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Redescription of Cheilinus quinquecinctus Rüppell, 1835 (Pisces: Perciformes, Labridae), a valid endemic Red Sea wrasse

  • Faculty of Marine Sciences, King Abdulaziz University, Jeddah, Saudi Arabia

Abstract and Figures

The labrid fish Cheilinus quinquecinctus Rüppell, originally described from the Red Sea, has long been regarded as a junior synonym of C. fasciatus (Bloch). Herein, both nominal species are redescribed, based on examination of the types and additional material from the Red Sea (for C. quinquecinctus) and the Indo-West Pacific (for C. fasciatus). Rüppell's description of Cheilinus quinquecinctus was originally based on three syntypes, and the most representative adult specimen is designated as the lectotype. We show that Cheilinus quinquecinctus is restricted to the Red Sea and the Gulf of Aden, and it differs from the similar C. fasciatus in having modally fewer gill rakers on the first gill arch, a total of 13–16 (mean 13.9, usually 13 or 14) (vs. 13–16, mean 14.7, usually 14 or 15), in developing a ragged posterior margin of the caudal fin with age (versus only upper and lower caudal-fin lobes developing with age), and in its color pattern. The phy-logenetic analysis of the COI barcoding region accords with the species status of C. quinquecinctus with the placement of the two sister species in two divergent and reciprocally monophyletic evolutionary lineages. A full description of C. quin-quecinctus and diagnosis of C. fasciatus is provided here for comparison. In addition, the data include a table of the results of the meristic and morphological examination of type and additional specimens of both species from throughout their distribution ranges as well as a table of gill-raker counts of all examined specimens. Underwater color photographs are provided for comparison of juveniles, females and males of both species.
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Accepted by W. Holleman: 13 Jul. 2016; published: 31 Aug. 2016
ISSN 1175-5326 (print edition)
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4158 (4): 451
Redescription of Cheilinus quinquecinctus Rüppell, 1835
(Pisces: Perciformes, Labridae), a valid endemic Red Sea wrasse
Senckenberg Research Institute and Natural History Museum Frankfurt, Senckenberganlage 25, 60325 Frankfurt a.M., Germany.
Station of Naturalists, Omsk, Russia.
Marine Biology Department, Faculty of Marine Sciences, King Abdulaziz University, Jeddah, Saudi Arabia.
Corresponding author
The labrid fish Cheilinus quinquecinctus Rüppell, originally described from the Red Sea, has long been regarded as a ju-
nior synonym of C. fasciatus (Bloch). Herein, both nominal species are redescribed, based on examination of the types
and additional material from the Red Sea (for C. quinquecinctus) and the Indo-West Pacific (for C. fasciatus). Rüppell's
description of Cheilinus quinquecinctus was originally based on three syntypes, and the most representative adult speci-
men is designated as the lectotype. We show that Cheilinus quinquecinctus is restricted to the Red Sea and the Gulf of
Aden, and it differs from the similar C. fasciatus in having modally fewer gill rakers on the first gill arch, a total of 13–16
(mean 13.9, usually 13 or 14 ) (vs. 13–16, mean 14.7, usually 14 or 15), in developing a ragged posterior margin of the
caudal fin with age (versus only upper and lower caudal-fin lobes developing with age), and in its color pattern. The phy-
logenetic analysis of the COI barcoding region accords with the species status of C. quinquecinctus with the placement of
the two sister species in two divergent and reciprocally monophyletic evolutionary lineages. A full description of C. quin-
quecinctus and diagnosis of C. fasciatus is provided here for comparison. In addition, the data include a table of the results
of the meristic and morphological examination of type and additional specimens of both species from throughout their
distribution ranges as well as a table of gill-raker counts of all examined specimens. Underwater color photographs are
provided for comparison of juveniles, females and males of both species.
Key words: Cheilinus, taxonomy, phylogeography, mitochondrial COI, evolutionary divergence, endemism, Red Sea and
Indo-West Pacific
The labrid genus Cheilinus, proposed by Lacepède (1801) for C. trilobatus, presently contains eight valid Indo-
West Pacific species: C. abudjubbe Rüppell, 1835, C. chlorourus (Bloch, 1791), C. fasciatus (Bloch, 1791), C.
lunulatus (Forsskål, 1775), C. oxycephalus Bleeker, 1853, C. quinquecinctus Rüppell, 1835, C. trilobatus
Lacepède, 1801, and C. undulatus Rüppell, 1835 (Parenti & Randall 2000; present study). The genus Oxycheilinus
Gill, 1862 was placed in synonymy with Cheilinus by Dor (1984) and Randall (1986). In a phylogenetic study
based on morphological characters of labrid fishes of the tribe Cheilinini, Westneat (1993) recognized two lineages,
a “cheiline” lineage of five genera, with all but one with species known from the Indo-Pacific, and a
“pseudocheiline” lineage. Cheilinus forms part of the “cheiline” lineage, with the genera Cheilinus, Doratonotus
Günther, 1862, Epibulus Cuvier, 1815, Oxycheilinus, and Wetmorella Fowler & Bean, 1928. According to Westneat
(1993) Cheilinus and Oxycheilinus each form monophyletic groups based on a set of morphological characters.
Cheilinus fasciatus, and the closely related species C. quinquecinctus, are accordingly retained in Cheilinus.
Bloch (1791) described two species of the genus Cheilinus as Sparus chlorourus and S. fasciatus [in
Naturgeschichte der ausländischen Fische], both species from Japan. Descriptions of both species included
illustrations in color. Rüppell (1828) reported C. fasciatus from the Red Sea (Massawa, Eritrea), but later he (Rüppell,
Zootaxa 4158 (4) © 2016 Magnolia Press
FIGURE 1. Cheilinus fasciatus. A. ZMB 8577, lectotype, 250.0 mm, Japan; B. Drawing of Sparus fasciatus from Bloch 1791;
C. Paralectotype, ZMB 2651, 147.5 mm, “Ostindien”. Photos by S.V. Bogorodsky.
Zootaxa 4158 (4) © 2016 Magnolia Press
FIGURE 2. Cheilinus quinquecinctus. A. Lectotype, SMF 2701, 210.0 mm, Jeddah, Saudi Arabia; B. Paralectotype, SMF
2732, 226.0 mm, Jeddah, Saudi Arabia. Photos by S.V. Bogorodsky.
1835) recognized C. quinquecinctus as a new species from the Red Sea, similar to C. fasciatus, with type locality
Jeddah. Some authors treated C. quinquecinctus as a valid species (e.g., Klunzinger, 1871) but it has been treated as a
junior synonym of C. fasciatus by most authors (e.g., Roux-Estève & Fourmanoir, 1955; Randall, 1983; Dor, 1984;
Parenti & Randall, 2000; Randall, 2005; Allen & Erdmann, 2012). Westneat (1993) does not mention C.
quinquecinctus in his analysis. Kuiter (2002) and Lieske & Myers (2004), in their popular books, however, have
accepted C. quinquecinctus as a valid Red Sea species with only a note on differences in the shape of caudal fin. More
recently DiBattista et al. (2015) listed C. quinquecinctus as a Red Sea endemic species, but gave no information on
the validity/synonymy of the species, or the foundation of their acceptance of C. quinquecinctus as a valid species.
The lectotype of C. fasciatus (ZMB 8577), designated by Paepke (1999), is a dry specimen missing both
caudal-fin lobes (Fig. 1A). However, Bloch’s (1791) drawing of species showed well-developed upper and lower
Zootaxa 4158 (4) © 2016 Magnolia Press
lobes (reproduced here as Fig. 1B). The paralectotype (ZMB 2651), collected from “Ostindien”, is in good
condition with intact lobes (Fig. 1C). Dor (1984) listed specimen SMF 2701 as the holotype of Cheilinus
quinquecinctus, but Rüppell (1835) did not designate a holotype in his description, and the three specimens
collected by him in Jeddah are then considered as syntypes (all in the fish collection of the Senckenberg Research
Museum and Natural History Museum Frankfurt, SMF). One of them, SMF 2701, is labelled “L-Holotyp.”,
presumably short for “Lecto-Holotypus”, i.e. the lectotype. However, to our knowledge no lectotype designation
has been published to date. All three dry adult specimens with rays posteriorly in caudal fin free of membrane.
Differences in shape of caudal fin between the males of the two species are confirmed by examination of additional
museum material and underwater photographs. Moreover, examination of material of both species shows that C.
quinquecinctus has modally fewer gill rakers than C. fasciatus. A comparison of the color pattern in females and
males of both species shows that C. quinquecinctus can easily be distinguished from C. fasciatus.
The objective of this study is to resurrect C. quinquecinctus from synonymy with C. fasciatus and to redescribe
it as a valid species, known from the Red Sea and the Gulf of Aden. Our examination is based on the examination
of the types and additional material of both species, as well as by comparison of numerous color photographs for
distinction of color patterns in C. fasciatus and C. quinquecinctus. In order to fix the name-bearing type of C.
quinquecinctus a lectotype is designated (Fig. 2A). Furthermore, the evolutionary divergence of the two species is
shown by analysis of COI sequences from C. quinquecinctus from the Red Sea and C. fasciatus from the Western
Indian and western Pacific Oceans.
Materials and methods
Specimens from the following institutions were examined: Bernice P. Bishop Museum, Honolulu (BPBM); King
Abdulaziz University Marine Museum, Jeddah (KAUMM) (temporarily housed at SMF); Museum National
d’Histoire naturelle, Paris (MNHN); Senckenberg Research Institute and Natural History Museum, Frankfurt
(SMF); Smithsonian Institution National Museum of Natural History, Washington (USNM), and Museum für
Naturkunde, Zoologisches Museum, Berlin (ZMB).
The length of specimens is given as standard length (SL), measured from the front of the upper lip (or from
projecting canine tooth of upper jaw) to the base of the caudal fin (posterior end of the hypural plate); head length
(HL) from the same point to the posterior end of the opercular flap; snout length from the same point to the nearest
fleshy edge of the orbit; upper-jaw length from the same anterior point to the posterior end of the maxilla. Body
depth is the greatest depth from the ventral edge of the abdomen vertically to the base of the spinous portion of the
dorsal fin; body width is measured just posterior to the gill opening; orbit diameter is the greatest fleshy diameter,
and interorbital width is the least bony width; suborbital depth is taken from the fleshy edge of the orbit to the
nearest edge of the snout above the upper lip; caudal peduncle depth is the least vertical depth, and caudal peduncle
length is the horizontal distance between verticals at the posterior end of the anal-fin base and the caudal-fin base;
length of spines and rays is measured from their base to tip; caudal-fin length is measured horizontally from the fin
base to a vertical at the tip of the longest ray; pectoral-fin length is the length of the longest ray measured from the
base of the uppermost ray; pelvic-fin length is measured from the base of the pelvic spine to the tip of the longest
soft ray.
