: 538."]}

BEROE CUCUMIS FABRICIUS, 1780 The body shape is strait or slightly oval in the oral part and oval at the aboral side, and it is more flattened in the paragastral plane than B. pseudocucumis (Fig. 6C, D). The adult length varies from 50 to 150 mm. Its length to width ratio (l/w) ranges 1.6���2.2. This ratio is variable: we observed specimens, which were shorter and longer, wider or slenderer, but never less than for B. pseudocucumis. Juvenile individuals may be narrower in the oral part of the body. Similar individuals were illustrated by Mayer (1912). The meridional canals lie under eight rows of ciliary combs, which extend about three-quarters of the distance from the aboral pole towards the mouth or a bit longer, but not up to the mouth. Its meridional canals have numerous diverticulae, which may branch out in adult ctenophores, but do not anastomose with each other, and do not connect with paragastral canals. At the aboral end, two oval polar-plates (Fig. 6A) surround the sense organ at the oval aboral pole, and are fringed with a row of short, branched papillae. Macrociliaries have three-toothed macrociliary tips with a somewhat larger middle tooth (Tamm & Tamm, 1993). Geographical distribution: Beroe cucumis was previously believed to be a widespread species, continuously distributed from the Arctic to the Antarctic (Pages & Orejas, 1999), but according to our genetic and morphological studies, and by comparison of published data, this species has a bipolar distribution, inhabiting cold polar and temperate waters, while being absent from tropical and subtropical zones. Distribution in the Arctic: all Eurasian seas (Sirenko, 2001), including the Barents Sea (Manko et al., 2015; Bandara et al., 2016; this study), the White Sea (Kosobokova & Pertsova, 2018), the Kara Sea (Dvoretsky & Dvoretsky, 2017), the Laptev Sea (Abramova & Tuschling, 2005), the East-Siberian Sea and the Chukchi Sea (Ershova et al., 2015); Canada Basin (Raskoff et al., 2005; Purcell et al., 2010). Distribution in the Atlantic Ocean: individuals of Beroe cucumis were sampled from the North of Norway to the south-east of the North Sea and analysed genetically (this research). In the Atlantic Ocean Beroe cucumis individuals were sampled from the northern part of Norway to the south-east of the North Sea and analysed genetically (this research). Also, B. cucumis is known to occur in the northern and north-eastern Atlantic (Fabricius, 1780; Van Soest, 1973; Granhag et al., 2012; Licandro et al., 2015; Knutsen et al., 2018), and along the eastern coasts of Canada and the USA (Mayer, 1912; Harbison et al., 1978). In the Mediterranean Sea, and the subtropical, tropical and equatorial regions of the Atlantic Ocean, it is replaced by B. pseudocucumis. Additional genetic studies are required to clarify the boundaries of the B. cucumis occurrence off the coast of South America, but presumably B. cucumis occurs also in the temperate and subpolar waters of the South Atlantic and Antarctic (Siegel & Harm, 1996; Mianzan 1999; Mianzan & Guererro, 2000; Pakhomov et al., 2000; Flores et al., 2010, 2011). There is evidence of B. cucumis occurring in the Benguela Current near South Africa (Gibbons et al., 1992). In the Pacific Ocean, Beroe cucumis occurs in cold waters: north-west Pacific (Kasuya et al., 2000; Napazakov & Chuchukalo 2011); north-east Pacific, eastern Bering Sea (Hoff et al., 2011); and the southeast Pacific off Chile (Pages & Orejas, 1999; Oliveira et al., 2016). It frequently occurs in southern Australia (Stiasny, 1931; Edgar, 1997; Gershwin et al., 2010) and in New Zealand waters (Ralph, 1950). Now that we have genetic confirmation of the presence of two species with a similar morphotype (Beroe cucumis and B. pseudocucumis), it is necessary to conduct a detailed analysis of the ecology and distribution of these two species. It is possible that there are areas of overlap in their habitat and geography. According to previous records, B. cucumis occurs in the areas between temperate and subtropical climatic zones, like the east coast of the USA (Mayer, 1912; Harbison et al., 1978), the west coast of the USA (Wrobel & Mills, 1998) and the Yellow and East China seas (Liu, 2013; Yin et al., 2017; Wang & Cheng, 2019). However, additional ecological and genetic studies of individuals in these areas are required to clarify the species identification and distribution. Habitat: Cold-water boreal species, has bipolar distribution and inhabits cold polar and temperate waters, while absent in tropical and subtropical zones., Published as part of Shiganova, Tamara A. & Abyzova, Galina A., 2022, Revision of Beroidae (Ctenophora) in the southern seas of Europe: systematics and distribution based on genetics and morphology, pp. 297-322 in Zoological Journal of the Linnean Society 194 on page 308, DOI: 10.1093/zoolinnean/zlab021, http://zenodo.org/record/5799206, {"references":["Fabricius, O. 1780. Fauna Groenlandica. Copenhagen and Leipzig: J. G. Rothe, 450.","Mayer AG. 1912. Ctenophores of the Atlantic coast of North America. Carnegie Institution of Washington Publication 162: 1 - 58.","Tamm SL, Tamm S. 1993. Diversity of macrociliary size, tooth patterns, and distribution in Beroe (Ctenophora). Zoomorphology 113: 79 - 89.","Pages F, Orejas C. 1999. Medusae, siphonophores and ctenophores of the Magellan region. Scientia Marina 63: 51 - 57.","Sirenko BI. 2001. List of species of free-living invertebrates of Eurasian Arctic seas and adjacent deep waters. Moscow: Russian Academy of Science, Zoological Institute.","Manko MK, Panasiuk-Chodnicka AA, Zmijewska MI. 2015. Pelagic coelenterates in the Atlantic sector of the Arctic Ocean-species diversity and distribution as water mass indicators. Oceanological and Hydrobiological Studies 44: 466 - 479.","Bandara K, Varpe O, Soreide JE, Wallenschus J, Berge J, Eiane K. 2016. Seasonal vertical strategies in a high-Arctic coastal zooplankton community. Marine Ecology Progress Series 555: 49 - 64.","Kosobokova KN, Pertsova NM. 2018. Zooplankton of the White Sea: communities, structure, seasonal dynamics, spatial distribution, and ecology. In: Lisitzin AP, Gordeev