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A review of the triplefin fish genus
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the western Indian Ocean...
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SMITHIANA
Publications in Aquatic Biodiversity
Bulletin 5
August 2005
A review of the triplefin fish genus Enneapterygius
(Blennioidei: Tripterygiidae) in the western Indian Ocean,
with descriptions of four new species
Wouter Holleman
Published by the South African Institute for Aquatic Biodiversity
Margaret Mary Smith (1916 - 1987),
James Leonard Brierley Smith (1897 - 1968)
with their dog Marlin
The publication series (Monographs, Bulletins & Special Publications) of the SAIAB
(formerly the JLB Smith Institute of Ichthyology), in its new format honours James
Leonard Brierley Smith and Margaret Mary Smith with the name Smithiana, in recognition of their many years of devoted service to African aquatic biology. Their life’s work,
a team effort, established modern ichthyology in southern Africa and laid the groundwork for the expansion of aquatic biology throughout the region.
© 2005, The South African Institute for Aquatic Biodiversity, Grahamstown, South Africa
Front cover photograph: Scales of a preserved coelacanth specimen by James Stapley. © James Stapley, 2002
A review of the triplefin fish genus Enneapterygius
(Blennioidei: Tripterygiidae) in the western Indian Ocean,
with descriptions of four new species
Wouter Holleman 1
ABSTRACT
The fishes of the tripterygiid genus Enneapterygius of the western Indian Ocean (excluding Sri
Lanka) are reviewed. Four new species, E. elaine, E. gruschkai, E. genamaculatus and E. kosiensis are
described and several species are redescribed. Enneapterygius elaine is known only from Rodrigues,
E. gruschkai, a medium-sized species, is known from the Comoro Islands, Mauritius, St Brandon
Shoals and the Chagos Archipelago; E. genamaculatus is known only from St Brandon Shoals and
E. kosiensis is known only from the northern KwaZulu-Natal coast. The first and last two species
are small species, less than 23 mm SL. A key is provided for the 17 species recognised in the region.
1
South African Institute for Aquatic Biodiversity, Private Bag 1015, Somerset Street, Grahamstown, 6140, South
Africa. E-mail: W.Holleman@ru.ac.za.
i
CONTENTS
Introduction ............................................................................................................................................... 1
Methods and materials ............................................................................................................................. 1
Taxonomic accounts ................................................................................................................................. 3
Genus Enneapterygius Rüppell ........................................................................................................... 3
Key to the western Indian Ocean species of Enneapterygius ................................................................ 3
Species accounts
Enneapterygius abeli (Klausewitz) ...................................................................................................... 4
E. clarkae Holleman ............................................................................................................................. 6
E. destai Clark ....................................................................................................................................... 6
E. elaine sp. nov. ................................................................................................................................... 7
E. elegans (Peters) ................................................................................................................................. 8
E. fasciatus (Weber) ............................................................................................................................... 9
E. genamaculatus sp. nov. ................................................................................................................... 11
E. gruschkai sp. nov. ........................................................................................................................... 11
E. hollemani Randall .......................................................................................................................... 13
E. kosiensis sp. nov. ............................................................................................................................ 13
E. melanospilos Randall ..................................................................................................................... 14
E. obscurus Clark ................................................................................................................................. 15
E. pallidus Clark ................................................................................................................................. 16
E. philippinus (Peters) ........................................................................................................................ 17
E. pusillus Rüppell ............................................................................................................................. 19
E. tutuilae Jordan & Seale .................................................................................................................. 20
E. ventermaculus Holleman ............................................................................................................... 21
Comments on Indian and Sri Lankan species .................................................................................... 23
Discussion on colour patterns .............................................................................................................. 23
Acknowledgements ................................................................................................................................ 24
Literature cited ........................................................................................................................................ 24
ii
A review of the triplefin fish genus Enneapterygius
(Blennioidei: Tripterygiidae) in the western Indian Ocean,
with descriptions of four new species.
Wouter Holleman
Holleman, 1982 & 1986) have not been included here:
measurements of snout and maxilla length in such small
species using a standard dial caliper are too inaccurate
and variable to be of statistical significance or taxonomic
value. Snout profile was also not measured, again
because the method I devised is too inaccurate for fishes
this small. Similarly, vertebral counts have not been made,
except for the newly described species, because they are
not readily available to those who would identify the
species.
Pectoral-fin ray counts are given as total number:
(from the top) number of simple rays, number of divided
rays, number of simple rays. In all tripterygiids (except
the free-swimming Obliquichthys) the lowermost 5-7
pectoral-fin rays are thickened and used as support
additional to the pelvic fins. The caudal fin of tripterygiids
has 7 dorsal and 6 ventral segmented, principal caudalfin rays, of which the uppermost and lowermost 2 or 3
are simple and the remainder bifurcate, and between 3
and 8 procurrent rays dorsally and ventrally.
In a few specimens one (seldom two) notched scales
occur anterior to the beginning of the notched segment
of the lateral line, just below the end of the anterior, pored
segment, and are not included in the count. Including
these and the two or three un-notched scales would result
in very variable counts. In most species the scales are
highly deciduous and, unless specimens are collected
with care, most of the scales are lost in collecting nets.
Total lateral scales are counted from the last notched
lateral line scale to the pectoral-fin axil. Transverse scale
counts are taken from the base of the anterior half of the
second dorsal fin to the pored segment of the lateral line,
and below it to the base of the anal fin.
Hansen (1986) included mandibular pore patterns
in her Helcogramma species descriptions, finding that they
were often diagnostic for species. Williams & McCormick
(1990) followed suit and pore patterns are used in this
revision as well. They are given as the number of pores
on the one side + the number at the symphysis + the
number on the other side. Two pores at the symphysis
may in some individuals join to form a single, wide oval
pore rather than two distinct pores. Where this is
relatively common in a species the symphysial pore is
indicated 1/2.
Live coloration is usually diagnostic for tripterygiid
species. In all species that I am aware of and of which
the live colours are known, males are more brightly
coloured than females, very often with much black on
the head, chest and pectoral-fin bases. In preserved
specimens melanophore patterns are often distinctive
and diagnostic, but these too tend to fade after extended
periods in alcohol, particularly where specimens are
exposed to light.
INTRODUCTION
Fishes of the genus Enneapterygius are characteristically
small, most species being less than 25 mm long. They
are generally coral reef inhabitants, their colouring and
small size rendering them cryptic. The advent of scuba,
ichthyocides and more sophisticated underwater
photographic equipment and techniques have resulted
in more of these small fishes being collected and
described. However, their colours fade rapidly in
preservative, leaving bottles of small straw-coloured
fishes with little to distinguish one species from the next.
Meristically and morphometrically many species are
very similar, and consequently long-preserved specimens
are difficult to identify and often relegated to the back of
shelves.
The taxonomic history of the family - and hence the
genus - has been checkered. Rosenblatt, in an
unpublished thesis, made the first - and to date only revision of the family in 1959. Some of the species
descriptions took decades to appear in print, often under
someone else’s name. Clark (1980) made the first
substantial regional revision, the tripterygiid fishes of
the Red Sea; Hansen (1986) revised Helcogramma; Hardy
addressed the taxonomy of the diverse New Zealand
fauna (e.g. Hardy 1984, 1986, 1987), but, it was not until
1994 that the next major revisions saw the light of day:
Shen & Wu’s revision of the tripterygiid fishes of Taiwan
and Fricke’s Tripterygiid fishes of Australia, New Zealand
and the southwestern Pacific Ocean. The most recent is
Fricke’s (1997) revision of the tripterygiid fishes of the
western and central Pacific Ocean.
This review of western Indian Ocean (WIO) species
of Enneapterygius is not comprehensive in that much
available material has not been examined. Its main
purpose is to provide information to identify newly
collected specimens, as well as preserved material that
has lost all coloration. The revision is based on material
from the St Brandon Shoals (collections of Springer in
1976), the Chagos Archipelago (collections of
Winterbottom and Emery in 1979), from the Comoro
Islands (collections of Winterbottom et al. in 1988) and
material available at the South African Institute for
Aquatic Biodiversity. This revision has also benefited
considerably from Fricke’s 1994 and 1997 revisions and
from Randall’s more recent work in Oman and the
Arabian Gulf (Randall, 1995).
METHODS AND MATERIALS
Measurements were taken as in Hubbs and Lagler (1958);
fin element counts follow Springer (1968). Several
measurements used by me in earlier papers (e.g.
1
Table 1. Selected characters of WIO species of Enneapterygius; usual counts are given in parentheses.
* Counts from Randall, 1995.
^ Counts from Clark, 1979.
The western Indian Ocean as dealt with in this
revision includes the Red Sea, but excludes Sri Lanka.
Sufficient material for this island was not available at
this time.
CAS - California Academy of Sciences, San Francisco
LACM - Los Angeles County Museum, California
HUJ - Hebrew University, Jerusalem
PMBC - Phuket Marine Biological Center, Thailand
ROM - Royal Ontario Museum, Toronto
SAIAB - South African Institute for Aquatic
Biodiversity, Grahamstown (formerly the J.L.B. Smith
Institute of Ichthyology, previously with the institutional
abbreviation of RUSI.)
USNM - National Museum of Natural History,
Smithsonian Institution, USA
Table 1 summarises morphometric and meristic data for
the 17 species.
The following institutional abbreviations are used:
AMS - Australian Museum, Sydney
BMNH - Natural History Museum, London
BPBM - Bishop Museum, Honolulu
2
TAXONOMIC ACCOUNTS
KEY TO THE WESTERN INDIAN
OCEAN SPECIES OF ENNEAPTERYGIUS
Genus Enneapterygius Rüppell, 1835
1a.
1b.
Enneapterygius Rüppell, 1835: 2; type species E. pusillus
Rüppell, 1835, by monotypy.
2a. Pectoral-fin rays 14; belly naked; black spot on
margin of second dorsal fin; banded green or brown in
life; adults >20 mm SL ........................... E. tutuilae (WIO)
2b. Pectoral-fin rays 13; belly with thin cycloid scales;
cluster of melanophores on midside, beneath pectoral
fin; adults <20mm SLE.kosiensis sp. nov. (KwaZulu-Natal)
DESCRIPTION. Small fishes with fusiform bodies, less
than 25 mm SL (14 of the 16 WIO species - there are
Pacific species that are considerably larger). First dorsal
fin with 3 spines; second with 11-14 spines; third with
8-11 simple rays, except the 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.
Height of first dorsal fin is about the same height as
second or considerably taller. Pelvic fins with one short,
hidden spine and two slender, simple rays which may
be united by a membrane for part of the length of the
shorter ray.
3a. Caudal peduncle with distinct dark bar or marking;
body with well defined vertical or oblique dark bars
(except E. elegans); anal fin with or without dark bars ..
............................................................................................ 4
3b. Caudal peduncle without distinctive bar or
marking; body without well defined bars ..................... 9
The supratemporal sensory canal is either U-shaped
with a pore at each posteromedial corner of the canal,
curving around the first dorsal fin, or variously curved
(Fig. 1). The mandibular canal has either one or two pores
at the jaw symphysis and 3-5 pores either side (except
for 2 species). Exposed edge of posttemporal bones with
fine serrations. Body with ctenoid scales; nape naked or
scaled; abdomen naked or with thin cycloid scales
(which are difficult to see) in three species. Lateral line
discontinuous, an anterior segment of 8-18 pored scales
ending below second dorsal fin, and a posterior segment
of 16-28 notched scales, starting 1-2 scale rows below
end of the pored segment, and continuing to base of
caudal fin. Orbital and anterior nasal cirri simple, the
nasal cirrus on the posterior margin of a short tube.
Teeth in jaws conical and slightly recurved, a row of
larger teeth in front followed by a variable-width band
of smaller teeth behind. Vomer always with a single row
of conical teeth; palatines edentate or with few small
teeth.
A
B
Pectoral-fin rays 13-14, all unbranched ............... 2
Pectoral-fin rays 13-16, at least some branched .. 3
4a. Caudal peduncle bar/marking not continuous, but
broken in some way ......................................................... 5
4b. Caudal peduncle bar continuous, encircling the
peduncle ........................................................................... 7
5a. Pored lateral-line scales 14-18; anal-fin rays 16
(rarely)-19 .......................................................................... 6
5b. Pored lateral-line scales 8-12 (usually 10-11); analfin rays 15-16 (usually 16); caudal peduncle mark two
round black spots, one above the other; body with oblique
black bars, pink in life ........................ E. destai (Red Sea)
6a. Pored lateral-line scales 16-18 (usually 17); notched
lateral-line scales 16-18 (usually 17); anal-fin rays 16-17
(usually 17); caudal peduncle mark two squarish black
spots, one above the other; body with indistinct darker
bars, red in life; males with lower half of head black . E.
elegans (WIO)
6b. Pored lateral-line scales 14-16 (usually 15); notched
lateral-line scales 19-23 (usually 20-21); anal-fin rays
17-19 (usually 18-19); caudal peduncle mark often
darker dorsally, often with small, clear window on
midline; small, dark saddle mark at base of last dorsal
fin rays; body cream-coloured in life with dark brown
bars which may divide ventrally .....................................
.......................... E. gruschkai sp.nov. (WIO islands)
7a. Pored lateral-line scales 15-16 (rarely 13-14); analfin rays 17-18 .................................................................... 8
7b. Pored lateral line scales 11-12 (usually 12); analfin rays 16-17 (usually 16); body with vertical dark bars,
often dividing ventrally; black bar encircling caudal
peduncle; belly with thin cycloid scales; pectoral-fin base
with single row of cycloid scales .......... E. clarkae (WIO)
C
Figure 1. Diagram of supratemporal sensory canals of
Enneapterygius destai, E. obscurus and E.
pusillus, showing the open ‘C’, deep ‘C’ and ‘U’shaped forms, respectively.
8a. Body with 6 distinct vertical dark bars; second
dorsal fin with black spot basally, capped by a bright
orange arc in life; caudal peduncle bar broad with
white band at base of caudal fin .....................................
