Additional Specimens of the Poorly Known Anchovy Stolephorus Multibranchus (Clupeiformes: Engraulidae) from Kosrae, Caroline Islands

Home , Stolephorus, Stolephorus commersonnii

Harutaka Hata and Hiroyuki Motomura Biogeography 20. 78–84. Sep. 20, 2018

Additional specimens of the poorly known anchovy Stolephorus multibranchus (Clupeiformes: Engraulidae) from Kosrae, Caroline Islands

Harutaka Hata1* and Hiroyuki Motomura2

1The United Graduate School of Agricultural Sciences, Kagoshima University, 1-21-24 Korimoto, Kagoshima 890-0065, Japan 2The Kagoshima University Museum, 1-21-30 Korimoto, Kagoshima 890-0065, Japan *Corresponding author: [email protected]

Abstract: Fourteen specimens of Stolephorus multibranchus Wongratana, 1987 (previously known only from the type specimens), collected from Pohnpei and Kosrae, Micronesia were located in museum collections. The Kosrae specimens represent the second locality for and easternmost record of the species. The newly-located specimens are fully described and comparisons made of S. multibranchus with S. commersonii Lacepéde, 1803.

Key words: anchovies, distribution, fish fauna, new record, taxonomy, morphology

Introduction et al., 1999), fourteen non-type specimens from Pohnpei and Kosrae were recently located in the The anchovy genus Stolephorus includes 20 valid Museum Support Center of the National Museum of species (Whitehead et al., 1988; Kimura et al., 2009; Natural History, Smithsonian Institution and Bernice Hata and Motomura, 2018), some of which, such as Papuahi Bishop Museum, Department of Zoology, Stolephorus commersonnii Lacepéde, 1803, Stole- USA. The Kosrae specimens represent the second phorus indicus (van Hasselt, 1823) and Stolephorus locality for and easternmost record of the species. A waitei Jordan & Seale, 1926, are abundantly caught full description of the specimens is given. and an important fisheries resource in the tropical Indo-Pacific (Baldwin, 1977; Ishikawa, 2013; Hata, Materials and Methods 2018). On the other hand, some species are poorly-known, including Stolephorus advenus Wongrata- Methods for counts and proportional measure- na, 1987, Stolephorus brachycephalus Wongratana, ments, shown in Tables 1 and 2, followed Hata & 1983, Stolephorus pacificus Baldwin, 1983 and Motomura (2017). Standard and head lengths are Stolephorus multibranchus Wongratana, 1987, all abbreviated as SL and HL, respectively. All measure- known only from their type specimens (Baldwin, ments were made with digital calipers to the nearest 1983; Wongratana, 1983, 1987a; Whitehead et al., 0.1 mm. Institutional codes follow Sabaj (2016). 1988; Wongratana et al., 1999). In addition to the nine type specimens of S. multibranchus collected Stolephorus multibranchus Wongratana, 1987 from Ponape (now Pohnpei), Caroline Islands (Won- (Fig. 1; Tables 1–2) gratana, 1987b; Whitehead et al., 1988; Wongratana ——————————————————————— ?Stolephorus commersonnii (not of Lacepéde): *Corresponding author:[email protected] Lee, 1973: 28 (Fiji).

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Fig. 1. (A) Lateral view of body and (B) dorsal view of head of preserved specimen of Stolephorus multibranchus (USNM 327513, 71.2 mm SL, Kosrae, Caroline Islands, Micronesia); (C) holotype of Stolephorus multibranchus, BMNH 1974.9.25.7, 57.0 mm SL, Pohnpei, Caroline Islands, Micronesia.

