First Records of the Luminous Deepwater Cardinalfish (Perciformes: Epigonidae), Epigonus Macrops (Brauer, 1906), from the Southeastern Atlantic and Timor Sea

Species Diversity 21: 79–83 25 May 2016 DOI: 10.12782/sd.21.1.079

First Records of the Luminous Deepwater Cardinalfish (Perciformes: Epigonidae), Epigonus macrops (Brauer, 1906), from the Southeastern Atlantic and Timor Sea

Makoto Okamoto1,4 and Naohide Nakayama2,3 1 Seikai National Fisheries Research Institute, 1551-8 Taira-machi, Nagasaki 851-2213, Japan E-mail: [email protected] 2 Laboratory of Marine Biology, Faculty of Science, Kochi University, 2-5-1 Akebono-cho, Kochi 780-8520, Japan 3 Present address: The Kyoto University Museum, Kyoto University, Yoshida, Sakyo, Kyoto 606-8501, Japan 4 Corresponding author (Received 16 February 2016; Accepted 18 April 2016)

The luminous deepwater cardinalfish, Epigonus macrops (Brauer, 1906), is recorded for the first time from the southeastern Atlantic (a single specimen, 121.0 mm SL) and the Timor Sea (two specimens, 128.6 and 150.8 mm SL). These specimens are described and compared with other specimens of the species from Western Australia, Indonesia, the western North Atlantic, and the Galápagos Islands. The present study reviews previous reports on the distribution of E. macrops in the world. Key Words: Percoidei, deep-sea fish, distribution, new record, luminous organ, Indo-Pacific.

Introduction Materials and Methods

The perciform epigonid fish Epigonus macrops (Brauer, Meristic and morphometric methods followed Mayer 1906) was originally described based on two specimens (1974) and Okamoto (2011). Missing lateral-line scales were from the western coast of Sumatra, Indonesia, eastern In- estimated by counting scale pockets. The number of pored dian Ocean, at a depth of 903 m (Brauer 1906). This species lateral-line scales on the caudal fin is represented as “+n”. has a ventral luminous organ as a unique character among The term “mustache-like maxillary process” is used for a lat- the species of this genus and belongs to the “E. telesco- eral process on the head of maxilla (see Mayer 1974; Oka- pus group” (Mayer 1974; Okamoto et al. 2012). Except for moto 2012). The definition of the first caudal vertebra fol- an accidental collection from the surface after a submarine lows Okamoto and Motomura (2011). The terminology and volcanic eruption in the Galápagos Islands [as E. merleni formula for the supraneural bones follow Mabee (1988) and McCosker and Long, 1997, relegated to a junior synonym of Ahlstrom et al. (1976) respectively. Counts of supraneurals, E. macrops by Okamoto et al. (2012)], adults of E. macrops vertebrae, and ribs were taken from radiographs. The num- have been collected exclusively by bottom trawls between ber of pyloric caeca, presence of a ventral luminous organ, 120 m and 1100 m in the Indian Ocean (off tropical east and sex were confirmed by dissection of the right side of Africa, Walters Shoals, Sumatra, and Western Australia), the abdomen. Standard length is abbreviated as SL. The the South China Sea (off Vietnam), and the western North specimens examined in the present study are deposited at Atlantic (Gulf of Mexico, Caribbean Sea, Bahamas, and off the Australian Museum, Sydney (AMS); Laboratory of Ma- French Guiana and Suriname) (Brauer 1906; Mayer 1974; rine Biology, Faculty of Science, Kochi University, Kochi Fujii 1983; Allen and Cross 1989; Abramov 1992; Williams (BSKU); California Academy of Sciences, San Francisco et al. 1996; McEachran and Fechhelm 2005). (CAS); CSIRO Marine and Atmospheric Research, Tasma- In an on-going study of the taxonomy of the genus (e.g., nia (CSIRO); Harvard University, Museum of Comparative Okamoto 2012, 2015), the first author (MO) examined a Zoology, Cambridge (MCZ); Texas Cooperative Wildlife single specimen of E. macrops from off Angola deposited Collection, Texas (TCWC); and Ichthyology, Zoologisches in the California Academy of Sciences (CAS). Additionally, Museum, Museum für Naturkunde, Berlin (ZMB). the second author (NN) found two specimens of this species from the Timor Sea deposited in the Laboratory of Marine Epigonus macrops (Brauer, 1906) Biology, Faculty of Science, Kochi University (BSKU). These [English name: Luminous Deepwater Cardinalfish] specimens represent the first confirmed records of E. mac- [Japanese name: Kushiba-yasemutsu] rops from the southeastern Atlantic and the Timor Sea. (Fig. 1)

