Enneapterygius altipinnis Clark, 1980

Holleman, Wouter & Bogorodsky, Sergey V., 2012, A review of the blennioid fish family Tripterygiidae (Perciformes) in the Red Sea, with description of Enneapterygius qirmiz, and reinstatement of Enneapterygius altipinnis Clark, 1980, Zootaxa 3152, pp. 36-60 : 41-46

Enneapterygius altipinnis Clark, 1980

Figs. 2 B, 3A & B, 4A & B; table 1 & 2

Enneapterygius altipinnis Clark, 1980: 99 –101, Figs. 3 b, 6c & 11 (Gulf of Aqaba and Ethiopia, Red Sea).

Enneapterygius tutuilae (non Jordan & Seale, 1906): Fricke, 1994: 285 ff, Figs. 58 & 59; Randall, Allen & Steene, 1997: 364; Holleman, 2005: 20 –21, Fig. 18, Pl. 2 I & J. Fricke, 1999: 468; Golani & Bogorodsky 2010: 44.

Enneapterygius pusillus (non Rüppell, 1835): Holleman, 1986: 757, pl. 116, fig. 236: 5 (Great Barrier Reef, Australia).

Diagnosis. Dorsal fins III + XII–XIII + 8–10 (usually III + XII + 8 or 9, rarely 10); anal fin I, 15–18 (rarely 15, usually 17); pectoral-fin 14, all rays simple. LL with 10–12 (usually 12) pored scales and 21–23 (usually 21) notched scales; total lateral scales 27 or 28; dentary pores 2 + 1 or 2 + 2, usually 2 + 2 + 2 ( Fig. 2 B). Head 3.3–4.0 [3.5] in SL; eye 2.6–3.2 [3.0] in head length.

Head, chest and pectoral-fin base naked; nape scaled; abdomen naked, but scales extend to base of dorsal and anal fins; single row of scales at base of caudal fin; supratemporal sensory canal crescent-shaped; supraorbital cirrus a rounded, flat ‘paddle’, its length about equal to pupil diameter; first dorsal fin higher than second in adult males and equal to slightly higher in females and in juveniles; longest pectoral-fin ray reaches junction between second and third dorsal fins; longest pelvic-fin ray reaches first anal-fin ray in some individuals.

Fresh colour. Males: body cream above, white ventrally, many scales edged in orange, with five irregular patches of olive-brown with melanophores, first from front of second dorsal fin and below pectoral fin, second from mid-second dorsal fin, third from junction of second and third dorsal fins, fourth at mid-third dorsal fin and last across peduncle at base of caudal fin; head cream with pale green markings, eye greenish with small white spots; first dorsal fin mottled cream and green, and with red patch at base of membrane between spine 3 and dorsum, spine 1 with alternating dark brown and white bands; second dorsal fin red anteriorly with 2 white bands, first near base of spines 1 and 2, second from tip of spine 2 to base of spine 4, middle of fin with a broad band of melanophores on hyaline membrane, followed by band of cream spots along posterior margin; third dorsal fin similar, but paler and without the red; caudal fin hyaline with white flecks on rays; anal with alternating bands of white, clear and black; pectoral fins with irregular bands of brown, white and green, colour on rays only; anal fin with 6 or 7 black subcutaneous spots at base and black pre-anal spot; pelvic fins whitish.

Females: very similar to males, but without red on dorsal fins.

Colour underwater. Individuals cryptic on sand, body with pale and olive-green bands; long first spine of the first dorsal fin banded black and white, membranes white, second dorsal fin with white band close to posterior half of margin and other white marks, third dorsal fin similar.

Key features. First dorsal fin tall; crescent-shaped supraoccipital sensory canal; underwater – tall first dorsal fin; white to yellowish band across the posterior portion of second dorsal fin, band continuing to LL.

Distribution. Egypt, Gulf Aqaba to Eritrea, but not recorded outside the Red Sea.

Habitat. In shallow water, collected material was taken from depth of 2–3 m, on dead corals near the base of reefs. Reaches 19.7 mm SL.

Remarks. The species was considered synonymous with Enneapterygius tutuilae Jordan & Seale by Fricke (1994: 285) and Holleman (2005: 20). In the Indian Ocean “ E. tutuilae ” — as then considered by these authors — is known from the coast of East Africa, Madagascar, and the Mascarene Islands, but not from the Maldives, India or Sri Lanka. Individuals of “ E. tutuilae ” have been recorded from the Cocos-Keeling and Christmas Islands, throughout Indonesia, tropical Australia, and in the Western Pacific to Fiji and Rotuma (see Fricke 1997: 339–349).

As in the species of the Helcogramma steinitzi complex ( Holleman, 2006: 91), there is little difference in morphometric counts between species of the Enneapterygius tutuilae complex distributed over this wide area (as shown in Table 2 below). There are, however, pigmentation differences suggesting that the various ‘forms’ are part of a widespread species complex, similar to those already described for the Helcogramma fuscopinna and H. steinitzi complexes (see also Williams & McCormick, 1990; Williams & Howe, 2003).

E. altipinnis from the Red Sea has no divided pectoral-fin rays (and Williams confirmed this for the types) and were incorrectly recorded by both Clark (1980: 100) and Fricke (1997: 338). All the Western Indian Ocean specimens of “ E. tutuilae ” examined by the first author have only simple pectoral-fin rays (see Holleman, 2005). Winterbottom (pers. comm.) confirmed that specimens from Raja Ampat also have only simple pectoral-fin rays. Photographs of a similar species Enneapterygius from the Ogasawara Islands, Japan, also show only simple pectoral-fin rays. This suggests simple pectoral-fin rays is a shared character for this species complex; E. kosiensis Holleman, 2005 is the only other Enneapterygius species known with only simple pectoral-fin rays.