...................................................... E. melanospilos (Oman)
3
8b.
Body bars oblique; second dorsal fin without black
spot; caudal peduncle bar narrow, lying at base of caudal
peduncle; anal fin with 6-7 dark basal spots, which are
dark blue in life .............. E. fasciatus (WIO to Indonesia)
pectoral fin base; pored lateral-line scales 13-15 (usually
15); ...................... ..................... E. genamaculatus sp. nov.
(St Brandon Shoals)
16a. Black on head of males to nape, pectoral fin base
and base of pelvic fins; entire body yellow to yellowgreen in life; females with line of melanophores from eye
onto upper lip; second dorsal fin 11-12 (usually 12); anal
fin rays 17-18 (usually 18)..... ..................... E. abeli (WIO)
16b ... . Black on head of males only to posterior margins
of orbit and preopercle; yellow on body confined to
anterior third; females with line of colour from eye to
upper lip in life only; second dorsal fin 12-13 (usually
13); anal fin rays 17-19 (usually 19) ................................
............................................. E. elaine sp. nov. (Rodrigues)
9a.
Supratemporal sensory canal U-shaped, curving
around first 1-2 dorsal-fin spines ................................ 10
9b.
Supratemporal sensory canal C-shaped, running
in a curve in front of the first dorsal-fin spine ........... 13
10a. .. Anal fin with 5-7 oblique black bars, basal spots
present or absent; mandibular pores 3+1+3 or 3+2+3 ..
.......................................................................................... 11
10b. Anal fin without bars or basal spots; mandibular
pores 2+2+2; body without distinguishing marks,
probably bright green in life .......... E. pallidus (Red Sea)
Enneapterygius abeli (Klausewitz)
(Fig. 2; Pl.2)
11a. Anal fin basal spots present; black preanal spot
present or absent; body with broad, diffuse dark bars
containing 3-4 pale (white in life) blotches along midside; white line at base of caudal fin; anal-fin rays 19
............................................................................... 12
11b. Anal fin basal spots absent; black preanal spot
present; body without bars; first dorsal fin of males higher
than second; anal-fin rays 19 (Red Sea) or 20-21 (East
coast of Africa); multicoloured in life ..............................
............................. E. pusillus (Red Sea; E coast of Africa)
Tripterygion abeli Klausewitz, 1960: 11, figs. 1-2 (Al
Ghardaqa, Red Sea).
Helcogramma abeli: Lal Mohan, 1971: 222
Enneapterygius abeli: Clark 1980: 97, figs. 2a, 2b, 4c & 10;
Holleman, 1986: 756, figs. 236.2.
DESCRIPTION. Dorsal fins III+XI-XIII+9-10 (usually
III+XII+10); anal fin I,17-18 (usually 18); pectoral fins
15: 2-4+4-6+7 (usually 3+5+7). Lateral line with 12-14
pored scales and 19-23 (usually 21- east coast of South
Africa - or 22 - Red Sea) notched scales. Notched segment
starts one scale row below end of pored segment (Figure
2c); total lateral scales 31; transverse scales 3/6. Vertebrae
10+24-26. Mandibular pores 3+1+3 (Fig. 2d). Head
length 3.2-3.8, body depth 4.5-5.2 in SL; eye 2.9-3.7 in
head length.
Nape scaled, base of first dorsal fin and belly naked.
Pelvic fin rays united by membrane for about half length
of shorter ray; longest ray reaching mid-vent. First dorsal
fin equal in height to second in males, lower in females;
second dorsal fin about 60% of body depth.
Supratemporal sensory canal crescent-shaped; mouth
slightly down-turned, reaching vertical through anterior
of pupil; orbital cirrus small and pointed.
Live colour. Adult males more darkly pigmented than
females or juveniles, with entire head, nape, pre-pectoral
area, pelvic fin bases and proximal third of pelvic fins
black. Body deep yellow (females) to yellow-green with
irregular black markings (males). Females and immature
individuals (over 14 mm SL) from Red Sea and Indian
Ocean islands with oblique line of melanophores from
either side of upper lip to anterior margin of eye, absent
in specimens from KwaZulu-Natal, South Africa. Dorsal
and caudal fins pinkish with black on margin of first
dorsal fin. Clark (1980) records Red Sea male specimens
with irregular vertical bars across body, a dark margin
to the second dorsal fin and incomplete, slender stripes
on third dorsal and caudal fins.
Colour in alcohol. Specimens straw-coloured except
for black head of males, black margin of first dorsal fin
and nasal stripes of females. (See Comparisons below.)
12a. .. Pectoral-fin rays 15; second dorsal-fin spines 13;
first dorsal fin not higher than second in males, white in
life; caudal peduncle with quadrangular dusky mark .
............................................................ E. hollemani (Oman)
12b. Pectoral-fin rays 14; second dorsal-fin spines
usually 12; first dorsal fin higher than second in males,
cream-yellow in life; caudal peduncle with triangular
dusky mark, apex anterior ......... E. ventermaculus (WIO)
13a. Pectoral-fin rays 15-16; single symphyseal
mandibular pore present; no labial folds ................... 14
13b. Pectoral-fin rays 14; mandibular pores 2+2+2; nape
scaled; labial folds prominent; no distinguishing marks
except for faint oblique bands on the anal fin; crimson in
life, first dorsal fin yellow and white ..............................
................................................................. E. obscurus (WIO)
14a.
Body with or without scattered melanophores;
head of males black or with cluster of melanophores on
cheek ............................................................................... 15
14b. Body, head and fins of males densely and evenly
covered with melanophores, lower half of head generally
darker than upper; of immature males and females less
densely covered with melanophores sometimes showing
indistinct oblique bars on body and anal fin
...................................... E. philippinus (WIO to W Pacific)
15a. Males with black head and throat, females with
line from eye onto upper lip in life; body of males with
orange-yellow to yellow-green in life; pored lateral-line
scales 12-15 (usually 13-14) ......................................... 16
15b. Males with cluster of melanophores on cheek; both
sexes with small black spot at upper and lower ends of
4
A
C
B
D
Figure2.
Enneapterygius abeli. A: male, 21.3mm SL; KwaZulu-Natal, South Africa. B: female, 24.0mm SL; KwaZuluNatal. C: junction between pored and notched segments of lateral line. D: mandibular pores.
31510 (17.5 mm). Kenya: SAIAB 31506 (9: 17.8-19.2 mm),
Shimoni. Mozambique: SAIAB 31507 (3: 16.0-18.5 mm),
Ibo Island; SAIAB 31505 (5: 16.5-20.2 mm), Pinda; SAIAB
31513 (19.3 mm), Inhaca Island. South Africa: SAIAB
DISTRIBUTION. (Fig. 3). Common in Red Sea, along
East African coast to southern Kwazulu-Natal, at
Mauritius, Seychelles (Randall & van Egmond, 1994),
Comoro Islands and St Brandon Shoals; not recorded
from Gulf of Oman (Randall, 1995).
COMPARISONS. Alive or freshly dead the bright
yellow body colour and the black head and chest of the
males is distinctive. Without any pigment (females in
particular) it is difficult to separate specimens of E. abeli
from E. elaine, E. obscurus and E. genamaculatus. In body
form and morphometrics the four species are very similar:
E. obscurus has higher dorsal fins than E. abeli, the length
of the longest spine being sub-equal to body depth,
whereas in E. abeli the longest spine is only about 60% of
body depth. Enneapterygius abeli may be separated from
E. elaine and E. genamaculatus by dorsal- and anal-fin
counts. Two of the species also differ in mandibular pore
patterns: E. abeli - 3+1+3 and E. obscurus - 2+2+2, while
E. abeli and E. elaine have the same pattern. See also under
E. elaine.
MATERIAL EXAMINED. Red Sea: BPBM 19869 (18.5
mm), Gulf of Aqaba; BPBM 35712 (15 & 16 mm), Hanish
Group, southern Red Sea. Comoro Ids: ROM 5470 (12:
11.8-18.6 mm); ROM 5474 (15.3 & 19.1 mm); ROM 67483
(10: 10.3-19.2 mm); ROM 67482 (15: 13.8-18.7 mm); ROM
67480 (17 mm); ROM 67481 (17.5 mm). Mauritius: SAIAB
31508 (19 mm); SAIAB 31511 (18.1 & 18.3 mm); SAIAB
31509 (3: 12.2-17.5 mm); SAIAB 60677 (42: 12.6-23.6 mm);
SAIAB 60681 (26:18.5-21.2 mm). St Brandon Shoals: SAIAB
Figure 3. Distribution of E. abeli (filled star), E. clarkae
(open star), E. elaine (star in square) E. elegans
(open star in roundel), E. destai (filled roundel)
and E. fasciatus (filled triangle).
5
B
A
Figure 4
C
Enneapterygius clarkae. A: holotype male, 23.5mm SL; Inhaca Island, Mozambique, SAIAB14175. B: junction
between pored and notched segments of lateral line. C: mandibular pores.
32418 (67: 10-24 mm); SAIAB 31514 (85:10-23 mm);
SAIAB 32415 (33: 9-24 mm); SAIAB 7934 (4: 15.3-21.1
mm); SAIAB 7936 (21.3 & 21.4 mm); SAIAB 9612 (18.3 &
19.5 mm); SAIAB 12362 (17.2 & 17.7mm).
portion of head. First dorsal fin dusky; second dorsal fin
with irregular dusky bars and one or two black marks
on the margin; third dorsal and pectoral fins with
irregular dusky markings. Little sexual dichromatism
was noted, except that males have red on first and second
dorsal fins and are generally more darkly marked than
females.
Colour in alcohol. Reds and brown fade and specimens
retain the bars and spotting as described above.
Enneapterygius clarkae Holleman
(Fig. 4, Pl. 1)
Enneapterygius clarkae Holleman, 1982: 121, fig. 6 (Inhaca
Island, Mozambique).
Enneapterygius n.sp. 2 Clark, 1980: 104, figs. 4a & 14
DISTRIBUTION. (Fig. 3) Red Sea, east coast of Africa to
northern KwaZulu-Natal, South Africa, the Comoro
Islands and St Brandon Shoals; not recorded from
Mauritius or Gulf of Oman.
DESCRIPTION. Dorsal fins III+XI-XII+8-10 (usually
III+XII+9); anal fin I,16-17 (usually 16 rays); pectoral fins
15: 2-3+5+6+7 (usually 2+6+7). Lateral line with 11-12
(usually 12) pored scales and 20-22 (usually 22) notched
scales, notched segment starting next scale row below
end of pored segment (Fig. 4b). Total lateral scales 29 30; transverse scales 3/6. Vertebrae 10+22-23.
Mandibular pores 3+2+3 (Fig. 4c). 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. Pelvic-fin rays
united for less than half their length, the longest ray
reaching almost to vent. First dorsal fin equal in height
to second in females, slightly higher (about 10%) in males.
Mouth slightly down-turned and reaches vertical
through anterior of pupil. Orbital cirrus length about
half pupil diameter, with serrate end; upper and posterior
margins of orbits with fine serrations. Supratemporal
sensory canal crescent-shaped.
Live colour. Body of males and females cream with 45 dark brown transverse bars, which continue across
the anal fin, darkest across caudal peduncle, least
conspicuous under pectoral fin. Bars may be partially
divided, forming Hs or inverted Ys; anal fin bars form
basal, subcutaneous black spots; prominent pre-anal
spot present. Anterior half of body dusted with
melanophores which may form irregular bars on lower
COMPARISONS. The species is very similar to E. destai
and E. ventermaculus. Fricke (1997) synonymised E. clarkae
and E. destai. The bars across the body of E. destai are less
well-defined than in E. clarkae, except for the caudal
peduncle bar of E. destai which has a distinctive
hourglass-shape; that of E. clarkae is solid. The live
colours of the two species are also different: bright pink
for E. destai and cream and dark brown for E. clarkae.
Body bars are absent in E. ventermaculus. Enneapterygius
clarkae also has a scaled nape and belly, naked in the
other two species.
MATERIAL EXAMINED, additional to that listed in
Holleman, 1982. Comoro Islands: ROM 67752 (16.5 mm);
ROM 67474 (16.4 mm); ROM 67472 (17.0 & 20.5 mm). St
Brandon Shoals: USNM 357598 (3: 18.1-19.5 mm); USNM
357609 (20.2 mm).
Enneapterygius destai Clark
(Fig. 5)
Enneapterygius destai Clark, 1980: 102, figs. 4b & 13 (Red
Sea).
6
A
Figure 5.
B
Enneapterygius destai. A: female, 20.2mm SL; Eritrea, Red Sea (from Clark, 1980). B: mandibular pores.
COMPARISONS. See under E. clarkae above.
DESCRIPTION (based partly on Clark, 1980;
measurements of examined material). Dorsal fins III+XIXIII+8-9 (usually III+XII+8-9); anal fin I,15-17 (usually
16 rays); pectoral fins usually 15: 3,5,7. Lateral line with
8-12 (usually 10-11) pored scales and 21-23 notched
scales, notched segment starting second scale row below
end of pored segment. Total lateral scales 28; transverse
scales 2/5. Vertebrae 10+21-22. Mandibular pores 3+2+3
(Fig. 5b). Head length 3.1-3.3, body depth 4.8-5.1 in SL;
eye 2.5-3.0 in head length.
Body robust. Scales large, nape scaled, scales to bases
of dorsal and anal fins. Two rows of scales at base of
caudal fin. Pelvic fins not united by membrane. First
dorsal fin lower than second, about 60% of height of
second. Pectoral fin of males long, longest ray extending
to 2nd or 3rd dorsal-fin ray. Mouth extending to vertical
through anterior margin of pupil. Orbital cirrus a
rounded flap, about half pupil diameter. Supratemporal
sensory canal crescent-shaped.