Oxyodon macrops Brauer, 1906: 288, fig. 172 (original de-

Fig. 1. Two specimens of Epigonus macrops. A, 150.8 mm SL, BSKU 16702, Timor Sea; B, 121.0 mm SL, CAS 234746, off Angola.

scription; type locality: west coast of Sumatra, Indonesia); lected by T. Iwamoto. Timor Sea: BSKU 16702, 150.8 mm Jordan 1920: 517 (list); Schultz 1940: 412 (list); Weber SL, female, 9°27.0′S, 127°58.6′E–9°28.5′S, 127°56.1′E, 610– and de Beaufort 1929: 351, fig. 81 (description, based on 690 m depth, 18–19 June 1972, beam trawl, R/V Hakuho- Brauer, 1906, west coast of Sumatra, Indonesia); Norman maru, collected by O. Okamura; BSKU 16703, 128.6 mm SL, 1939: 60 (list). male, same data as BSKU 16702. Epigonus macrops: Mayer 1974: 159, fig. 4 (description, In- Other material examined. 13 specimens, 103.7– dian Ocean, Gulf of Mexico, Caribbean Sea, and north- 202.5 mm SL. Australia: AMS I.22814-018, 131.6 mm SL, western Atlantic); Fujii 1983: 323, unnumbered fig. (de- northwest of Port Headland, Western Australia, Indian scription, Suriname and French Guiana); Shcherbachev Ocean, 18°48′00″S, 116°60′00″E, 704 m depth, 6 April 1982; 1987: 42 (list, Indian Ocean); Allen and Cross 1989: 553 AMS I.31161-004, 170.5 mm SL, off Cape Cuvier, Western (list, Western Australia); Abramov 1992: 98, table 1 [key Australia, Indian Ocean, 24°53′67″S, 111°80′83″E, 901 m and distribution, western North Atlantic (from Florida depth, 28 January 1991; CSIRO H 2562-01, 178.4 mm SL, to Guiana), Indian Ocean (from Tanzania to Sumatra), west of Quobba Point, Western Australia, Indian Ocean, and South China Sea (off southern Vietnam)]; Williams 24°28′S, 111°51′E, 905 m depth, 28 January 1991; CSIRO H et al. 1996: 153, appendix 1 (list, Western Australia); Gon 6575-08, 126.8 mm SL, northwest of Cape Leveque, Western 1999: 2613 (key and brief description, off southern Viet- Australia, Indian Ocean, 14°35′S, 121°21′E, 709 m depth, nam); Gon 2000: 614 (list, South China Sea, off southern 26 June 2007. Indonesia: ZMB 17678, syntype of Oxyodon Vietnam); Hutchins 2001: 32 (list, Western Australia); macrops, 202.5 mm SL, western coast of Sumatra, Indone- Gon 2003: 1394 (key and list, western North Atlantic); sia, eastern Indian Ocean, 03°22′01″S, 101°11′05″E, 903 m McEachran and Fechhelm 2005: 235 (list, Gulf of Mexi- depth, 21 January 1899. Galápagos Islands: CAS 86581, ho- co); Hoese et al. 2006: 1114 (list, Western Australia). lotype of Epigonus merleni, 139.1 mm SL, surface offshore of Epigonus merleni McCosker and Long, 1997: 126, fig. 1 (type Cabo Hammond, Isla Fernandina, Galápagos Islands, east- locality: Galápagos Islands). ern Pacific, 00°28′S, 91°37′W, February 1995. Northwest- Epigonus atherinoides (not of Gilbert, 1905): Horikoshi et al. ern Atlantic: MCZ 48827, 148.7 mm SL, Caribbean Sea, 1983: 117 (list, Timor Sea, present specimens). 11°54′N, 69°18′W, 910 m depth, 4 October 1963; TCWC 6372.09, 4 specimens, 103.7–145.6 mm SL, Gulf of Mexico, Material examined. 3 specimens, 121.0–150.8 mm 27°64′N, 91°53′W, 731 m depth, 3 March 1986; TCWC SL. Southeastern Atlantic: CAS 234746, 121.0 mm SL, fe- 7003.10, 2 specimens, 120.6–156.7 mm SL, Gulf of Mexico, male, off Angola, 9°54.09′S, 12°44.47′E, 620–630 m depth, 27°14′N, 93°39′W, 792–864 m depth, 8 April 1986. 10 March 2007, bottom trawl, R/V Dr. Fridtjof Nansen, col- Diagnosis. A species of Epigonus with the following Records of Epigonus macrops 81 combination of characters: pungent opercular spine absent; elliptical and without dermal flap. Eye large, round, orbital dorsal-fin rays VII-I-I, 10 or rarely VIII-I, 10; total gill rak- diameter greater than postorbital length; bony rim of orbit ers 18–20; vertebrae 10+15; pyloric caeca 8; pored lateral- raised above dorsal profile; interorbital region concave in line scales 45–51+3–5; mustache-like maxillary processes middle. Mouth large, terminal; gape oblique; posterior mar- absent; ribs presence on last abdominal vertebra; and ven- gin of maxilla extending to vertical drawn approximately tral luminous organ present. through center of pupil; lower jaw slightly projecting when Description. Counts and proportional measurements mouth closed; anteriorly projecting teeth or nub-like struc- as given in Table 1. Body elongate, laterally compressed, tures absent on symphysis of lower jaw. Curved conical deepest at pectoral-fin base; nape not humped. Head large, teeth arranged in single row on maxilla and dentary. Small slightly compressed. Mustache-like maxillary process ab- number of minute teeth present on vomer and palatine. Ba- sent. Snout short and round, its length less than interorbital sihyal toothless. Opercular spine weak and flat, not pungent, width; two nostrils closely set at level of upper edge of pupil, forming low ridge; preopercular edges smooth. Origin of anterior nostril without membranous tube, posterior nostril first dorsal fin above anterior part of pectoral fin; first- dor