Morphometric data are given as ratio of SL in the text, and as percentages of SL in the Table 2. Proportional
measurements in the text are rounded to the nearest 0.5 mm. Lateral-line scale counts (LL) include the last pored
scale that overlaps the end of the hypural plate; scales above and below the lateral line are counted in oblique rows
to the origin of the dorsal and anal fins, and do not include the very small scales present at the base of fins; the
count of gill rakers is made on the first gill arch and includes rudiments.
A phylogenetic analysis of the barcoding portion of the mitochondrial COI gene (Folmer et al., 1994, Hebert et
al., 2003) of Cheilinus quinquecinctus and C. fasciatus by maximum likelihood (ML) was performed, in order to
assess the evolutionary divergence of the two species. Genomic DNA was isolated either with a DNeasy tissue kit
(Qiagen, Hilden, Germany) or according to the protocol developed by the Canadian Centre for DNA Barcoding
(Ivanova et al., 2006) from tissue samples of specimens of C. quinquecinctus that were stored in undiluted analysis
grade EtOH at –26°C. Amplification of a 652 bp sequence of the mtCOI gene was carried out with the universal,
M13-tailed primer set COI-3 from Ivanova et al. (2007; partly taken from Ward et al., 2005) according to the PCR
Zootaxa 4158 (4) © 2016 Magnolia Press
protocol in that study or the modified protocol from Geiger et al. (2014). Amplicons were Sanger sequenced from
both ends with primers M13F (-21) and M13R (-27) (Messing, 1983) and contigs were assembled in Geneious Pro
5.4.4 (Biomatters, Aukland, New Zealand). An alignment of COI sequences obtained in this study, sequences of C.
fasciatus available from GenBank, and sequence of C. trilobatus and C. undulatus as outgroup was constructed in
MAFFT v7.017 (Katoh et al., 2013). A maximum likelihood (ML) phylogenetic tree was then inferred in PhyML
3.0 (Guindon et al., 2010; Guindon & Gascuel, 2003) under the TPM3uf+Γ model, the best fitting model of
nucleotide substitution as estimated according to AIC scores in jModelTest (Posada, 2008; Guindon & Gascuel,
2003). Reliability of branch support was assessed by 1.000 bootstrap replicates. Uncorrected pairwise distances
and pairwise K2P-distances for all aligned sequences were estimated in PAUP* (Swofford, 1998), and the
computer program Species Delimitation (Masters et al., 2011) was used to calculate distances among C. fasciatus
and C. quinquecinctus sequences, respectively, as well as to assess closest pairwise distances between the two
species. Computational analyses were conducted via respective software plugins in Genious Pro. Material for the
genetic analysis is listed in Table 1.
Cheilinus fasciatus (Bloch, 1791)
Redbreasted Wrasse
Figures 1, 3–8; Tables 1–3.
Sparus fasciatus Bloch, 1791: 18 (Japan).
Sparus bandatus Perry, 1810: unnumbered page (“Eastern Ocean”).
Labrus enneacanthus Lacepède, 1801: 433 (No locality given, probably western Indian Ocean).
Cheilinus fasciatus: Günther, 1862: 129 (Indian Seas); Playfair & Günther, 1867: 89 (Zanzibar); Klunzinger, 1871: 555 (East
Africa, Indian Ocean); Day, 1877: 394 (“seas of India”); Fowler & Bean, 1928: 245 (Philippines, Halmahera, Okinawa);
de Beaufort, 1940: 81 (Indo-Australian Archipelago); Smith, 1957: 109 (Indo-Pacific to Mozambique); Jones & Kumaran,
1980: 483 (Laccadives); Winterbottom et al., 1989: 52 (Chagos Archipelago); Randall & Anderson, 1993: 33 (Maldives);
Randall et al., 1997: 322 (Great Barrier Reef); Myers, 1999: 189 (throughout Micronesia); Fricke, 1999: 401 (Mascarene
Ids.); Parenti & Randall, 2000: 8 (Mozambique to Samoa and Marshall Ids.); Westneat, 2001: 3414 (Indo-Pacific); Kuiter,
2002: 64 (Indo-Pacific); Randall et al., 2003: 21 (Tonga); Randall, 2005: 396 (east coast of Africa to Samoa and
Micronesia); Satapoomin in Kimura et al., 2009: 219 (Andaman Sea); Nishiyama & Motomura, 2012: 240 (Japan); Allen
& Erdmann, 2012: 642 (throughout East Indian region).
Diagnosis. A species of Cheilinus with IX spines and 10 soft rays in dorsal fin; 13–16 (usually 14 or 15) gill rakers;
body depth 2.3–2.7 in SL; upper and lower lobes of caudal fin prolonged in males; juveniles (Fig. 3) brown with
five white bars across body, first bar broadest and brightest, below third dorsal-fin spine, second bar indistinct, on
ventral half of body, fifth bar faint, anteriorly on caudal peduncle; all bars but the second extend onto dorsal and
anal fins; three faint, short, greenish bars on nape and interorbital space; short, oblique, white or yellowish band
from eye across preopercle; narrow white bar at base of caudal fin; large, dark blue spot, surrounded dorsally with
orange, anteriorly in dorsal fin; subadults and females (Figs. 4 and 5) with similar white bars on nape and body as
juveniles, but second bar on lower body becoming more distinct and nearly reaching dorsal fin; few scales behind
eyes and many scales on body with vertical indistinct dark streak; orange area from behind eye and nape to
pectoral-fin base; humeral area with two (sometimes third above) double, rounded to nearly quadrangular, dark
blue or black spots; head becoming olive with short orange-red lines radiating from eye; lower body, dorsal and
anal fins, and posterior half of caudal fin with small dark orange to red spots, only a few spots in fins in some
individuals, sometimes median fins also with short red lines, similar to those radiating from eye; caudal fin white
with black bar in centre (bar not reaching upper and lower margins) and black posterior margin; males (Figs. 6–8)
with similar color pattern but the suffusion of orange becoming bright orange-red, covering postorbital part of
head, anterior of body (including anterior abdomen and chest), and pectoral-fin base, the area restricted posteriorly
to the first white bar, not enclosing it; second white bar across the body reaching dorsal fin; black streak on scales
becoming broader and well-defined. Reaches about 36 cm.
Zootaxa 4158 (4) © 2016 Magnolia Press
TABLE 1. Information on specimens of Cheilinus included in the phylogenetic analysis of the barcoding region of the COI gene.
Species name Catalog number of
voucher specimen
Locality GenBank
Cheilinus fasciatus HLC-13075 western Pacific Ocean (Vietnam, Thanh Pho-Ho Chi
FJ583119 Steinke et al. (2009)
SMRF6 western Pacific Ocean (Samar, Philippines) KF809396 Norcio et al. (unpublished)
UG0029 western Pacific Ocean (Australia, Lizard Island) KP194074 Steinke et al. (unpublished)
UG0047 western Pacific Ocean (Australia, Lizard Island) KP194008 Steinke et al. (unpublished)
UG0048 western Pacific Ocean (Australia, Lizard Island) KP194612 Steinke et al. (unpublished)
UG0540 western Pacific Ocean (Australia, Lizard Island) KP193981 Steinke et al. (unpublished)
KUT 6997 Indian Ocean (Seychelles, Mahe) KF929729 Bentley & Wiley (unpublished)
NBE0203 Indian Ocean (Madagascar, Nosy Vorona) FJ583120 Hubert et al. (2012)
NBE0478 Indian Ocean (Madagascar, Bay d`Ambanoro) JF458020 Hubert et al. (2012)
NBE0479 Indian Ocean (Madagascar, Bay d`Ambanoro) JF434854 Hubert et al. (2012)
NBE0480 Indian Ocean (Madagascar, Bay d`Ambanoro) JF458018 Hubert et al. (2012)
NBE0625 Indian Ocean (Madagascar, Nosy Tanikely) JF434852 Hubert et al. (2012)
NBE0626 Indian Ocean (Madagascar, Nosy Tanikely) JF458016 Hubert et al. (2012)
NBE0627 Indian Ocean (Madagascar, Nosy Tanikely-Sud) JQ349878 Hubert et al. (2012)
NBE1017 Indian Ocean (Madagascar, Ampasindava) JF457349 Hubert et al. (2012)
Cheilinus quinquecinctus HLC-11559 Red Sea (locality not known) FJ583120 Steinke et al. (2009)
KAUMM 384 Red Sea (Saudi Arabia, Al Qunfidah) KU841539 this study
SMF 33635 Red Sea (Saudi Arabia, Rabigh) KU841540 this study
SMF 35787 Red Sea (Saudi Arabia, Farasan Island) KU841541 this study
SMF 35784 Red Sea (Saudi Arabia, Al Lith) KU841542 this study
SMF 35785 Red Sea (Saudi Arabia, Jeddah) KU841543 this study
KAUMM 386 Red Sea (Saudi Arabia, Jeddah) KU841544 this study
KAUMM 387 Red Sea (Saudi Arabia, Jeddah) KU841545 this study
KAUMM 385 Red Sea (Saudi Arabia, Al Lith) KU841546 this study
SMF 35786 Red Sea (Saudi Arabia, Al Lith) KU841547 this study
Cheilinus trilobatus NBE0652 Indian Ocean (Madagascar, Nosy Tanikely-Sud) JQ349881 Hubert et al. (2012)
Cheilinus undulatus CSIRO H 4877-03 western Pacific Ocean (Australia, Queensland) EF609322 Ward & Holmes (2007)
Zootaxa 4158 (4) © 2016 Magnolia Press
Cheilinus fasciatus Cheilinus quinquecinctus
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3UHSHOYLFOHQJWK  ± ±  ± ±
%DVHRIGRUVDOILQ  ± ±  ± ±
)LUVWGRUVDOILQVSLQH  ± ±  ± ±
1LQWKGRUVDOILQVSLQH  ± ±  ± ±
/RQJHVWGRUVDOILQUD\  ± ±  ± ±
%DVHRIDQDOILQ  ± ±  ± ±
)LUVWDQDOILQVSLQH  ± ± ± ±
6HFRQGDQDOILQVSLQH  ± ±  ± ±
7KLUGDQDOILQVSLQH  ± ±  ± ±
6XERUELWDOGHSWK  ± ±  ± ±
/RQJHVWDQDOILQUD\  ± ±  ± ±
3HFWRUDOILQOHQJWK  ± ±  ± ±
3HOYLFVSLQHOHQJWK  ± ±  ± ±
3HOYLFILQOHQJWK  ± ±  ± ±
&DXGDOILQOHQJWK  ± ±  ± ±
  ±  ± ±
//VFDOHV  ±± ±±  ±± ±±
Zootaxa 4158 (4) © 2016 Magnolia Press
FIGURE 3. Cheilinus fasciatus, juvenile, Bali, Indonesia. Photo by A. Ryanskiy.