Stolephorus multibranchus Wongratna, 1987b: mens, 45.9–66.4 mm SL, Pohnpei, Caroline Islands. 3, fig. 1 (type locality: Metalanim Harbour, Pohn- Diagnosis. A species of Stolephorus with the fol- pei, Caroline Islands); Whitehead et al., 1988: 415, lowing combination of characters: gill rakers 24–27 unnumbered fig. (Pohnpei, Caroline Islands); Won- (modally 26) in upper series on first gill arch, 32–36 gratana et al., 1999, 1737, unnumbered fig. (Pohnpei, (33) in lower series, 57–64 (59) in total; gill rak- Caroline Islands); Myers, 1999: 62, fig. 1g [Pohnpei, ers 18–20 (18) in upper series on second gill arch, Caroline Islands, Micronesia]; Kimura et al., 2009: 30–34 (30, 32) in lower series, 47–53 (48, 49, 50, 295, fig. 1c (Metalanim Harbour, Pohnpei, Caroline 51) in total; gill rakers 13–16 (15) in upper series on Islands). third gill arch, 17–19 (18, 19) in lower series, 31–35 Materials examined. 14 specimens (57.8–71.2 (34) in total; gill rakers 11–13 (12) in upper series on mm SL): BPBM 28277, 5 specimens, 57.8–69.1 mm fourth gill arch, 14–15 (15) in lower series, 25–28 SL, mouth of Tofol Stream, Lalu, Kosrae (brackish (27) in total; prepelvic scutes 2–4 (4); predorsal water), Caroline Islands, 2 feet (0.6 m ca.) depth, 5 scales 15–17 (16, 17); long upper jaw, posterior tip Aug. 1981; USNM 327513, 5 specimens, 60.6–71.2 just reaching or just short of posterior border of pre- mm SL, Lele Harbor, Kosrae, Caroline Islands, 0–3 opercle; no spine-like scute on dorsal-fin origin and m depth, 24 June 1971; USNM 330894, 4 speci- pelvic scute; posterior border of preopercle convex,

− 79 − Additional specimens of the poorly known anchovy Stolephorus Harutaka Hata and Hiroyuki Motomura multibranchus (Clupeiformes: Engraulidae) from Kosrae, Caroline Islands rounded; a pair of dark patches behind occiput with covered by two to four prepelvic scutes. Pelvic scute a following pair of dark lines; no black spots below without a spine-like scute. Anus just anterior to anal- eye and on lower-jaw tip. fin origin. Caudal peduncle relatively compressed; Description. Counts and measurements, expressed depth slightly greater than orbit diameter. Snout tip as percentages of SL or HL, given in Tables 1 and 2. rounded; snout length less than eye diameter. Mouth Body compressed laterally, elongate, deepest at large, inferior, ventral to body axis, extending back- dorsal-fin origin. Dorsal profile of body slightly ward beyond posterior margin of eye. Lower jaw elevated from snout tip to dorsal-fin origin, straight slender, shorter than upper jaw. Maxilla rather long, along dorsal-fin base. Dorsal profile from dorsal-fin its posterior tip pointed, just reaching or just short of base end to uppermost point of caudal peduncle posterior border of preopercle. Single row of conical gently lowering. Ventral profile of body lowering teeth on each jaw and palatines. Small conical teeth slightly from lower-jaw tip to pelvic-fin insertion, in patch on vomer. Eye large, round, covered with thereafter straight and parallel to body axis until adipose eyelid, positioned laterally on head dorsal anal-fin origin, slightly elevated thereafter to lower- to horizontal through pectoral-fin insertion, visible most point of caudal peduncle. Abdomen rounded, in dorsal view. Pupil round. Orbit elliptical. Nostrils

Table 1. Meristics of specimens of Stolephorus multibranchus, Caroline Islands, Micronesia. Non-types Holotype Paratypes Kosrae Pohnpei Pohnpei Modes BMNH n = 10 n = 4 n = 2 1974.9.25.7 Standard length (SL; mm) 57.8–71.2 45.9–66.4 57.0 53.3–57.2 Dorsal-fin rays (unbranched) 3 3 3 3 3 Dorsal-fin rays (branched) 12–13 13 13 13 13 Anal-fin rays (unbranched) 3 3 3 3 3 Anal-fin rays (branched) 17–19 18–19 19 18 18 Pectoral-fin rays (unbranched) 1 1 1 1 1 Pectoral-fin rays (branched) 12–13 12–13 13 13 13 Pelvic-fin rays (unbranched) 1 1 1 1 1 Pelvic-fin rays (branched) 6 6 6 6 6 Caudal-fin rays (upper + lower) 10 + 9 10 + 9 10 + 9 10 + 9 10 + 9 Gill rakers on 1st gill arch (upper) 24–27 25–28 26 25–26 26 Gill rakers on 1st gill arch (lower) 33–34 33–36 35 32 33 Gill rakers on 1st gill arch (total) 57–61 58–64 61 57–58 59 Gill rakers on 2nd gill arch (upper) 18–19 18–20 19 18 18 Gill rakers on 2nd gill arch (lower) 29–34 30–34 31 29–30 30, 32 Gill rakers on 2nd gill arch (total) 47–53 48–52 50 47–48 48, 49, 50, 51 Gill rakers on 3rd gill arch (upper) 13–16 14–16 15 14 15 Gill rakers on 3rd gill arch (lower) 17–19 18–19 18 17 18, 19 Gill rakers on 3rd gill arch (total) 31–35 32–35 33 31 34 Gill rakers on 4th gill arch (upper) 11–13 12–13 12 11 12 Gill rakers on 4th gill arch (lower) 14–15 15 15 14 15 Gill rakers on 4th gill arch (total) 25–27 27–28 27 25 27 Gill rakers on posterior face of 3rd gill arch 5–8 7–8 8 7 7 Prepelvic scutes 2–4 3–4 4 2–3 4 Scale rows in longitudinal series 35–36 35–36 36 35–36 36 Predorsal scales 15–17 15–17 16 16–17 16, 17 Transverse scales 8 8 8 8 8 Pseudobranch filaments 17–20 17–19 20 17 17, 18, 19

