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

Abstract

The fishes of the blennioid family Tripterygiidae of the Red Sea are reviewed. Twelve species are recognised: Norfolkia brachylepis, Helcogramma obtusirostris, H. steinitzi and nine species of Enneapterygius. Enneapterygius qirmiz sp. nov. is described for a species originally described, but not named, by Clark (1980), of which the specimens on which she had based her description had become lost. Clark’s (1980) Enneapterygius altipinnis is re-instated as a valid species, and its inclusion as one of a complex of several species is discussed. Enneapterygius ventermaculus reported as first record for
the Red Sea. Detailed colour descriptions are given for all species, including their colour underwater where possible, as many of the species are cryptic and difficult to identify.

Full text

36 Accepted by M. Craig: 2 Nov. 2011; published: 6 Jan. 2012
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2012 · Magnolia Press
Zootaxa 3152: 36–60 (2012)
www.mapress.com/zootaxa/Article
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
WOUTER HOLLEMAN1 & SERGEY V. BOGORODSKY2
1South African Institute for Aquatic Biodiversity, Grahamstown, South Africa. E-mail w.holleman@saiab.ac.za
2Station of Naturalists, Omsk, Russia. E-mail ic187196@yandex.ru
Abstract
The fishes of the blennioid family Tripterygiidae of the Red Sea are reviewed. Twelve species are recognised: Norfolkia
brachylepis, Helcogramma obtusirostris, H. steinitzi and nine species of Enneapterygius. Enneapterygius qirmiz sp. nov.
is described for a species originally described, but not named, by Clark (1980), of which the specimens on which she had
based her description had become lost. Clark’s (1980) Enneapterygius altipinnis is re-instated as a valid species, and its
inclusion as one of a complex of several species is discussed. Enneapterygius ventermaculus reported as first record for
the Red Sea. Detailed colour descriptions are given for all species, including their colour underwater where possible, as
many of the species are cryptic and difficult to identify.
Key words: Red Sea, Tripterygiidae, Enneapterygius altipinnis, Enneapterygius qirmiz sp. nov.
Introduction
The colour patterns of many fish species are described from specimens ‘out of water’. This can raise several prob-
lems: some specimens loose colour quite fast; some species change colour when under stress; water selectively fil-
ters out longer wavelengths of light, and, while some species may seem very ‘obvious’ out of water, their colour
patterns render them cryptic under water. This last is particularly so of small species such as Tripterygiidae, many
of which are semi-translucent, and the authors experienced some difficulty identifying species of the family that
had been photographed in the Red Sea. One of the purposes of this review is to provide both colour descriptions
and photographs of Red Sea tripterygiid species both ‘out of the water’ and underwater, where these are available.
There is also considerable variation in colour in many species of tripterygiids. Males and females of most species
have different colour patterns, and more intense colours of some males being a mark of sexual maturity / reproduc-
tive activity (e.g. Wirtz, 1978; Clements, 2006).
The first tripterygiid was described from the Mediterranean Sea, Blennius tripteronotus (Risso, 1810) and Trip-
terygion nasus (Risso 1827), giving the family its name. The first tripterygiid from the Red Sea, Enneapterygius
pusillus, was described shortly thereafter, by Rüppell in 1835. However, by 19 , when Clark reviewed the family
in the Red Sea, only one further species, Enneapterygius abeli Klausewitz 1960, had been described. Clark
described and named eight new species and listed two more, but without names as the types had become lost.
In 1982 Holleman described two new species of Enneapterygius from the Western Indian Ocean, one of them
being Clark’s unnamed sp.2. Holleman (1991) revised Norfolkia and synonymised Clark’s Norfolkia springeri with
N. brachylepis. Randall (1995) reviewed the tripterygiids of the Gulf of Oman, described two new species of Enne-
apterygius and extended the range of E. ventermaculus from the NW coast of India to the Gulf.
In a review of the Enneapterygius species of the Western Indian Ocean the first author identified colour photo-
graphs of a tripterygiid from the Red Sea made by Hackenberg (Debelius, 1998: 174) as E. obscurus (Holleman,
2005). Another photograph of the species, also taken by Hackenberg, was reproduced in Lieske & Myers (2004:
177). Subsequent photographs of the species made by the second author show the identification to be incorrect and

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REVIEW RED SEA TRIPTERYGIIDAE
that Hackenberg’s photographs are in fact of Clark’s unnamed sp.1, which is herein described and named. In that
review the first author — in agreement with Fricke (1994) — considered E. altipinnis Clark, 1980 a junior syn-
onym of E. tutuilae Jordan and Seale, 1906. Exploration of the Red Sea by the second author over the past few
years has produced a selection of photographs and collected specimens of tripterygiids from the area. Some of
these called into question the validity of this synonymy, as the Red Sea species has a different colour pattern com-
pared to specimens from Samoa, the type locality of E. tutuilae, and from specimens from Western Indian Ocean
islands.
Most Western Indian Ocean species of Enneapterygius (and from elsewhere — see Fricke, 1994 and 1997) dif-
fer very little in fin element counts, often by a single spine or ray, and with little variability, these counts invariably
overlap. Such counts are thus rarely of diagnostic value, and species identification relies on colour patterns, height
of the first dorsal fin, dentary pore patterns, form of the supraoccipital sensory canal, and sometimes lateral line
scale counts. Thus, while there are no differences in counts, pore patterns or fin heights between ‘tutuilae’ speci-
mens from the Red Sea, Western Indian Ocean Islands and elsewhere, the disjunct distributions and differences in
colour pattern lead us to consider E. altipinnis a distinct species, and one of a species complex which is distributed
throughout the Indo-Western Pacific Ocean.
Methods and materials
Only the following measurements were made: standard length (SL), head length and eye diameter, the horizontal
measure inside the bony orbit. Making other measurement on fishes this small with a dial caliper is inherently inac-
curate. Proportions of head length in SL and eye diameter in head length are given, means being shown in brackets.
Fin spine and ray counts, and vertebral counts were made directly from specimens and from radiographs. Pectoral-
fin ray counts are given as the total count on the left side, and, when branched rays are present, the number of
unbranched rays, the number of branched rays, and the number of branched rays, beginning with the dorsalmost
ray. Body scales are deciduous and often lost, particularly in specimens less than 20 mm SL; lateral line (LL) scales
were counted on the left side where scales were present, and scale pockets were included where these could be dis-
tinguished. The nape may be naked (no scales between the anteriormost lateral-line scales and the first dorsal fin);
partially scaled (at least one row of scales above the lateral line, but not to base of first dorsal fin or supraoccipital
sensory canal, or scaled, with scales to base of first dorsal fin and to immediately behind the supraoccipital sensory
canal. There are usually one or more rows of scales at the base of the caudal fin.
Dentary pore patterns have been found to be consistent for a species and are often species specific (e.g. Hansen
1986, but referred by her and others as ‘mandibular’ pores). These are most easily detected when stained with cya-
nine blue.
The underwater photographs were made by the second author, by John E. Randall, and by Richard Field. Juer-
gen Herler of the University of Vienna provided the tank photograph of Enneapterygius destai. No photographs of
Norfolkia brachylepis from the Red Sea was available and a photograph of the species from Bali by G. R Allen is
used. Depending on the water depth and source of light, species can look very different. The same is true of speci-
mens photographed against black or white backgrounds. Thus, identifying species from underwater photographs is
often not easy, and one is reliant on particular colour pattern characteristics. Thus for example, female E. abeli can
be distinguished by the dark line from the eye to the upper lip and the small orange bar below the eye.
All specimens were collected using rotenone or by hand net.
The following institutional abbreviations are used: BPBM – Bishop Museum, Honolulu; HUJ—Hebrew Uni-
versity, Jerusalem; SAIAB—South African Institute for Aquatic Biodiversity; SMF—Senckenberg Forschungsin-
stitut und Naturmuseum; SMNS—Staatliches Museum für Naturkinde Stuttgart; USNM—National Museum of
Natural History, Smithsonian Institution.
Table 1 gives a summary of selected characters of the species that occur in the Red Sea, Fig. 1 shows localities
referred to, and Fig. 2 shows the dentary pore patterns of the species of Enneapterygius.