FIGURE 4. Cheilinus fasciatus, sudadult, western Papua New Guinea. Photo by A. Ryanskiy.
Zootaxa 4158 (4) © 2016 Magnolia Press
FIGURE 5. Cheilinus fasciatus, female, Bali, Indonesia. Photo by A. Ryanskiy.
FIGURE 6. Cheilinus fasciatus, male, SAIAB 78100, 185 mm SL, Seychelles. Photo by P.C. Heemstra.
Zootaxa 4158 (4) © 2016 Magnolia Press
FIGURE 7. Cheilinus fasciatus, male, Anilao, Philippines. Photo by R. Myers.
FIGURE 8. Cheilinus fasciatus, male, Kwajalein Atoll, Marshall Islands. Photo by S. Johnson.
Distribution and ecology. Ranges from the east coast of Africa and islands of the western Indian Ocean to
Micronesia and Samoa and Tonga, in the western Pacific north to the Ryukyu Islands, Japan, and south to
Queensland, Australia. The species is still unknown from the Arabian Sea (Randall, 1995), the record from Socotra
(Kemp, 1998) represents either C. quinquecinctus or C. fasciatus, but no images from field surveys are available to
resolve the question (J. Kemp, pers. comm.).
Usually solitary, mainly on sand and rubble substrata near coral reefs, more common in protected than exposed
areas, generally at depths less than 20 m, juveniles secretive.
Zootaxa 4158 (4) © 2016 Magnolia Press
Remarks. The synonymy of Sparus bandatus and Labrus enneacanthus follows Parenti & Randall (2000). In
his description of L. enneacanthus Lacepède (1801) gave the color pattern of caudal fin as “deux autres bandes
transversales sur la caudale, qui est en croissant”, which well matches that for C. fasciatus (caudal fin with a broad
black middle bar and black margin), and we confirm the synonymy of Sparus bandatus and Labrus enneacanthus
with Cheilinus fasciatus.
Material examined. Lectotype, ZMB 8577, 250.0 mm, Japan; paralectotypes: ZMB 2651, 147.5 mm,
“Ostindien”; ZMB 8158, 187.5 mm, Java, Indonesia.
Madagascar: MNHN 1900-236, 170.0 mm; MNHN 1999-079, 141.0 mm, Diego-Suarez; MNHN 1999-149,
130.5 mm; SMF 1157, 135.0 mm. Seychelles: MNHN A-7474, 2: 122.0–136.5 mm; MNHN A-7475, 134.5 mm;
MNHN A-7478, 162.0 mm; SAIAB 78100, 185 mm, west of Cachee Island. Mauritius: MNHN A-7476.
Maldives: BPBM 32923, 103.0 mm; SMF 27244, 2: 153.0–181.5 mm + 7 heads; SMF 5267, 176.0 mm; SMF
5268, 2: 150.0–169.0 mm; SMF 5269, 5: 118.0–172.5 mm; SMF 5270, 151.0 mm. Sri Lanka: SMF 27245, 87.5
mm. Indonesia: MNHN A-7420, Djakarta; MNHN A-7473, 2: 120.0–141.5 mm, Djakarta; MNHN 0000-5862,
Djakarta; ZMB 21304, 103.0 mm, Java; MNHN A-9042, 170.0 mm, Makasar; MNHN A-7472, 2: 132.0–142.5
mm, Nias; ZMB 17758, 129.0 mm, Padang, Sumatra. Philippines: USNM 377929, 139.0 mm, Mindoro,
Maricaban Island; USNM 408892, 172.0 mm, Sorsogon province; USNM 431519, 180 mm, Palawan, Puerto
Princesa Bay; USNM 431533, 140 mm, Palawan, Honda Bay. Vietnam: MNHN 2005-2123, 147.0 mm. Japan:
BPBM 8687, 115.0 mm, Ishigaki Island. Marshall Islands: BPBM 8789, 2: 193.5–235.0 mm, Enewetak Atoll;
BPBM 37180, 154.5 mm, Enewetak Atoll. Caroline Islands: USNM 224016, 4: 81.5–128.5 mm, Pohnpei, Tanak
Island. Palau: BPBM 31408, 3: 75.0–93.0 mm. New Caledonia: MNHN A-7477, 2: 126.5–173.0 mm; MNHN
1980-763, 206.0 mm; MNHN 1980-918, 181.5 mm. Van u a t u : MNHN 1894-342, 163.0 mm.
Cheilinus quinquecinctus ppell, 1835
Whitebarred Wrasse
Figures 2, 9–13, Tables 1–3
Cheilinus quinquecinctus Rüppell, 1835: 19 (Jeddah).
Cheilinus quinquecinctus: Günther, 1862: 130 (Red Sea); Kuiter, 2002: 64 (Red Sea); Lieske & Myers, 2004: 154 (Mangrove
Bay, Red Sea).
Chilinus quinquecinctus: Klunzinger, 1871: 555 (Red Sea).
Cheilinus fasciatus quinquecinctus: Klausewitz, 1967: 59 (Sarso Id., Saudi Arabia, Red Sea).
Cheilinus fasciatus (non Bloch): Rüppell, 1828: 23 (Massawa, Eritrea, Red Sea); Playfair & Günther, 1867: 89 (Red Sea
quoted); Day, 1877: 394 (Red Sea quoted); Borsieri, 1904: 216 (Massawa); Fowler & Bean, 1928: 245 (Red Sea quoted);
de Beaufort, 1940: 81 (Red Sea quoted); Roux-Estève & Fourmanoir, 1955: 198 (Abu Latt, Saudi Arabia, Red Sea); Smith,
1957: 109 (Red Sea quoted); Randall, 1983: 113 (Red Sea); Tortonese, 1983: 108 (Jeddah, Saudi Arabia, Red Sea); Dor,
1984: 197 (Red Sea); Gomon & Randall, 1984: LABR Che 5 (western Indian Ocean including Red Sea); Goren & Dor,
1994: 53 (Red Sea); Parenti & Randall, 2000: 8 (Red Sea quoted); Westneat, 2001: 3414 (Red Sea quoted); Khalaf, 2004:
43 (Jordan); Randall, 2005: 396 (Red Sea quoted); Golani & Bogorodsky, 2010: 39 (Red Sea); Allen & Erdmann, 2012:
642 (Red Sea quoted).
Diagnosis. A species of Cheilinus with IX spines and 10 soft rays in dorsal fin; 13–16 (usually 13 or 14) gill rakers;
body depth 2.3–2.6 in SL; caudal-fin rays free of membrane posteriorly in adults; juveniles brown, usually with
five white bars across body, first bar broadest and brightest, bar on midside of abdomen when present indistinct,
fifth bar anteriorly on caudal peduncle and faint; all bars except the bar on midside of abdomen extend onto dorsal
and anal fins; three short faint bars on nape and interorbital space; short oblique white band from eye across
preopercle; narrow white bar at base of caudal fin; large dark blue spot, surrounded with orange dorsally, anteriorly
in dorsal fin; subadults and females with similar white bars as juveniles but second bar becoming distinct and
nearly reaching dorsal-fin base; scales on postorbital head, dorsum, and ventrally on body edged with black;
humeral area with three (rarely two) irregular, dark blue or black marks; postorbital head and anterior body
suffused with orange; caudal fin broadly white basally, followed by a slightly broader black zone, outer fifth pale
grey; males with much more black pigment on body scales and posteriorly on head, and the suffusion of orange
becoming bright orange-red area on postorbital head, anterior of body, abdomen, and pectoral-fin base, color
extending posteriorly nearly to second white bar, enclosing first bar in large males; head brown with a large
orangish area anteriorly around eye, on cheek and most of snout; caudal fin as in females. Reaches about 35 cm.
Zootaxa 4158 (4) © 2016 Magnolia Press
FIGURE 9. Cheilinus quinquecinctus, juvenile, Jeddah, Saudi Arabia. Photo by H. Sjoeholm.
FIGURE 10. Cheilinus quinquecinctus, subadult, Jeddah, Saudi Arabia. Photo by S.V. Bogorodsky.
Zootaxa 4158 (4) © 2016 Magnolia Press
FIGURE 11. Cheilinus quinquecinctus, female, Temples, Egypt. Free rays in caudal fin begin to develop. Photo by A.
FIGURE 12. Cheilinus quinquecinctus, male, Jeddah, Saudi Arabia. Photo by H. Sjoeholm.
Zootaxa 4158 (4) © 2016 Magnolia Press
FIGURE 13. Cheilinus quinquecinctus, male, Yanbu, Saudi Arabia. Photo by J.E. Randall.
Description. Dorsal-fin rays IX,10, spines progressively longer posteriorly; anal-fin rays III,8, spines stout,
third longest; all dorsal- and anal-fin rays branched; pectoral-fin rays 12, upper two rays unbranched; pelvic-fin
rays I,5; principal caudal-fin rays 13, the middle 11 branched, upper procurrent rays 5, lower 4; lateral line
interrupted below posterior portion of dorsal-fin, anterior lateral line following dorsal contour of body, posterior
part midlaterally on caudal peduncle; pored lateral-line scales 14–16 (rarely 16) + 9−10 (rarely 10); 2 pored scales
on caudal-fin base, the last very large and pointed; each lateral-line scale with a horizontal tubule ending in a single
pore; scales above lateral line to origin of dorsal fin 1.5; scales below lateral line to origin of anal fin 5; median
predorsal scales 5 and 2 large scales anteriorly; gill rakers 13–16 (rarely 15 or 16), including 2 or 3 rudiments on
lower limb; branchiostegal rays 5.
Body moderately deep, depth 2.3–2.6 in SL, and compressed, width 4.7–5.3 in SL; head large, length 2.4–2.6
in SL; dorsal profile of head of juveniles nearly straight, forming an angle of about 30° to horizontal axis of body;
head profile of adults straight to above eye at an angle of about 45°, then convex to origin of dorsal fin; snout
obtuse, its length 2.1–2.5 in HL; suborbital moderately deep, depth 2.9–4.5 in HL; eye moderately large in
juveniles, decreasing in size with growth, its diameter 3.7–4.7 in HL in juveniles, 4.8–6.3 in subadults and adults;
interorbital space strongly convex, least bony width 3.0–3.5 in HL; caudal peduncle moderately deep, the depth
0.5–0.6 in its length; caudal-peduncle depth 5.2–5.8 in SL; caudal-peduncle length 8.1–11.5 in SL.