Table 1. Counts and proportional measurements of Epigonus macrops.

off Angola Timor Sea Other specimens n=1 n=2 n=13 Standard length (mm) 121 128.6–150.8 103.7–202.5 Counts Dorsal-fin rays VII-I-I, 10 VII-I-I, 10 VII-I-I, 10 or VIII-I, 10 Anal-fin rays II, 9 II, 9 II, 9 Pectoral-fin rays 19 18 18–19 Pored lateral-line scales 47+5 47+3 45–51+3–5 Scales above lateral line 3 3 3 Scales below lateral line 8 8–9 8–9 Pyloric caeca 8 8 8 Gill rakers 4+14=18 4–5+14=18–19 4–5+13–15=18–20 Vertebrae 10+15 10+15 10+15 Measurements (% standard length) Head length 35.2 35.4–37.6 34.6–37.8 Head height 17.2 19.0–20.4 17.4–20.2 Body depth 21.3 21.0–24.9 20.8–24.3 Body width 15.6 15.6–18.0 14.0–17.7 Caudal-peduncle depth 10.0 10.7–11.1 8.9–10.9 Caudal-peduncle length 28.3 25.1–25.4 24.4–28.9 Orbital diameter 15.6 14.9–15.8 12.7–17.5 Interorbital width 10.7 11.0–11.1 10.2–12.0 Postorbital length 13.1 13.8–13.9 12.3–14.6 Upper-jaw length 13.0 14.4–14.5 13.3–14.8 Lower-jaw length 16.9 17.3–18.4 15.9–19.3 Snout length 7.4 7.5–7.6 6.7–9.5 Pre-1st dorsal-fin length 38.8 37.4–39.5 36.8–40.1 Pre-2nd dorsal-fin length 57.5 58.7–60.1 57.0–60.3 Pre-pectoral-fin length 35.1 35.1–37.9 35.2–39.5 Pre-pelvic-fin length 35.7 36.1–37.7 35.8–41.7 Pre-anus length 57.9 58.9–59.0 57.0–61.7 Pre-anal-fin length 65.8 66.4–67.3 64.3–69.4 1st spine length on 1st dorsal fin 2.4 2.1–2.3 1.2–2.8 2nd spine length on 1st dorsal fin 12.6 15.3 11.5–15.9 3rd spine length on 1st dorsal fin Tip broken 14.7–15.7 12.3–15.7 2nd dorsal-fin spine length 7.2 7.3 4.9–7.8 1st anal-fin spine length 1.9 1.7–2.3 1.2–2.7 2nd anal-fin spine length 7.7 8.1 4.8–8.8 Pelvic-fin spine length 12.7 14.1–14.6 11.7–14.8 1st dorsal-fin base length 11.9 11.3–13.5 10.8–16.0 2nd dorsal-fin base length 10.8 10.6–11.1 9.6–11.3 Anal-fin base length 9.4 9.2–9.7 9.3–10.9 Pectoral-fin length 21.6 20.8–22.3 16.7–21.3 Pelvic-fin length 15.7 15.1 14.9–16.0 82 M. Okamoto and N. Nakayama