HOLLEMAN & BOGORODSKY
38 · Zootaxa 3152 © 2012 Magnolia Press
TABLE 1. Comparative characters for Red Sea tripterygiid species.
All data taken from Holleman 1991, 2005 & 2007; Clark 1980: 101 (Enneapterygius n. sp. 1) records upper and lower 5 rays
unbranched.
Key to the males of Red Sea species of Tripterygiidae
1a. First dorsal fin with three spines; no scales on head. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
1b. First dorsal fin with four spines; head with scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Norfolkia brachylepis
2a. LL divided into anterior section of pored scales and posterior section of notched scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
2b. LL only of pored scales, ending below second or third dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
3a. Males red in life, head dark with broad, pale blue mark above lip and on lower preopercle; females translucent greenish; body
with indistinct brownish-red bars and scattered blotches; membrane of first dorsal fin of both sexes cream with reddish mark-
ings and many micro-melanophores; dentary pores 3 + 1–2 + 3; underwater males with indistinct brownish-red bars and scat-
tered blotches. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Helcogramma steinitzi
3b. Males dark grey to dark red in life; lower half of head with many melanophores and bright blue line from corner of mouth,
angling posteroventrally across cheek onto preopercle; both sexes with distinct, irregular bright red bars; first dorsal-fin mem-
brane translucent with some iridescence; dentary pores 4 + 1 + 4; underwater with distinct irregular, bright red bars in both
sexes, first dorsal-fin membrane translucent with some iridescence . . . . . . . . . . . . . . . . . . . . . . . . .Helcogramma obtusirostris
4a. Body and anal fin with bars. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5
4b. Body without bars; anal fin with narrow bars or without bars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
5a. Body with four dark bars usually divided below LL and continuing onto anal fin, and two paler bars below pectoral fin; nape
Species D2 & D3 A P LL Mandib
pores Ht D1 Nape/
belly
scales
Supra-
temp.
canal
E. abeli XI–XIII+9–10
(XII+10) I, 17–18
(I, 18) 15:
2-4+4+6-7 12–14+19–23
(13+21–22) 3+1+3 =D2 m
<D2 f scaled/
naked ‘C’
E. altipinnis XI–XII+9–10
(XII+10) I, 16–18
(I, 17) 14, all simple 11–12+20–23
(12+22) 2+1/2+2 >D2 m
=D2 f scaled/
naked ‘C’
E. clarkae XI–XII+8–10
(XII+9) I, 16–17
(I, 16) 15: 2+6+7 11–12+20–22
(12+22) 3+2+3 >D2 m
=D2 f scaled/
scaled ‘C’
E. destai XI–XIII+8–9
(XII+8–9) I, 15–17
(I, 16) 15: 3+5+7 8–12+21–23
(10–11+21–23) 3+2+3 <D2 scaled/
naked ‘C’
E. obscurus XII–XIV+9–10 I, 16–18
(I, 17–18) 14–15:
3–4+5+6 10–12+21–22 2+2+2 =D2 m
<D2 f scaled/
naked open
‘U’
E. pallidus XIII–XIV+10–
11 I, 21–22 13–14:
3+5–6+5 10–11+28 2+2+2 =D2 m
<D2 f naked/
naked ‘U’
E. pusillus XII–XIV+10–11
(XIII+10–11) I, 20–21
RS 17–21
(I, 19)
13–14:
3–4+3–4+
6–7
12+26–28 2+2+2 >D2 m
=D2 f naked/
naked ‘U’
E. qirmiz* XII–XIII+9–10 I, 16–18
(I, 17) 14–15: usually
2+6+6 9+ ~20 2+2+2 ~D2 m
<D2 f scaled/
naked ‘C’
E. venter-
maculus XI–XIII+8–10 1, 17–21
(I, 19) 14: usually
2+5+7 15+23
Oman 14+21–
22
3+1+3 >D2 m
<D2 f naked/
naked ‘U’
Total lat.
scales Mandib.
pores Nape
H. obtusiro-
stris XII–XIII+9–10
(XIII+10) I, 18–19
(I, 19) 16: 2+7+7 20–23
(21–22) 36–38
(37–38) 4+1+4 naked
H. steinitzi XII–XIV+10–12 I, 19–21
(20) 16: 2+7+7 21–27 37–42 3+1–2
+3 scaled
D1–D3
N. brachyle-
pis IV+XII–XV+9–
11
(IV+XIII+9)
II, 18–20
(II, 19) 16: 2+7+7 14–18+18–23
(16–17+18–23) 31–34 4+1+4

Zootaxa 3152 © 2012 Magnolia Press · 39
REVIEW RED SEA TRIPTERYGIIDAE
and belly scaled; yellow and brown in life . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Enneapterygius clarkae
5b. Body with dark bars, darkest across peduncle, with black spots above and often below; anal fin with five oblique black bars,
prominent subcutaneous black spots at base of anal fin, and black pre-anal spot; belly without scales; pinkish in life . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enneapterygius destai
6a. Anal fin with narrow bars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
6b. Anal fin without bars. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
7a. First dorsal fin of males higher than second . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7b. First dorsal fin subequal to second; head of males dark olive, body olive green with white blotches, anal fin barred black and
white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enneapterygius ventermaculus
8a. Third dorsal fin with 8–9 rays; supraoccipital sensory canal ‘C’-shaped (A); all pectoral-fin rays simple; in life base of ventral
fins and fins white, head and body speckled with orange, first dorsal fin with crimson . . . . . . . . . . . Enneapterygius altipinnis
8b. Third dorsal fin with 10–11 rays; supraoccipital sensory canal ‘U’-shaped (B); base of ventral fins and proximal part of rays
black in males; some pectoral-fin rays divided; in life face red, first dorsal fin yellow and sometimes very tall . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Enneapterygius pusillus
9a. Nape scaled, anal fin with fewer than 20 rays. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10
9b. Nape naked, anal fin with 21–22 rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Enneapterygius pallidus
10a. Dentary pores 2 + 2 + 2. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10b. Dentary pores 3 + 1 + 3, males bright yellow in life, head with many melanophores to solid black, females with body scales
edged in orange-brown, forming narrow irregular bars; orange bar below eye . . . . . . . . . . . . . . . . . . . . . .Enneapterygius abeli
11a. Small black spot on upper caudal peduncle; no patch of melanophores on dorsum; anterior LL scales 10–12; males translucent
and red in life, first dorsal fin white to cream, females translucent, scales edged in pink . . . . . . . . . . .Enneapterygius obscurus
11b. No black spot on upper caudal peduncle; small oval patch of melanophores on dorsum between 3rd and 4th dorsal-fin spines;
anterior LL scales 8–10; crimson in life, first dorsal fin crimson anteriorly with gold spots, and white posteriorly, males with
rectangular blotch of melanophores on midside, below second dorsal fin . . . . . . . . . . . . . . . . .Enneapterygius qirmiz sp. nov.
Species accounts
Genus Enneapterygius Rüppell, 1835
Enneapterygius Rüppell, 1835: 2; type species Enneapterygius pusillus Rüppell, 1835, by monotypy.
Diagnosis. Small fishes with fusiform bodies, generally less than 25 mm SL in the Western Indian Ocean. First
dorsal fin with III spines, second with XI–XIV spines, third with 8–11 simple rays, except for last which is divided
to its base; anal fin with a single short spine and 15–22 simple rays, except for the last which is divided to its base;
pelvic fins with one short, hidden spine and two simple rays, often partially joined by membrane. LL with an ante-
rior section of pored scales and a posterior section of scales with a notch in the posterior margin, starting from 1–2
scale rows below end of pored section and continuing to base of caudal fin. Head without scales; nape scaled, par-
tially scaled or naked, belly naked in most species. Supratemporal sensory canal either an open ‘C’-shape, ‘U’-
shape, or an open ‘U’-shape, curving around the first dorsal-fin spine (see figure in Key above). Orbital and ante-
rior nasal cirri simple, the nasal cirrus on the posterior edge of a short tube. Teeth fixed, conical and slightly
recurved, larger in front; vomer with a single row of teeth, palatines without. Currently 61 recognised species, nine
species in the Red Sea.

HOLLEMAN & BOGORODSKY
40 · Zootaxa 3152 © 2012 Magnolia Press
FIGURE 1. Map of the Red Sea showing localities mentioned in the text.
Enneapterygius abeli (Klausewitz, 1960)
Figs. 2A, 5A–C; table 1
Tripterygion abeli Klausewitz, 1960: 11, Figs. 1–2 (Al-Ghardaqa, Egypt, Red Sea).
Helcogramma abeli: Lal Mohan, 1971: 222.
Enneapterygius abeli: Clark, 1980: 97, Figs. 2a, 2b, 4c, & 10; Holleman, 1986: 756, Fig. 236.2; 2005: 4–5, Fig. 2; Fricke, 1999:
468; Golani & Bogorodsky, 2010: 44.
Diagnosis. Dorsal fins III + XI–XIII + 9–10 (usually III + XII + 10); anal fin I, 17–18 (usually 18 rays); pectoral-
fin 15: usually 3, 5, 7. LL 12–14 pored scales and 20–23 (usually 22) notched scales; longitudinal scale series 31;
dentary pores 3 + 1 + 3 (Fig. 2A). Head 3.2–3.8 in SL; eye 2.9–3.7 in head length. Nape partially scaled, abdomen
without scales, single row of scales at base of caudal fin; supratemporal sensory canal crescent-shaped; supraorbital
cirrus small and pointed; first dorsal fin about equal in height to second in males, lower in females.
Fresh colour. Males with semi-translucent, deep yellow body, scale margins orange-red; head, chest, and base
of pectoral and pelvic fins of males with numerous melanophores (entire head becomes black during courtship);
eye with alternating brown and greenish bands; first dorsal-fin membranes white, suffused with yellow and with
some melanophores along margin; second dorsal-fin membranes translucent with spines yellow and orange, some
small melanophores in patches along margin; third dorsal fin similar with patches of small melanophores about
half-way in from margin; caudal fin pale yellow and orange on rays; anal fin lemon yellow with orange spots at
base of alternate rays; pectoral-fin rays deep yellow, membranes pale yellow; pelvic fins yellow with some black
proximally and at base.
Females: body translucent with scale margins orange-brown, forming irregular bars, darkest at caudal-fin base;
head and nape brownish, snout and lower half of head white, dark brown stripe from upper lip to anterior margin of

