Beroe albens Forskål, 1775 Beroe ovata Bosc, 1802 Beroe ovata Chamisso & Eysenhardt, 1821 Beroe capensis Chamisso & Eysenhardt, 1821 Beroe punctata Chamisso & Eysenhardt, 1821 Beroe ovata Eschscholtz, 1829 Beroe punctata Idya mertensii Mertens, 1833 Idyiopsis clarkii L. Agassiz, 1860 Idyiopsis clarkii Idyiopsis affinis L. Agassiz, 1860 Beroe ovata Beroe ovata Mnemiopsis leidyi Bolinopsis vitrea Pelagic ctenophores from the São Sebastião Channel, southeastern Brazil Oliveira, Otto M. P. Migotto, Alvaro E. Zootaxa 2006 2006-04-24 1183 1 26 5WJTR (Chamisso & Eysenhardt, 1821) Chamisso & Eysenhardt 1821 [264,814,1228,1254] Nuda Beroidae Beroe Animalia Beroida 16 17 Ctenophora species ovata  ( Fig. 6, Table 3)  ?  Beroe albens Forskål, 1775. ?  Beroe ovata Bosc, 1802.   Beroe ovata Chamisso & Eysenhardt, 1821.   Beroe capensis Chamisso & Eysenhardt, 1821.   Beroe punctata Chamisso & Eysenhardt, 1821. Not  Beroe ovata Eschscholtz, 1829.   Beroe punctata— Eschscholtz, 1829; McCrady, 1859.   Idya mertensii Mertens, 1833.   Idyiopsis clarkii L. Agassiz, 1860.   Idyiopsis clarkii— L. Agassiz, 1865.   Idyiopsis affinis L. Agassiz, 1860.   Beroe ovata— Moser, 1903, 1910; Mayer, 1912[see for complete synonyms until 1912]; Pratt,1935; Kremer et al., 1986a; Sterrer, 1986; Mianzan, 1999; Shiganova et al., 2001b.    Beroe capensis— Chun, 1880.   Beroe clarkiiMayer, 1900. Not  Beroe ovata— Chun, 1880. Not  Beroe“ ovata”— Mills et al., 1996.   Examined material     22.ix.2003, Near Farol dos Moleques, Canal de São Sebastião, 23°49'27"S 45°24'42"W, A.A.S.Mouracoll., one juvenile (ca. 3 mm), reared to   13.x.2003, Fig. 6J–K, and one adult (ca. 70 mm), fixed in 2% formalin, Fig. 6E( MZUSP00001);      06.iv.2004, Baíado Saco Grande, Canal de São Sebastião, 23°49'35"S 45°25'26"W, bloom of adults (up to 70 mmin length), observed during snorkelling; O.M.P.Oliveiracoll., three adults ( 37–68 mmin length), one specimen preserved in 95% ethanol, other two preserved in 2% formalin ( MZUSP00002);     05.v.2004, Off Praia do Segredo, Canal de São Sebastião, 23°49'27"S 45°25'19"W, V. Radashevskycoll., two juveniles (ca. 2 mmin length), reared to   11.v.2004, preserved in 2% formalin ( MZUSP00003);     21.v.2004, Near Farol dos Moleques, Canal de São Sebastião, 23°49'27"S 45°24'42"W, O.M.P.Oliveiracoll., ten adult ( 10–50 mmin length), reared to   24.v.2004, seven specimens preserved in 2% formalin ( MZUSP00004);     18.viii.2004, Off Vila, Ilhabela, Canal de São Sebastião, 23°45'27"S 45°21'49"W, bloom of adults (up to 70 mmin length), observed during snorkelling;   31.viii.2004,  Baíado Saco Grande, Canal de São Sebastião, 23°49'35"S 45°25'26"W, O.M.P.Oliveiracoll., one adult (ca. 60 mmin length), reared to 01.ix.2004, preserved in 95% ethanol and two larvae released from reared specimen at 01.ix.2004(ca. 0.5 mmin length), reared to 02.ix.2004;     27.xi.2004, Baíado Segredo, Canal de São Sebastião, 23°49'27"S 45°25'19"W, O.M.P.Oliveiracoll., one adult (ca. 59 mmin length), preserved in 95% ethanol;     01.iv.2005, Off Praia do Curral, Ilhabela, Canal de São Sebastião, 23°50'54"S 45°25'59"W, one juvenile (ca. 7 mmin length), preserved in 95% ethanol;     05.iv.2005, Off Praia do Curral, Ilhabela, Canal de São Sebastião, 23°50'54"S 45°25'59"W, ten juveniles ( 3–12 mmin length), seven specimen preserved in 95% ethanol;     07.iv.2005, Near Ilha das Cabras, Canal de São Sebastião, 23°49'25"S 45°23'33"W, one juvenile (ca. 8 mmin length), preserved in 95% ethanol;     17.v.2005, Off Praia do Curral, Ilhabela, Canal de São Sebastião, 23°50'54"S 45°25'59"W, bloom of adults (up to 70 mmin length), observed during snorkelling; O.M.P. Oliveiracoll., 39 adults( 34–66 mmin length), one specimen measured at 19.v.2005and reared to   20.v.2005(table 3, Fig. 6 B–D, MZUSP00005), four specimens reared to   07.vi.2005( Fig. 6F–I), 5 specimenspreserved in 2% formalin ( MZUSP00006), 23 specimenspreserved in 95% ethanol.  