− 80 − Additional specimens of the poorly known anchovy Stolephorus Harutaka Hata and Hiroyuki Motomura multibranchus (Clupeiformes: Engraulidae) from Kosrae, Caroline Islands

close to each other, anterior to orbit. Posterior mar- of gill membranes. Urohyal hidden by isthmus mus- gins of preopercle and opercle smooth. Subopercle cle (not visible without dissection). Gill membrane with rounded posterior margin. Opercular membrane on each side joined distally, isthmus muscle mostly without serrations. Posterior margin of preopercle exposed (not covered by gill membrane). Scales rounded, not indented. Interorbital space flat. Inter- cycloid, thin, deciduous, absent on head and fins, orbital width less than eye diameter. Pseudobranchial except for broad triangular sheath on caudal fin. filaments present, length of longest filament less than Scales on lateral surface of body with approximately eye diameter. Gill rakers long, slender, rough, visi- seven striae bending anteriorly on central part of ble from side of head when mouth opened. Isthmus scale. Lateral line absent. Dorsal-fin origin posterior muscle long, reaching anteriorly to posterior margin to vertical through base of last pelvic-fin ray, slightly

Table 2. Morphometrics of specimens of Stolephorus multibranchus, Caroline Islands, Micronesia. Non-types Holotype Paratypes Kosrae Pohnpei Pohnpei Means BMNH n = 10 n = 4 n = 2 1974.9.25.7 Standard length (SL; mm) 57.8–71.2 45.9–66.4 57.0 53.3–57.2 As % SL Head Length (HL) 24.8–26.2 25.0–26.7 26.2 25.8–26.3 25.8 Body depth 19.2–20.3 19.5–20.9 19.2 19.6–20.4 19.9 Pre-dorsal-fin length 52.5–55.0 53.9–54.8 55.6 53.3–53.8 54.2 Snout tip to pectoral-fin insertion 27.1–29.1 27.3–29.2 28.8 27.5–28.1 28.1 Snout tip to pelvic-fin insertion 45.2–49.0 44.9–47.9 47.4 46.2–46.5 46.7 Snout to anal-fin origin 63.0–65.0 63.4–65.0 64.8 61.8–65.1 64.1 Length of dorsal-fin base 14.0–15.7 15.3–16.1 15.8 14.6–15.0 15.1 Length of anal-fin base 18.3–21.3 19.2–20.9 20.8 18.4–19.8 19.9 Length of caudal peduncle 14.5–18.6 15.2–18.2 15.3 17.1–18.0 16.6 Depth of caudal peduncle 8.8–10.5 8.5–9.4 10.0 9.1–9.6 9.4 Pectoral-fin length 15.5–17.3 broken 17.0 17.8 16.9 Pelvic-fin length 10.0–11.4 10.5–11.4 11.6 11.7–11.9 11.0 Interorbital width 5.9–6.6 5.6–6.5 6.3 5.9–6.7 6.2 Snout length 4.3–5.1 4.1–4.7 4.3 4.3–4.5 4.5 Upper-jaw length 18.5–21.1 19.0–20.6 18.0 18.4–20.3 19.8 Mandibular length 16.2–18.5 16.0–17.6 16.9 17.1–17.4 17.1 1st dorsal-fin ray length 0.8–1.8 1.3–2.0 1.4 0.9–1.2 1.3 2nd dorsal-fin ray length 7.2–9.1 8.6 broken broken 8.3 3rd dorsal-fin ray length 16.0–19.0 broken broken broken 17.7 1st anal-fin ray length 0.8–1.7 1.4–1.7 1.4 1.0–1.2 1.3 2nd anal-fin ray length 4.7–6.2 broken 5.1 5.3–5.8 5.5 3rd anal-fin ray length 12.6–13.8 broken broken broken 13.2 1st pectoral-fin ray length 15.5–17.3 broken broken broken 16.5 1st pelvic-fin ray length 10.0–11.4 11.2–11.4 10.2 11.7–11.9 11.0 As % HL Length of orbit 32.1–37.5 30.6–34.8 32.2 31.8–33.0 33.6 Length of eye 26.0–31.0 25.0–27.8 29.3 27.5–28.5 27.5 D–P1 129.7–142.9 128.5–140.8 133.2 130.0–131.3 135.6 D–P2 82.2–92.6 79.5–89.9 81.5 81.5–82.9 85.5 D–A 82.6–90.6 80.1–89.0 82.9 81.5–85.0 84.9 P1–P2 77.6–88.0 73.6–76.4 78.1 69.2–74.2 78.6 P2–A 62.0–75.8 63.9–74.8 67.1 68.4–72.8 69.2 Postorbital length of the head 48.0–51.6 47.3–49.9 51.0 48.9–50.7 49.6

