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Redescription of the genus Koumansetta (Teleostei: Gobiidae), with description of a new species from the Red Sea

August 2018
Zootaxa 4459(3):453

DOI:10.11646/zootaxa.4459.3.3

Authors:

Marcelo Kovačić

Natural History Museum Rijeka

Sergey V. Bogorodsky

Station of Naturalists

Ahmad Osman Mal

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

Tilman J. Alpermann

Senckenberg Society for Nature Research

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.

Gill rakers and structures of Koumansetta hoesei sp. nov.: A) gill rakers on lower limb, B) gill rakers on upper limb, C) gill structures on upper limb. Non-type material, SMF 35872, 32.4 + 8.3 mm, Al Lith, Saudi Arabia.

…

. Morphometric characters (as proportional measurements in %) of Koumansetta hoesei sp. nov. and K. hectori. Characters are sorted in alphabetic order.

…

. (Continued)

…

Koumansetta hoesei sp. nov. A: SMF 35723 [KAU14-796], holotype, female, 28.8 + 7.4 mm, freshly collected, Al Lith, Saudi Arabia, Red Sea; B: SMF 35724 [KAU14-797], paratype, male, 24.4 + 6.7 mm, freshly collected, Al Lith, Saudi Arabia, Red Sea; C: SMF 35723 [KAU14-796], holotype, preserved. D: SMF 35723, paratype, preserved. Photos by S.V. Bogorodsky (A & B) and M. Kovačić (C & D). Non-type material. MNHN 1977-0743 (2: 27.0-32.0 mm SL), Djibouti, Gulf of Tadjoura. All other material from Saudi Arabia: KAUMM 445 (2: 20.8 & 27.5 mm SL), Jeddah, Obhur, 3 July 2013, coll. T.J. Alpermann & S.V. Bogorodsky; KAUMM 446 [KAU14-846] (1: 34.5 mm SL), Al Lith, 16 November 2014, coll. T.J. Alpermann & S.V. Bogorodsky; KAUMM 447 (5: 23.5-29.6 mm SL), Al Lith, 16 November 2014, coll. T.J. Alpermann & S.V. Bogorodsky; KAUMM 448 (2: 24.6 & 30.5 mm SL), Al Lith, 19 November 2014, coll. T.J. Alpermann & S.V. Bogorodsky; KAUMM 449 (1: 24.9 mm SL), Al Lith, 5 March 2012, coll. T.J. Alpermann & S.V. Bogorodsky;

…

Cephalic sensory pores system and papillae of Koumansetta hoesei sp. nov. Paratype, PMR VP4048, male, 31.6 + 8.3 mm, Al Lith. Terminology in text. Drawing by M. Kovačić.

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Accepted by W. Holleman: 11 Jul. 2018; published: 16 Aug. 2018

ZOOTAXA

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(online edition)

Zootaxa 4459 (3): 453

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Article

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https://doi.org/10.11646/zootaxa.4459.3.3

http://zoobank.org/urn:lsid:zoobank.org:pub:E764FED3-CFC9-4120-8A72-917D46D1EDED

Redescription of the genus Koumansetta (Teleostei: Gobiidae),

with description of a new species from the Red Sea

MARCELO KOVAČIĆ

1,4

, SERGEY V. BOGORODSKY

2,3

, AHMAD O. MAL

& TILMAN J. ALPERMANN

Prirodoslovni muzej Rijeka, Lorenzov prolaz 1, HR–51000 Rijeka, Croatia. E-mail: marcelo@prirodoslovni.com

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

Abstract

The gobiid genus Koumansetta Whitley, placed in synonymy by some authors with the similar genus Amblygobius Bleek-

er, is redescribed and its validity based on an integrated morphological and molecular assessment is confirmed. The fol-

lowing 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 x

and x

; snout pointed, prom-

inent, 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 as-

signed 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 con-

fused 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 phylo-

genetic 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.