Zootaxa 3152 © 2012 Magnolia Press · 41
REVIEW RED SEA TRIPTERYGIIDAE
eye, vertical orange mark below eye, orange marks on opercle and pectoral-fin base; first dorsal fin greenish with
two orange bars on first spine; second and third dorsal fins with oblique banding, colour on rays only, anal and pec-
toral-fin rays pale yellow, all fin membranes transparent.
Colour underwater. Males semi-translucent greenish yellow, scale margins orange-brown, head with scat-
tered melanophores to black, eye with orange-brown and green radii, all fins transparent, spines and rays of dorsal
and caudal fins translucent, rays of anal and pectoral fins pale yellow, in moderate to faintly pigmented males a
brown stripe from corner of eye to upper lip; females paler than males, top of head greenish and without melano-
phores, brown stripe from corner of eye to upper lip, head below eye white with cluster of orange spots below eye.
Both males and females with vertebral column coloured in alternating brown and white, the white often quite
bright.
Key features. Low first dorsal fin, brown and green radii around eye, males with yellow body and melano-
phores on head to base of pectoral fins; females with yellow-green body, brown stripe on nose and orange bar
below eye.
Distribution. Known from the Red Sea, where it is common in the north, the east coast of Africa to as far south
as KwaZulu-Natal, the Seychelles, southern Madagascar, Réunion and Mauritius, St Brandon Shoals and the Cha-
gos Archipelago; common in some areas.
Habitat. Shallow, sheltered waters, on hard corals. E. abeli are found on the upper part of coral reefs, usually
along the reef margin in depths of about 1 m, often on corals covered by low algae. Found in bays or sheltered
lagoons. E. abeli attains 25 mm SL.
Material examined. Additional to that listed in Holleman 2005: SAIAB 88711 (5: 15–21 mm SL), Mangrove
Bay, El Quseir (25.52°N, 34.24°E), collected S. Bogorodsky, 8 &10 June 2008; SAIAB 88721 (18.1 mm SL),
Abingdon Reef, Sudan (20.50°N, 37.25°E), depth 14 m, collected S. Bogorodsky, 15 October 2009.
Enneapterygius altipinnis Clark, 1980
Figs. 2B, 3A & B, 4A & B; table 1 & 2
Enneapterygius altipinnis Clark, 1980: 99–101, Figs. 3b, 6c & 11 (Gulf of Aqaba and Ethiopia, Red Sea).
Enneapterygius tutuilae (non Jordan & Seale, 1906): Fricke, 1994: 285ff, 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. 2B). 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 cir-
rus 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 dor-
sum, 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 mel-
anophores 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.

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42 · Zootaxa 3152 © 2012 Magnolia Press
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.
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.
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

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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).
FIGURE 3. Enneapterygius altipinnis, M (A) grey form; (B) green form. Photos by S.V. Bogorodsky.
As in the species of the Helcogramma steinitzi complex (Holleman, 2006: 91), there is little difference in mor-
phometric 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. stein-
itzi 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 speci-
mens of “E. tutuilae” examined by the first author have only simple pectoral-fin rays (see Holleman, 2005). Win-
terbottom (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 pecto-
ral-fin rays. This suggests simple pectoral-fin rays is a shared character for this species complex; E. kosiensis Hol-
leman, 2005 is the only other Enneapterygius species known with only simple pectoral-fin rays.

HOLLEMAN & BOGORODSKY
44 · Zootaxa 3152 © 2012 Magnolia Press
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).
TABLE 2. Comparative counts for E. altipinnis sensu Clark and E. tutuilae sensu Jordan & Seale.
* 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 coloura-
tion, and species may be sexually dichromatic. Pertinent to this paper, however, is to distinguish between speci-
mens 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 com-
parison, 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 dor-
sum 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
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)

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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 recogn-
ised as a valid species, distinct from similar fish from the east coast of Africa and westernmost Western Indian
Ocean islands.
FIGURE 5. (A) Enneapterygius abeli, M, El Quseir; (B) Enneapterygius abeli, F, Marsa Alam; (C) Enneapterygius abeli, M,
El Quseir; (D) Enneapterygius clarkae, El Quseir; (E) Enneapterygius destai, M, Al Lith, Saudi Arabia; (F) Enneapterygius
destai, F, Gulf of Aqaba. Photos by J. Herler (F); S.V. Bogorodsky (the others).
Comparative material examined. Enneapterygius altipinnis: SAIAB 88749 (3: 16.8–20.3 mm SL), Shams
Alam Reef, S. Egypt, S. Bogorodsky, 08 June 2008; SAIAB 88750 (3: 18.2–19.7 mm SL), El Quseir, Egypt, S.
Bogorodsky, 08 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, 09 April 2011.

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46 · Zootaxa 3152 © 2012 Magnolia Press
Enneapterygius cf. tutuilae: SAIAB 31515 (3: 15.0–18.6 mm SL), Pinda Island, Mozambique; SAIAB 31520
(14.8 & 15.2 mm SL) Mauritius.
Enneapterygius clarkae Holleman, 1982
Figs. 2C, 5D; table 1
Enneapterygius clarkae Holleman, 1982: 121, Fig. 6 (Inhaca Island, Mozambique). Holleman 2006: 6, Fig. 4, Pl.1C. Golani &
Bogorodsky, 2010: 44.
Enneapterygius n. sp. 2: Clark, 1980: 104, Figs. 4a & 14.
Diagnosis. Dorsal fins III + XI–XII + 8–10 (usually III + XII + 9); anal fin I,16–17 (usually 16); pectoral fin 15:
usually 2, 6, 7. LL 11–12 (usually 12) pored scales and 20–22 (usually 22) notched scales, notched segment starting
next scale row below end of pored segment; total lateral scales 29 or 30; dentary pores 3 + 2 + 3 (Fig. 2C). Head
length 3.3–3.9, body depth 4.4–5.2 in SL; eye 2.8–3.6 in head length. Nape scaled; belly with thin cycloid scales;
single row of cycloid scales on pectoral fin base, parallel with margin of branchiostegal membrane. Supratemporal
sensory canal crescent-shaped. Orbital cirrus length about half pupil diameter and either rounded or divided; upper
and posterior margins of orbits with fine serrations. First dorsal fin equal in height to second in females, about 10%
higher in males. Pelvic-fin rays united for less than half their length, the longest ray reaching almost to vent. Mouth
slightly down-turned and reaches vertical through anterior of pupil.
Fresh colour. Males: body pale yellow with 5 or 6 clearly-defined dark brown bars, usually divided ventrally,
and continue across the anal fin, anterior-most two very close together and sometimes forms single, broad bar and
1st bar paler than others, last bar across peduncle; scale pockets anteriorly outlined in black; head yellowish with
brown markings between eyes, below eye and on opercle; first dorsal fin dusky; second with two dark red bands as
extension of body bars; third with faint black on rays, as extension of body bars; anal fin with 6 or 7 brown to black
bands (extensions of body bars), with subcutaneous black pots at base; pectoral fins hyaline with black banding,
colour on rays only; pelvic fins cream.
Females: paler versions of males and with little red on the dorsal fin.
Colour underwater. Body semi-iridescent yellowish-green with 5 or 6 brown bars; top of head dark brown,
opercle brown, head below eye white with two brown bars, sides of snout brown, interorbital green, extending in a
stripe to the tip of the mouth, orbital and nasal cirri green, eye mottled green with thin yellow and red ring around
the pupil. First dorsal fin spine banded brown and white; second and third dorsal and caudal-fin rays reddish, mem-
branes largely transparent; anal fin with 7 or 8 brown bars; pectoral fins with red and green on rays, and green
blotches with melanophores on the base.
Key features. Diagonal brown bars across body with reddish dorsal and caudal fins.
Distribution. Red Sea, east coast of Africa, to northern KwaZulu-Natal.
Remarks. E. clarkae may be confused with E. destai as preserved specimens. E. destai, however, lacks scales
on the belly and pectoral-fin base, has much darker anal fin bars and is a smaller species, with a maximum recorded
length of 21 mm SL, whereas E. clarkae attains 27 mm SL. Both these species live in close association with hard
corals with many nooks and crannies in sheltered lagoons and both are very secretive.
Material examined. Additional to that listed in Holleman, 2005: SAIAB 88712 (17.1 mm SL), Mangrove Bay,
El Quseir, Egypt (25.52°N, 34.24°E), collected S. Bogorodsky, 10 June 2008.
Enneapterygius destai Clark, 1980
Figs. 2D, 5E & F; table 1
Enneapterygius destai Clark, 1980: 102, Figs; 4b & 13 (northern end of Delemone Island, Eritrea, Red Sea). Holleman, 2005:
7; Golani & Bogorodsky, 2010: 44.
Diagnosis. Dorsal fins III + XI–XIII + 8–9; anal fin I, 15–17 (usually 16); pectoral-fin 15: 3, 5, 7; LL 8–12 pored
scales and 21–23 notched scales; total lateral scales 28–29; dentary pores 3 + 2 + 3 (Fig. 2D). Head length 3.1–3.4
[3.2], body depth 4.5–5.2 in SL; eye 2.5–3.0 [2.7] in head length. Nape scaled, belly without scales; supratemporal