− 81 − Additional specimens of the poorly known anchovy Stolephorus Harutaka Hata and Hiroyuki Motomura multibranchus (Clupeiformes: Engraulidae) from Kosrae, Caroline Islands posterior to middle of body. Dorsal-fin origin with- head et al. (1988) and Wongratana et al. (1999): a out a spine-like scute. Dorsal and anal fins each with pair of dark lines on dorsum from occiput to dor- a minute first ray, three anteriormost rays closely sal-fin origin; lower gill rakers on first gill arch more spaced and unbranched. Anal-fin origin just below than 33; no spine-like scute on dorsal-fin origin and base of ninth to eleventh dorsal-fin ray; posterior pelvic scute; posterior tip of maxilla just reaching or tip of depressed fin not reaching caudal-fin base. just short of posterior border of preopercle; posterior Uppermost pectoral-fin ray unbranched, inserted margin of preopercle rounded, not indented; predor- below midline of body; posterior tip of fin not reach- sal scales15–17. ing vertical through pelvic-fin insertion. Pelvic fin Although S. multibranchus closely resembles Sto- shorter than pectoral fin, insertion anterior to vertical lephorus commersonnii Lacepède, 1803 in having through dorsal-fin origin; posterior tip of depressed a rather long upper jaw, its posterior tip reaching fin reaching to vertical through origin of second to to the posterior margin of the preopercle, a paired fifth dorsal-fin ray. dark line on the dorsum from the occipital region to Coloration of preserved specimens. Body uni- the dorsal-fin origin, and a predorsal scute absent formly pale ivory with light beige longitudinal band, (Whitehead et al., 1988; Wongratana et al., 1999), its width slightly less than pupil diameter, from just the former has numerous lower gill rakers on the behind upper opercular margin to caudal-fin base. A first gill arch (32–35 vs. 23–28 in S. commersonnii) pair of dark patches behind occiput without a follow- and a lower count of predorsal scales (15–17 vs. ing pair of dark lines; no black spots below eye or 18–19; Wongratana, 1987b; Whitehead et al., 1988; on snout and lower-jaw tips. Pectoral and pelvic fins Wongratana et al., 1999). Although Wongratana translucent, without melanophores. Melanophores (1987b) showed that body depth also separated the scattered on dorsal and caudal fins, along bases of two species (18.6–19.2% SL in S. multibranchus vs. dorsal and anal fins. Scales on dorsum margined 19.2–22.6% in S. commersonnii), the body depth of by melanophores. Double paired black patches on S. multibranchus examined in this study was 19.2– occipital region, posterior patches indistinct; a pair 20.9% SL, making separation on this basis equiv- of dark obscure lines on dorsum from occiput to dor- ocal. Although Wongratana (1987), Whitehead et sal-fin origin. al. (1988) and Wongratana et al. (1999) showed the Distribution. Stolephorus multibranchus, former- count of lower gill rakers on first gill arch as 32–35, ly known only from the type specimens collected a specimen examined here had 36 lower gill rakers from Pohnpei, Caroline Islands (Wongratana 1987b; on the first gill arch, indicating a greater range than Whitehead et al., 1988; Wongratana et al., 1999; this previously known. study), is newly recorded from Kosrae Island in this Stolephorus multibranchus was described by study. A report of the species from Fiji is unsupport- Wongratana (1987b) based on nine specimens col- ed (see Remarks). lected from Metalanim Harbour, Pohnpei, Caroline Remarks. The specimens were assigned to the Islands. Thereafter, Whitehead et al. (1988) and genus Stolephorus, defined by Whitehead et al. Wongratana et al. (1999) followed the diagnostic (1988) and Wongratana et al. (1999) as having a characters of the species given in the original de- long isthmus muscle reaching anteriorly to the pos- scription. Although Myers (1999) stated that “this terior margin of the gill membrane, hidden urohyal anchovy is known only from the coastal waters and prepelvic scutes, and postpelvic scutes absent, of Pohnpei, he made no mention extra specimens, and closely matched the diagnostic features of S. instead (apparently) citing the original description multibranchus given by Wongratana (1987b), White- and reproducing the original figure. Subsequently,

