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Ophidion zavalai new species Figs. 1, 3, 5 and 6, Table 3 urn:lsid:zoobank.org:act: B9D7C4E8-5A9B-4402-A5FC-FD275639EF60 Ophidion holbrookii (not of Putnam 1874) — Menezes & Figueiredo 1980: 47 (guide of fishes of southeastern Brazil; short description; first report from western South Atlantic)— Robins & Ray 1986: 99 (in part; assigned to Southeastern Brazil}— Smith 1997: 361 (in part; assigned to Southeastern Brazil)— McEachran & Fechhelm 1998: 748 (in part; assigned to Southeastern Brazil)— Nielsen et al. 1999: 41 (in part; FAO Fisheries Synopses; short description; illustration; occurrence in Southeastern Brazil)— Nielsen & Robins 2002: 972 (in part; reported to Southeastern Brazil)—Menezes & Figueiredo in Menezes et al. 2003: 59 (catalog of marine fishes of Brazil; listed)— Lea & Robins 2003: 7 (in part; assigned to Southeastern Brazil)— Zavala-Camin & Rotundo 2011: 3–10 (anatomy of the head lateral line and hearing system) — Britski & Figueiredo 2019: 204 (plate 004 from Burkhardt 1865–1866; Rio de Janeiro, Brazil; regarded as new species of “ Ophidium ”). Doubtful references for this species Ophidion cf. holbrookii — Garcia Jr. et al. 2010: 47; 2015: 51 (fishes of Bacia Potiguar, coast of Rio Grande do Norte; Brazil. Short description; photograph). Holotype: MPEG 039113 (1, 271 mm SL, male), 24º14’59” S / 46º41’32” W, laje da Conceição, Itanhaém, São Paulo, Brazil Paratypes: MPEG 039114 (1, 250 mm SL, female), laje da Conceição, Itanhaém, São Paulo, Brazil; NPM 6885 (2, 195 mm SL, male, 294 mm SL, female), 24º14’59” S / 46º41’32” W, laje da Conceição, Itanhaém, São Paulo, Brazil; MZUSP 125946 (2, 208 mm SL, female, 271 mm SL, male), 24º14’59” S / 46º41’32” W, laje da Conceição, Itanhaém, São Paulo, Brazil; AZUSC 6916 (1, 218 mm SL, female), 24º14’59” S / 46º41’32” W, laje da Conceição, Itanhaém, São Paulo, Brazil; ZUEC 17383 (2, 190 mm SL, female, 296 mm SL, male), 26º15’23” S / 48º01’58” W, barra de S„o Francisco, S„o Francisco do Sul, Santa Catarina, Brazil; LPB 31046 (2, 188 mm SL, female, 278 mm SL, male) 26º15’23” S / 48º01’58” W, barra de São Francisco, São Francisco do Sul, Santa Catarina, Brazil; AZUSC 6917 (1, 281 mm SL, male), 26º15’23” S / 48º01’58” W, barra de São Francisco, São Francisco do Sul, Santa Catarina, Brazil. Non-type specimens: AZUSC 1616 (9, 250– 291 mm SL), 24º34’50” S / 46º39’01” W, Parcel Pedro II, Praia Grande, S„o Paulo, Brazil; AZUSC 1698 (5, 244– 285 mm SL), 25º15’45” S / 47º39’55” W, Ilha do Bom Abrigo, Cananéia, S „o Paulo, Brazil; AZUSC 6917.1 (15, 148– 217 mm SL, male), 26º15’23” S / 48º01’58” W, barra de São Francisco, S„o Francisco do Sul, Santa Catarina, Brazil; AZUSC 6685 (1, 255 mm SL), 26º38’16” S / 48º37’43” W, barra de Barra Velha, Barra Velha, Santa Catarina, Brazil. Diagnosis. Ophidion zavalai n. sp. is distinguished from its western Atlantic congeners as follows: from O. antipholus by the rear of skull not outlined in dark pigment (Fig. 5) (vs. rear of skull broadly outlined in dark pigment); lacking a dark streak along the lateral line (vs. may have a dark streak along the lateral line, Fig. 5); from O. dromio by having 63–65 vertebrae, rarely 67 or 68 (vs. 67–69, Table 3); 14–16 caudal vertebrae (vs. 17, rarely 16, Table 3); from O. grayi by lacking irregular rows on body (vs. 2 irregular rows of dark brown spots, upper row from top of body to lateral line, lower row from lateral line down, Fig. 5); lacking dark brown spots on head and dorsal fin (vs. top of head and dorsal fin with scattered dark brown spots, Fig. 5); from O. guianense by rocker bone present (vs. absent, Fig. 6); 7.9–11.4 body height at center of the eye (vs. 13–14, Table 3); from O. josephi by having 107–135 dorsal-fin rays (vs. 138–146, Table 3), 88–107 anal-fin rays (vs. 114–121, Table 3), lacking dark spots on body (vs. 3 rows of dark spots along body, Fig. 5); from O. lagochila by having 2 upper gill rakers (vs. 3), rear of top of head not outlined in black (vs. outlined in black, Fig. 5), juvenile without a dark streak along the lateral line (vs. with a dark streak along the lateral line, Fig. 5); from O. marginatum by having 107–135 dorsal-fin rays (vs. 147–158, Table 3), 88–107 anal-fin rays (vs. 118–124, Table 3), body and lateral line without dark stripes (vs. with 2–3 dark stripes along its body and one at lateral line, Fig. 5); from O. puck by having 63–68 vertebrae (vs. 70, Table 3), 14–16 preanal vertebrae (vs. 17–18, Table 3); from O. robinsi by lacking dark brown spots (vs. with dark brown spots, Fig. 5); from O. selenops by having 88–107 anal-fin rays (vs. 123–129, Table 3), lacking a dark line along top of back before dorsal fin (vs. with a dark line along top of back before dorsal fin, Fig. 5); from O. holbrookii by caudal-fin length 0.7–2.8 (vs. 2.9–7.7, Table 3); postorbital length 8.4–11.2 (vs. 11.7–15.3, Table 3) in specimens less than 230 mm SL. Description. Meristic and morphometric characters are shown in Table 3. Body moderately compressed, elongate with tapering tail. Head moderately broad, not depressed, bones firm in touch. Scales elongate, not overlapping, present on flanks and belly, 32 to 40 scales transversely across trunk above beginning of anal fin, 152 to 230 scales in transversal rows from pectoral base to caudal fin base. Pectoral fin pointed, shorter than postorbital distance; pelvic fin origin below anterior margin of orbit to mid-eye, longest ray not reaching pectoral fin base. Head without scales, except on predorsal region to supratemporal canal (about 27–41 predorsal scales). Anterior nostril a small tube over upper lip, posterior nostril slender, oval, over horizontal imaginary line through mid-eye. Snout rounded to slightly pointed, roughly equal to eye diameter. Eye large, oval, slightly elongated, lens well developed. Upper jaw ends just below mid-eye to posterior margin of orbit, rear upper jaw border partially covered by infraorbital regions in young specimens, exposed in adults; 3–4 rows of villiform teeth on jaws, larger on outer series, 3–4 rows of villiform teeth on vomer, palatines long with 3 rows of villiform teeth. Opercular and nasal spines absent. Upper branch of anterior gill arch with 2 knob–like rakers, lower branch with 4–7 short rakers. Head canals: supraorbital 2, infraorbital 3, supratemporal 4, preopercle to lower jaw 5. Rocker bone with sickle to semicircular, half-doughnut shape, dorsal profile semicircular, ventral profile with anterior arm moderately long, slightly triangular to rounded, rear arm with ventral margin truncate to slightly indented. Concavity between arms gentle to semicircular, more so on largest specimens. Swimbladder carrot-like, pointed on rear portion, with two small, fringed anterior horns, sulcus present on ventral portion. Rocker bone abuts anterior portion of the bladder. Color in life. Dark brown to tan on dorsum and flanks, whitish or light tan on belly, head tan, with whitish or silvery hue on opercle, eye lens dark, cornea dark brown with some golden hue. Pectoral fin dusky, pelvic fin whitish, dorsal and anal fins dusky with narrow black margin. Younger specimens pale or more yellow in ground color than adults. Morphological variation. Significant developmental morphological variation was identified in two meristic and six morphometric characters in specimens ranging from 148 to 325 mm SL (Table 3). Smaller specimens have fewer dorsal-fin and anal-fin rays (Table 3); a shorter postorbital distance (Fig. 3e), a shorter outer and inner pelvic ray (Figs. 3f,g), shallower body at dorsal- and anal-fin origin (Figs. 3h,i), and a narrower interorbital distance. In addition, head shape and coloration vary between young and adult specimens (Fig. 5, see Color in life). Females exhibit a cylindrical gas bladder, posterior portion narrow, anterior portion with 2 small lateral chambers, this condition does not vary during ontogenetic development (Fig. 6), they also lack of rocker bone and parapophyses modifications that compose sonic apparatus (Fig. 6). Adult males present a posterior portion of gas bladder distended and narrow (Fig. 6), a differentiated chamber in anterior portion of gas bladder articulating with the reniform rocker bone (shape of a human kidney) (Fig. 6), and parapophyses modified at first neural spine (Fig. 6). n males, the rocker bone and parapophyses exhibit evident ontogenetic differentiation, being reduced at juvenile stages when the rocker bone is less evident in lateral view radiographs and differentiated from parapophyses which are clearly distinguishable (Fig. 6). Sexual dimorphism. In a sample of 15 females (158–250 mm SL) and 35 males (148–296 mm SL), we observed a difference in the pattern related to dorsal outline between males and adult females (rectilinear and steeply inclined vs slightly rounded, Fig. 5). Examinations of gas bladder and x-ray reveal that males possess a rocker bone and modifications to the anterior vertebral parapophyses (Fig. 6). Distribution and habitat. Ophidion zavalai is found on soft bottom associated coastal marine habitats. It occurs in the western South Atlantic, off the eastern to southeastern coast of Brazil (Fig. 4), Etymology. The new species is named in honor of Dr. Luis Alberto Zavala-Camin, researcher at the Instituto de Pesca, S„o Paulo, Brazil, and professor at the Universidade Santa Cecília in Santos, Brazil, for his great contribution to the knowledge of marine fishes in Brazil. Remarks. The description of O. zavalai is based on the knowledge and account of variation associated with ontogenetic and sexual development, yet not described for gas bladder and sonic apparatus (Lea & Robins 2003). Also, structures used by males for sound production and display for females (Rose 1961; Marshal 1967; Courtenay 1971; Parmentier et al. 2006; Nguyen et al. 2008; Zavala-Camin & Rotundo 2011) are likely important in species recognition. The gap observed between the distribution of O. holbrookii and O. zavalai (Fig. 4) is supported by an extensive search of zoological collections and literature of the northeastern coast of Brazil. Material from Brazilian collections (Universidade Federal da Paraíba; Universidade Federal de Alagoas) regarding Ophidion were either based on identifications or specimens deposited were too small to be properly identified. Personal communication with colleagues (Dr. Claudio Sampaio, UFAL) support absence of these taxa from northeastern Brazil. Also, there is no mention of this species taxa in most of the known literature for the region (Eskinazi 1972; Araújo et al. 2004; Nobrega et al. 2009).A presumable report of Ophidion (as Ophidion cf. holbrookii) from Northeastern Brazil (Bacia Potiguar) has been made by Garcia Jr. et al. (2010, 2015), but regardless of the similarity of specimen depicted to O. zavalai, the description is rather short and with no meristic data, and material of this specimen is not available. Specimens from this area previously identified as O. holbrookii (MNHN 00005772) were found to be Lepophidium. Thus, the occurrence of Ophidion holbrookii and O. zavalai in northeastern Brazil has yet to be confirmed.