Live colour. Freshly dead adults with head, body and
first dorsal fin bright pink, with tints of pink on other
fins except pelvics. Five broad oblique dusky bands
across body present to some degree in adults, but absent
in juveniles. Two prominent black spots on caudal
peduncle form an hourglass-shape, upper usually darker
than lower. Anal fin with 5, rarely 6, oblique black bars.
Prominent black spot present in front of vent on ventral
midline. Second and third dorsal fins with irregular
dusky bars, second dorsal black in mature males; pectoral
fin with 4 vertical dusky bars; 2-3 dusky bars radiate
across the lower head from the lower posterior quadrant
of the eye. Females generally less intensely coloured than
males.
Colour in alcohol. Pink fades and the body becomes a
pale straw colour, retaining dusky bars on body and
fins, the black hour-glass mark across the caudal
peduncle and the black bars on the anal fin.
MATERIAL EXAMINED. SAIAB 69185 (17.3 mm);
SAIAB 69262 (15.0 mm); SAIAB 69204 (4: 15.4-17.3 mm),
all from Karaman Island, Uqban,Yemen.
Enneapterygius elaine sp.nov.
Fig. 6, Pl. 1
Holotype: SAIAB 65582, male 18.5 mm SL; Rodrigues,
Baladirou (ca 19°40' S, 63°26’E); bay with coral
rubble; collected P C Heemstra et al., 16 October 2001;
sta. ROD-34.
Paratypes (all from Rodrigues): AMS 41476-001 (19.6 &
21.0 mm), sta. ROD-37; BPBM 39012 (4: 20.3-21.9 mm),
sta. ROD-37; BMNH 2002-8-22:1-3 (3: 17.2-19.2 mm), sta.
ROD-34; ROM 73204 (4: 15.3-22.7 mm), sta. ROD-35;
SAIAB 65577 (3: 15.0-18.9 mm), sta. ROD-34; SAIAB
65578 (5: 14.6-20.3 mm), sta. ROD-43; SAIAB 65579 (17.4
& 17.6 mm), sta. ROD-04; SAIAB 65580 (3: 17.8-19.1 mm),
sta. ROD-27; SAIAB 65581 (3: 21.0-20.7 mm), sta.ROD31; SAIAB 65584 (5: 13.2-21.7 mm), sta. ROD-35; 65586
(5: 15.3-21.7 mm), sta. ROD-37; USNM 370527 (4: 14.120.2), sta. ROD-31.
DESCRIPTION. Dorsal fins III + XIII + 10 (rarely with
12 spines and 9-11 rays); anal fin I,17-19; pectoral fins
15: 3+5+7 (rarely 4+4+7); lateral line with 14-15 (usually
14) pored scales and 21-22 notched scales (rarely 20-23),
starting second scale row below end of pored segment
(Fig. 6b). Total lateral scales 33; transverse scales 2/5.
Vertebrae: 10 precaudal, 25 caudal. Mandibular pores
3+1+3 (Fig. 6c). Head 3.2-4.0 in SL; eye 2.4-3.0 in head
length.
Small fish, seldom longer than 20 mm SL. Body
slender, scales large, nape scaled, abdomen naked. Pelvic
fin rays slender, united by membrane for half the length
of the longer ray. First dorsal fin about half height of
second. Pectoral fin large, longest ray reaching to
DISTRIBUTION. The species is restricted to the Red
Sea, where in the south it replaces E. abeli as the most
abundant species.
7
B
A
Figure 6.
C
Enneapterygius elaine. A: holotype, male, 18.5mm SL; Rodrigues; SAIAB 65582. B: junction between pored
and notched segments of lateral line. C: mandibular pores.
penultimate spine of second dorsal fin. Mouth oblique,
small and pointed, maxilla reaching vertical through
anterior margin of orbit. Orbital cirrus small and pointed.
Live colour. Head of males densely covered with large
melanophores, colour dorsally to just behind orbits,
laterally to margin of preopercle and ventrally to base of
pelvic fins, spotting on lips and throat less dense than
elsewhere. Sometimes some brown on lower portion of
opercle. Body behind head and including the pectoralfin base bright yellow, colour extending to junction
between first and second dorsal fins and base of pectoral
fin rays, giving appearance of a fish wearing a yellow
waistcoat. Posterior to this body with light brown
reticulations colour generally on margins of scales.
Narrow brown bar across caudal peduncle. First dorsal
fin pale yellow with a few melanophores on membrane
between second and third spines and between third
spine and dorsum. Elements of all other median fins light
reddish-brown, membranes without colour; caudal fin
with narrow yellow-white band at base of rays. Pectoral
fin base with two light brown circular spots, one above
the other, rays pale brown, membranes immaculate.
Head of females forward of hind margin of orbit white
with two brown streaks running from eye to upper lip.
Head behind eyes brown, colour extending to nape and
posterior margin of opercle. Body with same light brown
reticulation as males. Median fins as in males, except
caudal fin which lacks yellow-white band at base.
Pectoral fin bases white, with two light brown circular
spots, one above the other, rays pale brown, membranes
immaculate.
Colour in alcohol. In both males and females all colour
fades except for the black of the melanophores on heads
of males and on margin of first dorsal fin.
currently known only from the lagoon around the island
of Rodrigues, western Indian Ocean.
COMPARISONS. At first glance Enneapterygius elaine
is very similar to E. abeli, but can be easily separated
both on colour pattern and counts. The black on the head
of male E. abeli extends to the base of the first dorsal fin
spine, the posterior margin of the opercles and onto the
belly whereas in E. elaine the black extends only to just
behind the eyes and to the posterior margin of the
preopercle. Enneapterygius elaine usually has 13 spines
in the second dorsal fin (usually 12 for E. abeli), usually
19 anal-fin rays (17-18 for E. abeli) and 14-15 pored lateral
line scales, compared to a norm of 13 for E. abeli. Female
E. elaine also lack the two nasal stripes of preserved female
E. abeli specimens.
Enneapterygius elegans (Peters)
(Fig. 7, Pl. 1)
Tripterygium elegans Peters, 1876: 441 (Mauritius)
(lectotype ZMB 9465, designated by Fricke, 1997: 182).
Enneapterygius elegans: Holleman, 1986: 756, pl. 116,
figure 236.4 ; Fricke & Randall, 1992: 3 (Maldives);.
Fricke, 1994: 203.
DESCRIPTION. Dorsal fins III+XI-XIII+8-10 (usually
III+XII+9); anal fin I,16-17 (usually 17 rays); pectoral fins
16: 2-3+6-7+7. Lateral line with 16-18 pored scales and
16-18 notched scales, notched segment starting one scale
row below end of pored segment, overlapping by 2-3
scales (Figure 7c).Total lateral scaled 30-31; transverse
scales 3/5. Mandibular pores 6-9+1+6-9 (Figure 7d).
Head length 3.2-3.6, body depth 3.8-4.6 in SL; eye 3.03.7 in head length.
Body robust. Nape scaled; belly with thin cycloid
scales, entire in males, posterior half only in females.
Pelvic-fin rays united by membrane for half length of the
shorter ray, longer ray extending three-quarters distance
to vent. First dorsal fin about two-thirds height of second
dorsal fin, height of second half to two-thirds body depth.
Mouth slightly oblique, extending to vertical through
ETYMOLOGY. The species is named for Elaine
Heemstra, in recognition of her considerable and
excellent contribution to the illustration of Indo-Pacific
fishes, including several in this paper. The epithet is to
be used as a noun in apposition.
DISTRIBUTION (Fig. 3). Enneapterygius elaine is
8
C
A
D
B
Figure 7.
Enneapterygius elegans. A: male, 30.3mm SL; St. Brandon Shoals, Mauritius, SAIAB 949, B: female, 35mm
SL; St. Brandon Shoals, Mauritius, SAIAB 949, C: junction between pored and notched segments of lateral
line. D: mandibular pores.
along the east coast of Africa from Kenya to Bazaruto,
Mozambique. It is also widely distributed through the
Indo-West Pacific Ocean (see Fricke, 1994).
anterior margin of pupil. Orbital cirrus small and simple.
Exposed posterior margins of median and lateral
extrascapulars finely serrate. Supratemporal sensory
canal crescent-shaped.
Live colour. Males with brick-red body and lighter,
irregular, narrow bars. Head from below middle of eye,
including upper lip, densely spotted with
melanophores, extending onto throat, pectoral fin bases
and anterior of belly; nasal area above upper lip not
spotted. Caudal peduncle distinctly marked with two
squarish, black spots separated by a mid-lateral
unpigmented space, often giving the impression of a
dumbbell or hourglass; bright yellow spot on the
peduncle just posterior to the last dorsal-fin ray. First
and second dorsal fins stippled with red and black,
second dorsal darker along margin and base, third
dorsal with broken orangey bars and black stipples.
Caudal fin with a narrow red stripe at its base; anal fin
red; pectoral fins red with creamy blotch at base of central
rays; pelvics stippled with red.
Females less intensely marked: body with indistinct
bands with light spots between; no black on lower part
of head and throat and fewer melanophores on head
and fins. Belly and pelvic fins white; stipples along the
base and margin of second dorsal fin indistinct.
Colour in alcohol. All colour fades and specimens are
pale brown, sometimes with indistinct banding on body
and always with twin black blotches on caudal peduncle
and black on head and throat of males.
COMPARISONS. Small Enneapterygius elegans may be
confused with specimens of E. destai or E. gruschkai, both
of which have broken bars across the caudal peduncle,
that of E. destai also two spots, one above the other.
Enneapterygius destai is however confined to the Red Sea.
Enneapterygius elegans can be readily separated from E.
gruschkai, with which it occurs sympatrically in the
western Indian Ocean, by lateral line count: usually
17+17 for E. elegans vs. usually 15+20-21 for E. gruschkai.
MATERIAL EXAMINED. Mauritius: BPBM 20171 (25
& 27 mm); SAIAB 61342 (19: 17.9-25.4 mm); SAIAB 60614
(4: 10.5-20.8 mm). Seychelles: SAIAB 7710 (10: 14.3-27.3
mm), Mahé; SAIAB 950 (41: 19.5-29.5 mm), La Digue.
Comoro Ids: ROM 67466 (4: 16.3-20.5 mm). St Brandon
Shoals: USNM 218483 (27.0 & 29.7 mm); USNM 218764
(4); USNM 218765 (26.5 & 29.7 mm); USNM 218766 (22:
18-23.8 mm); USNM 218767 (17: 19.4-30.0 mm). Kenya:
SAIAB 951 (6: 19.2-20.7 mm), Shimoni. Mozambique:
SAIAB 7700 (26.1 mm), Pinda; SAIAB 7702 (23.5 & 28.5
mm), Pinda; SAIAB 7699 (4:13.8-28.2 mm), Ibo; SAIAB
7698 (27.4 & 28.3 mm), Bazaruto.
Enneapterygius fasciatus (Weber)
(Fig. 8)
DISTRIBUTION (Fig. 3). Mauritius, Seychelles, Comoro
Islands, St Brandon Shoals and Maldives as well as
9
Tripterygium fasciatum Weber, 1909: 148 (Savu [Sawu],
southern Indonesia and the Karakelang [Karakelong]
Islands, northern Indonesia).
B
A
Figure 8.
C
Enneapterygius fasciatus, A: male, 26mm SL; Ibo Island, Mozambique, SAIAB 60609. B: junction between
pored and notched segments of lateral line. C: mandibular pores.
bands of melanophores and 6-8 distinctive basal analfin spots. Even scatter of melanophores cover head to
pectoral fin bases and anterior portion of belly. Dorsal
and anal fins with light scatter of melanophores. Females
lack the dark head, show more obvious banding and
also carry the basal anal fin spots.
Tripterygion (Enneapterygius) fasciatum: Mukerji, 1935: 272
Tripterygion fasciatum: Herre, 1939: 351
Tripterygion fasciatus: Lal Mohan, 1968: 116, figure 2
DESCRIPTION. Dorsal fins III+XI-XIII+9-11 (usually
III+XII+9-10); anal fin I,17; pectoral fins15-16: 3-4+4-5+78. Lateral line with 13-16 (usually 15-16) pored and 1821 notched scales, notched segment starting one scale
row below end of pored segment and overlapping it by 2
-3 scales (Fig. 8b). Total lateral scales 30-32; transverse
scales 3/6. Mandibular pores 3+1+3 (Figure 8c). Head
length 3.3-3.5, body depth 3.9-4.6 in SL; eye 2.9-3.2 in
head length.
Body robust. Nape scaled, belly naked. Pelvic fins
united by membrane for two-thirds of length of shorter
ray, longest ray reaching anus. First dorsal fin lower than
second, second about 70% of body depth. Supratemporal
sensory canal crescent-shaped. Mouth reaching vertical
through anterior margin of pupil; orbital cirrus small
and lobate.
Live colour. Weber’s original description of the species
gives live colours of a yellowish body with seven brown
crossbars, head with brown flecks and bands, a dark
base to the first and second dorsal fins and seven barlike flecks on the anal fin. Herre (1939: 397) gives a colour
description that closely agrees with that of Weber. A more
recent description by Lal Mohan (1968: 117), based on a
single specimen, reads: “Cheeks, snout, anterior part of
belly mottled with blue chromatophores, a dark blotch
on first dorsal between first and second spines; five
vertical irregular greenish bands on body, a brown blotch
on lower part of opercle, a reddish blotch on base of
pectoral peduncle and ventral fin, six deep blue spots
on base of anal fin.” Herre however noted a fairly
distinctive dark band at the base of the caudal fin not
mentioned by Lal Mohan. His illustration, as does that
of Herre, shows the body bars as oblique, which is in
agreement with the specimens I have seen. The 6-8 black
(blue) spots at the base of the anal fin of both sexes are
noteworthy.
Colour in alcohol. Body of males light brown with faint
DISTRIBUTION (Fig. 3). East coast of Africa, the
Seychelles, the southeast coast of India, Sri Lanka, the
Andaman Islands and as far east as the Solomon Islands
(Fricke, 1997: 195).