Key words: Indo-West Pacific, endemic fish, phylogenetic analysis, taxonomy

Introduction

The gobiid genus Koumansetta, with its type species K. rainfordi, was described based on a single specimen from

Queensland, Australia, by Whitley (1940), who provided generic and species diagnoses without discussing its

affinities to other gobies. Another monotypic genus, with the type species Seychellea hectori, was described by

Smith (1957) based on specimens from Mahé and Aldabra, Seychelles. Smith (1957) noted that the new genus

differs from closely related Amblygobius by the absence of scales on the head and reduced dentition, but a

comparison with then monotypic Koumansetta (i.e. K. rainfordi) was not included. The gobiid genus Amblygobius

was created and briefly described by Bleeker (1874) for Gobius sphinx (correct name G. s p hy nx ) Valenciennes in

Cuvier & Valenciennes 1837. Characters of Amblygobius were extended to include more species in the short genus

description in Smith (1959), who considered Seychellea as a distinct valid genus. However, Seychellea was later

recognized as a junior synonym of Amblygobius by Hoese & Winterbottom (1979), followed by Masuda et al.

(1980) and Winterbottom & Emery (1986). In all three publications, however, this decision was not commented or

explained, and consequently the only species of the genus, S. hectori, was just placed in Amblygobius by these

authors. Additional characters for Amblygobius were provided by Hoese & Allen (1977) for comparison with their

new genus Signigobius. However, no revision and no redescription of the genus Amblygobius was later undertaken

KOVAČIĆ ET AL.

454

(Allen & Erdmann 2016), and the situation is still unclear, with 14 species divided into two species groups

according to Allen & Erdmann (2016), based on overall appearance and patterns of scalation, or with 15

Amblygobius species recognized by Eschmeyer et al. (2017). No relationship of S. hectori with Koumansetta was

established until it was placed in Koumansetta by Randall (2005), and therefore, according to this author, the genus

Seychellea needed to be considered a junior synonym of Koumansetta. Randall (2005) listed some characters that

distinguish Koumansetta from Amblygobius. Some later authors (e.g. Kottelat 2013) listed both genera

Koumansetta and Seychellea in synonymy of Amblygobius, again a taxonomical decision without any comment or

explanation, as previously for Seychellea. However, Eschmeyer et al. (2017) shared Randall’s opinion that K.

rainfordi Whitley 1940 and K. hectori (Smith 1957) belong to a valid genus Koumansetta, despite confusing data

from the same source on the genus Seychellea, for which the current status is given as a junior synonym of

Amblygobius. Consequentially, the second species originally assigned to Seychellea (i.e. S. tekomaji Smith 1959) is

placed in Amblygobius in Eschmeyer et al. (2017).

In the Red Sea Koumansetta hectori was first reported by Clark et al. (1968) as Seychellea hectori, later Goren

(1979) provided a description of specimens of S. hectori from different localities. The coloration and other

differences between specimens of “K. hectori” from the Red Sea and specimens from the Indo-West Pacific were

not commented by these authors. However, barcoding of the mitochondrial COI gene as well as comparison of

photographs from the entire distribution range of K. hectori indicated that the Red Sea population could represent

an undescribed species. Our investigations detailed herein, show that the new Red Sea species forms a clearly

divergent evolutionary lineage which differs from the true K. hectori by details of coloration and by morphological

characters.

The aim of this paper is to redescribe the genus Koumansetta, providing the distinguishing characters in

comparison with related genera, and to describe a new species of this genus from the Red Sea. We also summarize

our observations on morphological differences and subgeneric groupings within the remaining Amblygobius.

Materials and methods

Morphometric methods. Considering the absence of exact descriptions of methods of measurements in Whitley

(1940) and Smith (1957) we describe the measurements used herein in an attempt to standardize measures at least

for closely related and morphologically similar gobiid genera. Proportional measurements are made point to point

with a micrometer under a binocular microscope to the nearest 0.1 mm. Length of a specimen is presented as

standard length plus caudal-fin length (with exception of non-type material of K. hoesei sp. nov.). Morphometric

data presented are given as ratio to standard length and other suitable lengths in the text and percentages of

standard length in Table 1 for easier comparison with published data (e.g., Allen & Erdmann 2016). Standard

length (SL) is measured from the median anteriormost point of the head to the end of the vertebral column (i.e.