V, eds. Biogeochemistry of the atmosphere, ice and water of the White Sea. Cham: Springer, 223 - 266.","Dvoretsky VG, Dvoretsky AG. 2017. Macrozooplankton of the Arctic - the Kara Sea in relation to environmental conditions. Estuarine, Coastal and Shelf Science 188: 38 - 55.","Abramova E, Tuschling K. 2005. A 12 - year study of the seasonal and interannual dynamics of mesozooplankton in the Laptev Sea: significance of salinity regime and life cycle patterns. Global and Planetary Change 48: 141 - 164.","Ershova EA, Hopcroft RR, Kosobokova KN. 2015. Interannual variability of summer mesozooplankton communities of the western Chukchi Sea: 2004 - 2012. Polar Biology 38: 1461 - 1481.","Raskoff KA, Purcell JE, Hopcroft RR. 2005. Gelatinous zooplankton of the Arctic Ocean: in situ observations under the ice. Polar Biology 28: 207 - 217.","Purcell JE, Hopcroft, RR, Kosobokova, KN, Whitledge TE. 2010. Distribution, abundance, and predation effects of epipelagic ctenophores and jellyfish in the western Arctic Ocean. Deep Sea Research Part II: Topical Studies in Oceanography 57: 127 - 135.","Van Soest RWM. 1973. Planktonic coelenterates collected in the North Atlantic Ocean. Bijdragen tot de Dierkunde 43: 119 - 126.","Granhag L, Majaneva S, Friis Moller L. 2012. First recordings of the ctenophore Euplokamis sp. (Ctenophora, Cydippida) in Swedish coastal waters and molecular identification of this genus. Aquatic Invasions 7: 455 - 463.","Licandro P, Blackett M, Fischer A, Hosia A, Kennedy J, Kirby RR, Raab K, Stern R, Tranter P. 2015. Biogeography of jellyfish in the North Atlantic, by traditional and genomic methods. Earth System Science Data 7: 173 - 191.","Knutsen T, Hosia A, Falkenhaug T, Skern-Mauritzen R, Wiebe PH, Larsen RB, Aglen A, Berg E. 2018. Coincident mass occurrence of gelatinous zooplankton in northern Norway. Frontiers in Marine Science 5: 158.","Harbison GR, Madin LP, Swanberg NR. 1978. On the natural history and distribution of oceanic ctenophores. Deep Sea Research 25: 233 - 256.","Siegel V, Harm U. 1996. The composition, abundance, biomass and diversity of the epipelagic zooplankton communities of the southern Bellingshausen Sea (Antarctic) with special references to krill and salps. Archive of Fishery and Marine Research 44: 115 - 139.","Mianzan HW. 1999. Ctenophora. In: Boltovskoy D, ed. South Atlantic zooplankton. Leiden: Backhuys Publishers, 561 - 573.","Mianzan HW, Guererro R. 2000. Environmental patterns and biomass distribution of gelatinous macrozooplankton. Three study cases in the southwestern Atlantic Ocean. Scientia Marina 64: 215 - 224.","Pakhomov EA, Ansorge IJ, Froneman PW. 2000. Variability in the inter-island environment of the Prince Edward Islands (Southern Ocean). Polar Biology 23: 593 - 603.","Flores H, Van Franeker JA, Feij B, Meijboom A, Van Dorssen M. 2010. Macrozooplankton and micronekton in the surface layer and under sea ice. In: Bathmann U, ed. The expedition of the research vessel ' Polarstern' to the Antarctic in 2007 / 2008 (ANT-XXIV / 2). Berichte zur Polar-und Meeresforschung (Reports on Polar and Marine Research). Bremerhaven, Germany: Alfred Wegener Institute, Helmholtz Centre for Polar and Marine Research (AWI), 604.","Flores H, Van Franeker JA, Cisewski B, Leach H, Van de Putte AP, Meesters EH, Bathmann U, Wolff WJ. 2011. Macrofauna under sea ice and in the open surface layer of the Lazarev Sea, Southern Ocean. Deep Sea Research Part II: Topical Studies in Oceanography 58: 1948 - 1961.","Gibbons MJ, Stuart V, Verheye HM. 1992. Tropic ecology of carnivorous zooplankton in the Benguela. South African Journal of Marine Science 12: 421 - 437.","Kasuya T, Ishimaru T, Murano M. 2000. Seasonal variations in abundance and size composition of the lobate ctenophore Bolinopsis mikado (Moser) in Tokyo Bay, central Japan. Journal of Oceanography 56: 419 - 427.","Napazakov VV, Chuchukalo VI. 2011. Feeding of Albatrossia pectoralis (Macrouridae) on the continental slope of eastern Kamchatka and the Kurils. Journal of Ichthyology 51: 343 - 351.","Hoff GR, Britt LL. 2011. Results of the 2010 eastern Bering Sea upper continental slope survey of groundfish and invertebrate resources. US Department of Commerce, NOAA technical memorandum NMFS-AFSC- 224, 300.","Oliveira OM, Miranda TP, Araujo EM, Ayon P, Cedeno- Posso CM, Cepeda-Mercado AA, Cordova P, Cunha AF, Genzano GN, Haddad MA, Mianzan HW. 2016. Census of Cnidaria (Medusozoa) and Ctenophora from South American marine waters. Zootaxa 4194: 1 - 256.","Stiasny G. 1931. Uber einige coelenterata von Australien. Zoologische Mededelingen 14: 27 - 42.","Edgar GJ. 1997. A new genus and three new species of Apseudomorph tanaidacean (Crustacea) from the Darwin region, 279 - 299, In: Hanly JR, Caswell G, Megirian D, Larson HK, eds. Proceedings of the Sixth International Marine Biological Workshop. The Marine Flora and Fauna of Darwin Harbour, Northern Territory, Australia. Darwin, Australia: Museums and Art Galleries of the Northern Territory and the Australian Marine Science Association.","Gershwin, LA, Zeidler W, Davie PJ. 2010. Ctenophora of Australia. Memoirs of the Queensland Museum 54: 1 - 45.","Ralph PM. 1950. Ctenophores from the waters of Cook Strait and Wellington Harbour. Transactions of the Royal Society of New Zealand 78: 70 - 82.","Wrobel D, Mills C. 1998. Pacific coast pelagic invertebrates. A guide to the common gelatinous animals. Monterey: Sea Challengers & Monterey Bay Aquarium.","Liu JY. 2013. Status of marine biodiversity of the China Seas. PLoS One 8: e 50719.","Yin J, Zhang G, Li C, Wang S, Zhao Z, Wan A. 2017. Community composition, abundance and biomass of zooplankton in Zhangzi Island waters, northern Yellow Sea. Chinese Journal of Oceanology and Limnology 35: 1144 - 1151.","Wang M, Cheng F. 2019. The complete mitochondrial genome of the Ctenophore Beroe cucumis, a mitochondrial genome showing rapid evolutionary rates. Mitochondrial DNA Part B 4: 3774 - 3775."]}