* Data from Holleman 2005, from specimens from E. Africa, Comoro Islands, Mauritius, St Brandon Shoals, west Thailand, and a single specimen from Japan.

Specimens from East African and Mascarene Islands and from Indonesia eastwards appear to vary in colouration, and species may be sexually dichromatic. Pertinent to this paper, however, is to distinguish between specimens from the Red Sea and those from elsewhere in the Western Indian Ocean, and this distinction lies in colouration.

Presumed immature / sexually inactive males from the Comoro Islands have whitish bodies with dark brown to black markings, scale pockets edged in red, the first dorsal fin yellow ( Fig. 4 ). Mature / sexually active males have the head and anterior part of the body suffused with bright green, the colour extending to the base and lower part if the first dorsal fin and on the belly as far as the pre-anal spot. Scale pockets on the body tend to be more intensely coloured, as are the rays of the pectoral fins, and the black spot between spines 2 and 4 of the second dorsal fin, with its ‘trailing tail’ of melanophores, rather like a black comet and tail. The orbital cirri are deep yellow. By comparison, an intensely coloured male E. altipinnis from the Red Sea lacks the green on the anterior of the body, has less intensely coloured body bars, but with the membrane between the third spine of the first dorsal fin and the dorsum with crimson and the anterior portion of the second dorsal fin crimson, as a triangle with the first spine and the bases of first 7 spines forming two of the sides, the third side is an irregular, narrow cream band.

It is now well established that there are a number of tripterygiid species complexes, with closely related species often differing little in meristic counts and measurements, but in subtle ways in colour patterns. For example, the three currently recognised species of the Helcogramma steinitzi species complex share the micromelanophores of the first dorsal fin and an overall red body colour in males ( Holleman 2006; see also Williams & McCormick, 1990 and Williams & Howe, 2003). The species of the Enneapterygius tutuilae species complex share the following characters: only simple pectoral-fin rays, a characteristic black area on the second dorsal fin, followed by a pale to golden band that runs from the fin onto the body, to or just beyond the midline, and six or seven dark or black bands across the anal fin.

On the basis of the difference in colour pattern and distribution Enneapterygius altipinnis Clark and is recognised as a valid species, distinct from similar fish from the east coast of Africa and westernmost Western Indian Ocean islands.

Comparative material examined. Enneapterygius altipinnis : SAIAB 88749 (3: 16.8–20.3 mm SL), Shams Alam Reef, S. Egypt, S. Bogorodsky, 0 8 June 2008; SAIAB 88750 (3: 18.2–19.7 mm SL), El Quseir, Egypt, S. Bogorodsky, 0 8 June 2008; SAIAB 88751 (16.3 mm SL), Shaab Suadi, Sudan, S. Bogorodsky, 12 October 2009; SMF 33608 (15.1 mm SL), Rabigh, Saudi Arabia (23°02.839' N, 38°46.621' E), S. Bogorodsky, 0 9 April 2011.

Enneapterygius cf. tutuilae : SAIAB 31515 (3: 15.0– 18.6 mm SL), Pinda Island, Mozambique; SAIAB 31520 (14.8 & 15.2 mm SL) Mauritius.

Figures

FIGURE 2. Dentary pore patterns of Red Sea species of Enneapterygius: (A) E. abeli; (B) E. altipinnis; (C) E. clarkae; (D) E. destai; (E) E. obscurus; (F) E. pallidus (diagrammatic - J. T. Williams); (G) E. pusillus; (H) E. qirmiz; (I) E. ventermaculus.

FIGURE 3. Enneapterygius altipinnis, M (A) grey form; (B) green form. Photos by S. V. Bogorodsky.

FIGURE 4. (A, B) Enneapterygius altipinnis, M, Rabigh, Saudi Arabia; (B) Enneaterygius altipinnis, F, Marsa Alam; (C, D) Enneapterygius cf tutuilae, Comoros. Photos by S. Bogorodsky (A, B); R. Winterbottom (C, D).

Tables

TABLE 2. Comparative counts for E. altipinnis sensu Clark and E. tutuilae sensu Jordan & Seale.

	E. altipinnis	E. tutuilae
Dorsal fins	III + XII–XIII + 8–10, usually XII + 8 or 9, rarely 10 rays (Clark 1980; this study)	III + XII + 9 (Jordan & Seale 1905) III + XI–XII + 9–10, usually III + XII + 10 (Holleman 2005)*
Anal fin	I, 15–17, usually 17 rays, rarely 15 (Clark 1980); 17 or 18 (this study)	18 (I, 17?) (Jordan & Seale 1905) I, 15–17, usually 17 (Holleman 2005)
Pectoral fins	13–15, usually14–15, usually 14, all simple (not from Clark — see below)	14, all simple (Holleman 2005)
Lateral line	10–12 (usually 12) + 21–23 (usually 21) (Clark 1980)	33 lateral scales (Jordan & Seale 1905) 11–12 + 20–23 (usually 12 + 22) (Holleman 2005)
Dentary pores	2 + 2 + 2 (types, Williams pers. comm); occasionally 2 + 1 + 2 (this study)	2 + 1 / 2 + 2 (Holleman 2005)