Upeneus asymmetricus Lachner, 1954

Uiblein, Franz & White, William T., 2015, A new goatfish of the genus Upeneus (Mullidae) from Lombok, Indonesia and first verified record of U. asymmetricus for the Indian Ocean, Zootaxa 3980 (1), pp. 51-66 : 56-63

Upeneus asymmetricus Lachner, 1954

Asymmetrical goatfish

( Figs 1 , 2 ; Tables 1–2)

Upeneus asymmetricus Lachner, 1954 ; Ben Tuvia & Golani, 1989; Kim & Nakaya, 2001; Uiblein & Lisher, 2013

Material examined (8 specimens, 74–100 mm SL): Philippines: USNM 154659, holotype, 75 mm, Pandanon Island, between Cebu and Bohol; USNM 154660, 2 paratypes, 74–79 mm, same locality; USNM 154661, paratype, 100 mm, off Western Samar, Catbalogan; Indonesia, Eastern Indian Ocean: CSIRO H 7417-02, 93 mm SL, Lombok , Tanjung Luar, fish market; CSIRO H 7417-01, 2 specimens, 97–98 mm SL, same locality; MZB 22711, 95 mm SL, same locality.

Diagnosis. Dorsal fins VII + 9; pectoral fins 12–14; gill rakers 7–8 + 19–21 = 26–28; measurements in % SL: body depth at first dorsal-fin origin 22–24; body depth at anus 20–22; caudal-peduncle depth 8.5–9.9; maximum head depth 19–20; head depth through eye 15–16; interorbital length 8.0–9.0; head length 26–29; snout length 9.9– 11; postorbital length 11–13; orbit length 5.7–7.7; upper jaw length 8.7–11; barbel length 17–19; caudal-peduncle length 22–24; caudal-fin length 27–30; anal-fin height 15–16; pelvic-fin length 19–22; pectoral-fin length 18–21; first dorsal-fin height 19–21; second dorsal-fin height 15–17; upper lobe of caudal fin with 4–6 red-brown oblique bars including 1 bar on or close to tip; lower caudal-fin lobe crossed by 6–8 red-brown oblique bars, most prominent along middle of lobe and much weaker or absent on inner (dorsal) and outer (ventral) side of lobe, leaving the the outermost (ventral) 3 rays mostly unpigmented hyaline; barbels pale reddish; body and head dorsally grey-green darkened and white silvery below eys and along ventral mid-body and caudal peduncle, with a yellow-green mid-body stripe of about pupil width from behind eye to caudal-fin base, stripe accompanied by a series of ca. 10 dark grey dots below and behind first dorsal fin; behind second dorsal fin a saddle-like darkened area extending from mid-body stripe towards dorsal body margin; dorsal fins with 3 red-brown stripes, the lowest close to body, the distal-most close to tip; preserved fish dark brown dorsally and pale brown ventrally, bars on both upper and lower caudal-fin lobes retained.

Distribution. First record for the Indian Ocean and for Indonesia. Prior to this record, Upeneus asymmetricus was known only from Pandanon (east of Cebu) and western Samar, Philippines.

Comparisons Upeneus asymmetricus differs from all congeneric species in the following combination of characteristics: 7 dorsal-fin spines, 12–14 pectoral-fin rays, 26–28 total gill rakers (19–21 rakers on lower limb), body depth at anal-fin origin 20–22% SL, pectoral-fin length 18–21% SL, and lower caudal-fin lobe with 6–8 redbrown oblique bars crossing middle of lobe, the innermost (dorsal) part of lobe as well as the outermost (ventral) 3 rays mostly unpigmented.

Upeneus asymmetricus differs from the meristically most similar species of the japonicus group as follows (comparative data in Table 1): from U. francisi , in fewer gill rakers, smaller head, shorter dorsal-fin bases, lower anal fin, and shorter pectoral fin; it differs from U. japonicus in shorter barbels, shorter pectoral fins, lower first dorsal fin, and lower caudal-fin lobe with oblique bars vs. no bars; it differs from U. lombok in deeper body at analfin origin, longer snout, higher anal and second dorsal fins, and oblique bars on lower caudal fin present vs. absent; it differs from U. pori in slightly shorter snout, slightly more pectoral-fin rays, presence of a dark saddle behind second dorsal fin in fresh fish, and in both fresh and preserved fish by presence of oblique bars vs. a stripe along middle of lower caudal-fin lobe and prominent bars along inner and outer lobe margin absent vs. present; it differs from U. saiab in having fewer gill rakers, deeper body, shorter head, presence of a dark saddle behind second dorsal fin vs. absence in fresh fish, and oblique red-brown bars along middle of lower caudal-fin lobe vs. a broad red stripe; and it differs from U. seychellensis in more developed gill rakers on lower limb, deeper body at anal-fin origin, wider interorbital, shorter snout, and oblique bars on lower caudal fin present vs. bars absent.