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sensory canal crescent-shaped; orbital cirrus a small, rounded flap with a serrate distal margin. First dorsal fin
about 60% height of second in both males and females; pelvic-fin rays not united by membrane.
Fresh colour. Males: body of mature / sexually active males cream with 6 oblique, dark brown bars, most
prominent above LL, tending to break up below, last across caudal peduncle with two black spots on upper and
lower part, upper spot larger and darker than the lower; belly white with prominent, black pre-anal spot. Head dor-
sally brown, below eyes cream with irregular greenish brown stripe on side of snout and bar below eye, orbital cirri
brown. First dorsal fin crimson anteriorly, but grey between third spine and dorsum; second dorsal fin densely cov-
ered with melanophores, first spine banded brown and white, other spines reddish brown; third dorsal fin irregu-
larly banded in light red, grey, yellowish and transparent areas, parallel with margin; caudal fin with yellow and
orange, but with clear band at base; anal fin with 5 black bands, broadest and most intense anteriorly, narrowest and
palest posteriorly, rays pale yellow proximally, orange distally, most intense colour on posterior rays; pectoral-fin
base brown, membranes transparent, rays with alternating yellow and orange bands; pelvic fins white. Less darkly
coloured males lack the crimson on the first dorsal fin, the fin being mottled pink and white, with a pink flush
behind the head and on opercle and pectoral-fin base; lower black spot on caudal peduncle very small to absent.
Females: body bars brown anteriorly and yellow posteriorly; head pale with pink above and onto opercle, with
many small brown spots; first dorsal fin pink with first spine banded brown and white; second dorsal fin with broad
bands of brown spots, hyaline between and white along anterior ¾ of margin; third dorsal fin yellowish bands with
white patches; caudal-fin rays deep yellow with irregular narrow white band and with white line at base; anal fin
with some black at base, and with white, yellow and pink on rays; pectoral-fin rays mostly white, upper ones with
orange distally, base pink with brown spots; pelvic fins white.
Colour underwater. A single photograph taken by Juergen Herler of a male in an aquarium shows the body
barred in brown and white, the white most prominent above the midline; head with cream coloured with a brown
band below the eye and brown over a pink sheen on the opercle, the pink continuing onto the pectoral-fin base; first
dorsal-fin spine banded dark brown and white, the fin mostly pink; second and third dorsal fins hyaline except for
white markings and red and white on the spines and rays; caudal fins red with white spots, anal fin banded black
and white, and pectoral fins with four irregular whitish bars, the first at the base of the rays, rays pinkish.
Key features. Brown bars on upper half of body with one or two black spots on caudal peduncle, upper one
largest; five black blotches on anal fin, anteriormost largest and darkest; face pink with brown bar below eye, first
dorsal fin crimson in males.
Distribution. Restricted to the Red Sea, where in the south it replaces E. abeli as the most abundant species.
Remarks. Occurs in coral-rich areas of bays and lagoons from depths of 2–12 m; usually found on vertical
faces of Porites heads or in crevices close to this coral. Attains 21 mm SL.
Material examined. Additional to that listed in Holleman, 2005: SAIAB 88713 1 (3: 17.9–18.5 mm SL), Man-
grove Bay, El Quseir (25.52°N, 34.24°E, collected S. Bogorodsky, 8, 9 June 2008; SAIAB 88714 (12.3 mm SL),
Hanish Island, Yemen, collected S. Bogorodsky, 22 October 2009; SMF 9454 (13.8 mm SL), paratype, Sanganeb
Reef, north Port Sudan; SMF 16027 (14.2 mm SL), SMF 16030 (3: 14.4–15.3 mm SL) and SMF 16032 (6: 14.0–
18.5 mm SL), all from Aqaba, Red Sea; SMF 29588 (17.0 mm SL), Uqban, Kamaran Island, Yemen (15°31.245'N,
42°23.10'E); SMF 29501 (7: 13.6–18.4 mm SL), Uqban, Kamaran Island, Yemen (15°30.48'N, 42°23.30'E; SMF
29564 (17.6 mm SL); SMF ASE-3 (17.6 mm SL), Khebir Island, Kamaran Archipelago, Yemen; SMF 33539 (15.2
mm SL), Al Lith, Saudi Arabia (20°14.965' N, 39°59.621' E), S. Bogorodsky, 29 March 2011; SMF 33597 (12.2
mm SL), Al Lith, Saudi Arabia (20°14.965' N, 39°59.621' E), collected S. Bogorodsky, 29 March 2011; SMF
33598 (13.9 mm SL), Al Qunfidhah, Saudi Arabia (19°16.813' N, 40°53.265' E), collected S. Bogorodsky, 30
March 2011; SMF 33599 (16.5 mm SL), Farasan Islands, Saudi Arabia (16°54.933' N, 41°50.761' E), collected S.
Bogorodsky, 02 April 2011; SMF 33600 (15.1 mm SL), Rabigh, Saudi Arabia (23°02.839' N, 36°46.621' E), col-
lected S. Bogorodsky, 08 April 2011; SMF 33594 (16.9 mm SL), Al Lith, Saudi Arabia (20°14.965' N, 39°59.621'
E), collected S. Bogorodsky, 29 March 2011; SMF 33601 (16.8 mm SL) Farasan Islands, Saudi Arabia (16°54.933'
N, 41°50.761' E), collected S. Bogorodsky, 02 April 2011.

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48 · Zootaxa 3152 © 2012 Magnolia Press
Enneapterygius obscurus Clark, 1980
Figs. 2E, 8A; table 1
Enneapterygius obscurus Clark, 1980: 105, Figs. 4d & 15 (Gulf of Aqaba & Eritrea, Red Sea).
Fricke & Randall, 1992: 6, Fig. 3 (Maldives); Holleman, 2005: 15–16, Fig. 13, Pl. 2A (not of E. obscurus); Golani & Bogorod-
sky, 2010: 44.
Diagnosis. Dorsal fins III + XII–XIV + 9–10; anal fin I, 16–18; pectoral fin 14–15, some rays branched; LL 10–12
pored scales and 21–22 notched scales; dentary pores usually 2 + 2 + 2 (Fig. 2E). Body elongate, depth 6.0–6.5 in
SL, head 3.2–4.0 [3.6] in SL; eyes large, 2.3–2.6 [2.5] in head length. Nape scaled; abdomen naked; supratemporal
sensory canal crescent-shaped; supraorbital cirrus an elongate flap, about equal to pupil diameter; first dorsal fin
usually equal in height to second fin, in males the first spine slightly longer than first of second dorsal fin; longest
pectoral-fin ray to junction between second and third dorsal fins; pelvic-fin rays not united by membrane.
Fresh colour. Males: body transparent with a few scattered black speckles, scale edges crimson, with small,
intense black spot on upper caudal peduncle; pectoral-fin base and opercle with crimson blotch; head with crimson
markings dorsally, white ventrally, with brownish yellow stripe on side of snout and with small cluster of red spots
below eye and between eye and upper lip, lips yellowish with small brown spots; eye red with two spots dorsally
and white ring around pupil; first dorsal fin cream suffused with brownish red on first membrane and with alternat-
ing broad dark and narrow white bands on leading edge, spines 1 and 2 red-brown and white; second and third dor-
sal fins pinkish with clusters of melanophores and with area of white spots posteriorly; caudal fin pinkish red with
dark red line near base of rays; anal fin red with white spots along part of some rays; lower pectoral-fin base with
brown spots, lower rays pink; ventral fins without colour.
Females: body translucent with irregular white marks above and below midline, and brown spots below mid-
line, spine alternating white and yellow, viscera visible and yellow, chest with large white patch; head mostly trans-
lucent with white on opercle and cluster of brown spots below eye and on pectoral-fin base. First dorsal fin grey-
white, but membrane between spine 3 and dorsum transparent, spines alternating black and white; second dorsal fin
transparent with rows of white marks and golden yellow on spines in middle; third dorsal fin transparent with rows
of white marks; caudal fin with a white dumbbell mark at its base and a row of white marks on rays from top distal
edge to mid-lower edge; anal fin transparent with alternating subcutaneous white and brown marks at base and on
rays; pectoral fins with three rows of white marks.
Colour underwater. Males more pink than red, and eye more obviously red and white; females almost trans-
parent with white first dorsal fin, white chest and white and brownish marks on body and fins.
Key features. Translucent red in life with small, intense black spot on upper caudal peduncle; first dorsal fin
brownish anteriorly and with white spots posteriorly.
Distribution. Known from the Red Sea (type locality, El Himeira, Gulf of Aqaba); also reported from the Mal-
dives, but not confirmed; rare.
Remarks: Occurs in the vertical walls of seaward coral and rocky reefs of bays, lagoons from depths of 2–22
m. Attains 25 mm SL.
Material examined. Additional to that listed in Holleman, 2005: SAIAB 88715 (4: 17.8–19.4 mm SL), 5 km
south of Hamata, southern Egypt (24.13°N, 34.25°E), collected S. Bogorodsky, 21 & 23 November 2008; SAIAB
88716 (16.1 & 16.4 mm SL), Shaab Suadi, Sudan (20.12°N, 37.22°E), collected S. Bogorodsky, 12 October 2009;
SMF 33612 (19.2 mm SL), Farasan Islands (16°54.933' N, 41°50.761 E), collected S. Bogorodsky, 02 April 2011.
Enneapterygius pallidus Clark, 1980
Figs. 2F, 8B; table 1
Enneapterygius pallidus Clark, 1980: 107, Figs. 4e & 16 (Gulf of Aqaba and Eritrea, Red Sea).
Holleman, 2005: 16–17, Fig. 14, Pl. 2B; Golani & Bogorodsky, 2010: 44.
Diagnosis. Dorsal fins III + XIII–XIV + 10–11; anal fin I, 21–22; pectoral fin 13–14, some rays branched; LL 10–
11 pored scales and 26–27 notched scales; dentary pores 2 + 2 + 2 (Fig. 2F). Head in SL 3.4–4.7; eye diameter 3.0–