Mouth nearly horizontal, jaws prominent, lower jaw not projecting above upper, maxilla not reaching to below
anterior margin of eye; both jaws with one row of small conical teeth on each side and with pair of strong canine
teeth at front, upper pair slightly outflaring; no enlarged tooth at angle of mouth on upper jaw; lips thick but not
fleshy; gill opening extending forward to a vertical at rear edge of eye; gill membranes broadly attached to isthmus,
without free fold across; posterior margin of preopercle smooth; preopercular margin free to level just below eye
dorsally and to below anterior margin of pupil ventrally; ventral edge of preopercle free nearly to a vertical at
posterior end of maxilla; outer and ventral part of preopercle thin and membranous, with slight indentation in
margin above rounded corner; gill rakers short and simple; anterior nostril in short tube without flap about the orbit
diameter before upper part of eye; posterior nostril resembles small sensory pore dorsoposteriorly to anterior
nostril, distance between nostrils about one-third of eye diameter.
Zootaxa 4158 (4) © 2016 Magnolia Press
Scales large, cycloid, thin, and membranous; head mostly scaled except for anterior interorbital space, snout,
lower jaw, chin, and membranous flange of preopercle; predorsal scales extending forward to about anterior one-
third of orbit; cheek with 2 rows of scales; scales extending basally onto dorsal and anal fins; a slender axillary
scale above base of pelvic fin, partly covered by a large scale between bases of pelvic and pectoral fins; caudal fin
with scales basally.
Origin of dorsal fin on a vertical at upper base of pectoral fin, predorsal length 2.1–2.3 in SL; first dorsal-fin
spine 12.7–16.9 in SL in adults, ninth dorsal-fin spine longest, length 5.7–7.0 in SL; membranes of spinous portion
of dorsal fin incised in juveniles and small individuals, not incised in adults; each interspinous membrane of dorsal
and anal fins extending above spine tip; sixth to eighth dorsal-fin soft ray longest, 3.8–4.8 in SL; origin of anal fin
below base of eighth dorsal-fin spine, preanal-fin length 1.4–1.5 in SL; anal-fin spines reducing in length with
growth, first anal-fin spine 9.8–14.2 in SL in juveniles, 14.1–17.3 in SL in adults; second anal-fin spine 5.8–8.4 in
SL in juveniles, 8.1–12.0 in SL in adults; third anal-fin spine longest, the length 4.3–5.6 in SL in juveniles, 6.3–7.5
in SL in adults; sixth ray of anal fin longest, in juveniles and subadults 4.5–5.6 in SL, posterior rays of anal fin
prolonged in adults, fin angular in shape posteriorly, the length of lobe 2.8–4.5 in SL; base of dorsal fin 1.7–2.0 in
SL; base of anal fin 3.7–4.3 in SL; pectoral fins rounded, longest ray 1.5–1.9 in HL; origin of pelvic fins slightly
posterior to lower base of pectoral fins, prepelvic length 2.3–2.6 in SL; pelvic-fin spine 7.2–8.9 in SL; first pelvic
soft ray longest, not reaching anus, 4.4–5.4 in SL; caudal fin slightly rounded in juveniles, truncate in females, the
rays in adult males free of membranes posteriorly and long (fin looks ragged in large males), the length of fin 2.2
3.1 in SL; longest free ray of caudal fin varying in length: 2.4–8.2 % caudal-fin length in subadults, 8.1–33.3%
caudal-fin length in adults. Free rays of caudal fin subequal in length in smaller adults, and usually median rays
longest in large adults.
Color: juveniles (Fig. 9) brown usually with five white bars (bars edged with dark red in very small juveniles)
on the body: first bar broadest and brighter than other bars, beginning from tip of second dorsal-fin spine, slanting
to base of fin and curving across body ending slightly behind the pelvic-fin insertion and continues through
posterior margin of pelvic fin (inner part of pelvic fin dark copper); second bar short, indistinct, on ventral half of
abdomen (this bar sometimes not visible in some juveniles but appears with growth, see below); third bar
beginning from tip of seventh dorsal-fin spine across body, then curving below midline of body and ending at tip of
second anal-fin spine; fourth bar extending from tip of ninth dorsal-fin spine to anal-fin base; a faint fifth bar
anteriorly on caudal peduncle and a narrow white bar at base of caudal fin; all bars, except for the second bar when
visible on midside of abdomen, continuing onto adjacent fins; two faint, short, white bars on nape, one band across
interorbital space, and a short oblique white band from eye across preopercle; two small, black spots present
midlaterally, one behind second white bar, and another on caudal peduncle, spots disappear with growth;
postorbital area of larger juveniles with small black spots; first and second interspinous membrane of dorsal fin
with a large black or deep blue spot at base and small, triangular, orange spot above, tip of first membrane black;
caudal fin translucent with faint white submarginal bar.
Subadults and females (Figs. 10 and 11) with similar white bars as juveniles but second bar on abdomen
becoming more obvious, nearly reaching dorsal fin; scales on postorbital head, anterior body, lower abdomen and
along back edged with black; margin of opercle outlined with blue dorsoposteriorly in subadults; rest of body with
small dark brown spots aligned in row along edge of most scales in females, spots connected to each other with
growth to form black edge of scale; postorbital head and anterior body suffused with dark orange; several irregular
black spots posteriorly on head and short orange lines extending posteriorly from eye or few small orange spots
behind eye; oblique white band ventroposteriorly from eye, becoming a short dark green band in subadults and
disappearing in females; first bar on nape and band across interorbital space disappearing with growth; caudal fin
broadly white basally, followed by slightly broader black zone, the outer fifth pale grey; inner part of pelvic fins
black, outer part and pelvic-fin spine pale brown suffused with red.
Body of males dark brown; edge of scales on postorbital head, anterior body, lower abdomen, and along back
black in smaller males (Fig. 12), but black on edge becoming broader ventrally and posteriorly on body in large
males (Fig. 13); broad bright orange area anteriorly on body extending from level of upper margin of eye and
enclosing abdomen and pectoral-fin base; white bars on body and bar on nape retain in smaller males; bars become
diffuse, including first bar enclosed by orange area, in large males; bars extending onto fins basally, more obvious
on hind dorsal and anal fins; white bar on nape and first bar dorsally on body with orange hue; head brown suffused
with orangish anteriorly, enclosing eye, cheek, and most of snout; humeral area with three (rarely two) irregular
Zootaxa 4158 (4) © 2016 Magnolia Press
dark blue or black marks; dorsoposterior margin of opercle black; iris green with orange inner and outer ring;
dorsal fin suffused with blue, with two red streaks distally on each membrane of spinous portion and anterior half
of soft portion, forming two longitudinal rows; first or first two membranes with diagonal black streak at base;
anterior half of anal fin with two irregular, longitudinal, red lines distally; caudal fin broadly white basally,
followed by a broad black zone, with narrow pale grey margin; pectoral fins with translucent membranes (with
slight orange hue) and orange-yellow rays; pelvic fins mostly blackish with brownish red outer part and spine.
Distribution and habitat. This species is known from the Red Sea and the Gulf of Tadjoura. Kemp (1998)
listed C. fasciatus from Socotra, but no photographs or voucher specimens are available to determine positive
identification. Adults of C. quinquecinctus are generally found as solitary individuals over sand or sand-and-rubble
substrata adjacent to sheltered coral reefs from depth of 2–40 m; large males are usually seen swimming along the
reef margin; juveniles are more secretive and may be found at entrance of small caves or crevices, resemble and
often are confused with Wetmorella bifasciata Schultz & Marshall, 1954. Sometimes accompanies goatfishes or
even divers who may expose fossorial animals by their flippers. Feeds on benthic invertebrates (Gomon & Randall,
Remarks. Cheilinus quinquecinctus had previously been synonymized with the similar C. fasciatus (Parenti &
Randall, 2000), a species found elsewhere in the Indo-Pacific from the western Indian Ocean to the Samoan Islands
and Tonga. Kuiter (2002) and Lieske & Myers (2004) used the name C. quinquecinctus for the Red Sea species
with note that it differed mainly in the absence of ragged posterior margin of the caudal fin in C. fasciatus.
However, Cheilinus quinquecinctus can be distinguished from C. fasciatus by a combination of the following
characters: caudal fin with rays free of membrane posteriorly in males vs. caudal fin with well-developed upper and
lower lobes in males of C. fasciatus; total number of gill rakers 13–16 (usually 13 or 14) vs. 13–16 (usually 14 or
15) in C. fasciatus (count for examined specimens is given in the Table 3); broad orange or orange-red area
anteriorly on body and pectoral-fin base, dorsally from the level of upper margin of eye below to chest and
abdomen, zone extending forward to eye, posteriorly nearly to second white bar on body (first white bar enclosed
by red area in large males) vs. broad orange-red zone from nape to chest and abdomen, extending forward to eye,
posteriorly restricted by first white bar in C. fasciatus; no spots on body and fins vs. lower body, dorsal and anal
fins, and posterior part of caudal fin with small dark red spots in C. fasciatus; no lines on the head in males vs. head
with short orange lines radiating from eye in C. fasciatus; caudal fin broadly white basally, followed by slightly
broader black zone, posterior margin pale grey vs. caudal fin white with median black bar not connected with upper
and lower lobes, and black posterior margin in C. fasciatus.
Caudal-fin lobes begin to prolong from length of 93.0 mm SL in C. fasciatus, whereas two specimens of 128
mm SL without lobes in caudal fin; free caudal-fin rays develop in size of about 100 mm in C. quinquecinctus, in
examined material the specimen of 170. 5 mm (SMF 17837) has only upper third free ray longest but the specimen
of 181.5 mm (MNHN 1952-179) has developed, subequal in length, free rays.
TABLE 3. Gill-rakers count for specimens of Cheilinus fasciatus and C. quinquecinctus.
Material examined. Lectotype (designated herein), SMF 2701, 210.0 mm, Saudi Arabia, Jeddah;
paralectotypes: SMF 2732, 226.0 mm, Saudi Arabia, Jeddah; SMF 2740, 205.0 mm, Saudi Arabia, Jeddah.
Red Sea. Egypt: USNM 276830, 4: 53.0–99.0 mm, Strait of Jubal, Shaab Abu Qualawa; MNHN 1977-861,
163.5 mm, Ras Muhammad; SMF 5061, 194.5 mm, Gubal Island; SMF 12011, 147.0 mm; ZMB 2652, 177.5 mm.