Furthermore, Upeneus asymmetricus differs from 14 other at least potentially co-occurring species as follows: from U. australiae , U. guttatus , and U. torres in more gill rakers (gill rakers on lower limb 19–21 vs. 16–19; total gill rakers 26–28 vs. 22–26), less body depth at dorsal-fin origin (22–24 vs. 18–23% SL), and oblique red-brown bars along middle of lower caudal-fin lobe vs. the lobe either almost entirely covered by an orange or red stripe ( U. torres ), or crossed by oblique bars at outside (ventrally) that connect to a stripe at inner side (dorsally) ( U. guttatus ), or fully crossed by brown or black oblique bars ( U. australiae ); it differs from U. moluccensis , U. quadrilineatus , and U. sulphureus in 7 vs. 8 dorsal-fin spines, fewer lateral-line scales (28–31 vs. 33–37), less body depth at first dorsal-fin origin (22–24 vs. 24–33% SL); shallower head (maximum head depth 19–20 vs. 20–25% SL; head depth through eye 15–16 vs. 16–20% SL), shorter pectoral-fin length (18–21 vs. 21–27% SL), and oblique bars on lower caudal-fin lobe vs. no bars; it differs from U. stenopsis , U. subvittatus , and U. vittatus in 7 vs. 8 dorsal-fin spines, shallower head (maximum head depth 19–20 vs. 21–26% SL; head depth through eye 15–16 vs. 17–20% SL), shorter head length (26–29 vs. 29–34% SL), shorter pectoral-fin length (18–21 vs. 21–26% SL), and oblique bars on middle of lower caudal-fin lobe vs. lobe entirely crossed by bars; it differs from U. luzonius , U. margarethae , U. sundaicus and U. tragula in 7 vs. 8 dorsal-fin spines, more gill rakers (gill rakers on lower limb 19–21 vs. 13–18; total gill rakers 26–28 vs. 18–24), and oblique bars on middle of lower caudal-fin lobe vs. either lobe entirely crossed by bars ( U. tragula , U. margarethae ), or bars covered by a broad red stripe (Western Indian Ocean U. margarethae ), or no bars or stripe ( U. sundaicus ) (fresh colour of U. luzonius not known); and it differs from U. nigromarginatus in 7 vs. 8 dorsal-fin spines, fewer lateral-line scales (28–31 vs. 36–37), shallower body (body depth at first dorsal-fin origin 22–24 vs. 31–35% SL; body depth at anal-fin origin 20–22 vs. 27–30% SL), shorter upper-jaw length (8.7–11 vs. 11–12% SL), and oblique bars on lower caudal-fin lobe vs. no bars.

The four specimens of U. asymmetricus from Lombok differ from the four Philippine type specimens in having smaller eyes, shorter jaw, narrower snout, shorter barbels, longer caudal fin, and longer pectoral fins ( Figure 2 B, D, Table 2). Upeneus asymmetricus attains 100 mm SL.

Remarks. Upeneus asymmetricus is most similar to U. pori , from which it differs mainly in lower caudal-fin lobe colour pattern. This colour pattern resembles U. australiae , which may lead to misidentifications, especially when body form and meristic character are not taken into account. Recent detailed taxonomic studies of a large number of specimens originally identified as U. asymmetricus from around Australia (see also Gloerfelt-Tarp & Kailola 1984; Sainsbury et al., 1985) were re-identified as U. australiae ( Uiblein & Gledhill, 2015) . Upeneus asymmetricus has also been cited from the South China Sea ( Randall & Lim, 2000; Randall, 2001), but no verified record from that area exists. This species was not identified from a large sample of goatfishes from off Vietnam of which many were photographed fresh (the specimens are now curated at the HIFIRE collection, Institute of Marine Research, Bergen, Norway). Apart from the four types, this species was also not seen when examining a large number of collection material from the Coral Triangle area (lodged e.g. at CAS, RMNH, ZMUC, and USNM).

The intraspecific differences in several body-form characters between the two geographic groups of U. asymmetricus may indicate population divergence that would be worthy of further investigation when more comparative material, colour of fresh and/or live fish, and tissue samples become available. Geographic comparisons of fresh colour patterns were not possible, as the species description of U. asymmetricus by Lachner (1954) was based on preserved material only and no description of fresh colour of the type material was made.

TABLE 1. Measurements and counts of Upeneus lombok n. sp., U. asymmetricus , and five other species of the japonicus group with high gill-raker count; differences from U. lombok n.sp. indicated in bold. those from U. asymmetricus in italics.