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3.5 in head length; body depth 5.0–6.0 in standard length. Nape and abdomen naked; supratemporal sensory canal
deeply ‘U’-shaped; supraorbital cirrus small and lobate; height of first dorsal fin about equal to or lower than sec-
ond fin.
Fresh colour. Males with translucent body, shaded with greenish hue on caudal peduncle and ventrally on pos-
terior half of body, scale borders red, with faint hourglass-shaped red bars on body and grayish green bar on poste-
rior caudal peduncle; head greenish dorsally, whitish ventrally with brownish red stripe from upper lip to anterior
margin of eye and short red bar below eye; head and body with a few scattered white speckles; eye with alternating
brownish red and greenish bands; fins mostly translucent, first two membranes of first dorsal fin sometimes with
whitish patch of melanophores and second dorsal fin often with patches of whitish to greenish dots forming a faint
short narrow bands.
Colour underwater. Males with body translucent with greenish tinge, scale pockets outlined in red-brown,
with about 10 subcutaneous brown bars below midline to A base, belly white; top of head emerald green, white
below eye with brown line from eye to upper lip and cluster of brown spots below eye, iris with green and brown
‘spokes’; anterior of first dorsal fin with white and brown marks; second dorsal fin with pigment along margin third
dorsal fin, caudal and anal fins without pigment; pectoral-fin rays reddish with white bands, colour on rays only;
pelvic fins white. Females with paler green on head and less pigment on first dorsal fin.
Key features. Greenish anteriorly, remainder of body transparent with 8–9 narrow, faint orange bars; first dor-
sal fin of females lower than second, of males same height as second, and with melanophores on membrane
between first two spines.
Distribution. Known only from the Red Sea; common in some areas.
Habitat. Usually seen on outer reef flats of lagoons from depths of 0.5–5 m. Reaches 30 mm SL.
Enneapterygius pusillus Rüppell, 1835
Fig. 2G, 8C & D; table 1
Enneapterygius pusillus Rüppell, 1935: (original description, Massawa).
Tripterygium pusillum: Klunzinger, 1871: 497
Enneapterygius pusillus: Smith, 1949: 359; Holleman in Smith & Heemstra, 1986: Holleman, 2005: 19; Golani & Bogorodsky,
2010: 44.
Helcogramma pusillus: Lal Mohan, 1971: 222.
Diagnosis. Dorsal fins III + XII–XIV (usually XIII) + 10–11; anal fin I, 20–21; pectoral fin 13–15 (usually 14),
some rays branched; LL 9–12 pored scales and 25–28 notched scales; longitudinal scale series 29–30; nape and
abdomen naked; supratemporal sensory canal deeply ‘U’-shaped; dentary pores 2 + 2 + 2; eye diameter 3.0–3.4 in
head length; supraorbital cirrus small and lobate; body depth 5.3–6.0 in standard length; first dorsal fin about equal
in height to second fin in females and distinctly higher than second in males, sometimes more than twice height of
second dorsal fin.
Fresh colour. Another species in which mature / sexually active males are much more intensely coloured than
other males, coupled with a much taller first dorsal fin. Underwater or photographed against a white background
fish have largely transparent bodies and fins, colour being concentrated on the head and chest. Against a coloured
background these fish have a white body against which pigmentation patterns stand out more clearly. Females are
almost entirely transparent against a white background or underwater, with some pigment on the head, a little on
body and fins, and the viscera encased in a silver peritoneum.
Body of males translucent, pale cream with four irregular darkish bars; scales on upper half with scattered
orange and yellow-green spots and bordered with a thin, deep orange line. Lower half of body with up to 10 irreg-
ular orange blotches interspersed with silver-grey spots. Abdomen covered anteriorly with dark brown rosettes and
posteriorly with pale red blotches; dark brown or black pre-anal spot — often ‘V’-shaped — usually present. Top
of head laced with subcutaneous green and cream, with red surface spots. Upper margin of eye and orbital cirrus
speckled with crimson, iris golden-red. Snout and lips pale yellow with orange spots; cheeks with yellow, red and
black spots; iridescent blue spot below eye. First dorsal fin orange-yellow with many small pale red spots and mel-
anophores; second dorsal fin heavily marked with dark brown to black melanophores interspersed with red spots

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and with 3 silvery-transparent vertical bands, and red and orange marks along base of fin; third dorsal fin with three
reddish bands, colour on rays only and scattered melanophores on margin. Upper half of caudal fin pale orange,
lower half pale yellowish green. Anal fin with red and black spots, tending to form 6 or more indistinct bars. Pecto-
ral fins yellow distally, orange proximally, with large circular orange-red mark with green centre on fin base. Pelvic
fin bases and surrounding skin heavily mottled with deep red and black, rays black for proximal half, distally with
red stipples; membrane black.
Females less intensely pigmented and lack the melanin of large males, but with an olive streak on side of snout,
broadening onto lips; membrane between first two dorsal-fin spines olive green, the first spine banded in pale green
and orange, membrane between second and third spines whitish, and between third spine and dorsum transparent;
second and third dorsal fins transparent with some olive green spots anteriorly on the first dorsal fin, spines and
rays with orange bands; caudal-fin rays with pale orange bands; anal fin transparent, but with some rays orange
proximally, colour continuous with darker spots at base of rays; pectoral fins white at base and with band of white
marks on rays; pelvic fins without colour.
Colour underwater. Body of males (Fig. 8D) mottled dark green with splashes of brown and white, scales
edged in brown, all fin elements greenish gold, except first dorsal fin which is golden yellow with black stippling,
and pelvic fins which are black; head reddish with many melanophores, snout grey, eye ridges red.
Females (Fig. 8C) are almost invisible, with only white and green spots on the body, the banded first dorsal-fin
spine and white on the pectoral fins visible.
Key features. Very tall first dorsal fin, in males to twice height of second dorsal fin; body of both males and
females translucent, males with black ‘chest’ and pelvic fins, yellow first dorsal fin, red at corner of mouth and
deep blue spots below eye. Females with brown bar below eye and on pectoral-fin base, first dorsal fin equal in
height to second and yellowish.
Distribution. Restricted to the Western Indian Ocean, east to Persian Gulf and southern India, south to South
Africa, described from the Red Sea (type locality, Massawa, Eritrea); rare.
Remarks. Like E. altipinnis, E. pusillus can be distinguished by the very tall first dorsal fin. Both species live
on a silty sand or mud substrate, close to the base of reefs, often on stones and dead corals of the same color. Found
in bays or sheltered lagoons. Attains 30 mm SL.
Enneapterygius qirmiz sp. nov.
Figs. 2H, 6 & 7; table 1
Enneapterygius sp.1 Clark, 1980: 101, Figs. 3c & 12, Gulf of Aqaba, Ras Muhammad and Ethiopia.
Enneapterygius sp.: Debelius, 1998: 174; Lieske & Myers, 2004: 177.
Enneapterygius obscurus non Clark, 1980: Holleman, 2005: 15, Fig. 13, Plate 2A.
Holotype. SMF 33093, male 16.9 mm SL; Uqban, Kamaran Island, Red Sea, Yemen (15°28.893' N, 42°23.141' E),
collected by P.C. Heemstra & Uwe Zajonz, 12 April 1998 (ex SMF 29594).
Paratypes. BPBM 35721 (17.8 mm SL), Parkin Rock, Hanish Group, southern Red Sea (13°37'12" N,
42°48'54" E), collected by J.E. Randall, 18 May 1993; SMF 29357 (11.1 & 15.6 mm SL), Uqban, east coast of
Kamaran Island, Red Sea, Yemen (15°31'20" N, 42°22'30" E), collected U. Zajonz, 1 April 1998; SMF 29487 (17.7
mm SL), same data as SMF 29357; SMF 29492 (15.5 & 16.5 mm SL); Kamaran Island, Yemen (15°29.077' N,
42°23'805" E), collected by M. Eisinger, 10 April 1998; SMF 29594 (17.5 mm SL), collected with holotype; SMF
33588 (25.3 mm SL), Al Lith, Saudi Arabia (20°14.965' N, 39°59.621' E), collected S. Bogorodsky, 20 March
2011; SMF 33589 (19.0 mm SL), Al Wajh. Saudi Arabia (26°10.552' N, 36°22.448' E), collected S. Bogorodsky,
10 April 2011; SMF 33590 (18.1 mm SL), Al Wajh, Saudi Arabia (26°10.552' N, 36°22.448' E), collected S.
Bogorodsky, 10 April 2011; SMF 33591 (16.9 mm SL), Al Wajh, Saudi Arabia (26°10.552' N, 36°22.448' E), col-
lected S. Bogorodsky, 10 April 2011; SMF 33592 (20.0 mm SL), Rabigh, Saudi Arabia (22°47.123' N, 38°57.507
E), collected S. Bogorodsky, 06 April 2011; NHCY-P1 (17.5 mm SL), Kadaman Isl, Kamaran Archipelago, Yemen
(15°34'13" N, 42°32'156" E), collected by U. Zajonz and P.C. Heemstra, 3 April 1998; SAIAB 88720 (14.9 & 16.4
mm SL), Shaab Rumi, Sudan (19°49' N, 37°21' E), depth 8 m, S. Bogorodsky, 10 October 2009; USNM 205805 (6:

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12.0–18,7 mm SL); Ras Muhammad, Sinai Peninsula, Red Sea, depth 0–9 m; collected V. G. Springer et al., 16
September 1969.
Non-type material. SAIAB 96002 (16.4 mm SL), Abingdon Reef, Sudan (20°50'N, 37°25'E), depth 14 m, col-
lected S. Bogorodsky, 15 October 2009.
Diagnosis. A small species of Enneapterygius with a maximum recorded SL of 25.3 mm. Dorsal fins III + XII
or XIII + 9, anal fin I, 17, pectoral fin 14; first dorsal fin about the same height as the second; nape scaled; both
sexes with characteristic oval patch of melanophores on dorsum between third and fourth dorsal-fin spines, males
with large rectangular blotch of melanophores and brown pigment on side below posterior of second dorsal fin; red
in life, with first dorsal of males crimson in front with golden yellow marks, white behind, base of fin a white patch
on dorsum.
Description. Dorsal fins III + XII or XIII + 9; anal fin I, 17 (one of 13 with 18 rays); pectoral fin 14 (rarely 15),
upper 2 and lower 6 simple, others divided once. (Clark recorded lower and upper 5 unbranched). LL with 8–10
pored scales followed two scale rows lower by 22 notched scales, some specimens with 8 pored scales followed by
2 notched scales in anterior series; total lateral scales about 33, with single row of scales at base of caudal fin;
transverse scales 2/5. Vertebrae 10 precaudal + 23 (2 of 13 with 24) caudal, including urostyle. Dentary pores 2 + 2
+ 2 (Fig. 2H). Head length 3.1–3.7 (3.4) in SL; eye 2.3–2.7 (2.6) in head length.
Nape scaled, belly naked; eye relatively large, orbital cirrus a rounded flap, its length equal to orbit diameter.
First dorsal fin of males about same height as second, a little lower than second in females. Pectoral fins long, lon-
gest ray reaching to first spine of third dorsal fin; pelvic-fin rays not united by membrane, longest ray reaching first
anal-fin ray. Supratemporal canal a broad, flat ‘U’-shape rather than a crescent. Mouth small and ‘pointed’, maxilla
reaching vertical though anterior margin of pupil.
FIGURE 6. Enneapterygius qirmiz sp. nov., M, Angarosh Reef. Photo by S.V. Bogorodsky.
Colour. Males (from underwater photographs by Helmut Debelius and the second author) body translucent,
scales with double, crimson-red line on edge, except for black borders to scales immediately below end of second
dorsal fin forming an indistinct narrow, oblique bar, anterior to which is large, rectangular patch of scattered mel-
anophores extending from just below the dorsal fin almost to the base of the anal fin. Some specimens with another
cluster of scattered melanophores anterior to main blotch. Nape and opercle deep crimson; deep crimson blotch on
mid-side below first two or three spines of second dorsal fin; crimson blotches along vertebral column and along

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base of anal fin; preanal triangular cluster of melanophores present in large specimens; head red above and pale
below, iris crimson, with a thin whitish ring around the iris, and with white patches behind and between eyes,
orbital cirri yellow, nasal cirri red, red on lips. First dorsal fin with a broad, inverted deep red triangle, apex at base
of first spine, with golden yellow marks, remainder of fin white as is base of fin and a narrow ‘saddle’ extending
from base of first spine nearly to base of fourth spine; second dorsal-fin membranes transparent with red on spines,
and with black and white spots along margin and at base, and a narrow line of melanophores along margin which
continues onto body as black spots on scale margins; third dorsal fin similar; caudal fin with crimson lines on edges
of rays; anal fin rays crimson and patches of black and red spots on membranes; pectoral-fin base with patch of
crimson spots anteriorly and white spots below, rays red with patches of bright white spots.
Live colour of females unknown, but probably also crimson, but without the blotch of melanophores on the
side and conspicuously coloured first dorsal fin.
Body of preserved fishes pale yellow, males with large blotch of melanophores on side below pectoral fin, line
of scale pockets outlined in black extending from posterior end of third dorsal fin to midline and oval patch of mel-
anophores on dorsum between third and fourth dorsal-fin spines, patch of melanophores on peduncle just behind
end of third dorsal fin; small pre-anal patch of melanophores; some clusters of melanophores on anal fin.
Key features. Translucent crimson; first dorsal fin lower than second, red anteriorly with yellow marks, white
posteriorly and with bright white patch on dorsum at base of fin; narrow line of melanophores along margin of sec-
ond dorsal fin which continues onto body as black spots on scale margins; males with broad patch of melanophores
on side below second dorsal fin.
Etymology. ‘Qirmiz’ is the Arabic word for deep red or crimson, and is given for the colour of the species. It is
used as a noun in apposition.
Comparisons. As can be seen from Table 1, there is little variation in morphometric characters between spe-
cies of Enneapterygius. Three species tend to have 17 anal-fin rays: E. altipinnis, E. obscurus and E. qirmiz. The
first has a tall first dorsal fin and the characteristic black blotch on the second dorsal fin; the last two species are
both red in colour, but males of E. qirmiz have a very characteristic red, white and yellow first dorsal fin, and E.
obscurus an intense black spot on the upper caudal peduncle. These two species can also be separated on LL scale
counts, E. qirmiz with 9 pored scales in the anterior series and E. obscurus with 10–12.
Distribution. Clark’s original specimens were collected in the Gulf of Aqaba, at Ras Mohammad and Ethiopia.
The types here described were collected in similar localities as well as off Yemen, suggesting the species is wide-
spread, though not common in the Red Sea.
Remarks. Debelius’ photographs in Red Sea Reef Guide (1998) were incorrectly identified as E. obscurus by
Holleman (2005). The species is found in environments very similar to those of E. obscurus. However, in compari-
son with other Enneapterygius spp. which are mostly known from shallow coastal waters, this species is surpris-
ingly common on some isolated reefs far from the coast, for example, on the isolated Abingdon Reef located about
20 km from the coast, where no other triplefins were seen.
FIGURE 7. Enneapterygius qirmiz sp. nov., SMF 33093, male 16.9 mm SL; Uqban, Kamaran Island, Yemen. Drawn by Elaine
Heemstra.

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Enneapterygius ventermaculus Holleman, 1982
Figs. 2I, 8E; table 1
Enneapterygius ventermaculus Holleman, 1982: 123, fig. 7 (northern KwaZulu-Natal, South Africa); Randall, 1995: 32, fig. 5
(Gulf of Oman); Holleman, 2005: 21, Fig. 19, Pl. 2; Golani & Bogorodsky 2010: 44.
Enneapterygius nasimae: Hoda, 1983: 116ff, figs. 1–3 (Karachi coast, Pakistan).
Diagnosis. Dorsal fins III + XI–XIII + 8–10 (usually III + XII + 10); anal fin I,17–20 (usually 19); pectoral fin 14:
1–3, 4–6, 6–8 (usually 2, 5, 7). LL 13–16 (usually 15) pored and 21–25 (usually 23) notched scales, notched series
starting second scale row below end of pored series; dentary pores 3 + 1 + 3 (Fig. 2I). Head 3.4–4.0 in SL. Nape,
base of first dorsal fin and belly naked. Orbital cirrus moderate and lobate; supratemporal sensory canal ‘U’-
shaped, embracing first dorsal fin spine. First dorsal fin slightly higher than second in males, slightly lower in
females.
Fresh colour. Males body olive green with darker scale margins, with 3 or 4 blotches of white spots along mid-
side, interspersed with smaller dark olive green marks, sometimes extending to anal-fin base, dark triangular mark
on peduncle at caudal-fin base, apex forward with two white streaks, one above the other, at base of fin, belly white
with black pre-anal spot, often a triangle, apex anterior; head dark olive green, with white spots, colour to pectoral-
fin base; first dorsal finely speckled with brown and orange, first spine banded, except on membrane between spine
3 and dorsum; second dorsal fin with hyaline band in centre and with melanophores and white on margin and at
base of spines; third dorsal fin similarly marked with white and black, but on rays only; caudal fin largely hyaline
but with dark and light banding along upper and lower rays in deeply pigmented males; anal fin alternately banded
black, white and hyaline, with 6–8 black subcutaneous basal spots; pelvic fins cream with lower rays ‘banded’ with
black on rays only; pelvic-fin base black, rays cream.
Colour underwater. Body translucent pale bottle-green with scale margins orange-brown, subcutaneous anal-
fin spots red-brown, white lines on peduncle very bright, as are alternating dark and white dashed lines along
length of vertebral column; head with alternating dark and white marks radiating out from eye, iris orange-brown
and green; first dorsal fin with white and banding on spine 1, second and third dorsal fins with ‘rows’ of black and
white marks; pectoral fins with ‘bands’ of white on rays.
Key features. Males with blackish head and yellow first dorsal fin, yellow on lower pectoral-fin rays and prox-
imal portion of the pelvic-fin rays; both males and females olive green with black and white barred anal fins.
Distribution. This species is known from the Red Sea, Aden, Oman, Pakistan and the east and southeast coast
of South Africa, from the Transkei to southern Mozambique. The record from the Red Sea is based on SMNS
21497 from the Kamaran Islands, Yemen (R. Fricke pers. comm.).
Discussion of the Enneapterygius spp.
The eight Enneapterygius species can be seen to fall into four ‘pair-groups’, which can be distinguished in colour,
morphology and habitat preferences, though the individual species are not necessarily sibling species. Within each
group the two species tend to be similar in appearance.
Group 1: E. altipinnis and E. pusillus, both with high first dorsal fins, though the form of the supraoccipital
sensory canal in the two is different. Both species live on silty sand or mud bottoms, close to the base of reefs, often
on stones or dead coral of the same colour, in bays or sheltered lagoons. E. altipinnis is however confined to the
Red Sea and is one of a number of species of the widespread Enneapterygius tutuilae species group. E. pusillus
also occurs along the coast of Oman and in the Persian Gulf, the west coast of Indian, and south to northern Kwa-
Zulu-Natal (Holleman, 2005). Both species occur throughout the Red Sea and E. pusillus is the most common
triplefin in Omani seas.
Group 2: E. obscurus and E. qirmiz, both slender-bodied, red in colour and with distinct first dorsal fin pig-
mentation. Both species are found on rocky reefs, often on vertical walls, on the seaward side of reefs. Both species
are only known from the Red Sea and are found both in the north (Eilat) and south (Hanish Islands, Yemen and
Shaab Suadi, Sudan)