Saudi Arabia: SMF 4328, 2: 182.5–199.0 mm, Farasan Archipelago, Sarso Island; SMF 27257, 151.5 mm,
Farasan Archipelago, Sarso Island; SMF 17838, 170.5 mm, Jeddah; SMF 35238, 40.0 mm, Jeddah, Obhur; SMF
35785 (KAU14-104), 62.0 mm, Jeddah, Obhur; SMF 35786 (KAU14-939), 139.0 mm, Al Lith; SMF 35787
(KAU12-394), 175.0 mm, Farasan Island; KAUMM 384 (KAU11-284), 176.0 mm, Al Qunfudhah; KAUMM 385
(KAU14-785), 103.5 mm, Al Lith; KAUMM 386 (KAU14-105), 56.0 mm, Jeddah, Obhur; MNHN 1952-175,
122.0 mm, Farasan Archipelago, Abu Latt Island; MNHN 1952-176, 136.0 mm, Farasan Archipelago, Abu Latt
Island; MNHN 1952-177, 169.0 mm, Farasan Archipelago, Abu Latt Island; MNHN 1952-178, 137.0 mm, Farasan
Number of gill rakers 13 14 15 16 mean
C. fasciatus 11419414.7
C. quinquecinctus 8113113.9
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Archipelago, Abu Latt Island; MNHN 1952-179, 181.5 mm, Farasan Archipelago, Abu Latt Island; MNHN 1952-
180, 154.0 mm, Farasan Archipelago, Abu Latt Island; MNHN 1952-181, 189.0 mm, Farasan Archipelago, Abu
Latt Island; MNHN 1952-182, 106.0 mm, Farasan Archipelago, Abu Latt Island. Gulf of Tadjoura: MNHN 1977-
666, 64.5 mm.
Molecular analysis
Cheilinus fasciatus and C. quinquecinctus formed reciprocally monophyletic clades with high bootstrap support in
the gene tree resulting from maximum likelihood analysis of partial COI gene sequences (Fig. 14). Sequence
variability within the two clades was low with an average pairwise K2P-distance of 0.2% (uncorrected p-distance:
0.2) in C. quinquecinctus and of 0.6 (0.6) in C. fasciatus. The closest pairwise distance between the two clades with
a value of 4.4% (4.2%) was considerably higher than the average pairwise distance within the clades.
FIGURE 14. Maximum likelihood (ML) gene tree of partial COI sequences (652 bp) of Cheilinus fasciatus and C.
quinquecinctus from various parts of the distribution range of the two species aligned with one sequences of C. trilobatus and
C. undulatus, with the latter defined as the outgroup. Values on branches indicate percent bootstrap support from 1.000
replicate analyses, if higher than 80. The scale bar represents the average number of nucleotide substitutions in relation to
branch length.
The number of diagnostic sites in the partial COI gene alignment (652 bp) that differed between all specimens
of C. fasciatus and C. quinquecinctus, respectively, was 24. Within C. fasciatus, two sub-clades were formed
containing specimens from the western Pacific (i.e., eastern Australia, Vietnam, and the Philippines), or the
western Indian Ocean (i.e., Madagascar, Seychelles), respectively. The closest pairwise distance between these two
subclades was relatively low with a value of 0.9% (0.9%) when compared with distance values within clades
(western Pacific: 0.3% (0.3%), Western Indian Ocean: 0.2% (0.2%)). Only three sites of the entire alignment were
diagnostic for the differentiation between the two geographically defined sub-clades.
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Of the syntypes of Cheilinus quinquecinctus specimen SMF 2701 is in the best condition and has the specific well-
developed ragged posterior margin of the caudal fin. We therefore designate this specimen as the lectotype of
Cheilinus quinquecinctus (Fig. 2A). Of the two paralectotypes, one is illustrated here (SMF 2732 in Fig. 2B).
As various authors have previously treated C. quinquecinctus as a junior synonym of C. fasciatus (e.g. Randall
& Parenti, 2000), clarification of the status of C. quinquecinctus is required. Such a step is also a prerequisite
before interpretations of the geographic distribution of Cheilinus species and the evolution of endemism in the Red
Sea can be undertaken. While previous studies of the status of C. quinquecinctus lacked detailed comparative
analysis of types and other specimens of C. fasciatus and C. quinquecinctus, the morphological examination and
assessment of molecular divergence provided in this study unambiguously establish C. quinquecinctus as a valid
species endemic to the Red Sea and Gulf of Aden. The species is readily distinguished from its widely-distributed
sibling species C. fasciatus by its color pattern (see remarks for C. quinquecinctus above). The most diagnostic
morphological character for these two very similar species is the development with age of a ragged posterior
margin of the caudal fin in C. quinquecinctus, whereas in C. fasciatus only the upper and lower caudal-fin lobes
become extended with age.
As a result of his morphological analysis, Westneat (1993) proposed C. fasciatus as sister group to other
Cheilinus species by having a unique synapomorphy: the central portion of the lateral edge of the hyomandibula
has a sharp dorsal process whereas in other species of the genus this process is a prominent bony ridge — the same
might be presumed to be the case for C. quinquecinctus. Two representatives of other species pairs, C. abudjubbe
(sister species of C. chlorourus) and C. lunulatus (sister species of C. trilobatus), are sympatric with C.
quinquecinctus, and both also possess a ragged posterior margin of the caudal fin. We assume that this feature is a
homoplastic character and has developed independently in these three species. Other species of the genus
distributed in Indo-West Pacific possess a rounded caudal fin with developed upper and lower lobes or upper lobe
only; only some large males of C. trilobatus may have an irregular posterior margin of the caudal fin.
Body proportions, as assessed in this study, do not allow to distinguish C. quinquecinctus and C. fasciatus
except for length of soft rays in the dorsal fin (Fig. 15). With regards to meristic characters, specimens of both
species can also not readily be distinguished. For example the number of gill rakers on the first gill arch, although
modally distinct in the material examined herein (mean 13.9 vs. mean 14.7 in C. quinquecinctus and C. fasciatus,
respectively), is not helpful in identifying specimens. As the numbers of spines and segmented fin-rays are
identical in all but one species of Cheilinus, these characters were not further investigated in this study. The
residual counts taken in this study, such as pored lateral line scales, also did not allow distinction between the two
species. This apparently low level of gross morphological differentiation between sister species, in spite of good
support for long lasting evolution along separate evolutionary trajectories (see below), seems to be not uncommon
in labrid evolution. In other labrid genera similar situations have been observed, where closely related species can
almost solely be distinguished on differences in coloration, but not by any morphological characters (e.g.
Hemigymnus (Randall, 2013), Paracheilinus (Allen et al., 2016) or Pseudocoris (Randall et al., 2015)). These
examples suggest that initiation of species divergence is not necessarily driven by selection for adaptive
morphological characters, but might also result from other adaptive or non-adaptive processes. In these cases,
morphological differentiation (of diagnostic value) might evolve a considerable time after reproductive
incompatibilities between allopatric populations of one ancestral species have come into effect. The molecular
divergence of C. quinquecinctus and C. fasciatus with an average percentage pairwise K2P-distance of 4.69
together with high interspecific genetic divergence compared with low intraspecific genetic divergence is
indicative of complete reproductive isolation and lineage separation. Although simple approaches for species
delimitation such as a general threshold for interspecific divergence (e.g., a 3% pairwise COI divergence threshold)
or for identifying sibling species according to their mean interspecific vs. their intraspecific variation (e.g., a 10x
threshold) may not be generally applicable, e.g., due to differences in traits such as generation time or dispersal
regime (see e.g., Hebert et al., 2004 and references therein), the fulfillment of both these criteria is supportive for
species level evolutionary divergence of Cheilinus fasciatus and C. quinquecinctus.
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FIGURE 15. Length of longest ray in dorsal fin vs. standard length in Cheilinus quinquecinctus (filled circles) and C. fasciatus
(open circles).
Assuming that rates of molecular evolution of the COI gene in the genus Cheilinus are similar to those that
have been reported for other tropical reef fishes, reproductive connectivity between the two sister species ceased to
exist at least two to three million years before present rather than only during the previous hundreds of thousands of
years (see e.g., the compilation in Lessios (2008) for geminate species pairs across the isthmus of Panama with 1 to
2 % K2P-distance divergence in COI per million years). During the Pleistocene, the Red Sea was repeatedly
effectively isolated from the Indian Ocean by low sea level stands during glacial periods (see Klausewitz, 1989 and
references therein). Hence, the existence of glacial refugia for C. quinquecinctus (either inside the Red Sea and/or
outside the Red Sea in the wider Gulf of Aden area) needs to be presumed throughout all glacial periods that
followed the divergence of the recent most common ancestor of C. quinquecinctus and C. fasciatus. Considering
the geological and zoogeographic history of the Arabian Seas region, the evolution of the two species was most
likely initiated in allopatry as a consequence of cessation of gene flow between Red Sea and Indian Ocean
populations with low sea levels during one of the earliest Pleistocene glacial periods. The finding of relatively low
differentiation between specimens of C. fasciatus from the western Pacific and the western Indian Ocean agrees
with frequent findings of population level divergence in reef fishes and other marine organisms, which presumably
has as its cause the separation of Pacific and Indian Ocean populations during more recent glacial sea-level
lowstands (see e.g., Williams et al., 2002 and references therein). In addition, no differences in morphology and
coloration were found between these populations, thus data for the species is given for its distributional range.
However, a more complete picture of the present day population genetic structure and the connectivity among
Indian Ocean and Pacific populations of C. fasciatus can only be obtained by a more complete geographic
sampling including the Indo-Malay Archipelago (see e.g., Timm & Kochzius, 2008).
Among eight species of the widespread Indo-West Pacific genus Cheilinus, four occur in the Red Sea: C.
abudjubbe, C. lunulatus, C. quinquecinctus, and C. undulatus. Only C. undulatus is broadly distributed from the
Red Sea and east coast of Africa to Hawaiian and Pitcairn Islands, whereas the other three species have restricted
Zootaxa 4158 (4) © 2016 Magnolia Press
distributions and are known only from the northwestern Indian Ocean, i.e., from the Red Sea to the either the Gulf
of Aden (C. abudjubbe and C. quinquecinctus) or to the Arabian Gulf (C. lunulatus) (Randall, 1995). Similar
evolutionary histories might be considered for these species and their widespread Indo-Pacific sister species.
Following Westneat (1993) and the present study, the genus Cheilinus contains eight valid species. With their
phylogenetic analysis of labrid tribes, however, Westneat & Alfaro (2005) cast some doubt on the monophyly of
the genus as presently defined, as the phylogeny inferred from mitochondrial and nuclear genes—including
representatives of other cheiline genera (Cheilinus, Epibulus, Oxycheilinus, and Wetmorella)—showed Cheilinus to
be polyphyletic. Although it is beyond the objectives of this study to resolve the boundaries of the genus Cheilinus,
this task, as well as the exploration of evolutionary trajectories of sister species in a common morphological and
phylogenetic framework appear to be rewarding.