Figure 9. Distribution of E. genamaculatus (circle/star), E.
gruschkai (closed star), E. hollemani, (open star),
E. kosiensis (open star), E. melanospilos (closed
triangle), E. obscurus (closed roundel) and E.
philippinus (square/star).
10
B
A
Figure10.
C
Enneapterygius genamaculatus, paratype, A: male, 21.0mm SL; St Brandon Shoals; SAIAB 60503.
junction between pored and notched segments of lateral line. C: mandibular pores.
B:
for entire length of shorter ray, longer ray reaching analfin spine. First dorsal fin lower than second; maximum
height of second dorsal fin about 70% of body depth.
Supratemporal sensory canal crescent-shaped. Orbital
cirrus small and rounded. Tip of tongue narrow and
rounded; palatines with teeth; maxilla reaching vertical
through anterior margin of pupil.
Live colour. The live colour of the species is not known.
Colour in alcohol. Specimens straw-coloured. Males
with large cluster of melanophores below and slightly
posterior to the eye. Large specimens may have some
pigment on the snout, in the interorbital area, on top of
the head and on the membrane across the throat. Two
small clusters of melanophores at upper and lower ends
of pectoral-fin base and small black spots on membrane
between third dorsal-fin spine and dorsum. Females with
only the two clusters of melanophores at the corners of
the pectoral-fin base and a few melanophores on the first
dorsal-fin membrane.
COMPARISONS. Enneapterygius fasciatus is similar in
size and robustness to E. elegans, but lacks the scaled
abdomen and conspicuous peduncular mark of that
species (note: not the band at the base of the caudal fin).
Their live colours also separate them: greenish with
brown banding for E. fasciatus and brick-red for E .elegans.
MATERIAL EXAMINED. Kenya: SAIAB 60612 (7: 17.522.2 mm), Malindi; SAIAB 60682 (23.0 mm), Shimoni;
LACM 31617-27 (25.8 mm), Manda Island. Tanzania:
SAIAB 60683 (28.3 & 29.5 mm), Zanzibar. Mozambique:
SAIAB 60609 (25.3 mm), Ibo; SAIAB 60610 (28.1 mm)
Mocimboa da Praia. Comoro Ids: ROM 5474 (15.3 & 20.3
mm).
Enneapterygius genamaculatus sp. nov.
(Fig. 10)
Holotype : USNM 357600, male 22.3 mm SL; St Brandon
Shoals, SE side of Grande Passe (ca 16°28' S, 59°40'
E); dead and rocky reef channels and ledges; collected
V G Springer et al. 5 April 1976; sta.VGS 76-9.
DISTRIBUTION (Fig. 9). The only known specimens
are those collected by Springer et al. from St Brandon
Shoals.
Paratypes (all St Brandon Shoals): SAIAB 60503 (6: 19.321.4 mm), same collection as USNM 357602; USNM
357597 (17.8 & 21.0 mm), sta.VGS 76-22; USNM 357606
(17.5 & 19.7 mm), sta. VGS 76-7; USNM 357602 (12: 19.522.8 mm), sta.VGS 76-10; USNM 357599 (29: 17.6-22.9
mm), sta.VGS 76-9.
ETYMOLOGY. The name is taken from the Latin gena, a
cheek, and macula, a spot, and is given for the spots on
the cheeks of the males of the species.
COMPARISONS. Morphometrically the species is
similar to E. gruschkai, but lacks the body markings of
that species. Unmarked specimens would be impossible
to separate except on mandibular pore patterns: 3+1+3
for E. genamaculatus and 3+2+3 for E. gruschkai.
DESCRIPTION. Dorsal fins III+XII-XIII+9-11 (usually
III+XIII+10); anal fin I,18-19 (usually 19 rays); pectoral
fins 15-16: 3-4+5-6+7-8. Lateral line with 13-15 pored
and 21-23 notched scales, notched segment starting one
scale row below end of pored segment (Fig. 10b). Total
lateral scales 32-33; transverse scales 2/4. Vertebrae
10+24. Mandibular pores 3+1+3 (Fig. 10c). Head length
3.5-3.9, body depth 4.6-5.4 in SL; eye 2.7-3.1 in head
length.
Small (less than 25 mm SL), slender fishes; nape
scaled, belly naked. Pelvic-fin rays united by membrane
Enneapterygius gruschkai sp.nov.
(Fig. 11; Pl. 1)
Holotype: SAIAB 61281; male, 24.3 mm SL; Passe
dAmbulante, Mauritius (20°26’10"S, 57°17’40"E); depth
6-8m, spur and groove formation; collected P. C. Heemstra
et al., 18 May 1995; sta. PCH 95-M30.
11
B
A
Figure 11.
C
Enneapterygius gruschkai, paratype, A: male, 27.2mm SL; Mauritius; SAIAB 61282. B: junction between
pored and notched segments of lateral line. C: mandibular pores.
brown body with 5 narrow creamy-white bars, the last
at the base of the caudal fin. Darkest bar across caudal
peduncle with small, clear window mid-laterally;
peduncle cream or yellow either side of bar. Penultimate
two bars darkest dorsally, forming a small dark saddles
at base of last two dorsal-fin rays and on caudal
peduncle. Scale-pocket margins with brown and orange
spots. Head markings dark brown to black, the face
almost tiger-striped; top of head deep orange with black
stippling. First dorsal fin with black stipples; second
with orange bands on spines and black stipples on
membranes; third same as second, but paler. Anal fin
stippled black except for tips of rays, which are
colourless. Pectoral fins with a black spot near dorsal
and ventral ends of base, a white blotch posterior to the
lower black spot, followed by another black spot and a
larger white blotch in the centre of the fin; rays orange
distally. Caudal fin deep orange to red proximally, with
narrow band of black stipples at its base, and dusky
distally.
Females and juveniles less heavily pigmented, body
bars and facial markings pale brown; dorsal fins with
yellow-orange pigment only and no black stippling; anal
fin pale yellow with some black stippling; caudal fin
with orange and black stippling at base of rays.
Colour in alcohol. All colour except black fades. Large
males with 5 dark bars across pale cream body, darkest
across caudal peduncle, with characteristic window at
centre, and dark saddle at base of last two dorsal-fin
rays. Face tiger-striped. Females and juveniles may lose
all colour except for the peduncular bar and saddle.
Juveniles, females and less heavily pigmented males may
have no markings except for the double saddle mark on
the caudal peduncle.
Paratypes: Seychelles: BPBM 35600 (28.1 mm). Comoro Ids:
CAS 33424 (23.3 mm); ROM 71834 (12.1 & 23.7 mm);
ROM 71833 (10: 10.4-25.2 mm). Chagos Archipelago: ROM
56795 (4: 17.8-23.1 mm); ROM 56796 (18.4 mm); ROM
56797 (21.8 mm); ROM 56798 (23.2 mm); ROM 56799
(20.5 & 22.0 mm); ROM 56800 (20.0 mm); ROM 56801 (5:
16.6-24.4 mm); ROM 56802 (6: 17.3-25.2 mm); USNM
357612 (25.8 mm). St Brandon Shoals: USNM 357601 (6:
22.6-29.2 mm); USNM 357607 (23.7 mm); USNM 357611
(22.3 & 24.5 mm); USNM 357604 (22.7 & 24.1 mm); USNM
357596 (5: 23.0-24.7 mm); USNM 357608 (8: 23.2-27.0
mm). Mauritius: SAIAB 61282 (11: 13.0-26.3 mm); SAIAB
61283 (11: 22.4-25.2 mm).
Non-types: Comoro Ids: CAS 34507 (21.3 mm). St Brandon
Shoals: USNM (VGS 76-18) (22.1 mm).
DESCRIPTION. Dorsal fin III+XII-XIII+8-11 (usually
III+XIII+9-10); anal fin I,17-19 (usually 18-19 rays);
pectoral fins 15: 2+7+6 (occasionally 3+6+6); lateral line
with 14-16 (usually 15) pored scales and 19-23 (usually
20-21) notched scales, notched segment starting next
scale row below end of pored segment, below mid-second
dorsal fin (Fig. 11b). Total lateral scales 33; transverse
scales 3/5. Vertebrae 10+25. Mandibular pores 3+2+3
(Fig. 11c). Head length 3.1-3.6, body depth 5.4-5.9 in SL;
eye 2.7-3.2 in head length.
A medium-sized species of Enneapterygius (less than
30 mm SL). Nape scaled, body scales continuing to bases
of dorsal and anal fins; belly naked. Pelvic-fin rays not
united by membrane. First dorsal fin equal in height to
second, second about 70% of body depth. Simple, lobate
orbital and anterior nasal cirri present. Supratemporal
sensory canal slightly curved. Tongue triangular and
sharply pointed; maxilla reaches vertical through
anterior margin of pupil; broad band of teeth in upper
and front of lower jaw; palatines without teeth. Spines
of first dorsal fin with small, laterally projecting spinelets.
Live colour. Colour of freshly caught males varies from
5 dark brown bars on creamy-yellow ground body colour,
with bars sometimes divided below midline, to a rich
ETYMOLOGY. The species is named for Victor
Gruschka Springer, doyen of blennioid systematics.
DISTRIBUTION (Fig. 9). The species seems to be
restricted to the islands of the western and central Indian
Ocean and has not been recorded from the east coast of
Africa.
12
First dorsal fin slightly dusky; second dorsal fin with
broad dusky band at base; third dorsal and caudal fin
pale. Anal fin with 7 black basal spots which continue
subcutaneously and as diagonal bands across the fin.
Prepectoral region dusky, darkest ventrally at base of
fin, continuing as a pointed dark blotch below
branchiostegal membranes. Pectoral fins pale with dark
brown spots on rays forming 6 irregular transverse
bands.
COMPARISONS. Enneapterygius gruschkai may be
mistaken for small specimens of E. elegans, but the former
species has 18-19 anal-fin rays compared to 16-17 for E.
elegans and 15 pored and 20-21 notched lateral-line scales
compared to 16-18+16-18 for E. elegans. Enneapterygius
gruschkai also lacks the scaled belly of E. elegans.
Enneapterygius hollemani Randall
(Pl. 1)
DISTRIBUTION (Fig. 9). Described from the Gulf of
Oman where 3 specimens were collected from shallow
tidepools along a rocky shore.
Enneapterygius hollemani Randall, 1995: 27, figure 1 (Gulf
of Oman).
DESCRIPTION (from Randall, 1995). Dorsal fins
III+XIII+10; anal fin I,19; pectoral fins 15: 2-3+6+6-7.
Lateral line with 16 pored and 23-24, notched scales
starting second scale row below end of pored segment
and overlapping it by a single scale. Mandibular pores
3+1+3. Head length 3.3 in SL; eye 3.2-3.4 in head length;
body depth 4.6-5.2 in SL.
Body moderately elongate. Nape scaled except
adjacent to anterior of first dorsal fin, belly naked. Pelvicfin rays united by membrane for half of length of shorter
ray, longest ray reaching anus. First dorsal fin equal in
height to second, about 65% of body depth.
Supratemporal sensory canal U-shaped. Mouth small,
reaching vertical through anterior of orbit. Orbital cirrus
very small and lobate.
Live colour. Body of males pale green, edges of scales
with red and dark brown spots forming 3 broad dusky
bars which divide ventrally, 2 below second dorsal fin
and one below third dorsal fin. Bars darker ventrally
than dorsally and separated by clusters of white flecks.
Quadrangular dark mark on caudal peduncle, its rear
margin a narrow dark brown bar at base of caudal fin.
Belly white, anus edged in black; black preanal spot may
or may not be present. Head a mixture of pale green, red
and brown, becoming pale green ventrally. Upper lip
dusky; orbital cirrus white. First dorsal fin creamy-white
with red tip to first spine; second dorsal fin with red
cross-bands on spines, 3 oblique white bands and a
dusky band at its base; third dorsal fin similar but paler.
Anal fin with 7 black spots basally, which continue
internally and diagonally across fin with white in
between; rays with white and dark red spots. Pectoral
fin rays white with red edges and 6 dark brown spots
forming 6 irregular transverse bars, lower rays orangeyellow distally; membranes pale. Pelvic fins white.
Caudal fin with narrow white band at base, transparent
membranes and bright green rays.
Colour in alcohol. Body tan with melanophores along
scale margins except ventrally, forming bands as
described above. Quadrangular dusky mark on
peduncle with narrow dark band at base of caudal fin;
head below eyes dusky, cheeks paler. Small dark brown
spot just anterior to origin of lateral line and a second
spot of same size just postero-ventral to the first. Anus
edged in black, black preanal spot present in 2 of the 3
specimens on which the original description is based.
COMPARISONS. Enneapterygius hollemani is similar in
colour to E. ventermaculus, but differs in the shape of the
peduncular mark (quadrangular in E. hollemani,
triangular in E. ventermaculus), the colour of the first
dorsal fin (brown and orange in E. ventermaculus; white
in E. hollemani), and pelvic fins (yellowish in E.
ventermaculus, white in E. hollemani) and on the head
(bottle green with more red and orange in E.
ventermaculus, greenish in E. hollemani). Enneapterygius
hollemani also has 15 pectoral fin rays vs. 13 (rarely) or
14 for E. ventermaculus; 16 pored lateral line scales vs. a
mode of 15 for E. ventermaculus and 13 spines in the second
dorsal fin vs. a mode of 12 for E. ventermaculus. Other
differences are detailed in Randall (1995: 29).
Enneapterygius kosiensis sp. nov.
(Fig. 12)
Holotype. SAIAB 59086 (19.4 mm SL), male, Sodwana
Bay, KwaZulu-Natal, South Africa, coll. P. C. Heemstra
et al. 10 August 1998; sta. FW 98-12.
Paratypes. SAIAB, 40265 (9: 14.5-17.1 mm SL ), reef 2 km
south of Kosi Mouth, KwaZulu-Natal, South Africa, coll.