Rotundo, Matheus Marcos, Caires, Rodrigo Antunes, Oliveira, Claudio, Kuranaka, Mariana, Figueiredo-Filho, Jessé Miranda De, Marceniuk, Alexandre Pires (2023): Taxonomic revision of the Ophidion holbrookii Putnam, 1874 (Ophidiiformes: Ophidiidae) species complex, with description of a new species from Brazil. Zootaxa 5318 (2): 237-252, DOI: 10.11646/zootaxa.5318.2.4, URL: http://dx.doi.org/10.11646/zootaxa.5318.2.4

Ophidion holbrookii Putnam, 1874 Figs. 1–4 and Table 3 Ophidium holbrookii Putnam, 1874: 340 (original description; United States, Florida, Key West; holotype unknown). Ophidion holbrookii. — Jordan & Gilbert, 1883: 793 (fish synopsis; listed).— Rose 1961: 280–308 (anatomy of swim bladder and vertebrae; sexual dimorphism).— Cohen & Nielsen 1978: 16 (NOAA Technical Reports: listed).— Robins & Ray, 1986: 99 (Atlantic coast fishes; plate 16; short description).—Valdez & Aguillera, 1987: 62 (Fishes of Venezuela; short description and illustration; perla).–Gomez-Soto & Contreras—Balderas, 1988: 12 (fauna del laguna Madre de Tamaulipas, Mexico, first record; listed).– Smith, 1997: 361 (short description; distribution).— McEachran & Fechhelm, 1998: 748 (Fishes of Gulf of Mexico; description; drawing).— Castro-Aguirre et al. (1999): 164 (fishes of Mexico; listed). — Nielsen et al., 1999: 41 (FAO Fisheries Synopses; short description; illustration; senior synonymy of O. beani Jordan & Gilbert, 1883).— Leopold, 2004: 87 (Fishes of Guyana; brotule de banc; short description).— Nielsen & Robins, 2002: 972 (FAO Species Identification Guide, Living Marine Species of Western Central Atlantic; diagnosis; short description; drawing; senior synonymy of O. beani Jordan & Gilbert, 1883). — Lea & Robins, 2003: 7 (description of four new western Atlantic Ophidion species; comparison). —Garrido-Linares, & Acero. 2006: 297 (Ophidiiformes of Colombian Caribbean; listed).— Caires et al. in Marceniuk et al., 2021: 198 (Fishes of Northern Brazilian coast; description; photograph). Ophidium graellsi (not of Poey, 1861).— Jordan & Gilbert, 1882: 301 (fishes from Pensacola).— Jordan & Gilbert 1883: 953 (fish synopsis; listed). Ophidium beani Jordan & Gilbert, 1883: 143 (original description; type locality, United States, Florida, off Pensacola; Syntypes: USNM 30868).— Jordan, 1885: 126 (Catalog of fishes on North America; listed).— Jordan & Evermann, 1898: 2487 (fishes of North and Middle America; description). Ophidion sp. —Arai in Uyeno et al., 1983: 223 (Fishes trawled off Suriname and French Guiana; description; photograph). — Lea & Robins, 2003: 7 (comment of Arai’s image; re-identified as O. holbrookii). Holotype: MCZ 34010, formerly BSNH 1117, (138 mm SL), Key West, Florida, U.S.A., 1872, coll. C.J. Maynard. Non type specimens. United States: UF 124058 (1, 188.6 mm SL, female), 34°6’0.000” N / 77°46’12.000” W, North Carolina, 02 Sep. 1959, coll. R / V Silver Bay; UF 28320 (1, 200.2 mm SL), 33°27’36.000” N / 78°6’36.000” W, North Carolina, 22 Jun. 1968, coll. J. Gillikin et al.; UF 180111 (1, 159.0 mm SL, female), 31°10’48.000” N / 81°20’60.000” W, Georgia, 30 May 1961, coll. F. Blossoms; UF 37849 (7, 269.3, 247.2, 243.4 mm SL, all male, 229.3, 222.9, 199.2, 205.6 mm SL, all female), 29°32’24.000” N / 87°39’36.000” W, Alabama, 30 Jun. 1983, coll. G. Burgess; CAS 216613 (4), 30°15’57.600” N / 81°15’0.000” W, Florida, 29 Oct. 1973, coll. R / V Dolphin; UF 37487 (1, 131 mm SL), 24°37’60.000” N / 82°28’0.000” W, Florida, 18 Apr 1980, coll. G. Burgess; UF 185151 (1, 207.4 mm SL, female), Gulf of Mexico, Florida, 29 Jan. 1963, no collection data. Mexico: UF 232191 (1, 246.4 mm SL, male), 22°15’36.000” N / 89°2’60.000” W, Yucatan, 13 Dec. 1952, coll. R / V Oregon. Puerto Rico: UF 221855 (1, 214.7 mm SL, male), 18°6’0.000” N / 67°22’48.000” W. Colombia: UF 221832 (9, 218.4 mm SL, male, 192.5, 193.8, 160.8, 171.5, 190.8 mm SL, all female, 186.7, 173.9, 177.3, unsexed), 11°57’36.000” N / 72°28’48.000” W. Guyana: UF 205082 (1, 191.5 mm SL, female), 7°39’36.000” N / 57°33’36.000” W, 31 Aug. 1958, coll. R / V Oregon. Suriname: UF 167743 (1, 198.8 mm SL, male), 6°57’36.000” N / 55°52’48.000” W, 26 Jun. 1972, coll. R / V Oregon II; UF 232193 (1, 205.1 mm SL, male), 6°55’48.000” N / 56°18’36.000” W, 29 Jan. 1974, coll. R / V Oregon II. Diagnosis. Ophidion holbrookii is distinguished from its western Atlantic congeners as follows: from O. antipholus by the rear of skull not outlined in dark pigment (Fig. 2) (vs. rear of skull broadly outlined in dark pigment), the absence of a dark streak along the lateral line (Fig. 2) (vs. may have a dark streak along the lateral line); from O. dromio by having 59–66 vertebrae (vs. 67–69, Table 3); from O. grayi by the absence of irregular rows on body (vs. 2 irregular rows of dark brown spots, upper row from top of the body to lateral line, lower row from lateral line down, Fig. 2), the absence of dark brown spots on head and dorsal fin (vs. top of head and dorsal fin with scattered dark brown spots; from O. guianense by rocker bone present (vs. absent); 97–107 anal-fin rays, rarely 96 (vs. 91–96, Table 3), 4 rakers on lower gill arch (vs. 5–6, Table 3); from O. josephi by having 116–137 dorsal-fin rays (vs. 138–146, Table 3), 96–107 anal-fin rays (vs. 114–121, Table 2), without dark spots on body (Fig. 2) (vs. 3 rows of dark spots along body); from O. lagochila by having 2 rudiments on upper gill arch (vs. 3), rear of top of the head not outlined in black (Fig. 2) (vs. outlined in black), juvenile without a dark streak along the lateral line (vs. with a dark streak along the lateral line); from O. marginatum by having 116–137 dorsal-fin rays (vs. 147–158, Table 3), 96–107 anal-fin rays (vs. 118–124, Table 3), body and lateral line without dark stripes (Fig. 2) (vs. with 2–3 dark stripes along its body and one at lateral line); from O. puck by having 59–66 vertebrae (vs. 70, Table 3), 13–16 preanal vertebrae (vs. 17–18, Table 3); from O. robinsi by lacking dark brown spots (Fig. 2) (vs. with dark brown spots); from O. selenops by having 96–107 anal-fin rays (vs. 123–129, Table 3), the absence of a dark line along top of back before dorsal fin (Fig. 2) (vs. with a dark line along top of back before dorsal fin); from O. zavalai n. sp. by caudal-fin length 2.9–7.7 (vs. 0.7–2.8, Table 3); and 11.7–15.3 of postorbital length (vs. 8.4–11.2, Table 3) in specimens less than 230 mm SL. Description. Meristic and morphometric characters given in Table 3. Body moderately compressed, elongate with tapering tail. Head moderately broad, not depressed, bones firm in touch. Scales elongate, not overlapping, present on flanks and belly, about 30 to 40 scales transversely across trunk above beginning of anal fin, about 150 to 170 scales in transversal rows from pectoral base to caudal fin base. Pectoral fin variably long and pointed, usually shorter than postorbital distance; pelvic-fin origin below orbit anterior margin, longest ray not reaching