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(A, B) Upeneus alama: Philippines, Manila Bay, Cavite; (C) U. brevianalis, Thailand, Andaman Sea, Phuket; (D) U. doriae: Persian/Arabian Gulf, Bahrain; (F, G) U. nigromarginatus: (F) Philippines, Negros Oriental, Dumaguete Citz; (G) Philippines, Panay Island, Tigbauan market near Iloilo City.
Upeneus sulphureus Cuvier, 1829. (A, B) Thailand, Gulf of Thailand, Prachuab Khiri Khan Province; (C) Philippines, Metro Manila; (D) Vietnam, Ha Long Bay.
in Uiblein, Motomura, Møller, Sabaj, Boonphienphol, Hoang et Parkinson, 2025. |
Abstract
For the goatfish genus Upeneus, the new sulphureus-species group is established, which includes five species, U. doriae, U. nigromarginatus, U. sulphureus, and the two new species, U. alama and U. brevianalis. This taxonomic group differs from the other seven species groups and four ungrouped species of this genus in the following combination of characters: 8 dorsal-fin spines, 14–17 pectoral-fin rays, 25–33 total gill rakers, 32–37 lateral-line scales, and lack of bars on caudal fin. Upeneus sulphureus is redescribed based on examination of a large set of morphometric, meristic, and color characters in 211 specimens, which revealed a bimodal distribution in barbel length prompting separation into short- and long-barbel morphs. This bimodality was neither sex-, size-, nor population-dependent, but found in co-occurring individuals of U. sulphureus in many areas of its distributional range from East Africa to Japan. The two new species differ from all other congeners primarily in U. alama, new species, having longer barbels (28–30% SL) and U. brevianalis, new species, having a shorter and shallower anal fin (8.1 and 12% SL, respectively) as well as a broad, dark-brown vertical band along the anterior third of the first dorsal fin. For U. nigromarginatus, new records from Papua New Guinea, Indonesia (West Papua), and three areas of the Philippines north of Mindanao are reported. For U. doriae, U. sulphureus, and Mulloides pinnivittatus, the latter a junior synonym of U. sulphureus, lectotypes are designated. An updated account for the genus Upeneus is provided. The barbel-length data of all 53 species of Upeneus are reviewed, and the finding of a wide range combined with a distinct dimorphism of this character in U. sulphureus is discussed.
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Upeneus sulphureus.
(A, B) KAUM-I.33094, SBM, 119 mm SL, Thailand, Gulf of Thailand, Prachuab Khiri Khan Province (H. Motomura); (C) USNM 403094, SBM, 105 mm SL, Philippines, Metro Manila (copyright J. T. Williams); (D) KAUM-I.67460, SBM, 90 mm SL, Vietnam, Ha Long Bay (H. Motomura);
(E, F): SAIAB 82237, LBM, 106 mm SL, Mozambique (E: copyright O. Alvheim, F: copyright P. C. Heemstra);
(G, H) KAUM-I.57209, LBM, 91 mm SL, Philippines, Panay Island, Iloilo City (H. Motomura); (I, J) KAUM-I.123161, LBM, 99 mm SL, Japan, Kagoshima, NE of Matsushima (H. Motomura). |
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(A, B) Upeneus alama: (A) USNM 466258, HT, 117 mm SL, Philippines, Manila Bay, Cavite (copyright D. Pitassy); (B) USNM 466258, PT, 125 mm SL, same locality (copyright D. Pitassy);
(C) U. brevianalis, ZMUC P49375, HT, 137 mm SL, Thailand, Andaman Sea, Phuket (F. Uiblein);
(D, E) U. doriae: (D): USNM FIN 31104 (unvouchered photograph), 106 mm SL, Persian/Arabian Gulf, Bahrain (copyright J. E. Randall); (E) ZMB 7056, LT, 104 mm SL, Iran, Persian/Arabian Gulf, Bandar Abbas (copyright E. Aßel);
(F, G, H) U. nigromarginatus: (F) USNM 403319, 159 mm SL, Philippines, Negros Oriental, Dumaguete Citz (copyright J. T. Williams); (G) VNMN-I 1986 (unvouchered image), ca. 70 mm SL, Philippines, Panay Island, Tigbauan market near Iloilo City (F. Uiblein); (H) AMS I.18495-001, 88 mm SL, Papua New Guinea, Madang Province, Broken Water Bay (K. Parkinson). |
Upeneus alama Uiblein and Motomura, new species
Cavite Goatfish
Diagnosis.—Dorsal fins VIII + 9; pectoral fins 16; gill rakers 8–9 upper + 20–22 lower = 28–31 total; lateral-line scales 33; body moderately deep; measurements as % SL: body depth at anal-fin origin 25–28; maximum head depth 26; head depth through eye 22; suborbital depth 14; head length 34; snout length 13; postorbital length 14; upper-jaw length 15; barbel length 28–30; barbel width 1.6–1.7; caudal-peduncle length 21–22; second dorsal-fin base length 14; caudal-fin length 28–32; length of anal-fin base 9.8–9.9; anal-fin height 15–17; pelvic-fin length 20; pectoral-fin length 28; pectoral-fin width 5.5–5.9; first dorsal-fin height 24–25; first two or three long dorsal-fin spines protruding; second dorsal-fin height 17; caudal fin without bars; first dorsal fin with black tip, retained in preservative; preserved fish with silvery-whitish head, pale brown body ventrally and dark brown dorsally; fins except for first dorsal-fin tip pale brown, partly hyaline; barbels very long, pale.
Etymology.—The new species name “alama” is used as a noun in apposition and honors the late Mr. Ulysses Banga Alama (1959–2023), who was employed at the College of Fisheries and Ocean Sciences, University of the Philippines Visayas, Miagao, Iloilo, Philippines, and manager of the UPVM fish collection, to commemorate his contributions to marine ichthyology.
Upeneus brevianalis Uiblein and Boonphienphol, new species
Phuket Goatfish
Diagnosis.—Dorsal fins VIII + 9; pectoral fins 16; gill rakers 8 upper + 20 lower = 28 total; lateral-line scales 31–32; body moderately deep; measurements as % SL: body depth at anal-fin origin 23; maximum head depth 27; head depth through eye 21; suborbital depth 14; head length 33; snout length 13; postorbital length 14; upper-jaw length 14; barbel length 20; barbel width 1.5; caudal-peduncle length 19; second dorsal-fin base length 15; caudal-fin length 31; length of anal-fin base 8.1; anal-fin height 12; pelvic-fin length 20; pectoral-fin length 26; pectoral-fin width 6.0; first dorsal-fin height 23, first five long dorsal-fin spines slightly protruding; second dorsal-fin height 17; caudal fin without bars; anterior third of first dorsal fin forming a broad, dark-brown vertically oriented band, becoming slightly wider dorsally; head and body pale brown to brown, belly lighter; paired and anal fins pale hyaline; barbels pale to pale brown.
Etymology.—The new species name “brevianalis” refers to the short anal fin, an important diagnostic character of this species.
Franz Uiblein, Hiroyuki Motomura, Peter R. Møller, Mark Sabaj, Supasit Boonphienphol, Tuan A. Hoang and Kerryn Parkinson. 2025. Two New Goatfish Species of the
Upeneus sulphureus Species Group (Mullidae), with a Redescription of
U. sulphureus Cuvier, 1829 and a Review of Barbel Length in the Genus.
Ichthyology & Herpetology. 113(3); 453-486. DOI:
doi.org/10.1643/i2024094 (22 August 2025)