C. D. Buxton et al., 9 August 1992; sta. 1-92-11. SAIAB
9897 (18.1 mm SL) Sodwana Bay, KwaZulu-Natal, coll.
P. C. Heemstra et al., 22 April 1979; sta. PCH 79-27.
DESCRIPTION. Dorsal fins III+XII+9; anal fin I,16-17
(usually 17 rays); pectoral fins 13, all rays simple; lateral
line with 12 pored scales and 21-22 notched scales,
notched segment starting second scale row below end of
pored segment, overlapping by one scale (Fig. 12b). Total
lateral scales 32; transverse scales 1/4. Vertebrae 10+22.
Mandibular pores 2+1/2+2 (Fig. 12c). Head 3.2-3.5, body
depth 5.1-5.3 in SL; eye 3.1-3.7 in head length; maxilla
2.6-2.9 in head length.
Small fish, less than 20 mm SL. Body slender; scales
large; nape scaled; belly with smaller, thin cycloid scales.
Pelvic fins slender, rays united by a thin, fragile
membrane for about one third length of shorter ray. First
dorsal fin slightly higher than second, second 75-90% of
body depth. Longest pectoral-fin ray reaches nearly to
base of first ray of third dorsal fin. Supratemporal sensory
canal crescent-shaped. Mouth oblique, reaching vertical
through anterior margin of pupil; interorbital concave
13
B
C
A
Figure 12.
Enneapterygius kosiensis holotype, A: male, 19.4mm SL; KwaZulu-Natal, South Africa; SAIAB 59086.
B: junction between pored and notched segments of lateral line. C: mandibular pores.
Enneapterygius melanospilos Randall, 1995: 29, fig. 2
(Arabian Sea, central coast of Oman).
lateral line with 14-15 pored and 21-22 notched scales,
notched segment starting one scale row below end of
pored segment. Total lateral scales 34. Mandibular pores
3+2+3. Head 3.2 in SL; eye 2.7-3.0 in head length; depth
4.8-5.5 in SL.
Body moderately elongate. Nape scaled, belly naked.
Pelvic-fin rays scarcely united by membrane. First dorsal
fin higher than second in males (by about 20%) and about
the same height as second in females, at about 75% of
body depth. Numerous small spinules along posterior
edge of supratemporal sensory canal and on upper
posterior edge of orbit. Supratemporal sensory canal
crescent-shaped. Mouth not large, reaching vertical
though anterior of orbit. Orbital cirrus bilobed or trilobed,
equal in length to pupil diameter.
Live colour. Body whitish with 6 orange-brown bars,
becoming progressively darker posteriorly, the darkest
across the caudal peduncle. Width of last 4 bars about
equal to white interspaces. Front, top of head and nape
brown mottled with orange; lower half of head pale with
brown-edged orange stripe below eye. Opercle dark
orangey-brown shading to yellowish posteriorly. First
dorsal fin pinkish; second with jet-black spot as large as
eye basally between 7th and 11th spines, fin white above
spot with bright orange arc on margin, remainder of fin
transparent with irregular oblique dusky bars with
orange on spines; third dorsal fin with 2 brownishorange bands, continuous with 4th and 5th body bars;
anal fin whitish with 4 broad dusky zones; caudal fin
with white band at base, membranes clear and rays pink,
becoming whitish at tips; pectoral fins with clear
membranes, upper rays pale yellowish, lower rays
yellow; pelvic fins white.
Colour in alcohol. Orange and yellow colours fade and
body becomes whitish with 6 dusky bands, head mottled,
stripe below eye, distinctive mark on second dorsal fin
and banding on other fins.
DESCRIPTION. (from Randall, 1995). Dorsal fins
III+XII+10; anal fin I,18; pectoral fins 15-16: 2+5-6+8;
DISTRIBUTION (Fig. 9). Described from three
specimens taken in the Arabian Sea off the coast of Oman.
and broad, about equal to pupil diameter; orbital cirrus
a simple rounded flap and about half pupil diameter in
length.
Live colour. Not known.
Colour in alcohol. Entire fish a pale straw colour. Males
with a few melanophores scattered on body and a cluster
of large melanophores on midside, beneath pectoral fin,
and a cluster of melanophores on the caudal peduncle
below the midline. Margin of first dorsal fin with cluster
of large melanophores, a band of melanophores parallel
to margin of second dorsal fin and 6 irregular black bars
on anal fin. Basal spots may be present subcutaneously
on anal fin and a preanal spot may be present. Females
lack the fin markings and have fewer melanophores on
the body.
ETYMOLOGY. The species in named after Kosi Bay,
the area off which specimens were first found.
DISTRIBUTION (Fig. 9). Currently known only from
northern KwaZulu-Natal, South Africa (Sodwana Bay
and Kosi Bay), taken on flat reef at 27-30m depth.
COMPARISONS. Preserved specimens of
Enneapterygius kosiensis may be mistaken for small
specimens of E. ventermaculus; E. kosiensis, however, has
a scaled belly and 2+1/2+2 mandibular pores, compared
to 3+1+3 for E. ventermaculus and 13 simple pectoral-fin
rays (vs. 14 for E. ventermaculus, of which 4-6 are divided).
Only one other western Indian Ocean species, E.tutuilae,
has all pectoral fin rays simple, 14 in number.
Enneapterygius melanospilos Randall
(Plate 1)
14
base of the caudal peduncle, which shows as deep red.
While the cluster of black spots on the lower base of the
pectoral fins is not visible, the other features confirm
identification of the Debelius illustrations as those of E.
obscurus.
Body of males transparent with irregular crimson
markings, scale margins lined with crimson; large
crimson blotch below pectoral fin, and two small deep
crimson spots at top and bottom of the caudal peduncle.
Vertical line of scales above lateral line, between the
second and third dorsal fins, edged in black. Opercles
crimson; upper portion of head crimson with some white
marks; eye crimson, except for a fine yellow line around
the pupil. Orbital cirri yellow. First dorsal fin deep
crimson anteriorly with yellow markings, white
posteriorly with black spots. Elements of second and
third dorsal fins crimson with irregular bands of
melanophores on membranes. Anal, caudal and
pectoral-fin elements pale red, membranes transparent.
(Preserved specimens show melanophores on the
membranes of the anal fin, banded in females.) Females
are lighter in colour, with pale red, irregular bands on
the body, and lack the intense red on the head and
brightly coloured first dorsal fin of males.
Colour in alcohol. Specimens pale with no
conspicuous markings except a cluster of few
melanophores on lower pectoral-fin base, a small black
spot on upper caudal peduncle and a vertical line of a
few black-edged scales on the body below the end of the
second dorsal fin. Few melanophores on head, just above
anterior part of pterotic sensory canal. Second and third
dorsal fins with irregular bands of melanophores; anal
fin with melanophores evenly scattered on membranes
in males, banded in females. Dorsal and anal fins of
females with weak oblique, dusky bands. Pectoral fins
with scattered melanophores on mid- and upper-base
and on membranes of proximal two-thirds of fin.
COMPARISONS. Enneapterygius melanospilos shares the
distinctive banded pattern on body with E. clarkae and
E. gruschkai and is similar in colour to E. gruschkai, which
also has an orange and brown colour pattern. The spot
on the second dorsal fin distinguishes E. melanospilos
from the other two species. Randall also notes the long
bi- or trilobed orbital cirrus, the spinules on the upper
posterior rim of the eye and along the posterior margin
of the supratemporal sensory canal as distinctive. The
long orbital cirrus is distinctive, but the serrate margin
of the median extrascapulars is also found in E. elegans
and the spinules on the rim of the eye in E. clarkae.
Enneapterygius clarkae has fewer pored lateral-line scales
than E. melanospilos (12 vs. 14-15) and E. gruschkai
generally has 13 spines in the second dorsal fin (vs. 12
for E. melanospilos) and the distinctive window in the bar
across the caudal peduncle.
Enneapterygius obscurus Clark
(Fig. 13, Pl. 2)
Enneapterygius obscurus Clark, 1980: 105, figs. 4d & 15
(Aqaba & Eritrea, Red Sea); Fricke & Randall, 1992:
6, fig. 3 (Maldives).
DESCRIPTION (partly from Clark 1980). Dorsal fins
III+XII-XIV+9-10, usually III+XIII+9-10; anal fin I,16-18
(usually 17-18 rays); pectoral fins 14-15: 3-4+5+6; lateral
line with 10-12 pored and 21-22 notched scales, pored
segment starting 2 scale rows below end of pored
segment (Fig. 13b). (In some of the Red Sea types
examined there were only 7-8 pored scales, followed by
2 notched scales - Fig. 13c.) Transverse scales 1/4.
Mandibular pores 2+2+2 (Fig. 13d). Head 3.2-4.0, body
depth 6.5 in SL; eye 2.4-2.6 in head length.
Body moderately slender. Nape scaled, belly naked.
First dorsal fin equal to or a little lower than second,
second subequal to body depth. Pelvic-fin rays united
by membrane for half length of shorter ray. Profile
rounded; mouth low on head, cleft nearly horizontal,
reaching vertical through anterior margin of pupil.
Prominent labial folds extend from lower lip, under the
jaw, in a crescent to the mandibular sensory canal (Fig.
13d). Supratemporal sensory canal crescent-shaped.
Orbital cirrus bilobed and moderately long, about half
pupil diameter.
Live colour. There are three transparencies, one taken
by Randall in the southern Red Sea and two in Debelius’
Red Sea Reef Guide (1998: 174). The Randall specimen
was captured and identified as E. obscurus, while the
specimens illustrated in Debelius were not collected or
identified. The fin element counts that can be made from
the Debelius illustrations agree with those of E. obscurus
and not with those of E. pallidus, another Red Sea species
for which the live colour is unknown. The Debelius
illustrations also show the irregular bands of
melanophores on the second dorsal fin, the black-edged
scales above the lateral line, between the second and
third dorsal fins and a small, black spot at the upper
DISTRIBUTION (Figs. 9, 17). Enneapterygius obscurus
was originally described from the Red Sea (Aqaba and
Eritrea) and was presumed endemic to the area. Randall
(1995) does not record the species from Oman, where the
species may reasonably be expected to occur if it does
occur outside the Red Sea. Fricke & Randall (1992) do,
however, record a single specimen from the Maldives
which agrees morphometrically with Clark’s (1980)
description, but differs considerably in pigmentation,
having 6 narrow bars across the body. Randall collected
a specimen in the Seychelles that has similar
pigmentation. I have seen a single specimen from
Malindi, Kenya that can only be identified as E. obscurus,
except that it has a single symphyseal pore. It is then
possible that the species is more widespread in the
western Indian Ocean.
COMPARISONS. There seems little to distinguish
preserved material of E. obscurus from that of several other
species, other that the lack of any obvious pigmentation
except the cluster of melanophores at the pectoral-fin
base and the spot on the upper caudal peduncle, the
15
A
B
Figure 13.
C
D
Enneapterygius obscurus, A: male, 18.9mm SL, Malindi, Kenya; composite, based on Clark, 1980, figure 15
and SAIAB 30422. B: junction between pored and notched segments of lateral line - normal condition; C:
exceptional condition (HUJ 5486). D: diagram of mandibular pores.
Head 3.4-4.7, body depth 6.9 in SL.
Body very slender. Nape and belly naked. First dorsal
fin equal to or lower than second, second about equal to
body depth. Pelvic-fin rays united by membrane for about
half of length of shorter ray. Supratemporal sensory canal
deeply U-shaped. Orbital cirrus small and lobate.
Live colour. Recent underwater photographs taken by
Richard Field at Farasan Island in the southern Red Sea
of an unidentified tripterygiid may possibly be of E.
pallidus. With the identification of Debelius’ photographs
as E. obscurus (see above) and Randall’s photographs, E.
pallidus remains the only Red Sea species for which the
live colour is unknown. Thus, unless Field’s photograph
represents an undescribed species, by a process of
elimination, it is of E. pallidus.
Body translucent with faint red bars, the most
prominent across the caudal peduncle. Top of head
opercle and anterior of body bright green; iris green and
brown; fins translucent with a few dark spots on
membrane between first two dorsal-fin spines. The
apparent absence of melanophores suggests that this is
E. pallidus, but will have to be confirmed.
Colour in alcohol. No sexual dichromatism evident.
There are no distinguishing marks. Few melanophores
over the brain area, a few on mid-opercle and a few on
pectoral-fin base.
height of the first dorsal fin and shape of the
supratemporal sensory canal. Morphometrically E.
obscurus is similar to E. abeli, E. clarkae, E. destai and E.
ventermaculus, except that E. obscurus is the only species
in this group with a 2+2+2 mandibular pore pattern.
The other three species usually have distinguishing
pigmentation, but female E. abeli specimens do not. The
height of the first dorsal fin of E. abeli is only about half
body depth compared to being subequal to body depth
in E. obscurus. The scarlet live colour of the species is,
however, very distinctive.
MATERIAL EXAMINED. Red Sea: BPBM 35721 (17.8
mm), southern Red Sea; HUJ 5486 (4: 14.5-16.2 mm),
paratypes, El Himera; HUJ 5483 (17.5 & 22.0 mm),
paratypes, El Himera. East Africa: SAIAB 30422 (24.3 mm),
Malindi, Kenya.
Enneapterygius pallidus Clark
(Fig. 14; Pl. 2)
Enneapterygius pallidus Clark, 1980: 107, Figures 4e & 16
(Aqaba & Eritrea, Red Sea).
DESCRIPTION (from Clark 1980). Dorsal fin III+XIIIXIV+10-11; anal fin I, 21-22; pectoral fins 13-14: 3+56+5. Lateral line with 10-11 pored scales and 28(?)
notched scales, notched segment starting second scale
row below end of pored segment. Mandibular pores
2+2+2; lower jaw with two large lip flaps (Figure 14b).
DISTRIBUTION (Fig. 15). The species was described
from four specimens from the Red Sea (Aqaba and
Eritrea).