Description Flattened body with widened oral extremity, compressed along tentacular plane, reaching about 7.0 cm in length ( Fig. 6A,B). Body surface covered by small red spots. Eight meridional canals extending from aboral region to a circular canal around mouth. Canals with lateral diverticula, mostly with blind ends, extending to inner gelatinous portion of body ( Fig. 6C). A variable number of diverticula from one meridional canal con­ necting with those of adjacent meridional canal, sometimes forming anastomoses ( Fig. 6E). Some diverticula connected with paragastric canals. Rows of ctene plates located above meridional canals, arising at aboral region, extending ca. 3/4 length of meridional canals to oral region. Mouth wide, occupying entire oral region, opening into a large pharynx that occupies most of central inner part of animal. Two opposing paragastric canals extending from aboral pole of pharynx to radial canal around mouth, penetrating jelly beneath diverticula at tentacular plane. Apical organ comprising statolith at centre of polar fields ( Fig. 6D). Aboral papillae projecting from margins of polar fields, forming a figure 8 orientated in stomodeal plane. Gonads formed at sides of meridional canals and diverticula ( Fig. 6F), testes at distal side and ova at proximal side of canals from tentacular plane in subtentaclar canals and opposite in substomodeal canals. Formation of temporary gonopores through rupture of epidermis near polar fields ( Fig. 6G). Gonopore at end of short cone; closing of gonopores leaving no scar on animal surface.   TABLE 3.—Measurements of a  Beroe ovata(Chamisso & Eysenhardt, 1821)specimen (MZUSP00005) with 33.2 mm in length, in stomodeal plane.    Total length of body  33.2 mm  Maximum width of body  29.3 mm  Length of subtentacular ctene rows 31.0 mm  Length of substomodeal ctene rows  30.9 mm  Maximum width between substomodeal and subtentacular ctene rows up to 8.0 mm  Maximum width between subtentacular ctene rows up to 9.0 mm  Width of mouth  24.5 mm  Number of ctene plates in subtentacular ctene rows 118–121  Number of ctene plates in substomodeal ctene rows 95–108  Number of connections between substomodeal and subtentacular meridi­ onal canals 0–7  Number of connections between subtentacular meridional canals 4–7  Length of polar field  2920 µm  Number of papillae 42  Length of papillae up to 220 µm  Diameter of genital pore  360 µm  Diameter of meridional canals up to 740 µm  Diameter of diverticula up to 240 µm  Diameter of paragastric canals up to 380 µm  Diameter of radial canal up to 875 µm  Width of ctene plates  180–725 µm  Distance between ctene plates  80–240 µm  Diameter of statocyst  40 µm   FIGURE 6.—  Beroe ovata(Chamisso & Eysenhardt, 1821). A, total view of an adult in stomodeal plane; B, total view of another adult in stomodeal plane; C, closed view of the diverticula pattern (few connections) of an adult in stomodeal plane; D, aboral view of an adult; E, closed view of the diverticula pattern (anastomosis) of another adult in stomodeal plane; F, closed view of a meridional canal and diverticula with male and female gametes; G, closed view of the aboral portion of the body presenting the temporary formation of a gonopore; H, closed view of anastomosed diverticula with a larva inside; I, two larvae in gelatinous egg inside diverticula of an adult specimen; J, closed view of the aboral portion of a larva; K, total view of a larva in stomodeal plane. Legend: ao, apical organ; di, diverticulum; fg, female gametes; ge, gelatinous egg; gp, gonopore; la, larva; mg, male gametes; mo, mouth; pa, papilla; pc, paragastric canal; pf, polar field; sscr, substomodeal ctene row; stcr, subtentacular ctene row. Larva similar to adult, but less compressed along tentacular plane and with no lateral diverticula along meridional canals ( Fig. 6K). Body surface covered by conspicuous small red spots ( Fig. 6J).  