BEROE FORSKALII MILNE EDWARDS, 1841 The body is strongly flattened. The aboral part is sharply conical, whereas the oral part is oval with a rounded, voluminous mouth, expanded on the entire oral part. The length is up to 150 mm (Fig. 9A���C). The length/width ratio is 1.75���2.00. The eight meridional canals lie under eight rows of dense ciliary combs plates. The elongate plates, covering them from the surface, extend up to three-quarters of the body length from the aboral pole towards the mouth. Diverticulae are connected to each other, forming dense anastomoses of fine networks. The colour is pink with a dark tinge along the ciliary comb rows, which is specific for B. forskalii (Mills & Haddock, 2007; Shiganova & Malej, 2009). There is an axial funnel-tube. The aboral pole is fringed with a row of long, branched papillae (Fig. 9D). The macrocilia are arranged in long, tapering stripes running from the lip edge to a considerable distance into the stomodaeal cavity. Each stomodaeal wall bears about 20 macrociliary stripes, which increase in length towards the middle of the stomodaeum (Tamm & Tamm, 1993). Geographical distribution: We presume the existence of two different species, listed under Beroe forskalii following the presumption of Johansson et al. (2018) and our morphological analyses. While there is no sufficient data on the genetic structure of individuals around the world, we only present generalized data on the species distribution. Beroe forskalii is widely distributed in the Atlantic: the Mediterranean Sea (Madin, 1991; Tamm & Tamm, 1993), off the Brazilian coast (Oliveira & Migotto, 2014), off the USA east coast (Podar et al., 2001); the Pacific: off the California coast (Wrobel & Mills, 1998; Podar et al., 2001; Stout et al., 2002; Schultz et al., 2020); the eastern Pacific off Mexico and near Guadalupe Island (Ruiz-Escobar et al., 2015); off Peru (Oliveira et al., 2016); off Japan (Kubota, 1997; Fujikura et al., 2012) and off Australia (Gershwin et al., 2014). It was recently discovered in the southern waters of the Atlantic (Oliveira & Migotto, 2014). A single specimen was collected in the S��o Sebasti��o Channel, in south-eastern Brazil (23.8��S, 45.4��W). The authors conducted a comparative morphological analysis and concluded that the individual belongs to the same species that lives in the Mediterranean Sea. Definitely belonging to this species, a recent specimen was recorded from the coastal waters of Pakistan (Gul & Jahangir, 2019; Shiganova identification from illustration). An individual of Beroe forskalii was found on the coast of Japan (Lindsay, 2006), but there is insufficient data on the morphology of this specimen, which made it difficult to compare it with the Australian or Mediterranean samples. All these records demonstrate that B. forskalii is a warm-water species (Chun, 1880; Mianzan, 1999). Individuals identified as B. forskalii, but probably belonging to another species, have been reported from cold Antarctic waters (Flores et al., 2010, 2011; Siegel & Harm, 1996). Habitat: Warm waters., Published as part of Shiganova, Tamara A. & Abyzova, Galina A., 2022, Revision of Beroidae (Ctenophora) in the southern seas of Europe: systematics and distribution based on genetics and morphology, pp. 297-322 in Zoological Journal of the Linnean Society 194 on pages 309-311, DOI: 10.1093/zoolinnean/zlab021, http://zenodo.org/record/5799206, {"references":["Chun C. 1880. Die Ctenophoren des Golfes von Neapel. Fauna und Flora des Golfes von Neapel vol. 1. Berlin: R. Friedlander & Sohn, 313.","Mills CE, Haddock SHD. 2007. Ctenophora. In: Carlton JT, ed. The Light and Smith manual: intertidal invertebrates from central California to Oregon. Berkeley and Los Angeles: University of California Press, 189 - 199.","Shiganova TA, Malej A. 2009. Native and non-native ctenophores in the Gulf of Trieste, Northern Adriatic Sea. Journal of Plankton Research 31: 61 - 71.","Tamm SL, Tamm S. 1993. Diversity of macrociliary size, tooth patterns, and distribution in Beroe (Ctenophora). Zoomorphology 113: 79 - 89.","Johansson ML, Shiganova TA, Ringvold H, Stupnikova AN, Heath DD, MacIsaac HJ. 2018. Molecular insights into the ctenophore genus Beroe in Europe: new species, spreading invaders. Journal of Heredity 109: 520 - 529.","Madin LP. 1991. Distribution and taxonomy of zooplankton in the Alboran Sea and adjacent western Mediterranean: a literature survey and field guide (No. WHOI- 91 - 26). Woods Hole: Woods Hole Oceanographic Institution.","Oliveira OM, Migotto AE. 2014. First occurrence of Beroe forskalii (Ctenophora) in South American Atlantic coastal waters, with notes on the use of macrociliary patterns for beroid identification. Zootaxa 3779: 470 - 476.","Podar M, Haddock SH, Sogin ML, Harbison GR. 2001. A molecular phylogenetic framework for the phylum Ctenophora using 18 S rRNA genes. Molecular Phylogenetics and Evolution 21: 218 - 230.","Wrobel D, Mills C. 1998. Pacific coast pelagic invertebrates. A guide to the common gelatinous animals. Monterey: Sea Challengers & Monterey Bay Aquarium.","Stout NJ, Kuhnz L, Lundsten L, Schlining K, von Thun S. 2002. Video Annotation and Reference System (VARS) database, Year 2000, Monterey Bay Aquarium Research Institute, Moss Landing, California USA, Database. Available at: www. mbari. org / vars.","Schultz DT, Eizenga JM, Corbett-Detig RB, Francis WR, Christianson LM, Haddock, SH. 2020. Conserved novel ORFs in the mitochondrial genome of the ctenophore Beroe forskalii. PeerJ 8: e 8356.","Ruiz-Escobar, F, Valadez-Vargas DK, Oliveira OM. 2015. Ctenophores from the Oaxaca coast, including a checklist of species from the Pacific coast of Mexico. Zootaxa 3936: 435 - 445.","Oliveira OM, Miranda TP, Araujo EM, Ayon P, Cedeno- Posso CM, Cepeda-Mercado AA, Cordova P, Cunha AF, Genzano GN, Haddad MA, Mianzan HW. 2016. Census of Cnidaria (Medusozoa) and Ctenophora from South American marine waters. Zootaxa 4194: 1 - 256.","Kubota S. 1997. Phylum Ctenophora. In: Chihara M, Murano M, eds. An illustrated guide to marine plankton in Japan. Tokyo: Tokai University Press, 555 - 566 [in Japanese]","Fujikura K, Okutani T, Maruyama T. 2012. Deep-sea life - biological observations using research submersibles: second edition. Kanagawa: Tokai University Press, 487 [in Japanese].","Gershwin L, Lewis M, Gowlett-Holmes K, Kloser R. 2014. The Ctenophores. In: Pelagic invertebrates