pectoral-fin base. Head without scales, except on predorsal region to supratemporal canal. Anterior nostril as small tube over upper lip, posterior nostril slender, oval, over horizontal imaginary line through mid-eye. Snout triangular, slightly shorter than eye diameter. Eye large, oval, slightly elongated, lens well developed. Upper jaw ends just below mid-eye to posterior margin of orbit, rear upper jaw border partially covered by infraorbital regions in young specimens, exposed in adults; 3–4 rows of villiform teeth on jaws, larger on outer series, 3–4 rows of villiform teeth on vomer, palatines long with 3 rows of villiform teeth. Opercular and nasal spines absent. Upper branch of first gill arch with 2 knob-like rakers, lower branch with 4–7 short rakers. Head canals: supraorbital 2, infraorbital 3, supratemporal 4, preopercle to lower jaw 5. Rocker bone with sickle to semicircular, half-doughnut shape, dorsal profile semicircular, ventral profile with anterior arm triangular or with ventral margin slightly blunt, rear arm with ventral margin rounded to truncate or slightly indented. Concavity between arms gentle to semicircular, more so on largest specimens. Swim bladder carrot-like, pointed on rear portion, with two small, fringed anterior horns, sulcus present on ventral portion. Rocker bone abuts anterior portion of the bladder. Color in preserved specimens. Tan to brownish on upper third and head, beige to light tan elsewhere, paler on belly, sometimes darker along lateral line and anal fin base; eye lens dark, cornea dusky, darker on upper margin. Pectoral fin dusky or yellow, pelvic fin dirty whitish or dusky, dorsal and anal fins dusky with slender black distal margins, sometimes with dark area following distal lines in caudal fin base. Young specimens lighter in color, with dark areas less evident. Specimens may be light tan due to preservation. Morphological variation. No significant variation was found in the development of Ophidion holbrookii from 131 to 269 mm SL (Table 3). Sexual dimorphism. Sexual dimorphism has been observed in the dorsal profile of the head in adult males and females of O. holbrookii. Males exhibit a straight, strongly inclined dorsal profile, whereas in females this profile is slightly rounded (see Fig. 2). Radiographs of examined specimens show variation for the rocker bone and parapophyses of anterior vertebrae described for males and females of O. zavalai n. sp. (see Zavala-Camin & Rotundo 2011), as described by Rose (1961) in specimens captured by shrimp trawlers in the Dry Tortugas west of Key West, Florida (USA). Distribution and habitat. Ophidion holbrookii is found in soft bottom associated coastal marine habitats. It can be found from the east coast of the United States to Suriname, including throughout Gulf of Mexico and Caribbean Sea (Fig. 4). Remarks. Until recently, Catalog of Fishes stated that whereabouts of the holotype of O. holbrookii were unknown (Fricke et al. 2022). The confusion regarding the status of holotype arose from the fact that Putnam deposited the type specimen in Boston Society of Natural History (catalog number 1117), but this collection was later merged to Museum of Comparative Zoology, Harvard University (MCZ), a condition confirmed by A. Wilkinson, MCZ. The catalog number of the holotype of O. holbrookii is MCZ 34010, and the type specimen is in good condition (Fig. 2a), although its swim bladder and rocker bone were extracted by Putnam and later included with the specimen. Ophidion holbrookii has been diagnosed by being tan without markings, except for a dark edge to edge to the dorsal and anal fins, by the presence of the rocker bone, the form the gas bladder, 4 developed rakers on the lower gill arch and long pelvic rays, the longer of the two extending to a point under or nearly under the pectoral-fin base, however, some of the characters listed may apply to more than one species sensu Arai (1983) and Lea & Robins (2003). Additionally, Lea & Robins (2003) diagnosed O. holbrookii with dorsal profile from the tip of snout to tip of nape straight rather than convex, however, this trait shows sexual variation, common in adult males (Fig. 2). The synonymy of Ophidion holbrookii and Ophidion beani Jordan & Gilbert, 1883 is supported by the shape of a long gas bladder (Lea & Robins 2003)., Published as part of Rotundo, Matheus Marcos, Caires, Rodrigo Antunes, Oliveira, Claudio, Kuranaka, Mariana, Figueiredo-Filho, Jessé Miranda De & Marceniuk, Alexandre Pires, 2023, Taxonomic revision of the Ophidion holbrookii Putnam, 1874 (Ophidiiformes: Ophidiidae) species complex, with description of a new species from Brazil, pp. 237-252 in Zootaxa 5318 (2) on pages 240-244, DOI: 10.11646/zootaxa.5318.2.4, http://zenodo.org/record/8169565, {"references":["Putnam, F. W. (1874) Notes on Ophidiidae and Fierasferidae, with description of new species from America and the Mediterranean. Proceedings of the Boston Society of Natural History, 16, 339 - 348.","Jordan, D. S. & Gilbert, C. H. (1883) Synopsis of the fishes of North America. 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Bony fishes part 1 (Acipenseridae to Grammatidae). FAO Species Identification Guide for Fishery Purposes and American Society of Ichthyologists and Herpetologists Special Publication No. 5. FAO, Rome, pp. 965 - 972.","Lea, R. N. & Robins, C. R. (2003) Four New Species of the Genus Ophidion (Pisces: Ophidiidae) from the Western Atlantic Ocean. Natural History Museum, The University of Kansas, Scientific Papers, 31, 1 - 9. https: // doi. org / 10.5962 / bhl. title. 8466","Jordan, D. S. (1885) A catalogue of the fishes known to inhabit the waters of North America, north of the Tropic of Cancer, with notes on species discovered in 1883 and 1884. United States Commission of Fish and Fisheries, Report of the Commissioner, 13, 789 - 973. https: // doi. org / 10.5962 / bhl. title. 8874","Jordan, D. S. & Evermann, B. W (1898) The fishes of North and Middle America: a descriptive catalogue of the species of fishlike vertebrates found in the waters of North America north of the Isthmus of Panama. Part III. Bulletin of the United States National Museum, 47, 2183 - 3136. https: // doi. org / 10.5962 / bhl. title. 39717","Uyeno, T., Matsuura, K. & Fujii, E. (1983) Fishes trawled off Suriname and French Guiana. Japan Marine Fishery Resource Research Center, Tokyo, 519 pp.","Zavala-Camin, L. A. & Rotundo, M. M. (2011) Gross anatomy of the head lateral line and hearing system of the Ophidiinae, Genypterus blacodes, Raneya brasiliensis and Ophidion holbrookii, of Southern Brazil. Bioikos, 25 (1), 3 - 10.","Fricke, R., Eschmeyer, W. N. & van der Laan R. (2022) Eschmeyer's catalog of fishes: genera, species, references [Internet]. San Francisco: California Academy of Science. Available from: http: // researcharchive. calacademy. org / research / ichthyology / catalog / fishcatmain. asp (accessed 27 June 2023)"]}

