Abstract
The new generic name Nemoossis is proposed for the albulid species Pterothrissus belloci Cadenat 1937, a species endemic to the eastern Atlantic Ocean. Nemoossis differs from other genera in the family Albulidae as follows: longer dorsal-fin base, absence of supraneural bones, dorsal-fin rays [51–57 (mode 52)], vertebrae [total number 88–92 (91)], pored lateral-line scales [79–89 (84)], pre-dorsal-fin scale rows [3–8 (8)], head length [24–32 % (mean 30 %) of SL], pectoral-fin length [16–21 % (19 %) of SL], pre-dorsal-fin length [27–34 % (30 %) of SL], dorsal-fin base length [53–64 % (60 %) of SL], postorbital length [13–16 % (14 %) of SL], and upper caudal-fin length [18–31 % (24 %) of SL]. MNHN 1938-0002, 192 mm SL, is designated as a lectotype for P. belloci, and Bathythrissa dorsalis Günther 1877 is shown to be a junior synonym of P. gissu.
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Introduction
The bonefish family Albulidae is currently considered to comprise two genera (Nelson 2006), Albula Scopoli 1777 and Pterothrissus Hilgendorf 1877, although the latter has been considered the sole member of the family Pterothrissidae (e.g., Smith 1986; Aizawa 1993). Easily distinguished from the circumglobal genus Albula by its greater dorsal-fin base length, Pterothrissus has long been considered to include two species, Pterothrissus gissu Hilgendorf 1877, from the northwestern Pacific, and P. belloci Cadenat 1937, from the eastern Atlantic (Uyeno 1984; Smith 1986).
The genus Pterothrissus (type species: P. gissu) was described on the basis of a single specimen (ZMB 9890) collected from Yedo (=Tokyo), Japan by Hilgendorf (1877; published 31 Aug.). In the same year, Günther (1877; published 1 Nov.) described Bathythrissa dorsalis from “off Inosima” (=Enoshima), Kanagawa Pref., Japan, also from a single specimen (BMNH 1879.5.14.532). Examination of these specimens confirmed their conspecificity (see below). Elsewhere, Cadenat (1937) described P. belloci from Senegal on the basis of three syntype specimens (MHNLR P. 386, MNHN 1938-0002 and MSNG 36335). However, significant differences in morphological characters from the Japanese species suggested that the former represented a new albulid genus, which is described herein. The type species of Pterothrissus and the new genus are redescribed.
Materials and methods
Counts and measurements followed Hubbs and Lagler (1958), with the following additions and modifications: body width, least distance between pectoral-fin bases; bony interorbital width, bony width across frontals above center of eye; interorbital width with membrane, broadest distance between eyes (including membrane and adipose tissue); pre-anus length, distance from snout tip to anteriormost point of anus; upper and lower caudal-fin lengths, distance from middle of posterior margin of hypural plate to posterior tip of longest caudal-fin ray of upper and lower lobes, respectively; jaw width, distance between posteriormost points of each jaw; and maxillary depth, distance between upper and lowermost points on posterior margin of maxilla. Terminology and formula of supraneural bones follow Mabee (1988) and Ahlstrom et al. (1976), respectively. Standard length is expressed as SL. In each species description, holotype or lectotype data are given in parentheses; characters given in the diagnosis are not repeated. Institutional codes follow Fricke and Eschmeyer (2016).
Nemoossis gen. nov.
Type species. Pterothrissus belloci Cadenat 1937: 443, figs. 8–9 (type locality: Senegal).
Diagnosis. A genus of the family Albulidae with the following combination of characters: supraneural bones absent (Fig. 1a); dorsal-fin rays 51–57 (mode 52; Table 3); total vertebrae 88–92 (91; Table 1); pored lateral-line scales 79–89 (84; Table 1); pre-dorsal scale rows 3–8 (8; Table 3); head length 24–32 % (mean 30 %) of SL (Fig. 4a); pectoral-fin length 16–21 % (19 %) of SL (Fig. 4b); pre-dorsal-fin length 27–34 % (30 %) of SL (Fig. 5a); dorsal-fin base length 53–64 % (60 %) of SL (Fig. 5b); postorbital length 13–16 % (14 %) of SL; upper caudal-fin length 18–31 % (24 %) of SL.