of south-eastern Australia: a field reference guide, v. 1.1. Hobart: CSIRO Marine and Atmospheric Research.","Gul S, Jahangir S. 2019. First occurrence of Pleurobrachia pileus (O. F. Muller, 1776) and sighting records of four other species of comb jellies (Ctenophora) from the coast of Pakistan (northern Arabian Sea): a brief account. International Journal of Biology and Biotechnology 16: 991 - 993.","Lindsay DJ. 2006. A checklist of midwater cnidarians and ctenophores from Sagami Bay - species sampled during submersible surveys from 1993 - 2004. Bulletin of the Plankton Society of Japan 53: 104 - 110.","Mianzan HW. 1999. Ctenophora. In: Boltovskoy D, ed. South Atlantic zooplankton. Leiden: Backhuys Publishers, 561 - 573.","Flores H, Van Franeker JA, Feij B, Meijboom A, Van Dorssen M. 2010. Macrozooplankton and micronekton in the surface layer and under sea ice. In: Bathmann U, ed. The expedition of the research vessel ' Polarstern' to the Antarctic in 2007 / 2008 (ANT-XXIV / 2). Berichte zur Polar-und Meeresforschung (Reports on Polar and Marine Research). Bremerhaven, Germany: Alfred Wegener Institute, Helmholtz Centre for Polar and Marine Research (AWI), 604.","Flores H, Van Franeker JA, Cisewski B, Leach H, Van de Putte AP, Meesters EH, Bathmann U, Wolff WJ. 2011. Macrofauna under sea ice and in the open surface layer of the Lazarev Sea, Southern Ocean. Deep Sea Research Part II: Topical Studies in Oceanography 58: 1948 - 1961.","Siegel V, Harm U. 1996. The composition, abundance, biomass and diversity of the epipelagic zooplankton communities of the southern Bellingshausen Sea (Antarctic) with special references to krill and salps. Archive of Fishery and Marine Research 44: 115 - 139.","Komai T. 1915. On ctenophores of the neighbourhood of Misaki. Annotationes Zoologicae Japonenses / Nihon Dobutsugaku Iho 9: 451 - 474."]}

GENUS: BEROE BROWNE, 1756 Representatives of Beroida and Cydipida are presumed to be the most primitive species among the Ctenophora because of the absence of a transmission stage, like true larva, which exist in the Orders Lobata, Cestida and Platyctenida (Aggasiz, 1865; Greve et al., 1976; Harbison 1985; Podar et al. 2001). The phylogeny of ctenophores is still controversial and Simion et al. (2015) have suggested that Beroe is perhaps not monophyletic. All representatives of Beroe have an oval or conical body shape beginning from the aboral part and a wide-flaring mouth with a large stomodeum. They have eight meridional and two paragastral canals that originate from the funnel and extend down to the middle of the broad sides of the body. Inside the mouth of the Beroida, macrocilia are located. They are finger-shaped, compound, ciliary organelles (Horridge, 1965; Harbison & Madin, 1982; Tamm, 1982). A single macrocilium contains hundreds or thousands (depending on the species) of hexagonally packed 9 + 2 axonemes, organized into many parallel rows (Tamm & Tamm, 1993). Most Beroe species are pink, especially along the meridional canals and comb-rows. All Beroe species are highly luminescent and flashing with blue-green light along the comb rows. Most of them are simultaneous hermaphrodites, capable of self-fertilization (Mianzan, 1999)., Published as part of Shiganova, Tamara A. & Abyzova, Galina A., 2022, Revision of Beroidae (Ctenophora) in the southern seas of Europe: systematics and distribution based on genetics and morphology, pp. 297-322 in Zoological Journal of the Linnean Society 194 on page 303, DOI: 10.1093/zoolinnean/zlab021, http://zenodo.org/record/5799206, {"references":["Browne P. 1756. The civil and natural history of Jamaica. London: printed for the author and sold by L. T. Osborne and J. Shipton, The author, 490.","Greve W, Stockner J, Fulton J. 1976. Towards a theory of speciation in Beroe. In: Mackie GO, ed. Coelenterate ecology and behavior. New York: Springer, 251 - 258.","Harbison GR. 1985. On the classification and evolution of the Ctenophora. In: Conway Morris S, ed. Origins and relationships of lower invertebrates. Oxford: Oxford University Press, 78 - 100.","Podar M, Haddock SH, Sogin ML, Harbison GR. 2001. A molecular phylogenetic framework for the phylum Ctenophora using 18 S rRNA genes. Molecular Phylogenetics and Evolution 21: 218 - 230.","Simion P, Bekkouche N, Jager M, Queinnec E, Manuel M. 2015. Exploring the potential of small RNA subunit and ITS sequences for resolving phylogenetic relationships within the phylum Ctenophora. Zoology 118: 102 - 114.","Horridge GA. 1965. Macrocilia with numerous shafts from the lips of the ctenophore Beroe. Proceedings of the Royal Society of London. Series B. Biological Sciences 162: 351 - 364.","Harbison GR, Madin LP. 1982. Ctenophora. In: Parker SP, ed. Synopsis and classification of living organisms, 1. New York: McGraw-Hill, 707 - 715.","Tamm SL. 1982. Ctenophores. In: Shelton GG, ed. Electrical conduction and behaviour in simple invertebrates. Oxford: Oxford University Press, 266 - 358.","Tamm SL, Tamm S. 1993. Diversity of macrociliary size, tooth patterns, and distribution in Beroe (Ctenophora). Zoomorphology 113: 79 - 89.","Mianzan HW. 1999. Ctenophora. In: Boltovskoy D, ed. South Atlantic zooplankton. Leiden: Backhuys Publishers, 561 - 573."]}

BEROE PSEUDOCUCUMIS SP. NOV. Zoobank registration: urn:lsid:zoobank. org:act: 87D103FA-F278-4D2F-B206- 0AAF96632F0C Synonyms: Beroe ovata of Eschscholtz, 1829); Beroe cucumis sensu Mayer (of Bayha et al., 2004) Etymology: From Latin cucumis, cucumber, and the Greek prefix ��ευ����-, false, because the name B. cucumis had been misapplied to this species in the past. The body is long-oval, from slightly flattened to nearly circular in cross-section, l/w: 2.0���3.0, with length up to three times longer than the width, which is specific for this species. The aboral end and oral part is rounded, which is specific for this species (Fig. 5). Adult length varies from 50 to 160 mm. The eight meridional canals lie under eight rows of ciliary combs, which extend for three-quarters of the length from aboral pole towards the mouth or, sometimes, further (Fig. 5A, B). Lateral branches of the meridional canals (diverticulae) are dense (which is