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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."]}

Although ctenophores can be conspicuous components of the plankton in coastal marine ecosystems, only six species have been formally described from around South Africa. Using photographs from local community scientists, we add a further three species (Cestum veneris, Beroe forskalii?,Ocyropsis maculata?) and six morphospecies to the regional fauna. These additions suggest that South Africa has a ctenophore fauna that is amongst the most diverse, globally; an observation in agreement with information from other taxa. Tips on how community scientists can improve their photographic contributions to understanding ctenophore diversity are provided.

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 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. Journal of Plankton Research, 31, 61 - 71. http: // dx. doi. org / 10.1093 / plankt / fbn 102","Forskal, P. (1775) Descriptiones Animalium, Avium, Amphibiorum, Piscium, Insectorum, Vermium; quae in itinere orientali observavit Petrus Forskal. Hauniae, ex officina Molleri, 20 + xxxiv + 164 pp., 1 map.","ICZN (1999) International Code of Zoological Nomenclature, 4 th edn. ITZN, London, 306 pp.","Moser, F. (1910) Die Ctenophoren der Deutschen Sudpolar-Expedition 1901 - 1903. Deutsche Sudpolar-Expedition, 11, Zoologie, 117 - 192."]}

Beroe forskalii Milne Edwards, 1841 is an oceanic ctenophore with a global distribution. The present study provides the first record of Beroe forskalii for the South American Atlantic coast, including a redescription of the species and a discussion on the utility of macrociliary patterns for the correct identification of at least some beroid species, exemplified by a comparison of the macrociliary patterns of Beroe forskalii and Beroe ovata (Chamisso & Eysenhardt, 1821).