Configuration of supraneural bones. a Nemoossis belloci, SAIAB 65836, 175 mm SL, Angola; b Pterothrissus gissu, HUMZ 71355, 193 mm SL, off Onahama, Iwaki, Fukushima Pref., Japan. Bars 10 mm
Etymology. The generic name “Nemoossis” is a combination of the Latin words “Nemo”, meaning absence, and “ossis”, meaning bones, in reference to the absence of supraneural bones.
Nemoossis belloci (Cadenat 1937)
(Figs. 1a, 2, 4–6; Tables 1, 3)
Nemoossis belloci. a lectotype, MNHN 1938-0002, 192 mm SL, Senegal; b paralectotype, MHNLR P. 386, 224 mm SL, Senegal; c paralectotype, MSNG 36335, 158 mm SL, Senegal
Pterothrissus belloci Cadenat 1937: 443, figs. 8–9 (type locality: Senegal); Smith 1986: 157 (West Africa); Whitehead 1990: 125 (eastern tropical Atlantic); Bianchi et al. 1993: 136 (Namibia); Heemstra and Heemstra 2004: 99 (southern Africa); Wirtz et al. 2013: 117 (Cape Verde Islands).
Lectotype. MNHN 1938-0002, 192 mm SL, Senegal.
Paralectotypes. MHNLR P. 386, 224 mm SL, Senegal; MSNG 36335, 158 mm SL, Senegal.
Other material ( n = 49, 129–278 mm SL). MHNLR 6501014, 4 specimens, 236–266 mm SL, eastern central Atlantic (ECA); MUFS 22667–22670, 4 specimens, 149–190 mm SL, off Republic of Equatorial Guinea; SAIAB 65836, 175 mm SL, Angola; SAIAB 65607, 4 specimens, 224–243 mm SL, Angola; SAIAB 48544, 208 mm SL, Namibia; SAIAB 10603, 6 specimens, 225–249 mm SL, ECA; SAIAB 3677, 2 specimens, 129–146 mm SL, no data; SAM 28239, 2 specimens, 239–256 mm SL, ECA; SAM 28836, 20 specimens, 211–257 mm SL, no data; ZMB 22010, 5 specimens, 183–229 mm SL, ECA.
Diagnosis. See generic diagnosis.
Description. Counts and proportional measurements of the lectotype, two paralectotypes and non-type specimens are given in Table 1. Data for the lectotype are given in parentheses if different.
Body elongate, subcylindrical; head large, its length nearly equal to pre-dorsal-fin length; snout conical, projecting slightly beyond lower jaw, its length nearly equal to upper-jaw length; mouth small, inferior; posterior margin of maxilla not reaching vertical through anterior margin of eye; eye moderate size, oblong, its diameter slightly less than bony interorbital width; small gular plate absent between lower jaws; suborbital bones well developed; teeth minute, conical and villiform on premaxillary and dentary, respectively; vomer and palatines toothless; gill rakers rudimentary; modified median row of enlarged scales in front of dorsal fin; all fins without spines; dorsal-fin base very long, about twice head length; longest dorsal-fin ray somewhat shorter than longest pectoral-fin ray (tip broken); anal fin located under posterior part of dorsal fin; longest anal-fin ray slightly shorter than caudal-peduncle length; pectoral fins low on side of body, near ventral outline; pelvic fins abdominal, located under central part of dorsal fin, length nearly equal to postorbital length; caudal fin deeply forked, upper lobe longer than lower; body covered with cycloid scales; head naked, cavernous; lateral line beginning at upper margin of opercle, running straight to caudal peduncle; intermuscular bone present.
Color of fresh specimens. Based on color transparencies of a freshly thawed specimen (MUFS 43595): Body bright silvery-white dorsally and ventrally; dark longitudinal lines between dorsal scale rows; head blackish; first dorsal-fin ray black; upper margin of dorsal-fin rays slightly black-edged, lower part yellowish-white; first pectoral-fin ray black, remaining rays dull yellow; pelvic-fin base grayish, other fin bases white; anal fin white; posterior margin of caudal-fin lobes black, lobes otherwise gray.
Color of preserved specimens. Based on the lectotype (MNHN 1938-0002, 192 mm SL) and non-type specimens: body black dorsally, brown ventrally; head blackish; first pectoral-fin ray blackish; other fins yellow; posterior margin of caudal-fin lobes blackish.
Distribution. Nemoossis belloci is a deep water (50–500 m depth, Whitehead 1990) eastern Atlantic species, ranging from Senegal to the west coast of South Africa (Fig. 6).