The rare deepsea ophidiid genus Leucicorus was described by Garman (1899) based on L. lusciosus Garman, 1899 caught in the East Pacific. Until 1973 only three additional specimens were caught of which two from the East Pacific belong to L. lusciosus and one from off Hawaiʽi is an undescribed species here described as L. lentibus n. sp. In 1973 a Soviet expedition to the Caribbean Sea trawled 18 specimens from abyssal and hadal depths and based on this material a second Leucicorus species was described, L. atlanticus Nielsen, 1975. Since then eight Leucicorus specimens from the Indo-Pacific and Atlantic Oceans have been caught of which two from the West Atlantic belong to a new species, L. gerringerae n. sp., herein described, four to L. atlanticus and two remain as Leucicorus sp. About 35 demersal Leucicorus specimens have been observed and photographed by ROV (remotely operated vehicle) at depths of 3804-5768 meters in the Pacific Ocean.

“aff. Glyptophidium ” stringeri sp. nov. urn:lsid:zoobank.org:act: B65B9A77-A733-41EB-9AEA-61F744665C4B Fig. 17G–N Diagnosis OL/OH = 1.11–1.32. Otoliths with a large protruding antero-dorsal expansion and a pointed posterior rim. Sulcus wide, oblong, and not divided into ostium and cauda; it is filled with a large colliculum. Etymology This species is dedicated to Gary L. Stringer (University of Louisiana at Monroe) for his many contributions to the knowledge of fossil otoliths from the Gulf Coast area. Material examined Holotype UNITED STATED OF AMERIA • Right otolith; Texas, Alabama Ferry; “ Stone City beds ”; Fig. 17G; IRSNB P 10066. Paratypes UNITED STATES OF AMERICA • 221 otoliths, of which five are figured: Fig. 17H–L; same collection data as for holotype; IRSNB P 10067–10071 • 543 otoliths, of which two are figured: Fig. 17M–N; Texas, Little Brazos River, Stenzel loc.; Wheelock Member; IRSNB P 10072–10073 • 756 otoliths; same locality as for preceding; Landrum Shale Member. Type locality and horizon United States of America, Alabama Ferry (Texas), “Stone City beds”. Dimensions of the holotype Length = 2.75 mm; height = 2.23 mm; thickness = 0.59 mm. Description The otoliths of this species are especially characterized by a large protruding antero-dorsal expansion, which is triangular and pointing upward, and by their acuminated posterior end. The anterior rim of the dorsal expansion can be smooth or undulated. The anterior rim of the otolith is almost vertical, forming a blunt rostrum. The posterior part of the otolith shows a triangular shape; it tapers strongly after a well-marked postero-dorsal angle. The ventral rim is smoothly curved and its deepest part is located in the middle of the otolith. This rim may be crenulate in smaller specimens (Fig. 17L). The inner face is slightly convex; the outer face is nearly flat, but may be slightly convex anteriorly (Fig. 17G). A long and wide sulcus occupies nearly the entire length of the inner face. It is located in the middle of that face, and is completely filled by a single colliculum. This sulcus is not clearly divided into ostial and caudal parts. The cristae are well-developed but not ridge-like. The dorsal and the ventral areas are almost equal in size. Remarks The otoliths of this new species are somewhat similar looking to those of the co-occurring Hoplobrotula melrosensis (Dante & Frizzell, 1965) and also to those of Bauzaia mucronata (Koken, 1888), but this similarity is only superficial and restricted to the outline of the otoliths. Firstly, the sulcus of both H. melrosensis and B. mucronata are clearly divided into an ostium and cauda, and secondly, their posterior end is much more extended and spine-like. The sulcus of “aff. G.” stringeri sp. nov. is straight, and the posterior tip is located in their lower part. Although the posterior part of the otoliths resembles that of the extant genus Glyptophidium, this genus has a sulcus type with distinct ostium and cauda. The lack of a clear division between ostium and cauda in the sulcus of “aff. G.” stringeri sp. nov. suggests affinities to Glyptophidium, but does not imply a certain attribution to that genus. Two related Eocene species with similar features are “aff. G.” biarritzense (Sulc, 1932) from the Aquitaine Basin and “aff. G.” pseudobiarritzense Nolf & Bajpai, 1992 from Java (Nolf & Bajpai 1992: pl. 3 fig. 11). Stratigraphic and geographic distribution Lutetian: “Stone City beds”, Texas; Dobys Bluff Tongue of Kosciusko Formation, Mississippi. Bartonian: Landrum Member of Cook Mountain Formation, Texas; “upper” Lisbon Formation, Alabama., Published as part of Lin, Chien-Hsiang & Nolf, Dirk, 2022, Middle and late Eocene fish otoliths from the eastern and southern USA, pp. 1-122 in European Journal of Taxonomy 814 on pages 52-54, DOI: 10.5852/ejt.2022.814.1745, http://zenodo.org/record/6484355, {"references":["Frizzell D. L. 1965. Otolith-based genera and lineages of fossil bonefishes (Clupeiformes, Albulidae). Senckenbergiana Lethaea 46 a: 85 - 110.","Koken E. 1888. Neue Untersuchungen an tertiaren Fisch-Otolithen. Zeitschrift der deutschen geologischen Gesellschaft 40: 274 - 305.","Nolf D. & Bajpai S. 1992. Marine middle Eocene fish otoliths from India and Java. Bulletin de l'Institut royal des Sciences naturelles de Belgique, Sciences de la Terre 62: 195 - 221."]}

Aequalobythites aequaloides (Nolf & Stringer, 2003) Fig. 38L–O “genus Neobythitinarum ” aequaloides Nolf & Stringer, 2003: 7, pl. 3 figs 11–14. “ Neobythitina ” aequaloides – Nolf 2013: 68, pl. 134. Remarks Otoliths of this species are only known by specimens from the Yazoo Clay (Priabonian) of Louisiana. They are closely related to those of the species originally described as Lepophidium aequalis Stinton & Nolf, 1970 from the Belgian Lede Formation (Lutetian), that Schwarzhans (1981b) placed in the extinct genus Aequalobythites. Other related species are A. regularis (Priem, 1911) from the Lutetian and Bartonian of the Paris Basin, Belgium and southern England, and A. hilgendorfi (Koken, 1891) from the German lower Oligocene. Stratigraphic and geographic distribution Priabonian: Yazoo Clay, Louisiana., Published as part of Lin, Chien-Hsiang & Nolf, Dirk, 2022, Middle and late Eocene fish otoliths from the eastern and southern USA, pp. 1-122 in European Journal of Taxonomy 814 on page 52, DOI: 10.5852/ejt.2022.814.1745, http://zenodo.org/record/6484355, {"references":["Nolf D. & Stringer G. L. 2003. Late Eocene (Priabonian) fish otoliths from the Yazoo Clay at Copenhagen, Louisiana. Louisiana Geological Survey Geological Pamphlet 13: 1 - 23.","Nolf D. 2013. The Diversity of Fish Otoliths, Past and Present. Royal Belgian Institute of Natural Sciences, Brussels.","Stinton F. C. & Nolf D. 1970. A teleost otolith fauna from the Sands of Lede, Belgium. Bulletin de la Societe belge de Geologie, de Paleontologie et d'Hydrologie 78: 219 - 234.","Schwarzhans W. 1981 b. Vergleichende morphologische Untersuchungen an rezenten und fossilen Otolithen der Ordnung Ophidiiformes. Berliner geowissenschaftliche Abhandlungen A 32: 63 - 122."]}