Susanne Dorow and Jennifer Steppler are gratefully thanked for technical assistance at SMF. Patrice Prouvost and
Gabsi Zora (MNHN) are thanked for providing the chance to examine material at MNHN and a loan of specimens.
We also thank Peter Bartsch and Christa Lamour (ZMB) for their help during visit in examination of type material
of Cheilinus fasciatus and other material of both species, and Loreen O’Hara (BPPM) and Jeffrey Williams
(USNM) who provided loans of specimens. Thanks are also due to Matthias Juhas and Stephanie Simon (SMF) for
assisting in molecular genetic analyses and we acknowledge the Grunelius-Möllgaard Laboratory at SMF for lab
support. The scientific research cooperation between King Abdulaziz University (KAU), Faculty of Marine
Sciences (FMS), Jeddah, Saudi Arabia, and the Senckenberg Research Institute (SRI), Frankfurt, Germany, in the
framework of the Red Sea Biodiversity Project, during which the present material was collected, was funded by
KAU GRANT NO. “D/1/432-DSR”. The authors acknowledge, with thanks, KAU and SRI for technical and
financial support as well as Ali Al-Aidaroos, Mohsen Al Sofiyani (KAU), Fareed Krupp (SRI and Qatar Natural
History Museum, Doha) for their help in the realization of the present study; also thanks to Scott Johnson, Robert
Myers, Andrey Ryanskiy, and Hans Sjoeholm for underwater photographs and Jerry Kemp for offering access to
his photo collection. John E. Randall is gratefully thanked for critical comments.
Allen, G.R. & Erdmann, M.V. (2012) Reef Fishes of the East Indies. Vol. 2. Tropical Reef Research, Perth, 425–856.
Allen, G.R., Erdmann, M.V. & Yusmalinda, N.L.A. (2016) Review of the Indo-Pacific flasherwasses of the
genus Paracheilinus (Perciformes: Labridae), with descripions of three new species. Journal of the Ocean Science
Foundation, 19, 18–90.
Bloch, M.E. (1785–1795) Naturgeschichte der ausländischen Fische. Vol. 5. Schlesinger, Berlin, viii + 152 pp.
Borsieri, C. (1904) Contribuzione alla conoscenza della fauna ittiologica della Colonia Eritrea. Annali del Museo Civico di
Storia Naturale di Genova, Series 3, 1 (41), 187–220.
Day, F. (1875–1878) The Fishes of India. Bernard Quaritch, London, xx + 778 pp.
De Beaufort, L.F. (1940) The Fishes of the Australian Archipelago. Vol. 8. E.J. Brill, Leiden, xv + 508 pp.
DiBattista, J.D., Roberts, M.B., Bouwmeester, J., Bowen, B.W., Coker, D.J., Lozano-Cortes, D.F., Choat, J.H., Gaither, M.R.,
Hobbs, J.-P.A., Khalil, M.T., Kochzius, M., Myers, R.F., Paulay, G., Robitzch, V.S.N., Saenz-Agudelo, P., Salas, E.,
Sinclair-Taylor, T.H., Toonen, R.J., Westneat, M.W., Williams, S.T. & Berumen, M.L. (2015) A review of contemporary
patterns of endemism for shallow water reef fauna in the Red Sea. Journal of Biogeography, 2015, 1–17.
Dor, M. (1984) Checklist of the Fishes of the Red Sea. The Israel Academy of Sciences and Humanities, Jerusalem, xxi + 427
Eschmeyer, W.N. (Ed.) Catalog of Fishes. Electronic version accessed April 06, 2015. California Academy of Sciences, San
Francisco. Available from: asp
Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial
cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3 (5),
Fowler, H.W. & Bean, B.A. (1928) Contributions to the biology of the Philippine Archipelago and adjacent regions. The fishes
of the families Pomacentridae, Labridae, and Callyodontidae, collected by the United States Bureau of Fisheries Steamer
Albatross” chiefly in Philippine seas and adjacent waters. Bulletin of the United States National Museum, Series 100, 7,
viii + 525 pp.
Zootaxa 4158 (4) © 2016 Magnolia Press
Fricke, R. (1999) Fishes of the Mascarene Islands (Réunion, Mauritius, Rodriguez). An annotated checklist with descriptions of
new species. Koeltz Scientific Books, Koenigstein, 759 pp.
Geiger, M.F., Herder, F., Monaghan, M.T., Almada, V., Barbieri, R., Bariche, M., Berrebi, P., Bohlen, J., Casal-Lopez, M.,
Delmastro, G.B., Denys, G.P.J., Dettai, A., Doadrio, I., Kalogianni, E., Kärst, H., Kottelat, M., Kovacic, M., Laporte, M.,
Lorenzoni, M., Marcic, Z., Özulug, M., Perdices, A., Perea, S., Persat, H., Porcelotti, S., Puzzi, C., Robalo, J., Šanda, R.,
Schneider, M., Šlechtová, V., Stoumboudi, M., Walter, S. & Freyhof, J. (2014) Spatial heterogeneity in the Mediterranean
Biodiversity Hotspot affects barcoding accuracy of its freshwater fishes. Molecular Ecology Resources, 14 (6), 1210–
Golani, D. & Bogorodsky, S.V. (2010) The Fishes of the Red Sea—Reappraisal and Updated Checklist. Zootaxa, 2463, 1–135.
Gomon, M.F. & Randall, J.E. (1984) Labridae. In: Fisher, W. & Bianchi, G. (Eds.), FAO Species Identification Sheets for
Fishery Purposes. Western Indian Ocean (Fishing Area 51). Vo l s . 1 6 . Food and Agriculture Organization of the United
Nations, Rome, without pagination.
Goren, M. & Dor, M. (1994) An Updated Checklist of the Fishes of the Red Sea; CLOFRES II. Israel Academy of Sciences and
Humanities, Jerusalem, XII + 120 pp.
Günther, A. (1862) Catalogue of the Fishes in the British Museum. Vol. 4. British Museum, London, xxii + 334 pp.
Guindon, S. & Gascuel, O. (2003) A simple, fast, and accurate algorithm to estimate large phylogenies by maximum
likelihood. Systematic Biology, 52 (5), 696–704.
Guindon, S., Dufayard, J., Lefort, V., Anisimova, M., Hordijk, W. & Gascuel, O. (2010) New Algorithms and Methods to
Estimate Maximum-Likelihood Phylogenies: Assessing the Performance of PhyML 3.0 Systematic Biology, 59 (3), 307–
Hebert, P.D.N., Stoeckle, M.Y., Zemlak, T.S. & Francis, C.M. (2004) Identification of birds through DNA barcodes. PLoS
Biology, 2 (10), 1657–1663.
Hubert, N., Meyer ,C.P., Bruggemann, H.J., Guerin, F., Komeno, R.J., Espiau, B., Causse, R., Williams, J.T. & Planes, S.
(2012) Cryptic Diversity in Indo-Pacific Coral-Reef Fishes Revealed by DNA-Barcoding Provides New Support to the
Centre-of-Overlap Hypothesis. PLoS ONE, 7 (3), E28987.
Ivanova, N.V., de Waard, J. & Hebert, P.D.N. (2006) An inexpensive, automation-friendly protocol for recovering high-quality
DNA. Molecular Ecology Notes, 6, 998–1002.
Ivanova, N.V., Zemlak, T.S., Hanner, R.H. & Hebert, P.D.N. (2007) Universal primer cocktails for fish DNA barcoding.
Molecular Ecology Notes, 7 (4), 544–548.
Jones, S. & Kumaran, M. (1980) Fishes of the Laccadive Archipelago. The Nature Conservation and Aquatic Sciences Service,
Trivandrum, Kerala, xii + 760 pp.
Katoh, K. & Standley, D.M. (2013) MAFFT Multiple Sequence Alignment Software Version 7: Improvements in Performance
and Usability. Molecular Biology and Evolution, 30 (4), 772–780.
Kemp, J.M. (1998) Zoogeography of the coral reef fishes of the Socotra Archipelago. Journal of Biogeography, 25 (5), 919–
Khalaf, M. (2004) Fish Fauna of the Jordanian Coast, Gulf of Aqaba, Red Sea. Marine Science, 15, 23–50.
Klausewitz, W. (1967) Die physiographische Zonierung der Saumriffe von Sarso. Meteor Forschungsergebnisse, Reihe D, 2,
Klausewitz, W. (1989) Evolutionary history and zoogeography of the Red Sea ichthyofauna. Fauna of Saudi Arabia, 10, 310–
Klunzinger, C.B. (1871) Synopsis der Fische des Rothen Meeres. II. Theil. Verh. K.-K. Verhandlungen der Zoologisch-
Botanischen Gesellschaft in Wien, 21, 441–688.
Kuiter, R.H. (2002) Fairy & Rainbow Wrasses and their Relatives. TMC Publishing, Chorleywood, 208 pp.
Lacepède, B.G.E. (1801) Histoire Naturelles des Poissons. Vol. 3. Chez Plassan, Paris, xlvi + 558 pp.
Lessios, H.A. (2008) The great American schism: Divergence of marine organisms after the rise of the Central American
Isthmus. Annual Review of Ecology Evolution and Systematics, 39, 63–91.
Lieske, E. & Myers, R.F. (2004) Coral Reef Guide Red Sea. Harper Collins Publishers Ltd, London, 384 pp.
Masters, B.C., Fan, V. & Ross, H.A. (2011) Species delimitation – a geneious plugin for the exploration of species boundaries.
Molecular Ecology Resources, 11 (1), 154–157.
Messing, J. (1983) New M13 vectors for cloning. Methods in Enzymology, 101, 20–78.
Myers, R.F. (1999) Micronesian Reef Fishes. 3
Edition. Coral Graphics, Guam vi + 330 pp.
Nishiyama, K. & Motomura, H. (2012) A Photographic Guide to Wrasses of Japan. Toho-shuppan, Osaka, 302 pp.
Zootaxa 4158 (4) © 2016 Magnolia Press
Paepke, H.-J. (1999) Bloch's fish collection in the Museum für Naturkunde der Humboldt Universität zu Berlin: an illustrated
catalog and historical account. Ruggell (Liechtenstein). Theses Zoologicae, 32, 1–216.
Parenti, P. & Randall, J.E. (2000) An annotated checklist of the species of the labroid fish families Labridae and Scaridae.
Ichthyological Bulletin of the J.L.B. Smith Institute of Ichthyology, 68, 1–97.
Playfair, R.L. & Günther, A. (1867) The Fishes of Zanzibar, with a List of the Fishes of the Whole East Coast of Africa. J. van
Voorst, London, xix + 153 pp.
Posada, D. (2008) jModelTest: Phylogenetic model averaging. Molecular Biology and Evolution, 25 (7), 1253–1256.
Randall, J.E. (1983) Red Sea Fishes. Immel Publishing, London, 192 pp.