16
A
Figure 14.
B
Enneapterygius pallidus, A: male, 20.8mm SL; Eritrea, Red Sea (from Clark, 1980). B: diagram of
mandibular pores.
Live colour. Fricke (1994: 245) quotes a description by
J. B. Hutchins of the colour of specimens after two weeks
in formalin: “Upper sides green-yellow, lower sides redyellow. All scales on sides profusely spotted in black,
also red spots on dorsal surface. Lower half of head with
larger black spots extending onto pectoral [fin base].
Breast and throat pale with black spots. Eye red. Caudal,
dorsal and anal fins spotted with black.”
Jordan and Seale’s (1906) description of E. tusitalae is
in reasonable agreement with Hutchins, but note that E.
pardochir (synonymised with E. minutus by Fricke, 1994)
varies from, “cherry red with darker bars, most distinct
and paired alongside, iris scarlet, fins all reddish, barred
with brown...” to “...bright green with bronze dots, sides
with large white spots, two white dots at base of caudal,
pectoral banded white with reddish brown...”
Underwater photographs of a male and a female in
Masuda & Kobayashi (1994: 310) show the following:
male with body, head and fins liberally covered with
densely packed melanophores with some white flecks
on body, first and second dorsal fin and head. Females
with less black, showing clearer banding on body and
fins. Photographs of a male and female by Winterbottom
of fresh specimens from Lizard Island, Australia, agree
with Hutchins description and a photograph by
Heemstra of a single male from Rodrigues (Plate 2) shows
the top of the head and anterior dorsum of the body as
dark yellowish-green.
Colour in alcohol. Body of mature males with
melanophores evenly and densely spread over body and
fins, the larger ones on cheeks, thorax and along the
dorsum, black in the centre surrounded by a brown halo.
Lower half of head darker and in large males becomes
uniformly brown-black. All fins densely covered with
melanophores; dorsal fins darker distally; anal, caudal
and pectoral fins very dark, particularly lower half of
caudal and pectorals, the latter with melanophores on
inner surface as well.
Immature males and females paler, with 4 oblique
black bars across the body, with white blotches between,
the bars dividing ventrally to continue as narrower bars
across the anal fin. First and second dorsal fins evenly
dark, pigment on spines; third dorsal without colour;
COMPARISONS. Fricke (1997) considers the species
synonymous with E. pusillus. It can however be
distinguished from all WIO tripterygiids by its high analfin ray count (21-22 rays), and that only the lower 5
pectoral rays are simple (for E. destai as well), compared
to 6-8 for other species. Clark (1980) also records that E.
pallidus has 11+26-28 vertebrae compared to 11-12+2225 for E. pusillus. Enneapterygius pusillus is further
distinguished from E. pallidus by the former’s very high
first dorsal fin.
Enneapterygius philippinus (Peters)
(Fig. 15; Pl. 2)
Tripterygium philippinum Peters, 1868: 269 (Luzon Island,
Philippines).
Tripterygium minutum: Günther 1877: 211(Samoa).
Enneapterygius tusitalae: Jordan & Seale 1906: 416, fig. 97
(Samoa).
Enneapterygius pardochir: Jordan & Seale 1906: 417, fig.
98 (Samoa).
Enneapterygius punctulatus: Herre 1935: 432 (New
Hebrides). (See Fricke 1997: 274 & 276 for a detailed
synonymy).
DESCRIPTION (based partially on Fricke, 1994 & 1997;
note that western Pacific Ocean specimens exhibit a
wider range of counts than specimens from the western
Indian Ocean - see Fricke 1994: 249; 1997: 287 & 288):
Dorsal fins III+X-XIII+8-9 (usually III+XI+9); anal fin I,1517, (usually 16 rays); pectoral fins 15: 3-4+4-5+7; lateral
line with 10-15 (usually 12-13) pored scales and 20-22
notched scales, notched segment starting second scale
row below end of pored segment (Figure 15c). Total lateral
scales 29-31; transverse scales 2/5. Mandibular pores 23+1+2-3 (Fig. 15d). Head 3.6-4.2, body depth 5.0-5.6 in
SL; eye 2.8-3.2 in head length.
Small, slender fishes, less than 22 mm SL. Nape, belly
and pectoral-fin bases naked. Pelvic-fin rays slender and
united by membrane for two-thirds of length of shorter
ray. First dorsal fin lower than second. Mouth slightly
oblique, reaching vertical through anterior margin of
orbit. Orbital cirrus small and lobate. Supratemporal
sensory canal crescent-shaped.
17
C
A
D
B
Figure 15.
Enneapterygius philippinus, A: male, 19.0mm SL; Mauritius; composite SAIAB 60676 and photograph by
R. Winterbottom. B: female, 21.0mm SL; Ibo Island, Mozambique (composite SAIAB 30157 and photograph by
RW.) C: junction between pored and notched segments of lateral line. D: mandibular pores.
the female of E. tusitalae (1912:518) but did not formally
synonymise the two species. Fowler (1958) synonymised
E. tusitalae and E. pardochir with E. philippinus, but did
not state whether he recognised Jordan & Seale’s species
as male and female of the same species. Schultz (1960)
did, but neither of these authors state whether they
examined Günther’s types. Günther’s description of E.
minutus is brief in the extreme and merely notes that the
two specimens on which the description is based were
reddish-white with indistinct brown transverse bands
on caudal and pectoral fins. His illustration of the species
does show that the first dorsal fin is lower than the second
(ibid: 213d). Fricke (1994) did examine Günther’s types
and found that they agreed with specimens of E.
pardochir, E. tusitalae, Tripterygion callionymi (Weber,
1909) and E. punctulatus (Herre, 1935) and synonymised
the four species with E. minutus. Fricke (1997: 279-281)
has a detailed analysis of the nomenclatural
complications of the species and synonymised E. minutus
with E. philippinus.
Schultz’s inclusion of E. tutuilae Jordan & Seale as
the same species as E. minutus is incorrect. The
pigmentation of E. tutuilae is quite different: the species
has a clearly banded anal fin, black mark in the margin
of the second dorsal fin, and first dorsal fin distinctly
higher that the second.
caudal and pectoral fins irregularly barred. Females have
no pigment on throat, belly or along the base of the anal
fin.
DISTRIBUTION (Fig. 9). The species is widely
distributed throughout the Indo-West Pacific Ocean. It
was originally described from the Philippines (Peters,
1868), Günther (1877) described the species from Samoa;
Herre (1935) from the New Hebrides; Schultz (1960) from
the Marshall Islands; Fowler (1958) from the Ryu Kyu
Islands, and Winterbottom collected material at Lizard
Island (Australia) in 1981. In the Indian Ocean the species
has been recorded from St Brandon Shoals, the Comoro
Islands, and several island localities along the East
African coast. There is a single collection of 16 specimens
from the Great Fish River Point in the Eastern Cape, South
Africa, made by J. L. B. and M. M. Smith in 1954, but the
locality is highly doubtful.
COMPARISONS. Male Enneapterygius philippinus, with
their distinctive dense covering of melanophores, are
easily distinguished from other species of the genus.
Females, sometimes with 5-6 bars across the anal fin,
may be confused with E. pusillus, E. tutuilae or E.
ventermaculus. Enneapterygius pusillus and E.
ventermaculus have distinctly U-shaped supratemporal
sensory canals compared with a crescent in E.
philippinus, and E. tutuilae has a high first dorsal fin,
whereas in E. philippinus it is lower than the second.
Enneapterygius philippinus also has 15 pectoral-fin rays
vs. 14 for E. tutuilae.
MATERIAL EXAMINED.
Pacific Ocean: New Hebrides: AMS I6317 (3); AMS I6418
(3); AMS I6451 (4); AMS I6582-3 (3). Lizard Island, Great
Barrier Reef: ROM 72516 (21.7 & 21.9 mm). Abaiang Atoll,
Gilbert & Ellis Islands: AMS I18049-011 (5). Indian Ocean:
Comoro Ids: CAS 33458 (6: 17.4-21.5 mm). Mauritius:
REMARKS. Snyder correctly recognised E. pardochir as
18
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. Snout
and lips pale yellow with orange spots; cheeks with
yellow, red and black spots. Pectoral 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. First dorsal fin with light red
spots and some melanophores; second dorsal fin heavily
marked with dark brown to black melanophores
interspersed with red spots and with 3 silverytransparent vertical bands; third dorsal fin pale orange
to transparent. Anal fin with red and black spots, the
melanophores tending to form 6 or more indistinct bars.
Upper half of caudal fin pale orange, lower half pale
yellowish-green. Females less intensely pigmented and
lack the melanin of large males. (Note: the colour
photograph in Holleman, 1986 (plate 116) is not of E.
pusillus but of E. tutuilae.)
Colour in alcohol. All colour except for the melanin
disappears, leaving bodies a pale straw colour, males
with irregular bars across body, black on lower portion
of head, thorax and extending to area around base of
pelvic fins and proximal third to half of fins; pre-anal
black spot present. Second and third dorsal and pectoral
fins with irregular dusky bands; anal fin with basal spots
and irregular black bars across fin. Females have the
basal spots and bars on the anal fin, but lack black on
the head, thorax and pelvics and dark body bars, having
more irregular markings.
SAIAB 30163 (18.0 mm). Mahé, Seychelles: SAIAB 30155
(19.0 & 19.4 mm). Nosi Bé, Madagascar: AMS I28108 (4).
Shimoni, Kenya: SAIAB 30158 (3: 15.2-17.8 mm). Ibo Island,
Moçambique: SAIAB 30162 (6: 14.0-20.9 mm) & SAIAB
30157 (6: 16.8-21.3 mm). Pinda, Mocambique: SAIAB 30161
(17.7 & 20.0 mm). Cabo Delgado, Moçambique: SAIAB 30156
(12: 16.5-20.2 mm) & SAIAB 30159 (19.2 & 19.8 mm).
Tekomasi Island, Moçambique: SAIAB 30154 (18.1 & 20.0
mm) & SAIAB 30160 (3: 18.0-20.9 mm).
Enneapterygius pusillus Rüppell
(Fig. 16; Pl. 2)
Enneapterygius pusillus Rüppell, 1835: 2, pl. 1, fig. 2
(Massawa, Red Sea); Clark, 1980: 108, Figs. 3a & 17
(Red Sea); Holleman, 1986: 757, Figure 236.5, non Pl.
116 (KwaZulu-Natal); Randall, 1995: 31 (Gulf of
Oman).
DESCRIPTION. Dorsal fins III+XII-XIV+10-11 (usually
III+XIII+10-11); anal fins I,20-21 (see Table 1 for counts
of Red Sea and Oman specimens); pectoral fins usually
13-14: 3-4+3-4+6-7 (usually 4+3+6). Lateral line with 912 pored scales and 25-28 notched scales, notched
segment starting second scale row below end of pored
segment (Fig. 16b). Total lateral scales 29-30; transverse
scales 2/4; mandibular pores 2+2+2 (Fig. 16c). Head
3.6-3.9, body depth 5.3-6.0 in SL; eye 3.0-3.4 in head
length.
Small (less than 25 mm SL), slender fishes. Nape,
belly and bases of dorsal and anal fins naked. First dorsal
fin distinctly higher than second in males and about
equal in height in females; second dorsal fin 70% of body
depth. Pelvic-fin rays united by membrane for about half
length of shorter ray. Maxilla reaching vertical through
anterior of eye. Orbital cirrus small and lobate;
supratemporal sensory canal deeply U-shaped.
Live colour. Body colour of males pale cream with 4
irregular dark bars; scales on upper half with scattered
orange and yellow-green spots and edged with a thin,
deep orange line. Lower half of body with series of up to
10 orange blotches interspersed with irregular, dark,
silver-grey spots. Abdomen covered anteriorly with dark
DISTRIBUTION (Fig. 17). Known from inshore
continental waters from the Red Sea south to KwaZuluNatal, South Africa, from the Persian Gulf and coast of
Oman (Randall, 1995), to the southeastern coast of India
(Gulf of Mannar - Lal Mohan, 1971).
COMPARISONS. Male E .pusillus are distinctive with
their very high first dorsal fin and black pelvic-fin base
B
A
C
Figure 16.
Enneapterygius pusillus, A: male, 21.7mm SL; KwaZulu-Natal, South Africa, SAIAB 6522. B: junction between
pored and notched segments of lateral line. C: mandibular pores.
19
transverse scales 2/4. Mandibular pores 2+1/2+2 (Fig.
18c). Head 3.3-3.6, body depth 4.4-5.3 in SL; eye 2.6-3.2
in head length.
Body moderately elongate. Nape scaled; belly naked.
First dorsal fin higher than second in males and about
the same height in females, second dorsal fin about 70%
of body depth. Pelvic-fin rays united by membrane for
about half length of shorter ray; longer ray reaches to
vent. Longest pectoral-fin ray reaches end of second
dorsal fin. Supratemporal sensory canal crescentshaped. Maxilla reaches vertical through anterior
margin of pupil. Orbital cirrus moderate and lobate.
Live colour. There are two colour forms, an apparently
more common green form and a brown form recorded
from Phuket, Thailand and Mauritius. Green form (from
transparencies made by R. Winterbottom in the Comoro
Islands, Fiji and on Escape Reef, Australia, and by G.
Allen [in: Randall et al., 1997] ) as follows: males with
bright green bodies with white blotches along mid-side,
a white saddle below rear half of second dorsal fin and
a white bar across body at end of third dorsal fin; analfin base with 6-8 black spots; black preanal spot present;
basal anal-fin spots and pre-anal mark absent in most St
Brandon Shoals specimens. Scale-pocket margins on
upper half of body lined with red dots. Snout, lower half
of head, chest and pectoral-fin bases with many red spots;
top of head green. First dorsal fin whitish-green; second
with many red spots anteriorly, with black spots on the
membranes between 4th and 9th spines and white band
posteriorly, the white continuing onto the body; third
dorsal fin transparent with some red basally and a white
band on the margin posteriorly. Caudal-fin rays lined
with red, membranes transparent; anal fin with
alternating blackish red and white bars; pelvic fins
spotted with red; pectoral fin rays lined in red but with 3
white bands, membranes transparent.