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Group 3: E. clarkae and E. destai, with robust bodies and low first dorsal fins, both with barred pigmentation.
These two species live in close association with hard corals, tend to be secretive and hide in crevices, and are found
in sheltered bays and sheltered lagoons. E. destai is however confined to the Red Sea, where it appears to be more
abundant in the north than in the south (Clark, 1980). Like E. pusillus, E. clarkae occurs outside the Red Sea as
well and is known from the east coast of Africa to northern KwaZulu-Natal, the Mascarene Islands (Holleman,
1982).
Group 4: E. ventermaculus and E. pallidus, both with low first dorsal fin and both translucent green in colour
and without distinct body markings. The first species is also found on the coast of Oman to the coast of Pakistan,
the west coast of Indian and has been recorded from South Africa and Mauritius (Holleman, 2005). E. pallidus is
endemic to the Red Sea and, again, is found throughout. E. ventermaculus occurs in tide pools and to a depth of
some 12 m (Randall, 1995), while E. pallidus is found on the upper part of reefs, on reef flats, often on corals cov-
ered in short algae, in fairly sheltered environments.
The odd species is E. abeli, in which the males are markedly sexually distinct with the prominent black head,
chest and pectoral-fin bases. This species too is widely distributed throughout the Red Sea and down the East Afri-
can coast to KwaZulu-Natal and Mascarene Islands. The species has a sibling species, E. elaine, which is endemic
to Rodrigues (Holleman 2005).
Genus Helcogramma McCulloch & Waite, 1918
Helcogramma McCulloch & Waite, 1918: 51; type species H. decurrens McCulloch & Waite, 1918, by original designation.
Diagnosis. Small to medium-sized tripterygiid fishes with fusiform bodies, ranging in length from 27 mm to 50
mm SL. First dorsal fin with 3 spines; anal fin with a single, short spine, usually less than half the length of the first
ray. Pelvic fins with one short, hidden spine and two simple, segmented rays, which may be joined by membrane
for part of their length. Dentary canals with 1 or (rarely) 2 pores at the symphysis, or 3–5 pores arranged symmetri-
cally about the symphysis, and 2–8 on either side along the dentaries. Exposed posterior margin of post-temporal
bones with fine serrations. Body with ctenoid scales; nape naked or scaled; abdomen and pectoral-fin bases always
naked. Head always without scales. LL continuous, of 14–35 pored scales, running in a shallow, downwardly con-
cave curve from the post-temporal to mid-body and extending to below the second dorsal fin or to the base of the
caudal fin. Base of caudal fin with 0–3 rows of scales. Orbital cirrus absent in some species, if present simple to
palmate. Anterior nasal cirri simple to palmate, on posterior margin of a short tube. Teeth in both jaws, conical and
slightly recurved, largest in single row in front; vomer with single row of similar teeth; palatines edentate.
The genus currently consists of 38 valid species, two in the Red Sea. Eschmeyer & Fricke (2011) lists only 37
species, the validity of Helcogramma ascensionis Lubbock, 1980 not being recognised, while it is a valid species
(Holleman, unpubl. data).
Helcogramma obtusirostris (Klunzinger, 1871)
Fig. 8F & G; table 1
Tripterygium obtusirostris Klunzinger, 1871: 498 (Red Sea). Golani & Bogorodsky, 2010: 45.
Helcogramma obtusirostre: Clark, 1980: 85, figs. 3e, 7 & plate 1; Holleman in: Smith & Heemstra, 1986: 757; Randall, 1995:
309, fig. 872; Holleman, 2007: 68, Fig. 1 & 10, Pl. 2.
Helcogramma obtusirostris: Hansen, 1986: 341 (in part: Red Sea).
Helcogramma trigloides (non Bleeker): Marshall, 1952: 242 (Gulf of Aqaba).
non Helcogramma shinglensis: Lal Mohan, 1971: 219, fig. 1 (Gulf of Mannar, India).
non Helcogramma ascensionis: Lubbock, 1980: 294, fig. 2 (Ascension Island, S Atlantic).

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FIGURE 8. (A) Enneapterygius obscurus, Marsa Alam; (B) Enneapterygius pallidus, Farasan Is.; (C) Enneapterygius pusillus,
M, Sharm el Sheik; (D) Enneapterygius pusillus, M, Oman; (E) Enneapterygius ventermaculus, Farasan Is.; (E) Helcogramma
obtusirostris, M, Sharm el Sheil; (F) Helcogramma obtusirostris, F, Sharm el Sheik; (G) Helcogramma steinitzi, F, Marsa
Alam; (H) Norfolkia brachylepis, Bali. Photos by R. Field (B, E), J.E. Randall (D); G.R. Allen (H); S.V. Bogorodsky (the
others).

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56 · Zootaxa 3152 © 2012 Magnolia Press
Diagnosis. Dorsal fins III + XII–XIII + 9–10 (rarely with XII spines or 9 rays); anal fin I, 18–19 (usually 19);
pectoral fin 16: usually 2, 7, 7. LL 20–23 (usually 21–22) tubed scales, ending below junction of second and third
dorsal fins; total lateral scales 36–38 (usually 37–38); transverse scales 7/5. Dentary pores 4 + 1 + 4. Head length
3.3–3.7 (3.5) in SL. Nape and belly naked; scales do not extend to bases of first dorsal and anterior of anal fins; 2
rows of scales on base of caudal fin. Pelvic–fin rays united by membrane for half shorter ray, longest ray reaching
vent. First dorsal fin half height of second. Maxilla reaches vertical through centre of pupil; orbital cirrus small and
simple.
Fresh colour. (From colour photograph by the second author and J. E. Randall.) Body of males mottled green-
ish with darker interconnecting areas, with 4 narrow, pale saddle marks extending from base of dorsal fins to mid-
line, 2 below second dorsal fin, 2 below third dorsal fin; series of paler spots along midline; dorso–anteriorly
suffused with dull red; belly pink. Head dark, black below eye and to branchiostegal membrane; bright blue stripe
extends from lower lip across cheek, broadly edged with black above and below. Pectoral–fin base with red spot
above and below, the lower one with blue above and anterior; pelvic fins dark pink proximally; first dorsal fin
dusky with some orange, darkest on the margin, and with small black spot at base of membrane connecting first
and second dorsal fins; second dorsal fin dusky with dull orange band basally; third dorsal fin dusky; caudal fin
dusky; anal fin dark, with an even cover of melanophores.
Females dull green, the dark areas forming a zig–zag pattern down the body, on a pale green background; body
pale ventrally. Head dorsally dark green, ventrally pale; median fins with light sprinkling of melanophores, first
dorsal fin with small black spots as in males, basal third of dorsal and entire anal fins white.
Colour underwater. Body of males translucent with reddish ‘H’-bars and streak of white marks from dorsal
fins onto body, belly and pelvic fins pinkish-red; head reddish above and with many melanophores below eyes,
extending onto base of pectoral fin; blue-white line from lower lip, through corner of mouth to preopercle, with
blue patch below; dorsal fins translucent with red and white marks, caudal fin pink, pectoral fins translucent with
red and white marks. Females paler versions of males and lack the distinctive pigmentation on the head and belly,
which are largely white.
Key features. Iridescent blue band from corner of mouth to opercle in males; males in ‘breeding dress’ nearly
entirely black with 3 or 4 white ‘slashes’ below second and third dorsal fins; females whitish with irregular brown-
ish bars.
Distribution. H. obtusirostris occurs in the Red Sea, Yemen and Oman.
Remarks. H. obtusirostris is one of a group of widespread Indo-west Pacific species with at least four sibling
species, which all have an iridescent blue line running from the lower lip to the corner of the mouth and onto the
opercle. It is easily distinguished from H. steinitzi, which is reddish in general colour, has a large head and lacks the
blue line from the corner of the mouth. Usually seen in pairs in coral and rocky areas of lagoons in depths of 0.5–8
m. Attains 38 mm SL.
the west coast of Indian and has been recorded from South Africa and Mauritius (Holleman, 2005). E. pallidus
is endemic to the Red Sea and, again, is found throughout. E. ventermaculus occurs in tide pools and to a depth of
some 12 m (Randall, 1995), while E. pallidus is found on the upper part of reefs, on reef flats, often on corals cov-
ered in short algae, in fairly sheltered environments.
The odd species is E. abeli, in which the males are markedly sexually distinct with the prominent black head,
chest and pectoral-fin bases. This species too is widely distributed throughout the Red Sea and down the East Afri-
can coast to KwaZulu-Natal and Mascarene Islands. The species has a sibling species, E. elaine, which is endemic
to Rodrigues (Holleman 2005).
Helcogramma steinitzi Clark, 1980
Fig. 8H; table 1
Helcogramma steinitzi Clark, 1980: 88, fig. 8, Pl. II–V (Gulf of Aqaba); Hansen, 1986: 347; Randall, 1995: 310; Holleman,
2006: 98, Figs. 5–8 and 2007: 75, Figs. 1 & 15, Pl. 2; Golani & Bogorodsky, 2010: 45.
Diagnosis (partially from Clark 1980 and Randall 1995). Dorsal fins III + XII–XIV + 10–12 (usually III + XIII +
10–11); anal fin I, 19–21 (usually 20); pectoral fins usually 15–17 (recorded by Clark and Randall; usually 16: 2, 7,
7). LL 21–27 tubed scales, ending below first five rays of third dorsal fin; total lateral scales 38–40 (37–41, from