Randall, J.E. (1986) Family Labridae. In: Smith, M.M. & Heemstra, P.C. (Eds.) Smiths’ Sea Fishes. Macmillan South Africa,
Johannesburg, pp. 1–1047.
Randall, J.E. (2005) Reef and shore fishes of the South Pacific. New Caledonia to Tahiti and the Pitcairn Islands. University
of Hawaii Press, Honolulu, 707 pp.
Randall, J.E. (1995) Coastal Fishes of Oman. University of Hawai’i Press, Honolulu, xiii + 439 pp.
Randall, J.E. (2013) Review of the Indo-Pacific labrid fish genus Hemigymnus. Journal of the Ocean Science Foundation, 6, 2–
Randall, J.E. & Anderson, R.C. (1993) Annotated checklist of the epipelagic and shore fishes of the Maldive Islands.
Ichthyological Bulletin of the J.L.B. Smith Institute of Ichthyology, 59, 1–47.
Randall, J.E., Connell, A.D. & Victor, B.C. (2015) Review of the labrid fishes of the Indo-Pacific genus Pseudocoris with a
description of two new species. Journal of the Ocean Science Foundation, 16, 1–55.
Randall, J.E., Allen, G.R. & Steene, R.C. (1997) Fishes of the Great Barrier Reef and Coral Sea. Crawford House Press,
Bathurst, N.S.W., xx + 557 pp.
Randall, J.E., Williams, J.T., Smith, D.G., Kulbicki, M., Tham, G.M., Labrosse, P., Kronen, M., Clua, E. & Mann, B.S. (2003)
Checklist of the shore and epipelagic fishes of Tonga. Atoll Research Bulletin, 502, 1–35.
Roux-Estève, R. & Fourmanoir, P. (1955) VII Poissons capturés par la mission de la “Calypso” en Mer Rouge. Annales de
l’Institut Oceanographique Monaco (New Series), 30, 195–203.
Rüppell, E. (1828–30) Atlas zu der Reise im nördlichen Afrika. Fische des Rothen Meers. Heinrich Ludwig Brönner, Frankfurt
am Main, 141 pp.
Rüppell, E. (1835–1838) Neue Wirbelthiere zu der Fauna von Abyssinien gehorig. Fische des Rothen Meeres. S. Schmerber,
Frankfurt am Main, 148 pp.
Satapoomin, U. (2009) Family Labridae. In: Kimura, S., Satapoomin, U. & Matsuura, K. (Eds.), Fishes of Andaman Sea.
National Museum of Nature and Science, Tokyo, vi + 346 pp.
Smith, J.L.B. (1957) List of the fishes of the family Labridae in the western Indian Ocean with new records and five new
species. Ichthyological Bulletin of the J.L.B. Smith Institute of Ichthyology, 7, 99–114.
Steinke, D., Zemlak, T.S. & Hebert, P.D. (2009) Barcoding Nemo: DNA-based identifications for the ornamental fish trade.
PLoS ONE, 4 (7), E6300.
Swofford, D.L. (1998) PAUP*. Phylogenetic analysis using Parsimony (* and other methods). Version 4. Sinauer Associates,
Sunderland, Massachusetts.
Timm, J. & Kochzius, M. (2008) Geological history and oceanography of the Indo-Malay Archipelago shape the genetic
population structure in the false clown anemonefish (Amphiprion ocellaris). Molecular Ecology, 17, 3999–4014.
Tortonese, E. (1983) List of fishes observed near Jeddah (Saudi Arabia). Journal of the Faculty of Marine Science, 3, 105–110.
Ward, R.D., Zemlak, T.S., Innes, B.H., Last, P.R. & Hebert, P.D.N. (2005) DNA barcoding Australia's fish species.
Philosophical Transactions of the Royal Society B-Biological Sciences, 360 (1462), 1847–1857.
Ward, R.D. & Holmes, B.H. (2007) An analysis of nucleotide and amino acid variability in the barcode region of cytochrome c
oxidase I (cox1) in fishes. Molecular Ecology Notes, 7 (6), 899–907.
Westneat, M.W. (1993) Phylogenetic relationships of the tribe Cheilini (Labridae: Perciformes). Bulletin of Marine Science, 52
(1), 351–394.
Westneat, M.W. (2001) Family Labridae. In: Carpenter, K.E. & Niem, V.H. (Eds.) The Living Marine Resources of the Western
Central Pacific. Volume 6. Bony fishes part 4 (Labridae to Latimeriidae). Food and Agriculture Organization of the United
Nations, Rome, pp. 3381–4218.
Westneat, M.W. & Alfaro, M.E. (2005) Phylogenetic relationships and evolutionary history of the reef fish family Labridae.
Molecular Phylogenetics and Evolution, 36, 370–390.
Williams, S.T., Jara, J., Gomez, E. & Knowlton, N. (2002) The Marine Indo-West Pacific Break: Contrasting the Resolving
Power of Mitochondrial and Nuclear Genes. Integrative & Comparative Biology, 42, 941–952.
Winterbottom, R., Emery, A.R. & Holm, E. (1989) An annotated checklist of the fishes of the Chagos Archipelago. Life Science
Contributions of the Royal Ontario Museum, 145, vi + 226 pp.
... In some taxa, such variation between Red Sea and Indian Ocean populations is of a subtle nature and as a consequence, it has only been detected by rigorous examination of a wide range of characters in a large sample of specimens. One recent example are the sibling labrid species Cheilinus quinquecinctus (Red Sea) and C. fasciatus (Indian Ocean) that long time have been considered one widespread species (see Bogorodsky et al. 2016). In such cases, the joint assessment of morphological variation together with evolutionary genetic divergence in an integrative taxonomical frame work can provide more profound evidence for species hypothesis than would be obtained by standard morphological analysis alone (see e.g., Dayrat 2005;Will et al. 2005). ...
... Integrative taxonomic study, based on a combination of morphological examination (including comparison of colorationespecially among alive and fresh specimens) and of molecular phylogenetic analyses, can help in distinguishing among closely related species of fish (see e.g. Bogorodsky et al. 2016;Bañón et al. 2016). Here, we compiled morphological and molecular data for two species of Crenidens -C. ...
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Additional data, resulting from examination of newly collected material from the Red Sea, east coast of Africa, Arabian Gulf, Pakistan, and western coast of India, and a phylogenetic analysis of the COI barcoding region, confirms Crenidens crenidens (Forsskål) and C. indicus Day as valid species. The latter species was earlier regarded as a subspecies of C. crenidens. In addition, the analyses herein show that specimens from the Red Sea form a distinct monophyletic sub-clade within C. crenidens, characterized by low genetic divergence from specimens from the southwestern Indian Ocean. Close comparison of 34 morphological characters showed that specimens from South Africa and Mozambique differ from Red Sea specimens only in having slightly longer pelvic fins [4.6–4.9 in standard length (SL) vs. 4.8–6.1 in SL]. Examination of additional specimens of both species provided more assessment of inter- and intraspecific variation in meristic and morphometric characters. A new set of characters that help to distinguish C. indicus from C. crenidens is proposed: the former species has a deeper body and caudal peduncle; more scales between fifth dorsal-fin spine and lateral line; scales on top of head extending forward to vertical through posterior margin of pupil; longer pelvic fins; lips with tiny cirri; caudal fin blackish distally; and usually with obvious black spot at pectoral-fin axil. Crenidens indicus was previously reported from central Oman, Arabian Gulf, to Pakistan; herein its presence from the western coast of India, Gujarat and Mumbai (= Bombay), is confirmed. Descriptions of C. crenidens and C. indicus based on material examined, photographs of alive and fresh fishes and an updated key to the three known species of Crenidens are provided.
... Meanwhile, the genus Cheilinus belongs to the family Labridae and contains wrasses (e.g. Cheilinus trilobatus and Cheilinus quinquecinctus) that are native to the Indian and Pacific Oceans and the Red Sea (Bogorodsky et al., 2016). ...
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Cheilinus trilobatus, Cheilinus quinquecinctus, and Chlorurus sordidus specimens from Saudi Arabia's Farasan Islands were collected and genotyped using inter simple sequence repeats (ISSRs) and start codon targeted (SCoT) primers. Mitochondrial cytochrome C oxidase subunit I (COI) gene fragments were used for DNA barcoding. ISSRs and SCoT primers showed moderate polymorphisms: expected heterozygosity (H exp) of 0.470 and 0.435 and average polymorphism information contents (PICs) of 0.359 and 0.339 for ISSRs and SCoT markers were observed, respectively. Cheilinus quinquecinctus had the highest genetic diversity from ISSRs (70%) and SCoT (73%). Chlorurus sordidus and C. trilobatus showed similar genetic diversities of 29% and 39% based on ISSRs, respectively, and 60.32% for both species based on SCoT. Cheilinus quinquecinctus had the lowest nucleotide diversity (π) of 0.003, while C. sordidus and C. trilobatus had π values of 0.065 and 0.103, respectively. Analysis of molecular variance (AMOVA) revealed greater genetic variation among species rather than within them using ISSRs (65% and 67%, respectively) and SCoT (35% and 33%, respectively). COI-based AMOVA showed similar genetic variation among (51.11%) and within (48.89%) species. The current study highlighted outperformance of COI compared to ISSRs and SCoT markers in differentiating among parrotfish species. Also, ISSR outperformed SCoT since it was able to clearly show three distinct groups in principal component analysis. This study also confirmed the presence of three distinct parrotfish species, which will provide an insight into parrotfish diversity. Moreover, the results will contribute to monitoring parrotfish migration between Farasan Islands and different geographic locations which significantly affect species conservation.
... Examples from various groups of fish have been identified to fall into the same pattern, including recent examples from moray eels (e.g., Smith et al. 2019), gobies (e.g., Kovačić et al. 2018), labrids (e.g. Bogorodsky et al. 2016), damselfishes (Randall & DiBattista 2013) and chaetodontids (DiBattista et al. 2018). As more data on the distribution of O. olivaceus and its congener O. lithinus become available, the reconstruction of the evolutionary process that led to formation of these two species might become feasible. ...
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A new species of snake eel Ophichthus olivaceus is described based on two specimens trawled from a depth of 35–63 m from a soft substratum off Jizan, Red Sea coast of southern Saudi Arabia. It differs from its congeners by the following combination of characters: vertebrae 141–145; tail moderately short (2.15 in TL); head short (9.6–11.1 in TL); uniserial teeth in jaws and on vomer; pectoral fins slightly elongate, not lanceolate, upper rays longer than the lower; dorsal-fin origin above middle of pectoral fin; and a generally uniform, dark tan body with an olivaceous hue shading to tan or pale orange ventrally, with two pale yellow blotches above pectoral-fin base, snout and lower jaw dark brown, and olivaceous median fins. Its divergence from other mitochondrial-analyzed species is shown by phylogenetic analysis of the mitochondrial COI barcoding region. A key to the Indian Ocean species is provided.