The brown form with brown and orange spots on
body and head and with narrow dark bar across caudal
peduncle; first dorsal fin orange and cream; second as
for the green form, but with orange and brown replacing
the red. The white bands of the green form are creamcoloured in the brown form; lower half of pectoral fins
red. Females of both forms less brightly coloured than
males.
Colour in alcohol. Males and females straw-coloured
with 6-8 black bars across anal fin, dark basal spots on
anal fin, prominent black pre-anal mark, and black blotch
on margin of the middle of second dorsal fin. Mature
males sometimes with stippling along inner pelvic fin
ray, on head, and show indistinct, irregular, dusky bars
across body. Immature males and females often show
little pigment except the basal spots on anal fin and some
black on second dorsal fin.
Figure 17.Distribution of original specimens of E. obscurus
(closed triangles), E. pallidus (open star), E.
pusillus (closedstar), E. tutuilae (closed roundel)
and E. ventermaculus (open star).
and proximal portion of the fins. Females from the Red
Sea may be confused with the sympatric E. pallidus, but
E. pusillus usually has fewer anal-fin rays than the latter
species: 17-21 (usually 19), vs. 21-22 (usually 22) for E.
pallidus. Another sympatric species which shares the
high first dorsal fin of male E. pusillus is E. tutuilae (= E.
altipinnis - see below), but that species lacks a U-shaped
supratemporal sensory canal and has fewer notched
scales in the lateral line: 20-23 vs. 25-28 for E. pusillus.
MATERIAL EXAMINED. SAIAB 7937 (21.7 mm);
SAIAB 7941 (21.6 & 24.5 mm); SAIAB 7942 (6: 19.5-22.5
mm); SAIAB 7940 (5: 17.3-22.9 mm), all from Sodwana
Bay, South Africa.
Enneapterygius tutuilae Jordan & Seale
(Fig. 18, Pl. 2)
Enneapterygius tutuilae Jordan & Seale, 1906: 418-419,
figure 99 (Pago Pago, Samoa); Fricke 1994: 285ff,
figures. 58 & 59; Randall, Allen & Steene, 1997: 364.
Enneapterygius altipinnis: Clark, 1980: 99-101, figs. 3b,
6c, 11 (Aqaba & Eritrea, Red Sea).
Enneapterygius pusillus: (non-Rüppell, 1835): Holleman,
1986: 757, pl. 116, fig. 236.5 (Australia).
DESCRIPTION. Dorsal fins III+XI-XII+9-10 (usually
III+XII+10); anal fin I,16-18 (usually 17 rays); pectoralfin rays 14, all simple. Lateral line with 11-12 (usually
12) pored scales and 20-23 (usually 22) notched scales,
notched segment starting second scale row below end of
the pored segment (Fig. 18b). Total lateral scales 29;
DISTRIBUTION (Fig. 17). Widely distributed
throughout the Indo-West Pacific Ocean, from East Africa
to Japan and Samoa.
COMPARISONS. Enneapterygius tutuilae, though similar
to E. pusillus with its high first dorsal fin, has more
20
prominent anal-fin bars, particularly the basal spots,
which are absent in E. pusillus (see also remark above
about St Brandon Shoals specimens), and lacks the deep
U-shaped supratemporal sensory canal of E. pusillus.
Enneapterygius ventermaculus shares the anal fin bars and
basal spots with E. tutuilae, but also has a U-shaped
supratemporal sensory canal and 21-25 notched lateral
line scales, vs. 20-23 for E. tutuilae.
Schultz (1960) considered E. tutuilae a synonym of E.
minutus (= philippinus), together with E. tusitalae and E.
pardochir. This is not so: E. tutuilae and E. philippinus are
quite distinct in colour and form of the first dorsal fin.
(See also under E. philippinus above).
Enneapterygius ventermaculus Holleman
(Fig. 19, Pl. 2)
Enneapterygius ventermaculus Holleman 1982: 123, fig. 7
(northern KwaZulu-Natal, South Africa); Randall
1995: 32, fig. 5 (Gulf of Oman).
Enneapterygius nasima: Hoda 1983: 116ff, figs. 1-3
(Karachi coast, Pakistan).
DESCRIPTION. Dorsal fin III+XI-XIII+8-10 (usually
III+XII+10); anal fin I,17-20 (usually 19 rays); pectoralfins rays 14: 1-3+4-6+6-8 (usually 2+5+7). Lateral line
with 13-16 (usually 15) pored and 21-25 (usually 23)
notched scales (see Table 1 for counts of Oman
specimens), notched series starting second scale row
below end of pored series (Fig. 19b); total lateral scales
31-34 (usually 32-33, but 35-36 for Oman specimens);
transverse scales 2/6. Mandibular pores 3+1+3 (Fig. 19c).
Head 3.4-4.0, body depth 4.8-5.2 in SL; eye 3.0-3.7 in head
length.
Body moderately robust; nape, base of first dorsal fin
and belly naked. Mouth small and nearly horizontal,
reaching vertical through anterior margin of orbit. 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; second dorsal fin about 70% of body
depth. Pelvic-fin rays united by membrane for about half
length of shorter ray, longer ray reaching as far as preanal mark.
Live colour (also from Randall 1995). Body of males
greenish, scale margins with brown stipples and orange
edge, except ventrally; black pre-anal mark present,
ranging from small spot to boomerang-shape half around
vent; three white blotches along midside, first under end
of pectoral fin, second below junction between second
and third dorsal fins, third below end of third dorsal fin;
caudal fin base with triangular dark area, apex anterior,
base darkest, followed by narrow white band at base of
caudal fin; head bottle-green above with orange-brown
markings, darkest on sides below eyes, on thorax and to
MATERIAL EXAMINED. Kenya: SAIAB 31521 (4:19.821.2 mm) Malindi. Mozambique: SAIAB 31522 (16.5 mm)
Cabo Delagado; SAIAB 31515; (3: 15.0-18.6 mm) & SAIAB
15024 (17.4 mm), Baixo Pinda;. Indian Ocean: Comoro Ids:
ROM 67490 (12.8 & 15.1 mm); ROM 67491 (13.6 & 15.1
mm); ROM 67493 (13.8 mm); ROM 67495 (6: 13.8-14.4
mm); ROM 67492 (13: 7.6-13.9 mm); ROM 67485 (14.6 15.2 mm); ROM 67486 (15.4 mm); ROM 67487 (11.6 &
16.0 mm); ROM 67497 (8: 9.4-15.7 mm); ROM 67498 (15.6
mm); ROM 67494 (3: 11.8-14.1 mm); ROM 67499 (12.8 &
16.4 mm); ROM 67489 (12.9 & 14.9 mm); ROM 67500 (3:
14.8-15.0 mm); ROM 67496 (3: 10.1-15.0 mm); ROM 67484
(8: 14.3-15.8 mm); ROM 67488 (14.4 & 15.7 mm).
Mauritius: SAIAB 31518 (14.7 mm); SAIAB 31519 (15.5
mm); SAIAB 31520 (14.8 & 15.2 mm). Chagos Archipelago:
ROM 67467 (21.6 mm). St Brandon Shoals: SAIAB 31516
(16.0 mm); SAIAB 31517 (17.1 mm); USNM 357610 (18.5
mm); USNM 360012 (5: 17.5-22.1 mm); USNM 359861
(24: 14.6-19.4 mm); 359859 (16: 17.2-18.7 mm); USNM
359856 (4: 14.1-19.5 mm); USNM 359857 (16.9 & 18.5
mm).Pacific Ocean: Thailand: CAS 48185 (24: 11.1-17.4
mm); PMBC 17468 (3); PMBC17473 (5); PMBC 17475 (2).
Japan: USNM 770713 (7: 9.3-20.4 mm).
B
A
Figure 18.
C
Enneapterygius tutuilae, A: male, 14.0mm SL; composite, based on SAIAB 31520, Mauritius, SAIAB 31515,
Baixo Pinda, Mozambique, and Holleman (1986), Plate 116, Fig. 236.5 B: junction between pored and
notched segments of lateral line. C: mandibular pores.
21
with triangular dark area, with densest stipples in band
at base of caudal fin; dorsal fins with scattered
melanophores, darkest between first two spines of first
dorsal fin; anal fin with 5-7 irregular black bars, the
colour on the rays only; pectoral fins irregularly marked
with melanophores.
Females irregularly pigmented with black and brown,
tending to form 4-5 irregular H- or inverse Y-shaped bars
along the sides. Prominent black preanal mark present,
frequently in shape of triangle or boomerang-shape, apex
anterior. Anal fin with 5-7 black basal spots which
continue as irregular bars across the fin. Caudal and
pectoral fins irregularly barred.
pelvic-fin bases and on pectoral-fin bases. First dorsal
fin finely speckled with brown and orange, but without
pigment on membrane between third spine and dorsum;
second dorsal spines orange with alternating areas of
orange and dense, dark brown stippling along fin margin
and band of stipples along base; third dorsal fin banded
with dark brown on the rays and orange stipples on
membranes anteriorly; anal fin with 5-7 black broken
bars with white and transparent areas between, and 57 black basal spots which continue subcutaneously;
caudal fin rays greenish, upper with some brownish
semi-bands, lowermost with black marks; pelvic fins
yellow; pectoral fins with white and dark brown, lower
rays with yellow, and two white spots between base of
pectoral fin and base of pelvic fin, which may be largely
obscured in darkly marked specimens.
Females dorsally translucent pale green with orange
margins to scales; no black or brown on ventral part of
head and thorax, but small clusters of melanophores
below midline more prominent than in males; a dark
streak in front of each pelvic fin, another on side of thorax
and a third from lower pectoral-fin base, latter two
continuing a little below branchiostegal membrane.
Colour in alcohol. The original description (Holleman
1982) noted no sexual dichromatism. However, Randall’s
(1995) description and material available from Sur,
Oman, shows males with densely clustered large
melanophores in interorbital area, on head below eyes,
opercles and lower portion of pectoral fin-bases, entire
thorax, base of pelvic fins and anterior of belly, including
proximal portion of fins themselves, the darkest a black
band extending from lower pectoral fin bases across chest
and pelvic fin bases. Body with irregular dusky markings
that may form indistinct bars ventrally; caudal peduncle
DISTRIBUTION (Fig. 17). Aden, Oman, Pakistan and
east coast of South Africa from northern KwaZulu-Natal
to Tshani, Eastern Cape Province, an apparently antitropical distribution.
COMPARISONS. Hoda (1983) differentiated
Enneapterygius nasimae from E. ventermaculus on the basis
of colour pattern; there is little to distinguish the two
species morphometrically, the counts of E. nasimae falling
within the range of those for E. ventermaculus. At the time
of description the colour of E. ventermaculus was not
known, but specimens from Pakistan were included in
the type series (Holleman, 1982: 125). Fresh material
subsequently available to me and Randall’s description
and photograph of E. ventermaculus (1995: 32, fig. 5) from
the coast of Oman agree with Hoda’s description of the
colour of E. nasimae. Hoda’s only distinction is that E.
nasimae has more regular blotches along the side than E.
ventermaculus, particularly that on the caudal peduncle.
His illustration (figs. 3a & b) suggests the triangular
B
A
D
C
Figure 19.
Enneapterygius ventermaculus: A: male, 26.6mm SL; Aliwal Shoal, South Africa, B: junction between pored
and notched segments of lateral line. C: mandibular pores. D: holotype, female, 24.8mm SL; Sodwana Bay,
South Africa.
22
caudal mark as described by Randall. The species is
distinctly not the same as E. hollemani (the differences
are dealt with in detail under that species) and the E.
nasimae is thus considered a junior synonym of E.
ventermaculus. (See also under E. hollemani and E. tutuilae.)
tripterygiids) males are more brightly and/or darkly
coloured than females, and a black or partially black
head, thorax, abdomen and prepectoral area is not
uncommon in males (E. abeli, E. elegans, E. minutus, E.
pusillus, E. tutuilae and E. ventermaculus), it is not known
whether the black coloration is sexual dichromatism or
whether it only represents nuptual/territorial coloration,
as it does in Tripterygion species.
Randall (1993) suggests that the change in colour of
large males of Helcogramma vulcana (and other species of
tripterygiids) to entirely black does not seem to be
associated with the attainment of sexual maturity,
because pale individuals with dark bars may have large
testes. One would not expect a fish to produce such an
extreme amount of melanin for a brief spawning period,
only to have it disappear after spawning. He records a
more likely Jeffrey Williams’ hypothesis that melanistic
males may be dominant in the local population (Randall
1993:31). This would seem to be in agreement with
Wirtz’s observations (1978:143) Furthermore, in
collections of Helcogramma obtusirostre from tide pools in
KwaZulu-Natal, South Africa, where the species is
abundant, few males are almost entirely black, while
most are darker versions of females, an observation that
may also corroborate Williams’ hypothesis. This degree
of melanism has only been observed in males of
Enneapterygius etheostomus (Masuda & Kobayashi
1994:310), E. kermadecensis, E. minutus and E. niger in
which the males are very extensively covered with
melanophores.
Melanophore patterns may be of further value, as
indicators of relationship. Fricke (1994:174)
distinguishes six species groups for Australian and
southwestern Pacific species of Enneapterygius, based on
the length of the pored (anterior) portion of the lateral
line, the amount of melanin in males and the height of
the first dorsal fin, in that order, arbitrarily considering
12 or fewer pored scales as a short anterior lateral line
and 13 or more as a long anterior lateral line. While this
may be useful in a key, to use the character to define
species groups implies phylogenetic relationship, for
which Fricke offers no other substantiation.