Remarks. Nemoossis belloci differs significantly from Pterothrissus gissu in lacking supraneural bones (Fig. 1), and in having different counts of dorsal-fin rays [51–57 (mode 52) vs. 54–65 (58) in P. gissu], total vertebrae [88–92 (91) vs. 105–107 (107)], pored lateral-line scales [79–89 (84) vs. 99–109 (104)], and pre-dorsal-fin scale rows [3–8 (8) vs. 13–20 (16)] (Tables 1, 3), and longer head [24–32 % (mean 30 %) of SL vs. 23–31 % (27 %)], pectoral fin [16–21 % (19 %) vs. 10–18 % (15 %)], pre-dorsal fin [27–34 % (30 %) vs. 31–39 % (35 %)] and dorsal-fin base [53–64 % (60 %) vs. 47–56 % (52 %)] (Figs. 4, 5).
A new genus “Nemoossis” is proposed for Pterothrissus belloci Cadenat 1937 because distinct differences of the skeletal system (no supraneural, more vertebrae, and fewer dorsal-fin rays) and many counts and proportional measurements (less lateral-line scales, fewer pre-dorsal-fin scale rows, and longer head length) are clearly recognized.
Nemoossis belloci was originally described (as Pterothrissus belloci) by Cadenat (1937) on the basis of three syntype specimens (MNHN 1938-0002, MHNLR P. 386, and MSNG 36335, Fig. 2a–c) from Senegal. We examined three syntype specimens and confirmed having counts and proportional measurement of the genus and species of N. belloci (Tables 1, 3 and Figs. 4, 5). One (MNHN 1938-0002, 192 mm SL) of the three syntype specimens is designated as the lectotype of P. belloci under Article 74.7.3 (ICZN 2003) because of avoiding future taxonomic confusion, while others automatically become the paralectotypes.
Pterothrissus Hilgendorf 1877
Type species. Pterothrissus gissu Hilgendorf 1877: 127 (type locality: Tokyo, Japan).
Diagnosis. A genus of the family Albulidae with the following combination of characters: 6–7 supraneural bones (Fig. 1b); dorsal-fin rays 54–65 (mode 58; Table 3); total vertebrae 105–107 (107; Table 1); pored lateral-line scales 99–109 (105; Table 1); pre-dorsal scale rows 13–20 (16; Table 3); head length 23–31 % (mean 27 %) of SL (Fig. 4a); pectoral-fin length 10–18 % (15 %) of SL (Fig. 4b); pre-dorsal-fin length 31–39 % (35 %) of SL (Fig. 5a); dorsal-fin base length 47–56 % (52 %) of SL (Fig. 5b); postorbital length 9–12 % (10 %) of SL; upper caudal-fin length 16–32 % (21 %) of SL.
Pterothrissus gissu Hilgendorf 1877
Pterothrissus gissu. a holotype, ZMB 9890, 319 mm SL, Tokyo, Japan; b holotype of Bathythrissa dorsalis, BMNH 1879.5.14.532, 365 mm SL, Enoshima, Kanagawa Pref., Japan
Pterothrissus gissu Hilgendorf 1877 (31 Aug.): 127 (type locality: Tokyo, Japan); Tameka 1982: 57, 316 (Kyushu-Palau Ridge, Japan); Shirai 1983: 65, 173 (north-eastern Sea of Japan); Machida 1984: 79, 314 (Okinawa Trough, Japan); Uyeno 1984: 21, pl. 23-H (Japan); Aizawa 2000: 189 (Japan); Sheiko and Fedorov 2000: 16 (Kamchatka, Russia); Shinohara et al. 2001: 291 (Pacific coast of northern Honshu, Japan); Aizawa 2002: 189 (Japan); Tsukamoto 2002: 267 (Japan); Shinohara et al. 2005: 399 (Ryukyu Islands); Shinohara et al. 2011: 39 (Sea of Japan); Aizawa and Doiuchi 2013: 190 (Japan).
Bathythrissa dorsalis Günther 1877 (1 Nov.): 443 [type locality: Enoshima, (originally given as “off Inosima”), Kanagawa Pref., Japan]: Günther 1887: 222, pl. 56-A.
Holotype. ZMB 9890, 319 mm SL, Tokyo, Japan (originally given as “Yedo”).