Genus Preophidion Frizzell & Dante, 1965 Remarks Preophidion is an extinct genus with otoliths resembling those of the extant Sirembo. We here place P. elevatus (Koken, 1888), P. granus (Müller, 1999), and P. petropolis (Dante & Frizzell, 1965) in this genus. “ Neobythites ” meyeri and “ N.” stintoni were initially also assigned to Preophidion by Frizzell & Dante (1965), and this was followed as such in Ebersole et al. (2019), but here we reassign the former two species to the Symmetrosulcus (see above). The most significant distinctive feature between otoliths of Symmetrusulcus and Preophidion consists in the cleavage between the ostial and caudal collicula, which is vertical and short in otoliths of Symmetrosulcus, whereas it is more oblique and inclined in an anterodorsal-posteroventral direction in the more Sirembo -like otoliths of Preophidion. Additionally, Sirembo -like otoliths usually have a wider sulcus, a sharply restricted collum on the crista inferior, a more considerable thickness and a more convex inner face., Published as part of Lin, Chien-Hsiang & Nolf, Dirk, 2022, Middle and late Eocene fish otoliths from the eastern and southern USA, pp. 1-122 in European Journal of Taxonomy 814 on page 58, DOI: 10.5852/ejt.2022.814.1745, http://zenodo.org/record/6484355, {"references":["Frizzell D. L. & Dante J. H. 1965. Otoliths of some early Cenozoic fishes of the Gulf Coast. Journal of Paleontology 39: 687 - 718.","Koken E. 1888. Neue Untersuchungen an tertiaren Fisch-Otolithen. Zeitschrift der deutschen geologischen Gesellschaft 40: 274 - 305.","Muller A. 1999. Ichthyofaunen aus dem atlantischen Tertiar der USA. Leipziger Geowissenschaften 9 / 10: 1 - 360.","Frizzell D. L. 1965. Otolith-based genera and lineages of fossil bonefishes (Clupeiformes, Albulidae). Senckenbergiana Lethaea 46 a: 85 - 110.","Ebersole J. A., Cicimurri D. J. & Stringer G. L. 2019. Taxonomy and biostratigraphy of the elasmobranchs and bony fishes (Chondrichthyes and Osteichthyes) of the lower - to - middle Eocene (Ypresian to Bartonian) Claiborne Group in Alabama, USA, including an analysis of otoliths. European Journal of"]}

Signata nicoli Dante & Frizzell, 1965 Fig. 38I Signata nicoli Dante & Frizzell in Frizzell & Dante, 1965: 709, pl. 88 figs 3–4, 21, 25. Signata nicoli – Schwarzhans 1981b: 113. — Ebersole et al. 2019: 200, fig. 70i–j. “genus Ophidiidarum ” nicoli – Nolf 1985: 67. “ Ophidiida ” nicoli – Nolf 2013: 65, pl. 124. Remarks Signata represents yet another extinct taxon within Ophidiidae. The otoliths have a very wide sulcus that is filled only with a very flat, undivided colliculum. This sulcus has a somewhat Sirembo -like aspect, but the stronger convexity of the inner face and the more extended and thinner posterior part are never seen in Sirembo or any other related otolith type. Also, in otoliths of Signata, the areas above and below the sulcus are very restricted. Otoliths of Signata are very well characterized by autapomorphic features, but do not provide any indication for affinities with other ophidiid groups. Two species are included in the genus: S. nicoli and S. stenzeli. The holotype of S. nicoli has smoothly curved cristae, with just an angle on the crista superior, while the otoliths of S. stenzeli have a more elevated angle on the crista superior and two notable undulations in their crista inferior. The holotype of S. nicoli is from the Weches Formation (Lutetian) of Texas, but we did not find additional specimens among our present collection. Ebersole et al. (2019), however, reported a specimen from the “upper” Lisbon Formation (Bartonian) of Alabama. Stratigraphic and geographic distribution Lutetian: Weches Formation, Texas. Bartonian: “upper” Lisbon Formation, Alabama.

Hoplobrotula melrosensis (Dante & Frizzell, 1965) Fig. 16A–E Bauzaia melrosensis Dante & Frizzell in Frizzell & Dante, 1965: 711, pl. 86 figs 7, 9. Hoplobrotula melrosensis – Schwarzhans 1981b: 113. — Nolf 1985: 66; 2013: 67, pl. 132. — Ebersole et al. 2019: 200, fig. 70k–l. Remarks Hoplobrotula melrosensis has a sulcus with a somewhat downward oriented cauda, a feature typical of the extant Hoplobrotula. Stratigraphic and geographic distribution Lutetian: Weches Formation and “Stone City beds”, Texas; “lower” Lisbon Formation, Alabama. Bartonian: “upper” Lisbon Formation, Alabama., Published as part of Lin, Chien-Hsiang & Nolf, Dirk, 2022, Middle and late Eocene fish otoliths from the eastern and southern USA, pp. 1-122 in European Journal of Taxonomy 814 on page 54, DOI: 10.5852/ejt.2022.814.1745, http://zenodo.org/record/6484355, {"references":["Frizzell D. L. 1965. Otolith-based genera and lineages of fossil bonefishes (Clupeiformes, Albulidae). Senckenbergiana Lethaea 46 a: 85 - 110.","Frizzell D. L. & Dante J. H. 1965. Otoliths of some early Cenozoic fishes of the Gulf Coast. Journal of Paleontology 39: 687 - 718.","Schwarzhans W. 1981 b. Vergleichende morphologische Untersuchungen an rezenten und fossilen Otolithen der Ordnung Ophidiiformes. Berliner geowissenschaftliche Abhandlungen A 32: 63 - 122.","Nolf D. 1985. Otolithi Piscium. In: Schultze H. P. (ed.) Handbook of Palaeoichthyology, Volume 10: 1 - 145. Gustav Fischer Verlag, Stuttgart.","Nolf D. 2013. The Diversity of Fish Otoliths, Past and Present. Royal Belgian Institute of Natural Sciences, Brussels.","Ebersole J. A., Cicimurri D. J. & Stringer G. L. 2019. Taxonomy and biostratigraphy of the elasmobranchs and bony fishes (Chondrichthyes and Osteichthyes) of the lower - to - middle Eocene (Ypresian to Bartonian) Claiborne Group in Alabama, USA, including an analysis of otoliths. European Journal of"]}

Genus incertae sedis Remarks “ Neobythites ” rotundus (Müller, 1999) and “ N.” virginicus Müller, 1999, two species that are known from the Atlantic Coastal Plain only (Piney Point Formation), are incertae sedis Neobythitinae that are not clairly related to other living or fossil genera of Neobythitinae. Otoliths of “ Neobythites ” virginicus have a deep ventral area and a small but well-marked antero-dorsal angle. Those of “ N. ” rotundus are easily recognizable by their height which is more considerable than their length, their important thickness, which is essentially located in their posterior portion, and by a salient antero-dorsal angle., Published as part of Lin, Chien-Hsiang & Nolf, Dirk, 2022, Middle and late Eocene fish otoliths from the eastern and southern USA, pp. 1-122 in European Journal of Taxonomy 814 on page 57, DOI: 10.5852/ejt.2022.814.1745, http://zenodo.org/record/6484355, {"references":["Muller A. 1999. Ichthyofaunen aus dem atlantischen Tertiar der USA. Leipziger Geowissenschaften 9 / 10: 1 - 360."]}

“ Neobythites ” virginicus Müller, 1999 Fig. 18L–M “genus Neobythitinarum ” virginicus Müller, 1999: 126, fig. 28/21–23. “ Neobythitina ” virginica – Nolf 2013: 69, pl. 138. Remarks See remarks under the genus. Stratigraphic and geographic distribution Lutetian: Piney Point Formation, Virginia., Published as part of Lin, Chien-Hsiang & Nolf, Dirk, 2022, Middle and late Eocene fish otoliths from the eastern and southern USA, pp. 1-122 in European Journal of Taxonomy 814 on page 58, DOI: 10.5852/ejt.2022.814.1745, http://zenodo.org/record/6484355, {"references":["Muller A. 1999. Ichthyofaunen aus dem atlantischen Tertiar der USA. Leipziger Geowissenschaften 9 / 10: 1 - 360.","Nolf D. 2013. The Diversity of Fish Otoliths, Past and Present. Royal Belgian Institute of Natural Sciences, Brussels."]}