In Helcogramma there are examples of colour pattern
indicating relationship as, for example, the H. fuscopinna
species group as defined by Williams & McCormick
(1990). The distinctive caudal peduncular bars of certain
Enneapterygius species (E. clarkae, E. destai, E. elegans and
E. gruschkai) may be indicative of relationship, as may
be the body bars of E. clarkae, E. gruschkai and E.
melanospilos, or the internally extending basal anal fin
spots of E. hollemani, E. kosiensis, E. tutuilae and E.
ventermaculus.
Certainly the form of the supratemporal sensory canal
with its associated osteological features, which may have
phylogenetic implications, must be taken into
consideration (Holleman, 1982). However, until study
of the entire genus is undertaken these issues must
remain speculative.
MATERIAL EXAMINED additional to that listed in
Holleman 1982. Oman - ROM 57548 (22.0 mm); ROM
57549 (11.2 & 20.0 mm); ROM 57537 (3: 18.5-19.2 mm);
ROM 57545 (4: 16.6-20.0 mm); ROM 57546 (3: 12.0-19.9
mm), all from Sur. South Africa - SAIAB 40336 (3: 21.027.6 mm) & SAIAB 40427 (25.8 & 26.6 mm), Park Rynie;
SAIAB 40506 (26.7 mm), SAIAB 60242 (28.7 mm), SAIAB
60303 (6: 17.3-29.0 mm), Aliwal Shoal; SAIAB 53511 (31.0
mm), Tshani, Transkei; SAIAB 43242 (20.0 & 21.5 mm),
Sodwana.
COMMENTS ON INDIAN
AND SRI LANKAN SPECIES
The most recent publications on tripterygiid fishes from
the coasts of India and Sri Lanka are those of Lal Mohan
(1968, 1971) and Hoda (1983). From those we know that
at least two species of Enneapterygius are found: E.
fasciatus and E. ventermaculus (E. nasimae). Fricke &
Randall (1992) record E elegans and E. obscurus from the
Maldives. It stands to reason that E. abeli, E. clarkae, E.
gruschkai, E. minutus and, perhaps, E. tutuilae also occur
in the Laccadive Islands and the islands around Sri
Lanka.
DISCUSSION ON COLOUR PATTERNS
With the exception of the New Zealand and southern
Australian species, there is still little record of colour
patterns of tripterygiids or information about associated
behaviour patterns. While the number of colour
photographs of species is increasing, including good
underwater photographs by J. Randall, G. Allen, H.
Debelius, R. Kuiter, H. Masuda, S. Dewa and others, few
of the probably more than 150 species have been
photographed or illustrated, and of only a handful of
species are there photographs that show changes in
colour patterns with growth, sexual dichromatism and
the nuptial/spawning colours of males.
While most species of Enneapterygius show sexual
dichromatism, it is not certain that all do. This
assumption was made in describing Enneapterygius
ventermaculus (Holleman, 1982) and was found to be
incorrect when specimens from Oman were examined.
Thresher (1984) records that during spawning male
tripterygiids generally develop some combination of
black and/or red on the head, body and caudal fin.
Zander & Heymer (1976) and Wirtz (1978:143) observed
a black head in territorial males of all three
Mediterranean species of Tripterygion, and both Shiogake
& Dotsu (1973: 36) and Masuda & Kobayashi (1994: 310)
figure a male Enneapterygius etheostomus in nuptial
colours where nearly the entire body is black.
While in all Enneapterygius spp. (probably in all
23
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Records of the Indian Museum 41(4):327-372.
HODA, S. & M. Shamsul. 1983. Enneapterygius nasimae, a
new species of tripterygiid fish from the Karachi
coast, northern Arabian Sea. Indian Journal of
Fisheries 30(1):116-123.
HOLLEMAN, W. 1982. Three new species and new
genus of tripterygiid fishes (Blennioidei) from the
Indo-West Pacific Ocean. Annals of the Cape
Provincial Museums (Natural History) 14(4):109-137.
HOLLEMAN, W. 1986. Tripterygiidae. In: Smith, M M &
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HUBBS, C. L. & K. F. Lagler. 1958. Fishes of the Great
Lakes Region. Bulletin of the Cranbrook Institute of
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Descriptions of the species found in the
archipelago with a provisional checklist of the
fishes of Oceania. Bulletin of the Bureau of Fisheries
25 (for 1905):175-455.
KLAUSEWITZ, W. 1960. Fische aus dem Roten Meeres.
III. Tripterygion abeli n.sp. (Pisces, Blennioidea,
Clinidae). Senckenbergiana Biologica 41(1-2):11-13.
LAL MOHAN, R. S. 1968. On the occurance of the
blennioid fishes Blennius semifasciatus Rüppell
(family: Blenniidae) and Tripterygion fasciatum
(Weber) (family: Clinidae) along the Indian coast.
Journal of the Marine Biological Association of India
10(1):114-117.
LAL MOHAN, R. S. 1971. Helcogramma shinglensis, a new
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with a key to the fishes of the family Tripterygiidae
of the eastern and central Indian Ocean.
Senckenbergiana Biologica 53(3/5):219-223.
MASUDA, H. & Y. Kobayashi. 1994. Grand Atlas of Fish
Life Modes. Tokai University Press, Tokyo, 465pp.
MUKERJI, D. D. 1935. Notes on some rare and interesting
fishes from the Andaman Islands, with
descriptions of two new freshwater gobies. Records
of the Indian Museum 37: 259-277.
PETERS, W. C. H. 1868. Über die von Hrn. Dr F. Jagor in
dem ostindischen Archipel gesammelten und dem
Königl. Zoologischen Museum übergebenen
Fische. Monatsberichte der Königlich Preussischen
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PETERS, W. C. H. 1876. Übersicht der von Dr K Möbius
in Mauritius und bei den Seychellen gesammelten
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RANDALL, J. E. 1993. Helcogramma vulcana, a new
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RANDALL, J. E. 1995. A review of the triplefin fishes
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ACKNOWLEDGEMENTS
I am most grateful for the loan of material from the
institutions listed, and particularly to Vic Springer and
Rick Winterbottom. Jeff Williams of the Smithsonian
Institution kindly provided data on Red Sea type
specimens and Daniel Golani provided some of the type
material of E. obscurus. Jeff Williams and Kendall
Clements reviewed this paper and provided valuable
comment. I also thank the Director of the South African
Institute of Aquatic Biodiversity, Prof. Paul Skelton, for
providing laboratory space and making the facilities of
the Institute available for my use.
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24
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25
PLATE 1
A.
Enneapterygius elaine, 17mm SL male, SAIAB 65579, Rodrigues. (PC Heemstra)
B.
Enneapterygius elaine, 20mm SL female, SAIAB 65581, Rodrigues. (PC Heemstra)
C.
Enneapterygius clarkae, male, Comoro Ids. (R Winterbottom)
D.
Enneapterygius elegans, 26mm SL male (above) and 25mm SL female (below), both
La Digue, Seychelles (JE Randall)
E.
Enneapterygius gruschkai, 24mm SL male, Mauritius. (JE Randall)
F.
Enneapterygius hollemani, 30mm SL male, Ras al Madrakah, Oman. (JE Randall)
G.
Enneapterygius hollemani, female, Red Sea. (R Field)
H.
Enneapterygius melanospilos, 26mm SL male, Masirah Is., Oman. (JE Randall)
I.
Enneapterygius melanospilos, Masirah Is., Oman. (JP Hoover)
PLATE 2
A.
Enneapterygius obscurus, Sinai, Red Sea. (H Debelius)
B.
Enneapterygius pallidus, Farasan Is., Red Sea. (R Field)
C.
Enneapterygius philippinus, 23mm SL male, Rodrigues. (PC Heemstra)
D.
Enneapterygius philippinus, 22.8mm SL female, ROM 47941, Dravuni, Fiji. (R Winterbottom)
E.
Enneapterygius pusillus, ~30mm male, Oman. (JE Randall)
F.
Enneapterygius ventermaculus, 22mm SL male, Aliwal Shoal, South Africa. (PC Heemstra)
G.
Enneapterygius abeli, 22mm SL male, Aliwal Shoal, South Africa. (PC Heemstra)
H.
Enneapterygius abeli, 19.5mm SL female, SAIAB 63076, Sodwana, South Africa. (PC Heemstra)
I.
Enneapterygius tutuilae, 20mm TL, green form, Queensland, Australia. (R Winterbottom)
J.
Enneapterygius tutuilae, 20mm SL, brown form, Gulf of Aqaba, Egypt. (JE Randall)
A. Enneapterygius elaine, male
C. Enneapterygius clarkae, male
P. C. H.
B. Enneapterygius elaine, female
P. C. H.
R. W.
D. Enneapterygius elegans, male (above)
and female (below)
J. E. R.
E. Enneapterygius gruschkai, male
J. E. R.
F. Enneapterygius hollemani, male
J. E. R.
G. Enneapterygius hollemani, female
R.F.
H. Enneapterygius melanospilos, male
J. E. R.
I.
J. H.
PLATE 1
Enneapterygius melanospilos
A. Enneapterygius obscurus
H. D.
B. Enneapterygius pallidus
R. F.
C. Enneapterygius philippinus, male
P. C. H.
D. Enneapterygius philippinus, female
E. Enneapterygius pusillus, male
J. E. R.
F.
G. Enneapterygius abeli, male
P. C. H.
H. Enneapterygius abeli, female
P. C. H.
J.
J. E. R.
I.
Enneapterygius tutuilae, green form
R. W.
PLATE 2
Enneapterygius ventermaculus, male
Enneapterygius tutuilae, brown form
R. W.
P. C. H.
SMITHIANA
SERIAL PUBLICATIONS FROM
THE SOUTH AFRICAN INSTITUTE FOR AQUATIC BIODIVERSITY
The Institute publishes original research on systematics, ecology, biology and conservation of fishes. Three series are
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Smithiana, JLB Smith Bulletins & Special Publications are available online at: http://www.bioline.org.br/special_journals and on
our web page at: http://www.saiab.ru.ac.za/smithiana
INSTRUCTIONS TO AUTHORS
Manuscripts shorter than 30 pages will generally be published in the Special Publications series; longer papers will be considered
for the Bulletin or Monographs series. Please follow the format of a recent Bulletin or Special Publication. The typescript must be
double-spaced throughout with 25 mm margins all round; two hard copies must be submitted to the Editor. Each table or figure
should be on a separate page and numbered with an Arabic numeral (not in sequence with text pages). All maps, graphs, charts,
drawings and photographs should be numbered as figures. If two or more illustrations are grouped as one figure, they must be
trimmed and spaced (but not mounted) as intended for final reproduction. Each part of a composite figure must be labelled with
a capital letter; typewriter lettering is not acceptable. Illustrations larger than 21 x 30 cm should be avoided, or sent as a reduced
bromide/line shot. Legends for figures should be on a separate page. A computer diskette/ CD, with the text in RTF, MSWord
or Corel WordPerfect format, must be submitted with the hard copies. The inclusion of digital copies of all images and figures
in acceptable formats (see web page) will expedite publication of the manuscript .
See extended Instructions to Authors at http://www.saiab.ru.ac.za/pubs.htm
STYLE OF THE HOUSE
Hyphens: Certain substantive compounds are hyphenated: gill-raker, soft-ray, type-species, type-locality, type-series, typespecimen. Other words often used together are not hyphenated unless they are used in adjectival expressions before a noun: anal
fin / anal-fin rays; lateral line / lateral-line scales; gill arch / gill-arch filaments, etc.
Word usage: Although the following word pairs are often used interchangeably, we believe that consistent use of the first
word as a noun and the second as an adjective will improve the precision of our writing: mucus / mucous; maxilla / maxillary;
opercle / opercular, operculum / opercular. The operculum (= gill cover) comprises (usually) four separate bones: opercle,
subopercle, preopercle and interopercle. The words preoperculum, suboperculum and interoperculum are unnecessary substitutes
and not to be used for preopercle, subopercle and interopercle. The plural of operculum is opercula.
Decimal comma versus decimal point: Contrary to most journals published in South Africa and some European countries, we
will not use a comma in place of a decimal point. Most computers do not read a comma as a decimal point. In addition, it is
common in ichthyological papers to give sequences of measurements that include decimal numbers, with each measurement
separated by a comma. If the comma is used to separate items in a series, as well as being used to indicate a decimal number, it
will cause considerable confusion.
Fin formulae: Fin formulae will be designated as follows: D XII,10-12 indicates on continuous fin with 12 spines and 10-12 soft
(segmented) rays; DX/I,10-12 indicates a fin divided to the base in front of the last spine; and D X+I,12 indicates two separate
dorsal fins, the first with 10 spines and the second with 1 spine and 12 soft rays. If it is necessary to differentiate branched and
unbranched soft-rays, lower-case Roman numerals will be used for unbranched rays and Arabic numerals for branched rays, e.g.
D iii,S. Principal caudal-fin rays are defined as those that touch the hypural bones. The number of principal caudal rays is usually
the number of branched rays plus two. If the principal caudal rays are in two separate groups, the number of rays in the dorsal
group is given first: thus, “principal caudal rays 8+7” means that there are 15 principal caudal rays, with 8 rays in the dorsal group
and 7 in the ventral group.
Abbreviations: Abbreviations normally end with a full stop: et al., e.g., etc., n.b., (note: these commonly used abbreviations
of Latin words are not italicized). Dr (Doctor) and Mr (Mister) and compass directions (north, west, northwest, etc.) are abbreviated
using capital letters without full stops: N, W, NW. We recommend the following abbreviations for ichthyological terms: SL standard length, TL - total length, FL - fork length, GR - gill-rakers, LL - lateral line.
M. E. Anderson, Editor
L-A. Fargher, Layout and production
South African Institute for Aquatic Biodiversity (formerly J.L.B. Smith Institute of Ichthyology),
Private Bag 1015, Grahamstown 6140, South Africa.
ISSN 1684-4130
Published by the South African Institute for Aquatic Biodiversity
Private Bag 1015, Grahamstown, South Africa, 6140
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