unique in this species) and do not anastomose with each other (Fig. 5C). Stomodaeum is voluminous and occupies most of the body (four-fifths). The colour is pink or translucent, being slightly pink along the ciliary comb rows. There is an axial funnel tube; the aboral pole is fringed with a row of long, branched papillae, which is also specific for this species (Fig. 5D). This new species differs from Beroe cucumis by its long body (up to three times longer compared to width), the roundish shape of the mouth, the dense diverticulae that do not anastomose with each other. Its stomodaeum is large and constitutes four-fifths of the width of the body. There is an axial funnel tube and an aboral pole with a row of long, branched, aboral papillae (Fig. 5). Reliable identification may only prove reliable thorough morphological examination of adult individuals and DNA analyses. Geographical distribution: This species occurs in warm waters. Genetically its occurrence was confirmed off Villefranche sur Mer (Ligurian Sea) and the Adriatic Sea (Mediterranean), the north-eastern Pacific off the Californian coast and in the north-west Atlantic at Woods Hole, Massachusetts (Podar et al., 2001; Bayha et al., 2004; Johansson et al., 2018; this study). The species is also known from morphological studies in the Mediterranean Sea (Chun, 1880; Mertens, 1833; Galil & Gevili, 2013; Shiganova & Malej, 2009). In an extensive study of warm-water ctenophores, Harbison et al. (1978) showed that Beroe cucumis is widespread also in the tropical and subtropical Atlantic. However, he highlighted (Harbison, 2001) that the species he called Beroe cucumis sensu Mayer after Harbison et al. (1978), is not the same as the one from Greenland described by Fabricius (1780). The same opinion was expressed by Wrobel & Mills (1998) after study of individuals from the Pacific coast of the USA. We suppose that all these individuals listed as B. cucumis, but found outside the cold-water areas, belong to Beroe pseudocucumis: Spanish coast, Madeira, the Canary Islands, the Caribbean and from the east coast of the USA to southern Brazil (Bayha et al., 2004; Oliveira & Migotto, 2006; Moss, 2009; Oliveira et al., 2016). A few studies mentioned similar individuals from the Indian Ocean and the Arabian Sea (Harbison et al., 1978; Gul & Jahangir, 2019). It is highly likely that these also are the warm-water Beroe pseudocucumis, also inhabiting the Indo-Pacific (von Drygalski, 1909; Bigelow, 1919). Habitat: Warm water, inshore and offshore (F. Lombard, pers. comm; Shiganova & Malej, 2009.); widespread in tropical and subtropical waters., Published as part of Shiganova, Tamara A. & Abyzova, Galina A., 2022, Revision of Beroidae (Ctenophora) in the southern seas of Europe: systematics and distribution based on genetics and morphology, pp. 297-322 in Zoological Journal of the Linnean Society 194 on pages 306-307, DOI: 10.1093/zoolinnean/zlab021, http://zenodo.org/record/5799206, {"references":["Eschscholtz R. 1829. System der Acalephen. Eine ausfuhrliche Beschreibung aller Medusenartigen Strahlthiere. Berlin: F. Dummler, 190.","Bayha KM, Harbison GR, Mcdonald, JH, Gaffney PM. 2004. Preliminary investigation on the molecular systematics of the invasive ctenophore Beroe ovata. In: Dumont H, Shiganova TA, Niermann U, eds. Aquatic invasions in the Black, Caspian, and Mediterranean Seas. Dordrecht: Springer, 167 - 175.","Shiganova TA, Malej A. 2009. Native and non-native ctenophores in the Gulf of Trieste, Northern Adriatic Sea. Journal of Plankton Research 31: 61 - 71.","Shiganova TA, Riisgard HU, Ghabooli S, Tendal OS. 2014 b. First report on Beroe ovata in an unusual mixture of ctenophores in the Great Belt (Denmark). Aquatic Invasions 9: 111 - 116.","Podar M, Haddock SH, Sogin ML, Harbison GR. 2001. A molecular phylogenetic framework for the phylum Ctenophora using 18 S rRNA genes. Molecular Phylogenetics and Evolution 21: 218 - 230.","Johansson ML, Shiganova TA, Ringvold H, Stupnikova AN, Heath DD, MacIsaac HJ. 2018. Molecular insights into the ctenophore genus Beroe in Europe: new species, spreading invaders. Journal of Heredity 109: 520 - 529.","Chun C. 1880. Die Ctenophoren des Golfes von Neapel. Fauna und Flora des Golfes von Neapel vol. 1. Berlin: R. Friedlander & Sohn, 313.","Mertens H. 1833. Beobachtungen und Untersuchungen uber die beroeartigen Akalephen. Memoires de l'Academie Imperiale des Sciences de St. - Petersbourg 2: 479 - 543.","Galil BS, Gevili R. 2013. A moveable feast: Beroe cucumis sensu Mayer, 1912 (Ctenophora; Beroida; Beroidae) preying on Mnemiopsis leidyi A. Agassiz, 1865 (Ctenophora; Lobata; Bolinopsidae) off the Mediterranean coast of Israel. BioInvasions Records 2: 191 - 194.","Harbison GR, Madin LP, Swanberg NR. 1978. On the natural history and distribution of oceanic ctenophores. Deep Sea Research 25: 233 - 256.","Harbison GR. 2001. A guide to American species of ctenophores introduced into the Black and Mediterranean Seas. Naples: CIESM Workshop.","Fabricius, O. 1780. Fauna Groenlandica. Copenhagen and Leipzig: J. G. Rothe, 450.","Wrobel D, Mills C. 1998. Pacific coast pelagic invertebrates. A guide to the common gelatinous animals. Monterey: Sea Challengers & Monterey Bay Aquarium.","Oliveira OM, Migotto AE. 2006. Pelagic ctenophores from the Sao Sebastiao Channel, southeastern Brazil. Zootaxa 1183: 1 - 26.","Moss AG. 2009. Ctenophora of the Gulf of Mexico. In: Felder DL, Camp DK, eds. Gulf of Mexico - origins, waters, and biota. Biodiversity. College Station, Texas: Texas A & M University Press, 403 - 411.","Oliveira OM, Miranda TP, Araujo EM, Ayon P, Cedeno- Posso CM, Cepeda-Mercado AA, Cordova P, Cunha AF, Genzano GN, Haddad MA, Mianzan HW. 2016. Census of Cnidaria (Medusozoa) and Ctenophora from South American marine waters. Zootaxa 4194: 1 - 256.","Gul S, Jahangir S. 2019. First occurrence of Pleurobrachia pileus (O. F. Muller, 1776) and sighting records of four other species of comb jellies (Ctenophora) from the coast of Pakistan (northern Arabian Sea): a brief account. International Journal of Biology and Biotechnology 16: 991 - 993.","von Drygalski E. 1909. Deutsche Sudpolar-Expedition, 1901 - 1903: im Auftrage des Reichsamtes des Innern 4. Berlin: G. Reimer.","Bigelow HB. 1919. Hydromedusae, siphonophores, and ctenophores of the ' Albatross' Philippine expedition. Washington, DC: US Government Printing Office."]}