The fossil otoliths of the southern USA have been known for more than 130 years and are among the richest assemblages worldwide. However, previous studies are often scattered and with outdated systematic scheme. A collection of over 25000 otoliths ranging in age from the Lutetian to the Priabonian from 47 sites in five states in the eastern and southern USA is analysed here. Combined with the earlier described material, at least 101 otolith-based taxa are documented, of which 83 are identified at species level. Fourteen of these are introduced as new species: Elopothrissus bernardlemorti sp. nov., “Muraenesox” barrytownensis sp. nov., Pseudophichthys texanus sp. nov., Paraconger wechesensis sp. nov., Neoopisthopterus weltoni sp. nov., “aff. Glyptophidium” stringeri sp. nov., Symmetrosulcus dockeryi sp. nov., Mene garviei sp. nov., “Citharus” varians sp. nov., Waitakia beelzebub sp. nov., Astroscopus compactus sp. nov., Parascombrops yanceyi sp. nov., Anisotremus rambo sp. nov., and Pagellus pamunkeyensis sp. nov. The assemblages are distinct fom contemporary European faunas by the complete lack of mesopelagic fish otoliths, and by the presence of sciaenids. Dominant taxa in the American Eocene are the Ophidiidae, Sciaenidae, Lactariidae, and Congridae. They indicate shallow-water environments for all the sampled sites. The notable abundance of those taxa suggests that they could have a higher turnover rate, and provided fundamental nutrition in the local Paleogene marine ecosystem. Further analyses of the species in the stratigraphic succession revealed that a faunal turnover between the Claiborne and Jackson seas was evident in teleosts, and it might be more widespread in other marine organisms in the region.

Brotula aquitanica Nolf, 1980 Fig. 38K Brotula aquitanica Nolf, 1980: 104, pl. 15 fig. 14. Brotula aquitanica – Nolf 1985: 64; 1988: 59, pl. 6 figs 6–7; 2013: 65, pl. 124. — Nolf & Stringer 2003: 5, pl. 7 fig. 12. Remarks As already mentioned by Nolf & Stringer (2003), two specimens from the Priabonian Yazoo Clay of Louisiana look almost identical to those of Brotula aquitanica from the Ypresian Gan Clay in Aquitaine, SW France (see Nolf 2013: pl. 124 for an iconography of the specimens from Aquitaine). Stratigraphic and geographic distribution Bartonian: Moodys Branch Formation, Louisiana. Priabonian: Yazoo Clay, Louisiana., Published as part of Lin, Chien-Hsiang & Nolf, Dirk, 2022, Middle and late Eocene fish otoliths from the eastern and southern USA, pp. 1-122 in European Journal of Taxonomy 814 on pages 48-49, DOI: 10.5852/ejt.2022.814.1745, http://zenodo.org/record/6484355, {"references":["Nolf D. 1980. Etude monographique des otolithes des Ophidiiformes actuels et revision des especes fossiles (Pisces, Teleostei). Mededelingen van de Werkgroep voor Tertiaire en Kwartaire Geologie 17 (2): 71 - 195.","Nolf D. 1985. Otolithi Piscium. In: Schultze H. P. (ed.) Handbook of Palaeoichthyology, Volume 10: 1 - 145. Gustav Fischer Verlag, Stuttgart.","Nolf D. 1988. Les otolithes de teleosteens eocenes d'Aquitaine (sud-ouest de la France) et leur interet stratigraphique. Memoire de l'Academie royale de Belgique, Classe des Sciences, 4 °, 2 e serie, 19 (2): 1 - 147.","Nolf D. & Stringer G. L. 2003. Late Eocene (Priabonian) fish otoliths from the Yazoo Clay at Copenhagen, Louisiana. Louisiana Geological Survey Geological Pamphlet 13: 1 - 23.","Nolf D. 2013. The Diversity of Fish Otoliths, Past and Present. Royal Belgian Institute of Natural Sciences, Brussels."]}

Signata stenzeli Dante & Frizzell, 1965 Fig. 17A–F Signata stenzeli Dante & Frizzell in Frizzell & Dante, 1965: 709, pl. 88 figs 12, 16–17. Signata stenzeli – Schwarzhans 1981b: 113. — Ebersole et al. 2019: 199, fig. 70g –h. “genus Ophidiidarum ” stenzeli – Nolf 1985: 67. “ Ophidiida ” stenzeli – Nolf 2013: 65, pl. 124. Remarks See under S. nicoli for the distinction of both species of Signata. Stratigraphic and geographic distribution Lutetian: “Stone City beds”, Texas; Dobys Bluff Tongue of Kosciusko Formation, Mississippi. Bartonian: Cook Mountain Formation, Texas; “upper” Lisbon Formation, Alabama., Published as part of Lin, Chien-Hsiang & Nolf, Dirk, 2022, Middle and late Eocene fish otoliths from the eastern and southern USA, pp. 1-122 in European Journal of Taxonomy 814 on page 51, DOI: 10.5852/ejt.2022.814.1745, http://zenodo.org/record/6484355, {"references":["Frizzell D. L. 1965. Otolith-based genera and lineages of fossil bonefishes (Clupeiformes, Albulidae). Senckenbergiana Lethaea 46 a: 85 - 110.","Frizzell D. L. & Dante J. H. 1965. Otoliths of some early Cenozoic fishes of the Gulf Coast. Journal of Paleontology 39: 687 - 718.","Schwarzhans W. 1981 b. Vergleichende morphologische Untersuchungen an rezenten und fossilen Otolithen der Ordnung Ophidiiformes. Berliner geowissenschaftliche Abhandlungen A 32: 63 - 122.","Ebersole J. A., Cicimurri D. J. & Stringer G. L. 2019. Taxonomy and biostratigraphy of the elasmobranchs and bony fishes (Chondrichthyes and Osteichthyes) of the lower - to - middle Eocene (Ypresian to Bartonian) Claiborne Group in Alabama, USA, including an analysis of otoliths. European Journal of","Nolf D. 1985. Otolithi Piscium. In: Schultze H. P. (ed.) Handbook of Palaeoichthyology, Volume 10: 1 - 145. Gustav Fischer Verlag, Stuttgart.","Nolf D. 2013. The Diversity of Fish Otoliths, Past and Present. Royal Belgian Institute of Natural Sciences, Brussels."]}