Upeneus lombok n.sp. U. asymmetricus U. francisi

Lombok Philippines and Indonesia Adult 4 subadults .. ....continued on the next page

TABLE 1. (Continued)

Upeneus lombok n.sp. U. asymmetricus U. francisi

Lombok Philippines and Indonesia Adult 4 subadults .. ....continued on the next page

TABLE 1. (Continued)

U. japonicus U. pori U. saiab U. seychellensis

E Malaysia to Korea Red Sea to Madagascar Mozambique Seychelles-Bank ......continued on the next page

TABLE 1. (Continued)

U. japonicus U. pori U. saiab U. seychellensis

E Malaysia to Korea Red Sea to Madagascar Mozambique Seychelles-Bank U. asymmetricus

Philippines (types) Lombok all ......continued on the next page

TABLE 2. (Continued)

U. asymmetricus

Philippines (types) Lombok all

Figures

FIGURE 1 (A) Upeneus lombok n. sp., paratype (MZB 22710), 51 mm SL, Lombok (W. T. White); (B) U. asymmetricus (CSIRO H 7417 - 02), 93 mm SL, Lombok (W. T. White); (C) Upeneus francisi (BPBM 34256), 56 mm SL, Norfolk Island (J. E. Randall); (D) U. pori, holotype (HUJ 13627), Eilat, Red Sea, 98 mm SL (D. Darom); (E) U. saiab, holotype (SAIAB 188298), 102 mm SL, Angoche, Mozambique (M. Lisher); (F) Upeneus seychellensis, paratype (SAIAB 84280), 102 mm SL, Seychelles Bank (O. Alvheim)

FIGURE 2. Relationships between SL and four body-form characters (A – D), between two body-form characters (E), and between a meristic and body-form character, respectively (F, G) in Upeneus lombok n. sp., U. asymmetricus, and five other species of the japonicus group with high gill-raker counts. The two geographic groups of U. asymmetricus from the Philippines and Lombok are indicated.

Tables

TABLE 2. Morphometric and meristic characters in types and newly recorded specimens of Upeneus asymmetricus; data revealing the most important population distinctions are emphasized in bold.

	Min	Mean	Max	n Min	Mean	Max	n Min	Mean	Max	n
SL (mm)	74	82.1	100	4 93	96.1	98	4 74	89.1	100	8
Body depth at first dorsal-fin origin	22	23.4	24	4 23	22.8	24	4 22	23.1	24	8
Body depth at anal-fin origin	20	20.8	22	4 20	19.9	20	4 20	20.4	22	8
Half body depth (from lateral line downwards) at first dorsal fin origin	19	19.5	20	4 19	19.2	20	4 19	19.3	20	8
Half body depth (from lateral line downwards) at anal fin origin	14	15.1	16	4 15	16.0	17	4 14	15.6	17	8
Caudal-peduncle depth	8.8	9.3	9.9	4 8.5	9.0	9.3	4 8.5	9.2	9.9	8
Caudal-peduncle width	3.7	4.1	4.8	4 3.9	4.5	5.0	4 3.7	4.3	5.0	8
Maximum head depth	20	20.1	20	4 19	19.3	20	4 19	19.7	20	8
Head depth across a vertical midline through eye	15	15.7	16	4 15	15.4	16	4 15	15.5	16	8
Suborbital depth	9.4	9.7	10	4 8.8	9.1	9.6	4 8.8	9.4	10	8
Interorbital length	8.0	8.6	9.0	4 8.2	8.6	9.0	4 8.0	8.6	9.0	8
Head length	27	27.7	29	4 26	26.5	28	4 26	27.1	29	8
Snout length	9.9	10.2	11	4 9.9	10.1	10	4 9.9	10.2	11	8
Postorbital length	11	11.5	13	4 11	11.3	12	4 11	11.4	13	8
Orbit length	7.0	7.4	7.7	4 5.7	6.1	6.4	4 5.7	6.7	7.7	8
Orbit depth	5.6	6.0	6.5	4 4.8	5.0	5.3	4 4.8	5.5	6.5	8
Upper-jaw length	9.6	10.2	11	4 8.7	9.1	9.6	4 8.7	9.6	11	8
Lower-jaw length	9.1	9.6	10	4 8.4	8.7	9.3	4 8.4	9.1	10	8
Snout width	8.1	8.8	9.3	4 7.6	7.7	7.9	4 7.6	8.2	9.3	8
Barbel length	18	18.6	19	4 17	17.3	18	4 17	17.9	19	8
Maximum barbel width	0.8	0.9	1.0	4 0.7	0.8	0.9	4 0.7	0.8	1.0	8
First pre-dorsal length	37	37.3	38	4 35	35.7	37	4 35	36.5	38	8
Second pre-dorsal length	64	64.9	66	4 62	63.7	64	4 62	64.3	66	8
Interdorsal distance	14	15.5	16	4 14	15.5	16	4 14	15.5	16	8