Other material ( n = 79, 61–406 mm SL, all from Japan, except where stated). ASIZP 60837, 325 mm SL, Dahsi fish market, Dahsi, Ylian, Taiwan; ASIZP 63258, 282 mm SL, Dahsi fish market, Dahsi, Ylian, Taiwan; BMNH 1879.5.14.532, 365 mm SL, off Inosima (=Enoshima, ca. 35°30’N, 139°50’E), Kanagawa Pref.; BSKU 4650, 307 mm SL, Suzaki, Kochi Pref.; BSKU 8986, 285 mm SL, Kochi Pref.; BSKU 18865–18880, 16 specimens, 63–113 mm SL, Sendai Bay, Miyagi Pref.; BSKU 33263, 386 mm SL, Okinawa Trough, East China Sea, ca. 29°N, 42°E; BSKU 53727–53728, 2 specimens, 319–331 mm SL, Kochi Pref.; FRLM 2343, 388 mm SL, Sea of Kumano, Wagu, Shima, Mie Pref.; FRLM 3463–3464, 3861, 3 specimens, 377–385 mm SL, Sea of Kumano, Shima, Mie Pref.; HUMZ 59213, 71231, 71355, 71421, 72132, 5 specimens, 151–248 mm SL, Onahama fish market, Iwaki, Fukushima Pref.; HUMZ 89817, 142 mm SL, off Muroran, Hokkaido; HUMZ 96624, 145 mm SL, southern part of Funka Bay, Hokkaido; OMNH 5865, 406 mm SL, Hyogo Pref.; OMNH 8158, 344 mm SL, Hyogo Pref.; OMNH 10083, 10415–10418, 10608–10609, 7 specimens, 286–383 mm SL, Suruga Bay, Shizuoka Pref.; NSMT-P 7506–7515, 10 specimens, 125–269 mm SL, Suruga Bay, Shizuoka Pref.; NSMT-P 12522, 2 specimens, 113–118 mm SL, off Fujigawa, Shizuoka Pref.; NSMT-P 12526, 151 mm SL, off Fujigawa, Shizuoka Pref.; NSMT-P 21812, 3 specimens, 88–121 mm SL, off Ida, Suruga Bay, Shizuoka Pref.; NSMT-P 46856, 277 mm SL, off Toda, Suruga Bay, Shizuoka Pref.; NSMT-P 48729, 5 specimens, 67–73 mm SL, Fukushima Pref.; NSMT-P 48909, 2 specimens, 70–71 mm SL, northeastern Pacific coast (37°N, 141°E); NSMT-P 49200, 2 specimens, 61–66 mm SL, Ibaraki Pref. (36°N, 140°E); NSMT-P 50031, 224 mm SL, off Choshi, Chiba Pref.; NSMT-P 53648, 2 specimens, 76–84 mm SL, off Miyagi Pref.; NSMT-P 65745, 65748–65749, 3 specimens, 73–88 mm SL, northeastern Pacific coast (37°N, 141°E), Japan; MUFS 498, 1174, 2 specimens, 199–254 mm SL, Owase market, Mie Pref.; MUFS 1601, 312 mm SL, off Atashika, Mie Pref.
Diagnosis. See generic diagnosis.
Description. Counts and proportional measurements of the holotype and non-type specimens of Pterothrissus gissu and holotype of Bathythrissa dorsalis are given in Table 2. Data for the holotype of P. gissu are given in parentheses if different.
Body elongate, subcylindrical; head moderate size, its length considerably less than pre-dorsal-fin length; snout conical, projecting slightly beyond lower jaw, its length equal to upper-jaw length; mouth small, inferior; posterior margin of maxilla not reaching vertical through anterior margin of eye; eye large, oblong, its diameter greater than bony interorbital width; small gular plate absent between lower jaws; suborbital bones well developed; teeth minute, conical and villiform on premaxillary and dentary, respectively; vomer and palatines toothless; gill rakers rudimentary; modified median row of enlarged scales in front of dorsal fin; all fins without spines; base of dorsal fin long, about twice head length; longest dorsal-fin ray shorter than longest pectoral-fin ray (tip broken); anal fin located under posterior part of dorsal fin; longest anal-fin ray slightly shorter than caudal-peduncle length (broken); pectoral fins low on side of body, near ventral outline; pelvic fins abdominal, located under central part of dorsal fin, length slightly greater than postorbital length; caudal fin forked, upper lobe slightly shorter than lower; body covered with cycloid scales; head naked, cavernous; lateral line beginning at upper margin of opercle, running straight to caudal peduncle; intermuscular bones present.