Fig. 2. Distributional records of Epigonus macrops (stars=present study; circles=previous studies). sal-fin spine minute; third dorsal-fin spine longest; isolated 1997: CAS 86581), these distributional records are located dorsal fin spine present between first and second dorsal- between 30°N and 35°S at depths greater than 120 m. Thus, fins, not connected to seventh dorsal-fin spine by fin mem- this species is distributed on continental slopes in subtropi- brane. Spine of second dorsal fin short, thicker than those cal and tropical regions. of first dorsal fin. Origin of anal fin below posterior portion No species of the genus Epigonus had previously been of second dorsal-fin base; first anal-fin spine minute; second reported from the Timor Sea. This is considered to be due anal-fin spine short, subequal in length to second dorsal-fin to the dearth of knowledge about the deep-sea fish fauna spine. Posterior tip of pectoral fin just reaching vertical line of the Timor Sea. On the other hand, six other species of drawn from anus. Caudal fin deeply forked. Anus located the genus have been recorded from around Angola (Mayer slightly posterior to vertical line through origin of second 1974; Abramov 1992; Okamoto et al. 2011), viz., E. affinis dorsal fin. Ribs present on last abdominal vertebra. Supra- Parin and Abramov, 1986, E. constanciae (Giglioli, 1880), E. neural bones three (0+0/0+2/1+1/1/). Scales deciduous, denticulatus Dieuzeide, 1950, E. mayeri Okamoto, 2011, E. weakly ctenoid, covering whole body except area anterior pandionis (Goode and Bean, 1881), and E. telescopus (Risso, to rim of orbit and surfaces of jaws; scales also present on 1810). bases of second dorsal, anal, and caudal fins; series of pored Epigonus macrops is unique in the genus in having a ven- lateral-line scales complete, 3–5 pored scales on caudal fin. tral luminous organ; one of the eight pyloric caeca appears Ventral luminous organ present; luminescent window locat- to have become modified into this organ. The lumines- ed mid-ventrally between anterior pelvic fins. cent window is located mid-ventrally between the anterior Color in alcohol (Fig. 1). Body and all fins uniformly light pelvic-fins and is covered by a single large scale (see Mayer brown; opercular region dark brown; luminous window, 1974: fig. 7; Okamoto et al. 2012: fig. 3). Also, the eighth tongue, and mouth cavity black. dorsal-fin spine is isolated between the first and second dor- Distribution (Fig. 2). Known from the western North sal fins or, rarely, is connected by a membrane to the sev- Atlantic [Gulf of Mexico and Caribbean Sea, Bahamas, off enth spine of the first dorsal fin (expressed as VII-I-I, 10 or French Guiana and Suriname (Mayer 1974; Fujii 1983; VIII-I, 10). This feature is shared with only four other spe- McEachran and Fechhelm 2005)]; off Angola, southeast- cies in the genus and is known as a unique diagnostic char- ern Atlantic (present study); off Vietnam, South China acter for the E. telescopus group (Abramov 1992; Okamoto Sea, (Abramov 1992); Galápagos Islands, eastern Pacific and Motomura 2011, 2013), which comprises E. macrops, (McCosker and Long 1997; Okamoto et al. 2012); and In- E. angustifrons Abramov and Manilo, 1987 (southwestern dian Ocean [Sumatra, Western Australia, Timor Sea, off Indian Ocean, southeastern Atlantic), E. notacanthus Parin tropical east Africa, Walters Shoals (Brauer 1906; Mayer and Abramov, 1986 (Nazca Ridge), and E. telescopus (In- 1974; Shcherbachev 1987; Allen and Cross 1989; Abramov dian Ocean, Atlantic, southwestern Pacific) (Mayer 1974; 1992; Williams et al. 1996; present study)]; at depths of 120– Abramov 1992; Okamoto et al. 2012; Okamoto 2016). 1100 m. In our examination of specimens of E. macrops, only one Remarks. There are no clear differences in the meristic specimen (BSKU 16702, 150.8 mm SL) was a female with and morphometric characters between the present speci- developed mature ova. This observation represents the first mens and previously recorded specimens, including a syn- confirmed information about sexual maturity in this species. type of Epigonus macrops (Table 1). The present study newly revealed that E. macrops is distributed in the southeastern Atlantic (off Angola) and the Timor Sea. Except for an ac- Acknowledgments cidental collection from the surface in the Galápagos Islands after a submarine volcanic eruption (McCosker and Long We are grateful to M. McGrouther and S. Reader (AMS), Records of Epigonus macrops 83

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