Leucicorus atlanticus Nielsen, 1975 Table 1, Figs. 1A, 2C, 3A���D, 5 Leucicorus atlanticus Nielsen, 1975: 113, fig. 1. Leucicorus atlanticus: Hureau & Nielsen 1981: 16; Nielsen & M��ller 2008: 29. Material examined (22 specimens, SL 48���240 mm). Holotype: ZMUC P 771813, 154 mm SL, male, Cayman Trench, 19��00���N, 80��38���W, R / V Akademik Kurchatov, cr. 14, st. 1243, Sigsbee trawl, 6400���6800 m, 16. Mar. 1973. Paratypes (17 specimens): UF 230276 (former UMML 30276), 205 mm SL, female, Puerto Rico Trench, 22��02���N, 65��10���W, R / V Pillsbury, st. 1374, 41��� otter trawl, 5541���5563 m, 2 July 1971. IORAS 02295, 110���151 mm SL and ZMUC P 77575, 154 mm SL, female, Cayman Trench, 18��59���N, 80��30���W, R / V Akademik Kurchatov, cr. 14, st. 1242-A, Sigsbee trawl, 6800 m, 20 Mar. 1973. IORAS 7 specimens not catalogued (specimens missing), 48���139 mm SL and ZMUC P 77585, 125 mm SL, Cayman Trench, 19��05���N, 80��29���N, R / V Akademik Kurchatov, st. cr. 14, 1259, Sigsbee trawl, 6800��� 6500 m, 22 Mar. 1973. IORAS 02296, 2 specimens, 79���152 mm SL and ZMUC P77586, 78 mm SL, Cayman Trench, 19��44���N, 76��12���W, R / V Akademik Kurchatov, cr. 14, st. 1266, Sigsbee trawl, 5900���6300 m, 24 Mar. 1973. IORAS 02297,120 + mm SL, Cayman Trench, 19��38���N, 76��40���W, R / V Akademik Kurchatov, cr. 14, st. 1267 Sigsbee trawl, 6740���6760 m, 25 Mar. 1973. IORAS not catalogued (specimen missing), 135 mm SL, Yucatan Trench, 20��08���N, 84��30���W, R / V Akademik Kurchatov, cr. 14, st. 1272, Sigsby trawl, 4580 m, 28 Mar. 1973. Additional material: MNHN 1979���689, 224 mm SL, Northeast Atlantic Ocean, 10��46.23���N, 42��40.41���W, Campagne Biovema, st. B, benthic trawl, 5100 m, 19 Nov. 1977. VIMS 6857, 183 mm SL, female, Silver Abyssal Plain, 23��20���22.5���N, 68��23���29.4���W, R / V Columbus Iselin, cr. CI-8123, st. D018, 45��� OT, 5303���5387 m, 25 May 1981. BMNH 2020.11.26.2, 240 mm SL, female, Northeast Atlantic Ocean, 38 �� 00���N, 11��32.7���W, R / V Discovery, st. 11911#7, OTSB, 5035 m, 26 Sep. 1989. ZMUC P771571, 87 mm SL, unripe, Solomon Sea, 7��49.4���S, 156��02.8���E, Galathea 3 exp., st. 061228-01, 1.2 m Agassiz trawl, 4350���4450 m, 2 Dec. 2006. Diagnosis. Dorsal fin rays 82���94, much reduced squamation on preopercle and opercle and occiput almost naked; 18���20 scale rows on trunk above beginning of anal fin. Black eye comparatively small (10���13.5% HL), without or minute lens Comparison. L. atlanticus is most similar to L. gerringerae by the number of vertebrae and anal fin rays. They differ by the reduced squamation of the occiput in L. atlanticus and in the anal-fin origin placed below dorsal fin rays 15���18 vs 19���21 in L. gerringerae. It further differs in the lower number of transversal scales (18���20 vs 25), more compressed otolith (OL:OH = 1.2���1.55 vs 1.62) with the inner face flat and the outer face strongly convex (vs inner face of otolith convex and outer face slightly convex or flat). It differs from L. lentibus by having fewer dorsal-fin (82���94 vs. 105) and anal-fin (68���80 vs. 87) rays, as well as minute or absent eye lenses (0���10% of black eye vs 57%). Description. The principal meristic and morphometric characters are shown in Table 1. Body compressed and elongate with tapering tail. Head broad and somewhat depressed with thin fragile bones. Scales often missing, but scale-pockets show a fully scaled body with about 18 to 20 scales transversely across trunk above beginning of anal fin. Head with scaleless occiput and a few scales on opercle and preopercle. Anterior nostril situated midway between upper lip and posterior nostril. Snout equal in length to horizontal orbit. Horizontal length of orbit twice as long as high; lens poorly developed (Distribution (Fig. 5). Known from eight localities in the West Indies (19 specimens from 4580���6800 m), two localities in the Northeast Atlantic Ocean (two specimens from 5035���5100 m) and one location from the Solomon Sea, SW Pacific (4350���4450 m). Leucicorus atlanticus is the deepest living species in the genus., Published as part of Schwarzhans, Werner W., Nielsen, J��rgen G. & Mundy, Bruce C., 2022, Revision of the circumglobal deep-sea genus Leucicorus (Teleostei, Ophidiidae) with two new species, pp. 541-558 in Zootaxa 5100 (4) on pages 543-544, DOI: 10.11646/zootaxa.5100.4.5, http://zenodo.org/record/6224876, {"references":["Nielsen, J. G. (1975) A review of the oviparous ophidioid fishes of the genus Leucicorus, with description of a new Atlantic species. Trudy Instituta Okeanologii, 101, 106 - 123. [translation No. 74 from NOAA, Washington, D. C.]","Hureau, J. - C. & Nielsen, J. G. (1981) Les poissons Ophidiiformes des campagnes du N. O. \" Jean Charcot \" dans l'Atlantique et la Mediterranee. Cybium, 5 (3), 3 - 27.","Nielsen, J. G. & Moller, P. R. (2008) New and rare deep-sea ophidiiform fishes from the Solomon Sea caught by the Danish Galathea 3 Expedition. Steenstrupia, 30 (1), 21 - 46."]}

Leucicorus lentibus n. sp. Table 1, Figs. 1C, 2D, 3G, 5 Holotype (and only specimen): ZMUC P 771812, 170 mm SL, female, off Hawaiʽi, 20°01’N, 154°05W, R / V Vityaz, cr. 29, st. 4313, Sigsbee trawl, 5385–5390 m, 1 Feb. 1959. Diagnosis. Dorsal fin with 105 rays, anal fin with 87 rays and 64 (14+50) vertebrae. Transversal scale rows above origin of anal fin 20 (up to 27 rows in presumable life photographs). Reduced squamation on occiput, preopercle and opercle. Diameter of black eye 13% head length with large eye lens of 57% of black eye diameter. Otolith with flat inner and convex outer face; OL:OH = 1.73. Comparison. L. lentibus seems most similar to L. atlanticus with the reduced head squamation and a comparable number of transversal scale rows; however, it differs from L. atlanticus by having more dorsal- (105 vs. 82–93) and anal-fin rays (87 vs. 68–80) and a more slender otolith (OL:OH = 1.73 vs 1.2–1.55). It differs from L. lusciosus by having reduced head squamation, fewer transversal scale rows (20 vs. 35–40) and fewer dorsal- (105 vs. 110–114) and anal-fin (87 vs. 96–99) rays. It differs from L. gerringerae for its reduced head squamation, and by having more dorsal- (105 vs 97) and anal-fin (87 vs 73–80) rays and the flat inner and convex outer face of the otolith (vs convex inner face and slightly convex to flat outer face). It differs from all the three congeners in the presence of a large eye lens (57% of black eye vs 0–10%). Description. The principal meristic and morphometric characters are shown in Table 1. Body elongate and compressed with tapering tail; head broad and flat with a slight concavity above the eyes; head bones thin and fragile. Diameter of black eye 13% head length; the distinct, light, oval lens is 57 % the diameter of the black eye. Opercular spine thin and broadly pointed. Scales and scale pockets show a fully scaled body; 20 transversal scale rows from anal-fin origin; head with reduced squamation on occiput, preopercle and opercle. Dorsal-fin origin above anterior part of pectoral fin; anal-fin origin distinctly closer to tip of snout than to base of caudal fin; pelvic fins each with two tightly joined rays. Mouth terminal with upper jaw ending below posterior part of orbit. Anterior gill arch with four knob-like rakers on upper branch, one long raker in the angle and lower branch with nine long and four knob-like rakers; longest gill raker 3–4 times the length of the longest gill filament; two pseudobranchial filaments. After sixty-two years of preservation in alcohol the specimen is brownish grey with blue eye balls. Axial skeleton (based on radiographs). Precaudal vertebrae 14, all with pointed neural spines. Anterior neural spine half the length of the second one. The six anterior neural spines slightly depressed. Basis of neural spines 4–14 enlarged. Parapophyses on precaudal vertebrae 8–14. Pleural ribs on precaudal vertebrae 3–14. Epineurals not observed. Dentition.All dentigerous bones with many close-set, pointed teeth in irregular rows. Premaxillae and dentaries with teeth along their full length. Vomer boomerang-shaped. Palatines spindle-shaped. A single long and narrow basibranchial tooth patch. Otolith (Fig. 3G). Large otolith of 8.2 mm in length (holotype) with slender, droplet-like shape (otolith length: height = 1.73). Ventral rim shallow; dorsal rim with broad, rounded, predorsal angle; anterior rim broadly rounded; posterior tip slightly tapering, rounded. Inner face nearly flat; outer face convex; otolith height: thickness = 1.9. Sulcus straight, shallow, narrow, with single well-defined colliculum; otolith length: sulcus length = 2.18; sulcus length: height = 4.4. Sulcus connected to the anterior-dorsal rim via collicular crest. Ventral furrow very close to ventral rim of otolith; dorsal depression small. Distribution (Fig. 5). Known from the holotype caught in a bottom working trawl at a depth 5385–5390 m off Hawaii. Etymology. From lentibus (Latin) = lens bearing, referring to the presence of a large eye lens.