... and K. hectori is in the range of other Red Sea-Indian Ocean or Red Sea-Indo-West Pacific pairs of sibling species, considering that among species from various higher reef fish taxa the estimated rates of nucleotide substitution in mitochondrial COI varied at least with a factor of 2 (see Lessios 2008). Molecular study of two species of Cheilinus (e.g., C. quinquecinctus and C. fasciatus), a pair of Red Sea-Indo-West Pacific Ocean sibling species from the family Labridae, accounted 24 diagnostic sites in the barcoding portion of mitochondrial COI (Bogorodsky et al. 2016). With only eight diagnostic sites within 652 bp of COI a considerably lower number was found for two divergent evolutionary lineages of C. crenidens (family Sparidae) from the Red Sea and the Western Indian Ocean, respectively (Bogorodsky et al. 2017). ...
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The gobiid genus Koumansetta Whitley, placed in synonymy by some authors with the similar genus Amblygobius Bleeker, is redescribed and its validity based on an integrated morphological and molecular assessment is confirmed. The following characters have been found that distinguish Koumansetta from any of 15 recognized valid species of Amblygobius: oculoscapular transverse rows trp and tra long, extending dorsally well above level of rows x1 and x2; snout pointed, prominent, longer than eye diameter, with gently sloping dorsal profile, overhanging mouth; mouth subterminal; the upper limb of first gill arch with 1–2 slender, weak and soft gill rakers anteriorly, followed by 1–5 short, also soft, broad structures; first two dorsal-fin spines elongate, remaining spines progressively shorter; pelvic frenum absent; body brown to brown-green in upper and lateral sides with narrow yellow or orange longitudinal stripes on body and head, black ocellated spot on the second dorsal fin, and another black spot dorsoposteriorly on caudal peduncle. The following three species are assigned to Koumansetta: K. rainfordi Whitley, the type species of the genus, known from the western Pacific Ocean; K. hectori (Smith), the most widespread species, known from islands of the western Indian Ocean to Micronesia and Fiji; and a new species, restricted to the Red Sea and the inner Gulf of Aden. Koumansetta hoesei sp. nov. has formerly been confused with similar K. hectori, but differs in various details of coloration, and in some morphological characters. Moreover, K. hoesei sp. nov. is evolutionary well divergent from K. rainfordi and K. hectori, its closest relative, as shown by phylogenetic analysis of the mitochondrial COI barcoding region. In addition to the description of the new species, brief species accounts are provided for K. hectori and K. rainfordi, and a key to the three species.
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The Socotra Archipelago, located in the eastern Gulf of Aden, has a unique marine environment which combines tropical and ‘pseudo-temperate’ elements. Studies on the fish biogeography of the archipelago, partially framed in regional studies, have substantially outpaced critical elementary research on the archipelago’s fish diversity. The present study seeks to close this gap and identifies the Socotra Archipelago as a major hotspot of coastal fish diversity in the Indian Ocean. The archipelago supports unique coastal fish assemblages which are predominantly composed of coral-associated (“reef”) species, in spite of the limited biogenic reef frameworks. A Preliminary Checklist comprises 682 species with confirmed records and a “Working List” includes an additional 51 records, totalling 733 faunal records in 108 families. The family Labridae is the most speciose, followed by Gobiidae, Pomacentridae, Serranidae and Chaetodontidae. The species richness of the archipelago is the highest when compared to adjacent Arabian ecoregions. The richness of the Acanthuridae, Chaetodontidae, Labridae, Pomacentridae and Pseudochromidae stand out as particularily high, and the richness of several families is as high as or higher than in the entire Red Sea. The total archipelagic richness is extrapolated at up to 875 species based on incidence-based richness models and expert opinion. Inshore fish inventories, covering 497 species, found between 14 and 132 species per site (x̄ = 66). Site diversity decreased across the archipelago from west to east and from north to south. Total fish diversity was highest around Socotra Island, followed by Abd al-Kuri & Kal Farun and Darsa & Samha. Occurrence frequencies were very unevenly distributed and dominated by Pomacentrus caeruleus and Thalassoma lunare, whilst many species were infrequent. The fish assemblages are dominated by species from the Indo-West Pacific and the north-western Indian Ocean. The assemblages are rich in rare species and hybrids, and include a low number of endemics (4–5), and a high number of species with far-reaching and Western Indian Ocean ranges.
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The Red Sea is characterised by a unique composition of species of fishes which, based on unpublished data of the present authors, currently consists of 1166 species from 159 families whose habitats range from shallow waters to the deep sea. There is a total of 1120 species in coastal waters of the Red Sea recorded within an overall depth range 0–200 m; among them, 165 species are exclusively endemics to the Red Sea, whilst another 51 species are restricted to the Red Sea and Gulf of Aden only, and 22 species living at depths greater than 200 m are endemic. As the westernmost peripheral area of the Indo-West Pacific region, the Red Sea is at the opposite end of the distributions of many widespread coral reef organisms that range to the easternmost regions, such as the Hawaiian Islands, Easter Island, and the Marquesas Islands. It is noted that these areas exhibit high percentages of endemism among coastal fishes. The Hawaiian archipelago has 30.7% of its fishes as endemic species; Easter Island has 21.7%, the Red Sea 14.7% (19.3% when combined with the Gulf of Aden), and the Marquesas Islands have 13.7% endemic fishes. The Red Sea is 2250 km in length and it is very deep, with an average depth of 490 m, and a maximum depth of 3040 m. As expected, the fish fauna is far from homogeneous. The most divergent sector is the Gulf of Aqaba. We have noted that its entrance to the rest of the Red Sea is shallow. It has a maximum width of only 24 km, but a maximum depth of 1850 m. The shore drops off quickly to deep water. The prevailing cross wind creates upwelling, resulting in surface sea temperature at least as low as 21 ℃. Twenty-two of 46 species of Red Sea fishes living at depths greater than 200 m in the Red Sea are endemic (48% endemism). The Gulf of Aqaba has 22 endemic coastal species of fishes and eight endemic deep-dwelling species. By contrast, the neighboring Gulf of Suez, with extensive sand flats and a maximum depth of 70 m, has only seven endemic species of fishes. Of the 165 endemic Red Sea species of fishes, only two are elasmobranchs. Twenty-three families of Red Sea fishes have more than 20% of endemic species with the highest rates of endemism occurring among the Pseudochromidae, Schindleriidae (83.3% and 100% respectively) and the family Gobiidae with the greatest number of endemic species (36 of 139 recorded species). A brief summary of the history of scientific research on Red Sea fishes is provided together with complete lists of endemic species for (i) the entire Red Sea (separately for coastal and deep-dwelling fishes); (ii) the Red Sea combined with the Gulf of Aden; (iii) the Gulf of Aqaba and the Gulf of Suez; and (iv) Lessepsian migrants. Ongoing research is likely to reveal additional endemic species in the region.
The Indo-Pacific labrid fish genus Paracheilinus now contains 20 species. Most of the currently known species inhabit the mega-diverse East Indian region including Paracheilinus angulatus, P. carpenteri, P. cyaneus, P. filamentosus, P. flavianalis, P. lineopunctatus, P. nursalim, P. rennyae, P. togeanensis, P. walton, as well as three recent discoveries described as new species herein. Five species are known from the Red Sea and Indian Ocean, including P. attenuatus (Seychelles and Kenya), P. hemitaeniatus (Madagascar and South Africa), P. mccoskeri (Kenya, Comoro Islands and Arabian Gulf to Andaman Sea), P. octotaenia (Red Sea), and P. piscilineatus (Mauritius). The remaining two species, P. bellae and P. rubricaudalis, are mainly confined to Micronesia/Marshall Islands and PNG/Fiji/Vanuatu, respectively. Members of the genus are typically distinguished on the basis of their caudal-fin and dorsal-fin shapes, the presence or absence of elongate filamentous dorsal-fin rays, and, in particular, the color of terminal-phase (TP) males, including their dramatic nuptial-display patterns. Paracheilinus paineorum n. sp. is described from 8 specimens, 43.1-70.0 mm SL, collected in Indonesia (southwestern Flores, Sulawesi, Nusa Penida, East Borneo, and Seribu Islands) in depths of 10-65 m. It is closely related to the allopatric P. filamentosus and P. xanthocirritus n. sp., differing mainly in coloration (particularly the bright red dorsal-fin markings) and larger maximum size (to at least 70 mm SL). Paracheilinus xanthocirritus n. sp. is described from 12 specimens, 33.9-49.3 mm SL, collected in the South China Sea at the Anambas Islands of Indonesia and Brunei in depths of 15-25 m. In contrast to the closely related P. paineorum n. sp., TP males of this species have a mostly yellow dorsal fin lacking red markings. The two new species further differ from P. filamentosus by having a narrower interorbital and a shorter caudal peduncle. A third new species, Paracheilinus alfiani, n. sp., is described on the basis of two specimens, 48.8 and 49.3 mm SL, from Lembata Island in the Lesser Sunda Islands of Indonesia. It is characterized by a rounded and relatively tall dorsal fin without elongate filamentous rays, a slightly rounded caudal fin, and distinctive TP male coloration. In addition to the new species descriptions, a diagnosis and color illustrations are included for all members of the genus. We also present a key to the species and a neighbor-joining tree of mitochondrial DNA sequences which clarifies the genetic relationships among species, revealing four discrete species complexes within the genus.
With a rather high percentage of endemic littoral fishes, the Red Sea, together with the Gulf of Aden and the Arabian Gulf, proves to be a separate zoogeographic unit (subprovince). Based on the deep-sea ichthyofauna, the Red Sea is recognized as a distinct province, while the Gulf of Aden forms part of the Indian Ocean. In addition to the oscillatory migrations to and from the Gulf of Aden during the glacial and postglacial phases, parts of the southern Red Sea may have acted as a refuge for tropical fishes. The bathyal fishes form a secondary deep-sea ichthyofauna with a number of endemics. As deep-sea inhabitants did not participate in southward migrations during the glacials, they might be indicators of the rather long existence of the Red Sea as a continuous habitat (at least since the last interglacial). -from Author
The Red Sea is characterised by a unique fauna and historical periods of desiccation, hypersalinity and intermittent isolation. The origin and contemporary composition of reef-associated taxa in this region can illuminate biogeographical principles about vicariance and the establishment (or local extirpation) of existing species. Here we aim to: (1) outline the distribution of shallow water fauna between the Red Sea and adjacent regions, (2) explore mechanisms for maintaining these distributions and (3) propose hypotheses to test these mechanisms.