− 82 − Additional specimens of the poorly known anchovy Stolephorus Harutaka Hata and Hiroyuki Motomura multibranchus (Clupeiformes: Engraulidae) from Kosrae, Caroline Islands

Kimura et al. (2009) compared the holotype and six viding a literature, and G. Hardy (Ngunguru, New paratypes of S. multibranchus deposited in BMNH Zealand), for checking an early version of the man- with their new species, Stolephorus teguhi Kimura, uscript. This study was supported in part by a Grant- Hori and Shibukawa, 2009. in-Aid from the Japan Society for the Promotion Although Lee (1973) reported that S. commerson- of Science for JSPS Fellows (DC2: 29-6652); the nii was used as baitfish in Fiji, that species, recorded Sasakawa Scientific Research Grant from the Japan from the Indo-West Pacific from the eastern coast Science Society (28-745); JSPS KAKENHI Grant of Africa to the Ryukyu Islands, Papua New Guinea Numbers JP19770067, JP26241027, JP24370041, and the northern coast of Australia, has not been re- JP23580259, and JP26450265; the “Biological Prop- ported from Fiji (Whitehead et al., 1988; Wongratana erties of Biodiversity Hotspots in Japan” project of et al., 1999; Kimura et al., 1999). Because S. com- the National Museum of Nature and Science, Tsuku- mersonnii resembles S. multibranchus, Lee’s (1973) ba, Japan; “Establishment of Research and Educa- “S. commersonnii” is likely to be the latter. Howev- tion Network on Biodiversity and Its Conservation er, no specimens appear to be available, and in the in the Satsunan Islands” project of Kagoshima Uni- absence of reports of additional material, following versity adopted by the Ministry of Education, Cul- the original description of the species, the specimens ture, Sports, Science and Technology, Japan; and the from Kosrae Island represent the second locality for “Island Research” project by Kagoshima University. and easternmost record of S. multibranchus. No information on biology and habitat of S. mul- Reference tibranchus have been reported. Some of present specimens (BPBM 28277) were collected from the Baldwin, W. J., 1977. A review of the use of live bait estuary of Tofol Stream, suggesting that S. multi- fishes to capture skipjack tuna, Katsuwonus pela- branchus at least sometimes occurs in brackish wa- mis, in the tropical Pacific Ocean with emphasis ters. on their behavior, survival and availability. NOAA Comparative material examined. Stolephorus Tech. Rep., NMFS Circ., 408: 8‒35. multibranchus (45.9–66.4 mm SL): holotype of Baldwin, W. J., 1984. Stolephorus pacificus, a new Stolephorus multibranchus: BMNH 1974.9.25.7, species of tropical anchovy (Engraulidae) from 57.0 mm SL, Metalanim Harbour, Ponape, Caroline the western Pacific Ocean. Micronesica, 19: 151– Islands, 19 Nov. 1971, T. S. Hida; paratypes of Sto- 156. lephorus multibranchus: AMNH 57158, 57.2 mm Hata, H., 2018. Stolephorus indicus (van Hasselt, SL, USNM 280364, 53.3 mm SL, collected with the 1823). In: Koeda, K., Hata, H., Yamada, M. & holotype. Motomura, H. (Eds.), Field guide to fishes landed at Uchinoura Fishing Port, Kagoshima, Japan: Acknowledgements 84. The Kagoshima University Museum, Kagoshi- ma. We thank R. Arrindell (AMNH), O. Crimmen and Hata, H. & Motomura, H., 2017. A new species of J. Maclaine (BMNH), A. Suzumoto (BPBM) and anchovy, Encrasicholina auster (Clupeiformes: J. Williams, K. Murphy, S. Raredon and D. Pitassy Engraulidae), from Fiji, southwestern Pacific. N. (USNM) for opportunities to examine specimens Z. J. Zool., doi: 10.1080/03014223.2016.1268177 of Stolephorus. We also thank Y. Haraguchi and (also appeared in N. Z. J. Zool., 44:122‒128). other volunteers, and students of KAUM for their Hata, H. & Motomura, H., 2018. Stolephorus con- curatorial assistance, K. Koeda (NMMB) for pro- tinentalis, a new anchovy from the northwestern