Key to species of Leucicorus 1a Extensive squamation on opercle, preopercle and occiput; transversal scale rows ���25; inner face of otolith more strongly convex than outer face.................................................................................... 2 1b Reduced squamation on opercle, preopercle and occiput; transversal scale rows ���21 (not considering ROV live photographs); inner face of otolith nearly flat, outer face distinctly convex................................................... 3 2a Dorsal fin rays 110���114; anal fin rays 96���99; vertebrae 68���69; transversal scale rows 35���40.................. lusciosus 2b Dorsal fin rays 97; anal fin rays 73���80; vertebrae 62; transversal scale rows 25...................... gerringerae n. sp. 3a Dorsal fin rays 88���94; anal fin rays 68���80; no eye lens or lens smaller than 10% of black eye; OL:OH = 1.2���1.55.................................................................................................... atlanticus 3b Dorsal fin rays 105; anal fin rays 87; lens at least 50 % of black eye; OL:OH = 1.73..................... lentibus n. sp., Published as part of Schwarzhans, Werner W., Nielsen, J��rgen G. & Mundy, Bruce C., 2022, Revision of the circumglobal deep-sea genus Leucicorus (Teleostei, Ophidiidae) with two new species, pp. 541-558 in Zootaxa 5100 (4) on page 543, DOI: 10.11646/zootaxa.5100.4.5, http://zenodo.org/record/6224876

Leucicorus Garman, 1899 Leucicorus Garman, 1899: 146 (type species by monotypy Leucicorus lusciosus Garman, 1899). Leucicorus: Nielsen 1974: 348; Nielsen 1975: 106; Nielsen et al. 1999: 73. Diagnosis. Elongate ophidiid with tapering tail. Scales on body, present or reduced on head. Median fins confluent. Pelvic fins with two rays in each. Mouth terminal. Infraorbital bones fragile and much expanded, almost covering maxillaries. Orbit large, equal in length to snout. Eye large, bulged, eye lens degenerated, except large in L. lentibus n. sp. Infraorbital, supraorbital and preopercular head canals extremely widened, covered with skin (when preserved), containing small pores. Opercular spine pointed, but weak. Posttemporal with a posterior, flap-like extension reaching beyond the opercle, bearing posterior part of supraorbital canal. Anterior nostril midway between upper lip and posterior nostril. Teeth small and pointed in irregular rows. No fangs. A single median basibranchial tooth-plate. 17���22 long gill rakers., Published as part of Schwarzhans, Werner W., Nielsen, J��rgen G. & Mundy, Bruce C., 2022, Revision of the circumglobal deep-sea genus Leucicorus (Teleostei, Ophidiidae) with two new species, pp. 541-558 in Zootaxa 5100 (4) on page 542, DOI: 10.11646/zootaxa.5100.4.5, http://zenodo.org/record/6224876, {"references":["Garman, S. (1899) The fishes. Memoirs of the Museum of Comparative Zoology, 24, 1 - 431.","Nielsen, J. G. (1974) List of ophidioid fishes from the 14 th cruise of the \" Akademik Kurchatov \" with a new species of Aphyonus. Trudy Instituta Okeanologii, 100, 348 - 353. [translation No. 72 from NOAA, Washington, D. C.]","Nielsen, J. G. (1975) A review of the oviparous ophidioid fishes of the genus Leucicorus, with description of a new Atlantic species. Trudy Instituta Okeanologii, 101, 106 - 123. [translation No. 74 from NOAA, Washington, D. C.]","Nielsen, J. G., Cohen, D. M., Markle, D. F. & Robins, C. R. (1999) Ophidiiform fishes of the world (order Ophidiiformes). FAO species catalogue, 18, 1 - 178."]}

Leucicorus gerringerae n.sp. Table 1, Figs. 1B, 2B, 3E–F, 5 Leucicorus atlanticus: Sêret & Andreata 1992: 92; Mincarone et al. 2008: 57. Material examined (2 specimens, SL 214–235 mm). Holotype: VIMS 42888, 235 mm SL, male, Silver Abyssal Plain, 23°20–22.5’ N, 68°23–29.4’ W, R / V Columbus Iselin, cr. CI-8113, st. D018, 45’ OT, 5303–5387 m, 25 May 1981. Paratype: MNRJ 52795, male, 214 mm SL, off Brazil, 21°39’S, 31°47’W, R / V Marion Dufresne, cr. MD-55, st. 26 CP 44, beam trawl, 4500–4510 m, 18 May 1987. Diagnosis. Dorsal fin with 97 rays, anal fin with 73–80 rays and 62 (13–14 + 48–49) vertebrae. Head squamation extensive on opercle, preopercle and occiput. 25 scales transversely across trunk above the anal-fin origin. Bluish eyes large (15–19% HL), without lens. Otolith elongate (OL:OH = 1.62), with inner and outer faces equally convex. C omparison. L. gerringerae is most similar to L. lusciosus with the extensive head squamation; however, it differs from L. lusciosus by having fewer dorsal-fin (97 vs. 110–114) and anal-fin (73–80 vs. 87–99) rays, larger head scales (2.0– 2.5 mm vs ~ 1 mm) and fewer scales across trunk above beginning of anal fin (25 vs. 35–40). It differs from L. lentibus by having an extensive head squamation, fewer rays in dorsal (82–94 vs. 105) and anal (73–80 vs. 87) fins and by lacking an eye lens. It differs from the sympatric L. atlanticus in having complete head squamation (vs reduced on occiput), higher number of dorsal-fin rays (97 vs 82–93), larger eye size (15–19% HL vs 10–13.5% HL except one eye in one case 18.4% HL), and the more elongate otolith (OL:OH = 1.62) with equally convex inner and outer faces (vs roundish otolith, OL:OH = 1.2–1.55, with flat inner face and strongly convex outer face). Description. The principal meristic and morphometric characters are shown in Table 1. The description is based on the holotype; characters in which the paratype differ from the holotype are given in brackets. Body elongate and compressed with tapering tail; head broad with dorsal profile straight except for a slight concavity above eyes; head bones thin and fragile. Anterior nostril placed midway between upper lip and posterior nostril and the larger posterior nostril close to orbit. Orbit equal in length to snout. Black eye about 15 to 19% of HL; no eye lens discernible. Opercular spine thin and pointed. Body fully scaled, with 25 scale rows transversely above beginning of anal fin, head with scales on opercle, preopercle and occiput. Dorsal-fin origin above anterior part of pectoral fin; anal-fin origin below dorsal fin ray 21(19), much closer to tip of snout than to base of caudal fin and with a preanal length ca. 40% SL. Pelvic fins each with two tightly joined rays. Pectoral fins placed on midbody. Mouth terminal; upper jaw ends below posterior edge of orbit; supramaxilla well developed. Anterior gill arch with four knob-like and one long raker on upper branch, one long raker in the angle between upper and lower branches and lower branch with nine(7) long and five(6) knob-like rakers; longest gill raker almost three times as long as longest gill filament. 2–4 pseudobranchial filaments. The color is brownish grey except for the bluish eyes. Axial skeleton (based on radiographs). Precaudal vertebrae 14(13) all with pointed neural spine tips. Anterior neural spine half the length of second spine whereafter the neural spines decrease in length. Six anterior neural spines somewhat depressed. Bases of neural spines 5–14(5–13) enlarged. Parapophyses on precaudal vertebrae 5–14(4–13). Pleural ribs on precaudal vertebrae 3–13(2–12). Epineurals not observed. Dentition. All dentigerous bones with many, small and pointed teeth arranged in irregular rows. Dentaries and premaxillae with teeth along their entire length with decreasing rows posteriad. Palatines with spindle shaped dentition and vomer ‘boomerang’-shaped with rather few teeth. One median basibranchial tooth patch, 6.5 mm long, 2.5 mm broad anteriorly and 0.8 mm broad posteriorly, with dentition similar to that on dentaries. Otolith (Fig 3E–F). Large otoliths up to 10.5 mm in length (holotype) with slender, droplet-like shape (otolith length: height = 1.62). Ventral rim shallow; dorsal rim with broad, obtuse, predorsal angle; anterior rim broadly rounded; posterior tip slightly tapering. Inner face about as convex as outer face; otolith height: thickness = 2.3. Sulcus slightly inclined, shallow, narrow, with single well-defined colliculum; otolith length: sulcus length = 2.28– 2.88; sulcus length: height = 3.3–4.5. Sulcus connected to anterior-dorsal rim via indistinct channel. Ventral furrow variably expressed; no distinct dorsal depression. Distribution (Fig. 5). Known from two localities (north of the Bahama Isl. and west of Rio de Janeiro) caught abyssally (4500 to 5303 m). Etymology. The species is named after Dr. Mackenzie Gerringer (State University of New York at Geneseo, USA) for her contribution to the knowledge of the biology of hadal fishes.