Beroe cucumis Fabricius, 1780 Remarks: Harbison et al. (1978) documented the presence of Beroe spp., ���most of them ressembling Beroe cucumis ��� off the northern Brazilian coast and Guianas. Distribution in South America: Pacific Ocean, Peru to Chile, from 3��S to 55��S (Palma 1971, 1994; Palma & Rosales 1995; Pag��s & Orejas 1999; Palma & Apablaza 2004; IMARPE database 2009; Pavez et al. 2010); Atlantic Ocean, Brazil to Argentina, from 30��S to 59��S (Mianzan 1999; Oliveira et al. 2007). Habitat: epipelagic species (Mayer 1912; Harbison et al. 1978)., Published as part of M. P. Oliveira 1,16, S P. Miranda 2, *,, Es W. Mianzan 10,, Ro E. Migotto 11,, Ne B. Nascimento 2,11, Eli Nogueira J��nior 12,, Er Qui��ones 13,, Izio Scarabino 14,, T��n Schiariti 10,, Io N. Stampar 15,, Tronolone 2, , Qu��ria B. & Onio C. Marques 2,11, 2016, Census of Cnidaria (Medusozoa) and Ctenophora from South American marine waters, pp. 1-256 in Zootaxa 4194 (1) on page 211, DOI: 10.11646/zootaxa.4194.1.1

Beroe ovata Chamisso & Eysenhardt, 1821 Remarks: ��� Beroe forskali Chun ��� and ���larves de Beroe forskali ��� recorded by Seguin (1965) are B. ovata. Beroe cucumis from Cartagena Bay (Moncaleano & Ni��o 1976) is also considered as B. ovata. Distribution in South America: Atlantic Ocean, Colombia to Argentina, from 10��N to 42��S (Domaneschi 1976; Moncaleano & Ni��o 1976; Genzano & Zamponi 1993; Mianzan 1986, 1999; Oliveira & Migotto 2006; Oliveira et al. 2007; Nogueira Jr. 2012; Nogueira Jr. et al. 2015b). Habitat: euryhaline and neritic species, found in coastal and estuarine waters (Mianzan 1986; Oliveira & Migotto 2006)., Published as part of M. P. Oliveira 1,16, S P. Miranda 2, *,, Es W. Mianzan 10,, Ro E. Migotto 11,, Ne B. Nascimento 2,11, Eli Nogueira J��nior 12,, Er Qui��ones 13,, Izio Scarabino 14,, T��n Schiariti 10,, Io N. Stampar 15,, Tronolone 2, , Qu��ria B. & Onio C. Marques 2,11, 2016, Census of Cnidaria (Medusozoa) and Ctenophora from South American marine waters, pp. 1-256 in Zootaxa 4194 (1) on pages 211-212, DOI: 10.11646/zootaxa.4194.1.1

Beroe forskalii Milne-Edwards, 1841 Distribution in South America: Pacific Ocean, Peru, at 13.8��S 76.26��W in Bahia de Paracas (specimens collected in this study); Atlantic Ocean, Brazil, at 23.8�� S, 45.4��W, in the S��o Sebasti��o Channel (Oliveira & Migotto, 2014). Habitat: epi-mesopelagic species (Lindsay & Hunt 2005)., Published as part of M. P. Oliveira 1,16, S P. Miranda 2, *,, Es W. Mianzan 10,, Ro E. Migotto 11,, Ne B. Nascimento 2,11, Eli Nogueira J��nior 12,, Er Qui��ones 13,, Izio Scarabino 14,, T��n Schiariti 10,, Io N. Stampar 15,, Tronolone 2, , Qu��ria B. & Onio C. Marques 2,11, 2016, Census of Cnidaria (Medusozoa) and Ctenophora from South American marine waters, pp. 1-256 in Zootaxa 4194 (1) on page 211, DOI: 10.11646/zootaxa.4194.1.1

Beroe gracilis K��nne, 1939 Remarks: The species assingned as Beroe sp. in Palma (1994) and Palma & Rosales 1995) are, in fact, B. gracilis. Distribution in South America: Pacific Ocean, Chile, from 32�� ��� 36��30���S, in Valpara��so Bay and off Dichato (Oliveira et al. 2014). Habitat: Epi-mesopelagic species (Wrobel & Mills, 2003)., Published as part of M. P. Oliveira 1,16, S P. Miranda 2, *,, Es W. Mianzan 10,, Ro E. Migotto 11,, Ne B. Nascimento 2,11, Eli Nogueira J��nior 12,, Er Qui��ones 13,, Izio Scarabino 14,, T��n Schiariti 10,, Io N. Stampar 15,, Tronolone 2, , Qu��ria B. & Onio C. Marques 2,11, 2016, Census of Cnidaria (Medusozoa) and Ctenophora from South American marine waters, pp. 1-256 in Zootaxa 4194 (1) on page 211, DOI: 10.11646/zootaxa.4194.1.1

We have compiled available records in the literature for medusozoan cnidarians and ctenophores of South America. New records of species are also included. Each entry (i.e., identified species or still as yet not determined species referred to as "sp." in the literature) includes a synonymy list for South America, taxonomical remarks, notes on habit, and information on geographical occurrence. We have listed 800 unique determined species, in 958 morphotype entries: 5 cubozoans, 905 hydrozoans, 25 scyphozoans, 3 staurozoans, and 20 ctenophores. Concerning nomenclatural and taxonomical decisions, two authors of this census (Miranda, T.P. & Marques, A.C.) propose Podocoryna quitus as a nomen novum for the junior homonym Hydractinia reticulata (Fraser, 1938a); Euphysa monotentaculata Zamponi, 1983b as a new junior synonym of Euphysa aurata Forbes, 1848; and Plumularia spiralis Milstein, 1976 as a new junior synonym of Plumularia setacea(Linnaeus, 1758). Finally, we also reassign Plumularia oligopyxis Kirchenpauer, 1876 as Kirchenpaueria oligopyxis(Kirchenpauer, 1876) and Sertularella margaritacea Allman, 1885 as Symplectoscyphus margaritaceus (Allman, 1885). Fil: Oliveira, Otto M. P.. Universidade Federal de Santa Catarina; Brasil Fil: Miranda, Thaís P.. Universidade de Sao Paulo; Brasil Fil: Araujo, Enilma M.. Faculdade Pitágoras de Feira de Santana; Brasil Fil: Ayón, Patricia. Instituto Mar del Peru; Perú Fil: Cedeño Posso, Cristina M.. Instituto de Investigaciones Marinas y Costeras; Colombia Fil: Cepeda Mercado, Amancay A.. ProCAT Colombia/Internacional; Colombia Fil: Córdova, Pablo. Pontificia Universidad Católica de Chile; Chile Fil: Cunha, Amanda F.. Universidade de Sao Paulo; Brasil Fil: Genzano, Gabriel Nestor. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Haddad, Maria Angélica. Universidade Federal do Paraná; Brasil Fil: Mianzan, Hermes Walter. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Subsede Instituto Nacional de Investigación y Desarrollo Pesquero; Argentina Fil: Migotto, Alvaro E.. Universidade de Sao Paulo; Brasil Fil: Miranda, Lucília S.. Universidade de Sao Paulo; Brasil Fil: Morandini, André C.. Universidade de Sao Paulo; Brasil Fil: Nagata, Renato M.. Universidade de Sao Paulo; Brasil Fil: Nascimento, Karine B.. Universidade de Sao Paulo; Brasil Fil: Nogueira, Miodeli. Universidade Federal da Paraíba; Brasil Fil: Palma, Sergio. Pontificia Universidad Católica de Chile; Chile Fil: Quiñones, Javier. Instituto Mar del Perú; Perú Fil: Rodriguez, Carolina Soledad. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Scarabino, Fabrizio. Instituto Nacional de Investigaciones y Desarrollo Pesquero; Argentina Fil: Schiariti, Agustin. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Subsede Instituto Nacional de Investigación y Desarrollo Pesquero; Argentina Fil: Stampar, Sergio. Universidade Estadual Paulista Julio de Mesquita Filho; Brasil Fil: Tronolone, Valquíria B.. Universidade de Sao Paulo; Brasil Fil: Marques, Antonio C.. Universidade de Sao Paulo; Brasil