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Clark). Dentary pores 3 + 1 + 3. Head length 3.0–3.5 in SL. Nape scaled, belly naked, scales do not extend to base
of first dorsal or anal fins; 1 or 2 rows of scales at base of caudal fin. Pelvic-fin rays united by membrane for half
length of shorter ray, longer ray reaching vent. Origin of first dorsal fin over posterior margin of preopercle, fin tri-
angular, in males first dorsal-fin spine about as long as longest spine of second dorsal fin, in females a little shorter;
first two spines set close together. Mouth large, maxilla reaching vertical through posterior border of pupil; orbital
cirrus minute.
Fresh colour. (based on Randall 1995 and photos by the second author and Randall). Body of males dark red
above LL, many scales outlined in black, paler red below LL, with white flecks and indistinct black markings
forming a reticulated pattern, and with 8 dark blotches interspersed with white blotches along mid–side below lat-
eral line; belly white. Head above lower level of eye dark red, below bluish–grey with many small melanophores,
and a bluish mark below eye and another on lower part of preopercle. First dorsal fin with red and black spots; sec-
ond and third with red basal band with black spots and pale red marginal band, hyaline band between, with white
on elements. Anal fin pale red, darker along margin; caudal fin pale red; pectoral fins red, bases dark grey with two
oval, bluish marks, one above the other; pelvic fins pink.
Females with translucent greenish body, irregularly spotted with white, with large interconnecting red blotches,
red and black spots on scale margins, forming a reticulated pattern; head greenish with numerous dark red spots
and short bands, the darkest a diagonal band on the side of the snout.
Colour underwater. Body of males mottled red-brown on white, head with many brown spots below eyes and
to base of pectoral fins, bluish mark below eye and on lower part of preopercle, and two bluish spots on pectoral fin
base. Median fins translucent with red and white, except for membrane between first two dorsal-fin spines white
with dark areas. Females semi-translucent with reddish-brown and subcutaneous white marks; median fins translu-
cent, fin elements with brown and pale ‘bands’, pectoral-fin rays with white marks and orange-pink distally.
Key features. Large, ‘crocodile-like’ head with thick lips; males red in life with lower part of head blackish
and many micromelanophores on membrane between first two dorsal-fin spines; females whitish with irregular
brown markings on body and lower part of head.
Distribution. Helcogramma steinitzi is known from the Red Sea, the coasts of Yemen and Oman and the Per-
sian Gulf.
Remarks. H. steinitzi is one of a complex of species of which there are three species in the Indian Ocean. The
males of all the species of the complex are red and have many micromelanophores on the membrane between the
first two dorsal fin spines. Occurs on corals and rocks in bays and lagoons in depths from 0.5 to at least 12 m, usu-
ally solitary. The species attains nearly 60 mm SL.
Material examined. Additional to that listed in Holleman 2006: SAIAB 88723 (27 mm SL), 5 km S of
Hamata, southern Egypt (24°13'N, 34°25'E), collected S. Bogorodsky, 18 November 2008; SAIAB 88724 (mm
SL), northern Hurghada (27°15'N, 33°49'E), collected S. Bogorodsky, 20 April 2009; SMF 33583 (17.8 mm SL),
Al Qunfidhah, Saudi Arabia (19°16.813' N, 40°53.265' E), collected S. Bogorodsky, 30 March 2011; SMF 33584
(18.0 mm SL), Rabigh, Saudi Arabia (22°47.123' N, 38°57.507' E), collected S. Bogorodsky, 06 April 2011; SMF
33585 (17.9 mm SL), Rabigh, Saudi Arabia (22°47.123' N, 38°57.507' E), collected S. Bogorodsky, 06 April 2011.
Genus Norfolkia Fowler, 1953
Norfolkia Fowler, 1953: 262. Type species N. lairdi Fowler, 1952 = Gillias squamiceps McCulloch & Waite, 1916: 449, by
original designation.
Diagnosis. Tripterygiid fishes with the following combination of characters: first dorsal fin with four spines; anal
fin with two spines; lateral line discontinuous, with an anterior series of pored scales ending below the second dor-
sal fin, and a posterior series of notched scales from below the end of the anterior series to base of caudal fin;
orbital and anterior nasal cirri present. Body and head below and behind eyes heavily scaled with ctenoid scales,
except for a few cycloid scales on pectoral-fin bases, abdomen and around vent. Margins of lateral ethmoids ‘cren-
ulate’ (see Holleman 1991); septal bone unossified to partially ossified; vomer with single row of coniform teeth,
palatines without teeth.
Four species, one in the Red Sea.

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Norfolkia brachylepis (Schultz, 1960)
Fig. 8I; table 1
Tripterygion brachylepis Schultz in Schultz et al., 1960: 291 (type locality Bikini Atoll, Marshall Islands).
Norfolkia springeri Clark, 1980: 95 (Aqaba, Red Sea).
Norfolkia brachylepis: Holleman, 1991: 233; Fricke, 1994: 465; 1997: 504; Golani & Bogorodsky, 2010: 45.
Diagnosis. Dorsal fins IV + XIV–XV + 10–11, usually IV + XV + 11; anal fin II, 20–21, usually 21, the spines less
than half the length of the rays; pectoral fin 16, upper 2–3 and lower 7 rays simple, remainder divided once; pelvic-
fin rays united by membrane for less than one-quarter of length of shorter ray. LL 21–24 (usually 22) pored scales
ending below junction of second and third dorsal fins, and 14–19 (usually 16) notched scales; total lateral scales
32–35, usually 34. Dentary pores 5–6 + 1 + 5–6. Orbital and nasal cirri large palmate, at least equal to pupil diam-
eter; cirri at tips of first dorsal-fin spines. Head 2.9–3.6 (3.3) in SL, eye 3.0–3.9 (3.3), maxilla 2.3–3.0 (2.6) and
snout 3.3–4.9 (3.8) in head length.
Fresh colour. (from photograph of Rick Winterbottom of specimen from Kwazulu-Natal, South Africa). No
sexual dichromatism present other than that males may be more darkly coloured than females. Body pale yellow,
scales outlined in yellow-ochre, with 5–6 dark brown blotches between dorsum and midline; head brownish-yellow
with darker brown bar below eye and pale area behind, iris yellow; first dorsal fin brown and yellow, second trans-
lucent with patches of light red and brown, third dorsal fin with irregular bars of brown-red and cream; caudal fin
with irregular brown and white barring, anal fin with yellow and black-brown bars, pelvic fins pale yellow and pec-
toral fins with irregular brown and yellow bars.
Colour underwater (from photograph of G.R. Allen of fresh specimen from Indonesia): body grayish with
golden-brown and gray-white blotches between dorsum and midline, blotched extending onto second and third dor-
sal fins; head grey with brown bar below eye, followed by cream bar and light brown preopercle, eye rim alternat-
ing brown and grey; first dorsal fin ‘dirty’ white, darker distally, second dorsal fin with white and red-brown
marks, the white extending as bands across the fin; third dorsal fin the same; caudal fin with alternating, irregular
white and brown bands; anal fin distinctly banded black and cream; pectoral fins irregularly banded with white.
Colour in preservative. Body pale gray with five, irregular dark bars along dorsum, which fade towards LL;
head gray with a tapering dark bar below the eye and another from the posterior margin of the orbit curving down
onto the preopercle. First dorsal fin distally dark gray, second and third dorsal fins with clusters of melanophores
basally, continuations of body bars, and which extend as irregular areas of spots across fin to margin; caudal fin
with several, narrow dark gray bars; anal fin with faint, diagonal bands of melanophores. Mature males are much
darker, with body and median fins dark gray and without any suggestion of bars.
Key features. Scaled head, palmate orbital and nasal cirri, four spines in the first dorsal fin and distinctive yel-
low and black barred anal fin.
Distribution. This is an apparently wide-spread species, currently known from the Red Sea, to South Africa,
from Sri Lanka, the east and west coast of Australia, Indonesia, southern Japan and east to the Marshall Islands,
New Caledonia and Fiji.
Remarks. Holleman (1991) revised the genus, recognising three species, N. brachylepis, with its very wide
distribution, N. squamiceps (southern Great Barrier Reef) and N. thomasi, from the Great Barrier Reef and southern
Pacific to Rapa Island. Photographs by J. E. Randall of specimens from Japan and Indonesia do not show the yel-
low and black anal fin. The colour described above is of a specimen from southern Madagascar. As the species was
recorded by Randall (1995) from the Gulf of Oman it is assumed here that the colour of the Madagascar specimen
is the same as for individuals from the Red Sea.
Acknowledgements
The authors are grateful for the loan of a large collection of material from the Red Sea from the Forschungsinstitut
und Naturmuseum Senckenberg (Uwe Zajonz and Tilman Alperman). Other specimens have been loaned over
some years by the Bishop Museum, Honolulu (Arnold Suzumoto) and the National Museum of Natural History,
Smithsonian Institution (Jeff Williams). Juergen Herler, Rick Winterbottom, Richard Field, Gerry Allen and Jack
Randall allowed us to use of their photographs. Elaine Heemstra made the excellent drawings of Enneapterygius

Zootaxa 3152 © 2012 Magnolia Press · 59
REVIEW RED SEA TRIPTERYGIIDAE
qirmiz. Jeff Williams made critical comments on the manuscript which improved it much. We also thanks Ronald
Fricke for his help in confirmation of identification and information regarding Red Sea record of E. ventermaculus.
SB thanks Friedhelm Krupp and KAU for the financial support and organization of field work in April 2011 in
Saudi Arabia, and also thanks Dmitry Pashkov and Captain Mauricio for their help in the organization of the diving
trip on the vessel Katharina from Sudan to Yemen in October 2009, and WH acknowledges the support and facili-
ties provided by the South Africa Institute for Aquatic Biodiversity, and the help of the Collections staff, particu-
larly Bernard Mackenzie and Bafo Konqobe.
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