Biological notes    Beroe ovatapreys on gelatinous zooplankton, being frequently associated with blooms of  Mnemiopsis leidyiand  Bolinopsis vitrea. Some authors (e.g. GESAMP 1997, Volovik & Korpakova 2004) believed that this species can be used as a biological­control agent in areas where gelatinous plankton blooms are not desirable, as in the case of the Black and the Caspian Seas ( Shiganova et al.2001b). The species is supposed to be oviparous, fertilization occurring outside the parental bodies ( Carré & Sardet 1984). Indeed, we observed spawning through the temporary gonopores, formed laterally to the apical organ. Male and female gametes were not released at the same time in reared specimens, what would be a way of avoiding self­fertilization outside the parental body. However, we also observed the presence of larvae inside the connected diverticula ( Fig. 6H–I), a strong indication that these larvae were the product of self­fertilization.  We collected and reared in the laboratory for several days specimens in several ontogenetic stages, from 3.0 mm larvae to adults. The smallest specimens were observed eating hydromedusae of about their size.   Remarks The genus name “  Beroe” was initially used in the descriptions of ctenophores in general, referring to species that are now within distinct families and genera ( e.g. Hormiphora cucumis,  Callianira compressa,  Euplokamis octoptera, and  Dryodora glandiformis, described in Mertens 1833, plates 8–11, as  Beroe cucumis,  B. compressa,  B. octoptera, and  B. glandiformis). This wide use of a genus name led to a great confusion around the concept of some species, as “  Beroe ovata” ( cf. Mills et al., 1996, p.161). This species name, with two distinctive origins, currently refers to two biological species ( Mills et al.1996) — “  Beroe ovata”, described by Chun (1880)and Mayer (1912, as  B. cucumis)as having a cylindrical body, and “  Beroe ovata” ( Mayer 1912), with a body compressed along the tentacular axis. Mayer (1912)used other characters, such as the connection between diverticula, to distinguish the species. Recently, in a biomolecular study, Bayha et al.(2004)distinguished two genetic pools of  B. ovata: one from the Mediterranean Sea, involving specimens morphologically similar to the ones described by Chun (1880)(=  B. cucumis sensu Mayer, 1912), and the other from the North American Atlantic coast and invasive in the Black Sea (=  B. ovataas described by Mayer, 1912). Our specimens are closer to those described as  B. ovataby Mayer (1912)and Mianzan (1999), except for the patterns of diverticula connections. The range from unconnected to anastomosed diverticula observed by us in specimens of the same population indicates that the anastomosis pattern is not an adequate character to distinguish species of the genus  Beroe. Furthermore, an extensive morphological revision, supported by molecular data and including specimens from South Atlantic and Caribbean, should be done to confirm the identity of these nominal species.   Distribution Species found on coastal waters of the western Atlantic, from USAto Argentina( Mayer 1912, Mianzan 1999). Recently Shiganova et al.(2001b)and Volovik & Korpakova (2004)reported its occurrence in Black and Caspian Seas, as an invader species ( cf. Bayha et al. 2004).  Acknowledgments The authors are grateful to CEBIMar­USP for logistics; to Dr. Hermez Mianzan (INIDEP­ Argentina) and Dr. G. Richard Harbison (WHOI­USA) for providing literature; to Dr. Keith M. Bayha (Dauphin Island Sea Lab­USA) for important suggestions; to Dr. George Matsumoto (MBARI­USA), Dr. Hermez Mianzan (INIDEP­Argentina), and Dr. Steven Haddock (MBARI­USA) for manuscript revisions. License for collecting specimens yielded by IBAMA/MMA (092/2004). Funding was provided by CAPES and FAPESP (Proc. 2004/15300­0) to Otto M. P. Oliveira and CNPq (Proc. 300194/1994–3) to A. E. Migotto.   References   Agassiz, A.( 1865)  Illustrated catalogue of the Museum of Comparative Zoology at Harvard College, Vol. 2, North American Acalephae. Sever and Francis, Cambridge, 234 pp.    Agassiz, L.( 1860)  Contributions to the natural history of the United States of America, Vol. 3. Little, Brown and Co., Boston, 301 pp.   Bayha, K.M.; Harbison, G.R.; McDonald, J.H.& Gaffney, P.M.( 2004) Preliminary investigation on the molecular systematics of the invasive ctenophore Beroe ovata. 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