Ruiz-Escobar, Fernando, Valadez-Vargas, Diana K., Oliveira, Otto M. P. (2015): Ctenophores from the Oaxaca coast, including a checklist of species from the Pacific coast of Mexico. Zootaxa 3936 (3): 435-445, DOI: http://dx.doi.org/10.11646/zootaxa.3936.3.9

Beroe forskalii Milne Edwards, 1841 (Fig. 4; Table 3) Material examined. Oaxaca, Mexico: Four specimens; one specimen 55 mm long at Puerto ��ngel Bay, 15 �� 39 ' 54.04 " N, 96 �� 29 ' 36.73 " W, 7 December 2013, surface, coll. A. Ramos-Silva, placed in 70 % ethanol, dissolved after 24 h; UMAR-CTEN-005, 2 specimens (Zipolite, 15 �� 38 ' 49 '' N, 96 �� 31 ' 46 '' W, 2 m, 25 January 2014, coll. F. Ruiz-Escobar); UMAR-CTEN-006, one specimen (Carrizalillo, 15 �� 51 ' 26.64 " N, 97 �� 4 ' 48.89 " W, surface, 15 February 2014, coll. F. Ruiz-Escobar). Description. Body long, very flattened at the tentacular plane, typically conical, at least in the aboral half, with a broad mouth (Fig. 4 B). Apical organ at the aboral extremity, surrounded by two sets of seven branched aboral papillae, of pink color, the five proximal longer, and two distal shorter. Subtentacular and substomodeal ctene rows about the same length, extending from near the aboral end to about 1 / 7 of the body distance to the mouth. Meridional canals covered by the ctene rows, with a small uncovered part, never reaching the circular canal around the mouth. Numerous anastomosed diverticulae connecting the meridional canals, and these with the circular canal. Paragastric canals running along both surfaces of the pharynx, also connecting with the anastomosed branches between every two subtentacular meridional canals. Gonads present along the meridional canals and nearest parts of the diverticulae; testes formed between subtentacular and substomodeal meridional canals; ovaries formed between subtentacular and substomodeal meridional canals. Remarks. The morphology of the studied specimens is similar to those described by Wrobel & Mills (2003). Although the macrociliary patterns were not checked, we had no doubts about species identification, since we found only large adult specimens with full morphological characteristics. It would be different if we had seen juveniles (viz Oliveira & Migotto 2014), whose morphology is sometimes less distinctive. Distribution. This species is widely distributed in the Atlantic and Pacific Oceans, as well as in the Mediterranean Sea (Wrobel & Mills 2003; Oliveira & Migotto 2014). In the Eastern Pacific, it occurs along most of the USA coast to Canada (Wrobel & Mills 2003) and south as far as the Peruvian coast (Oliveira et al., in press). Total length of the body 50.2 mm Maximum width of the body 22.2 mm Length of the subtentacular ctene rows 37.1 mm Length of the substomodeal ctene rows 38.2 mm Maximum width between substomodeal and subtentacular ctene rows up to 5.2 mm Maximum width between subtentacular ctene rows up to 7.6 mm Width of the mouth 17.5 mm Number of ctene plates in subtentacular ctene rows 97���107 Number of ctene plates in substomodeal ctene rows 92���105 Length of the polar field 2785 ��m Length of the papillae up to 823 ��m Diameter of the meridional canals up to 950 ��m Diameter of the diverticulae up to 253 ��m Diameter of the paragastric canals up to 654 ��m Diameter of the adradial canal up to 591 ��m Diameter of the radial canal up to 443 ��m Width of the ctene plates 190���1160 ��m Distance between ctene plates 169���401 ��m Diameter of the statocyst 189 ��m, Published as part of Ruiz-Escobar, Fernando, Valadez-Vargas, Diana K. & Oliveira, Otto M. P., 2015, Ctenophores from the Oaxaca coast, including a checklist of species from the Pacific coast of Mexico, pp. 435-445 in Zootaxa 3936 (3) on pages 441-443, DOI: 10.11646/zootaxa.3936.3.9, http://zenodo.org/record/245778, {"references":["Milne Edwards, M. H. (1841) Observations sur la structure et les fonctions de quelques Zoophytes, Mollusques et Crustaces des cotes de la France. Annales des Sciences Naturelles, Serie 2, 16, 193 - 232.","Wrobel, D. & Mills, C. (2003) Pacific Coast Pelagic Invertebrates - A Guide to the Common Gelatinous Animals. Sea Challengers and the Monterey Bay Aquarium, Monterey, California, 108 pp.","Oliveira, O. M. P & Migotto, A. E. (2014) First occurrence of Beroe forskalii (Ctenophora) in South American Atlantic coastal waters, with notes on the use of macrociliary patterns for beroid identification. Zootaxa, 3779 (4), 470 - 476. http: // dx. doi. org / 10.11646 / zootaxa. 3779.4.5"]}

Beroe forskalii Milne Edwards, 1841 (Figures 1���2; Table 1) Beroe forskalii Milne Edwards, 1841 ��� Chun 1880; Tamm & Tamm 1991, 1993; Haddock & Case 1999; Mianzan 1999; Wrobel & Mills 2003; Lindsay & Hunt 2005; Lindsay 2006; Shiganova & Malej 2009. Material examined. MZUSP 00012, a single specimen 2.5 cm long from Canal de S��o Sebasti��o, 23.8�� S, 45.4�� W, coll. A. Migotto 8 July 2010, reared to 10 July 2010 then fixed in 4% formalin. Description. Body sac-shaped, pinkish, with broad mouth; body width tapering from middle to aboral extremity of body (Fig. 1A); compressed along tentacular plane (Fig. 1C). Eight meridional canals extending from aboral region towards mouth and connected with a circular canal around mouth by a mesh of anastomosing canals; these with lateral diverticulae, mostly with blind ends in aboral half of body, but forming anastomoses along oral half (Fig. 1F). Diverticulae from subtentacular meridional canals connected with paragastric canal (Fig. 