Color of fresh specimens. Based on color transparencies (OMNH 5865): Body dark grayish dorsally, silvery-white ventrally; dark longitudinal lines between dorsal scale rows; head dark grayish; anterior part of dorsal fin blackish; upper margin of dorsal-fin rays black edged, lower part hyaline white; pectoral fin gray; pectoral-fin base blackish, other fin bases white; anal fin white; posterior margin of caudal-fin lobes black.
Color of preserved specimens. Based on the holotype (ZMB 9890, 319 mm SL) and a non-type specimen (NSMT-P 46856): body dark brown dorsally and ventrally; head blackish; fins yellow; posterior margin of caudal-fin lobes blackish.
Distribution. Pterothrissus gissu is a moderately deep water (200–1000 m depth, Machida 1984) northwestern Pacific species (Fig. 6).
Remarks. Examination of the holotypes of Pterothrissus gissu Hilgendorf (1877; published in 31 Aug.) and Bathythrissa dorsalis Günther (1877; published in 1 Nov.) showed them to be conspecific, having the same diagnostic features (Tables 2, 3, and Figs. 4, 5). Accordingly, B. dorsalis is recognized as a junior synonym of P. gissu.
Relationships of a head length, b longest pectoral-fin length, c pre-dorsal fin length, and d dorsal-fin base length to standard length in Nemoossis belloci (open circles) and Pterothrissus gissu (solid triangles)
Distributional record of Nemoossis belloci (solid circles) and Pterothrissus gissu (solid triangles), based on specimens examined in this study
The original description (Günther 1877) of B. dorsalis was written as only “Off Inosima”, although Fricke and Eschmeyer (2016) erroneously noted it as “Off Inoshima, Hiroshima Prefecture, Japan, Inland Sea, depth 345 fathoms?” However, Günther (1887) added a postscript as “Off Inosima; purchased from Japanese fishermen; depth, (?) 345 fathoms. One specimen, 15 inches long”. Furthermore in the same reference, the same locality species of Centrophorus squamulosus was mentioned in addition to the station number (station 232). The latitude and longitude data of station 232 in the Expedition of HMS Challenger were 35°11’N and 139°28’E, respectively (Brady 1884). These data clearly point to Sagami Bay, Kanagawa Pref., Japan. Accordingly, we presumed that “Inosima” is equal to Enoshima.
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Acknowledgments
We greatly appreciate the specimen loans from the following persons and institutions: K-T. Shao (ASIZP); J. Maclaine (BMNH); H. Endo (BSKU); T. Nakabo and Y. Kai (FAKU); S. Kimura (FRLM); M. Yabe, H. Imamura, and T. Kawai (HUMZ); C. Ferrara, R. Causse, Z. Gabsi, P. Pruvost, and G. Duhamel (MNHN); G. Shinohara and K. Matsuura (NSMT); K. Hatooka (OMNH); B. Mackenzie, R. Bills, K. Dubula, U. Lwana, E. Heemstra, M. Mwale, V. B. Kongobe, V. Hanisi, W. Holleman, M. E. Anderson, P. C. Heemstra, and P. Skelton (SAIAB); M. Bougaardt and L. J. V. Compagno (SAM); and P. Bartsch and H.-J. Paepke (ZMB). K. E. Carpenter (ODU), organizer of the 2004 FAO workshop (Tenerife, Canary Is., Spain), with the co-operation of the Spanish Government, also provided examples of Nemoossis belloci for which we were very grateful. G. S. Hardy, Ngunguru, New Zealand, read the initial manuscript and offered helpful comments. This study was financially supported in part by the Sasakawa Scientific Research Grant from the Japan Science Society (No. 15-258) and the Ito Grant for Ichthyology, Fujiwara Natural History Foundation, awarded to the first author.
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Hidaka, K., Tsukamoto, Y. & Iwatsuki, Y. Nemoossis, a new genus for the eastern Atlantic long-fin bonefish Pterothrissus belloci Cadenat 1937 and a redescription of P. gissu Hilgendorf 1877 from the northwestern Pacific. Ichthyol Res 64, 45–53 (2017). https://doi.org/10.1007/s10228-016-0536-5
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DOI: https://doi.org/10.1007/s10228-016-0536-5