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South China Sea and redescription of Stolephorus tutionalresource-collections-herpetology-ichthy- chinensis (Günther 1880) (Clupeiformes: Engraul- ology. Accessed 19 March 2018 idae). Ichthyol. Res., doi: 10.1007/s10228-018- Whitehead, P. J. P., Nelson, G. J. & Wongratana, T., 0621-z. 1988. FAO species catalogue, vol 7. Clupeoid Ishikawa, S., 2013. Stolephorus waitei Jordan & fishes of the world (suborder Clupeoidei). An an- Seale, 1926. In: Yoshida, T., Motomura, H., Musi- notated and illustrated catalogue of the herrings, kasinthorn, P. & Matsuura, K. (Eds.), Fishes of sardines, pilchards, sprats, shads, anchovies and northern Gulf of Thailand: 34. National Museum wolf-herrings. Pt 2 – Engraulidae. FAO Fish. Syn- of Nature and Science, Tsukuba, Research In- op., no 125, 7 (2): i–viii + 305–579. stitute for Humanity and Nature, Kyoto and Ka- Wongratana, T., 1983. Diagnoses of 24 new species goshima University Museum, Kagoshima. and proposal of a new name for a species of In- Kimura, S., Hori, K., & Shibukawa, K., 2009. A do-Pacific clupeoid fishes. Japan. J. Ichthyol., 29: new anchovy, Stolephorus teguhi (Clupeiformes: 385–407. Engrauliadae) from north Sulawesi, Indonesia. Wongratana, T., 1987a. Four new species of clupeoid Ichthyol. Res., 56: 292–295. fishes (Clupeidae and Engraulidae) from Austra- Kimura, S., Sado, T., Iwatsuki, Y. and Yoshino, T., lian waters. Proc. Biol. Soc. Washington, 100 (1) 1999. Record of an engraulid fish, Stolephorus 1987: 104–111. commersonnii from Ishigaki I., southern Japan. Wongratana, T., 1987b. Two new species of ancho- Japan. J. Ichthyol., 46: 45–50. vies of the genus Stolephorus (Engraulidae), with Lee, R., 1973. Live bait research. Skipjack tuna fish- a key to species of Engraulis, Encrasicholina, and ing project in Fiji. South Pac. Isl. Fish. Newsl., 9: Stolephorus. Am. Mus. Novit., 2876: 1–8. 26–30. Wongratana, T., Munroe, T. A., & Nizinski, M. S., Myers, R. F., 1999. Micronesian reef fishes. A com- 1999. Order Clupeiformes. Engraulidae. Ancho- prehensive guide to the coral reef fishes of Micro- vies. In: Carpenter, K. E.& Niem, V. H. (Eds.), nesia, 3rd revised edition. Coral Graphics, Guam. FAO species identification guide for fishery pur- Sabaj, M. H., 2016. Standard symbolic codes for in- poses. The living marine resources of the western stitutional resource collections in herpetology and central Pacific, vol 3. Batoid fishes, chimaeras ichthyology: an online reference. Version 6.5 (16 and bony fishes pt 1 (Elopidae to Linophrynidae): August 2016). American Society of Ichthyologists 1698–1753. FAO, Rome. and Herpetologists, Washington, DC. http://www. asih.org/resources/standard-symbolic-codes-insti- (Received June 30, 2018; Accepted July 30, 2018)

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