1E). Ctene rows located above meridional canals arising at aboral region. Substomodeal ctene rows extending for about 3/4 the length of the meridional canals to oral region. Subtentacular ctene rows limited to aboral 2/3 of body. Mouth wide, semicircular, occupying whole oral region, opening into large pharynx that occupies most of central inner part of animal. Two opposite paragastric canals, extending from aboral pole of pharynx to circular canal around mouth, crossing at tentacular plane. Aboral extremity of body mitriform (Fig. 1D). Apical organ including a statolith at center of polar fields. Aboral papillae prominent, branched (Fig. 1G), projecting from margins of polar fields, forming an ���hourglass��� figure, oriented in stomodeal plane when in aboral view (Fig. 1B). Each papilla with first- and sometimes second-order branches in a pinnate arrangement; branches curving toward exterior of polar field. The animal is a simultaneous hermaphrodite, with gonads formed at lateral walls of meridional canals and nearby diverticulae (Fig. 1E). Ovaries restricted to portion of meridional canals under ctene rows. Testes lying along all meridional canals and adjacent diverticula. Gonads of adjacent meridional canals facing those of same gender on the adjacent canal, i.e. female gonads face in stomodeal-stomodeal and tentacle-tentacle patterns; male gonads face in a stomodeal-tentacular arrangement. Macrocilia are robust (length 5���7 times width), with about 20 small teeth along edge (Fig. 2), covering extensive area of oral portion of stomodeal cavity, arranged in long tapering stripes that run from lip toward center of stomodeum. Remarks. The young individual on which the above description is based, was very active for the three days it was kept in the aquarium, swimming quickly, sometimes with the mouth open. It resembled Beroe mitrata Moser, 1907, except for the presence of anastomoses and the pointed end of the aboral portion (Wrobel & Mills, 2003). Based on the shape of their macrocilia, these two species form a distinct group within the genus Beroe (Tamm & Tamm 1993). In these two species, the macrocilia are very similar, differing only by their pattern of distribution along the stomodeal cavity, comprising a carpet covering all of the oral portion of the stomodeum in B. mitrata and long tapering stripes that run from the lip toward the center of the stomodeum in B. forskalii (Tamm & Tamm 1993). On this basis, inter alia, the specimen from S��o Sebasti��o Channel was certainly B. forskalii. The first description of what was probably this species was made by Forsk��l (1775), as Beroe rufescens. This name was not used by later researchers after Milne Edwards (1841) questioned its validity. The combination Beroe forskalii has been used in at least 20 publications in the last 50 years and, following the nomenclatural Principle of Stability (ICZN 1999), its use may continue. The author of the combination Beroe forskalli is taken as Milne Edwards (1841), not Chun (1880), following the Principle of Priority (ICZN 1999). Distribution. Beroe forskalii is distributed worldwide (e.g. Moser 1910; Mianzan 1999; Lindsay & Hunt 2005; Shiganova & Malej 2009; Oliveira et al. submitted)., Published as part of Oliveira, Otto M. P. & Migotto, Alvaro E., 2014, First occurrence of Beroe forskalii (Ctenophora) in South American Atlantic coastal waters, with notes on the use of macrociliary patterns for beroid identification, pp. 470-476 in Zootaxa 3779 (4) on pages 471-473, DOI: 10.11646/zootaxa.3779.4.5, http://zenodo.org/record/4910453, {"references":["Milne Edwards, M. H. (1841) Observations sur la structure et les fonctions de quelques Zoophytes, Mollusques et Crustaces des cotes de la France. Annales des Sciences Naturelles, ser. 2, 16, 193 - 232.","Chun, C. (1880) Die Ctenophoren des Golfes von Neapel und der angrenzenden Meeres-abschnitte. Eine Monographie. Fauna and flora des Golfes von Neapel, 1, Monographie, 18, 1 - 313, 18 pls.","Tamm, S. L. & Tamm, S. (1991) Reversible epithelial adhesion closes the mouth of Beroe, a carnivorous marine jelly. Biological Bulletin, 181, 463 - 474.","Tamm, S. L. & Tamm, S. (1993) Diversity of macrociliary size, tooth patterns, and distribution in Beroe (Ctenophora). Zoomorphology, 113, 79 - 89. http: // dx. doi. org / 10.1007 / bf 00403086","Haddock, S. H. D. & Case, J. F. (1999) Bioluminescence spectra of shallow and deep-sea gelatinous zooplankton: ctenophores, medusae and siphonophores. Marine Biology, 133, 571 - 582. http: // dx. doi. org / 10.1007 / s 002270050497","Mianzan, H. W. (1999) Ctenophora. In: Boltovskoy, D. (Ed.), South Atlantic Zooplankton. Backhuys Publishers, Leiden, pp. 561 - 573.","Wrobel, D. & Mills, C. (2003) Pacific Coast Pelagic Invertebrates - A Guide to the Common Gelatinous Animals. Second Printing with corrections. Sea Challengers and the Monterey Bay Aquarium, Monterey, California, iv + 108 pp.","Lindsay, D. J. & Hunt, J. C. (2005) Biodiversity in midwater cnidarians and ctenophores: submersible based results from deep- water bays in the Japan Sea and north-western Pacific. Journal of the Marine Biological Association of the United Kingdom, 85, 503 - 517. http: // dx. doi. org / 10.1017 / s 0025315405011434","Lindsay, D. J. (2006) A checklist of midwater cnidarians and ctenophores from Sagami Bay - species sampled during submersible surveys from 1993 - 2004. Bulletin of the Plankton Society of Japan, 53, 104 - 110.","Shiganova, T. & Malej, A. (2009) Native and non-native ctenophores in the Gulf of Trieste, Northern Adriatic Sea. 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Oliveira, Otto M. P., Migotto, Alvaro E. (2014): First occurrence of Beroe forskalii (Ctenophora) in South American Atlantic coastal waters, with notes on the use of macrociliary patterns for beroid identification. Zootaxa 3779 (4): 470-476, DOI: http://dx.doi.org/10.11646/zootaxa.3779.4.5

Oliveira, Otto M. P., Feliú, Guillermo, Palma, Sergio (2014): Beroe gracilis (Ctenophora) from the Humboldt Current System: first occurrence of this species in the southern hemisphere. Zootaxa 3827 (3): 397-400, DOI: http://dx.doi.org/10.11646/zootaxa.3827.3.10