posterior end of the hypural plate). Other measurements in alphabetic order are listed in Table 1: Anal-fin base is

measured between the insertions of the spine of the anal fin and its last ray. Anal-fin spine and ray lengths are

distances from the insertion of the spine/ray to its tip. Body depth and width at anal-fin origin are measured at the

level of the insertion of the first anal-fin spine. Body depth and width at pelvic-fin origin are measured at the level

of the insertion of the pelvic spine. Caudal-fin length is the distance from the base of the fin, i.e. the posterior end

of the hypural plate to the tip of the longest ray of the caudal fin. Caudal peduncle length is the oblique distance

from the fleshy insertion of the last ray of the anal fin to the caudal-fin base i.e. the posterior end of the hypural

plate. Caudal peduncle depth is the least depth of caudal peduncle. Cheek depth is the least distance from lower

margin of eye to the most-posterior fleshy point of the jaws angle and includes the scleral cartilage or ossicle ring

framing the eyes. Eye diameter is the largest orbit diameter excluding the scleral cartilage or ossicle ring around the

eye. First dorsal-fin base is measured between the insertions of the first and last spines of the dorsal fin. First

dorsal-fin spine lengths are distances from the insertion of the spines to their tips. Head depth is the maximum

depth in a vertical line at the mid-eye. Head length is measured from the anteriormost end of the upper lip to the

posterior end of the opercular membrane. Head width is the maximum width adjusting for any excessive flaring of

the operculum. Interorbital width is the minimum width including the scleral cartilage or ossicle ring framing the

eye. Pectoral-fin length is measured from the uppermost base of the pectoral-fin rays to the tip of the longest ray.

Pelvic-fin length is measured from the base of the pelvic spine to the tip of the longest soft ray. Pelvic to anus is the

distance from the base of the pelvic spine to the anterior margin of the anus. Postorbital length is measured from

455

NEW KOUMANSETTA FROM THE RED SEA

the posterior margin of the eyeball to the posterior end of the opercular membrane and includes the ligamentous

ring. Second dorsal-fin base is measured between the insertions of the spine of the second dorsal fin and its last ray.

Second dorsal-fin spine and ray lengths are distances from the insertion of the spine/ray to its tip. Snout length is

measured from the anteriormost end of the upper lip to the anterior margin of the orbit including the ligamentous

ring. Snout to origin of anal fin is the distance from the anteriormost end of the upper lip to the insertion of the

anal-fin spine. Snout to origin of anus is the distance from the anteriormost end of the upper lip to anterior anus

origin. Snout to origin of first dorsal-fin is the distance from the anteriormost end of the upper lip to the insertion of

the first dorsal-fin spine. Snout to origin of second dorsal fin is the distance from the anteriormost end of the upper

lip to the insertion of the second dorsal-fin spine. Snout to origin of pelvic fin is the distance from the anteriormost

end of the upper lip to the insertions of the pelvic-fin spines. Upper jaw length is the distance from the anteriormost

point of the premaxilla to the posteriormost point of the maxilla. Total dorsal bases is measured between the

insertion of the first spine of the first dorsal fin and the insertion of the last ray of the second dorsal fin.

Meristic methods. Scales in lateral series is taken from the upper attachment of the opercular membrane

backward to the caudal-fin base, with separate count of small scales on the caudal-fin base; scales in transverse

series is counted up and back from anal-fin origin to the second dorsal-fin base; circumpeduncular scales were

counted in a vertical zigzag row around the caudal peduncle, immediately anterior to the caudal-fin base; predorsal

scales were counted forward along mid-dorsal from the first dorsal fin to the anteriormost scale; gill rakers were

counted separately on the lower limb and on the upper limb of the first gill arch.

Terminology of the cephalic sensory canals and papillae follow Akihito (1984) and Miller (1986). The material

was stained in 2% solution of Cyanine Blue in distilled water (Saruwatari et al. 1997) for positive confirmation of

data on scales and sensory papillae rows. The type material has been deposited in SMF—Senckenberg Museum,

Frankfurt, KAUMM—King Abdulaziz University Marine Museum and PMR—Natural History Museum, Rijeka,

Croatia. The comparative material was loaned from the collections of BMBP—the Bernice P. Bishop Museum,

Honolulu, ROM—Royal Ontario Museum, Toronto, USNM—National Museum of Natural History, Smithsonian

Institution, Washington, D.C.

Molecular phylogenetics. In order to investigate the evolutionary divergence of Koumansetta from the Red

Sea a maximum likelihood phylogenetic analysis was performed on partial sequences of the mitochondrial COI

gene. Genomic DNA was isolated with a DNeasy tissue kit (Qiagen, Hilden, Germany) from ethanol-preserved

tissues samples of specimens collected in this study or that had generously been provided from other collections

(Table 2). Amplification of the 652 bp barcoding portion 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

protocol in this 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, Auckland, New Zealand). An alignment of COI sequences obtained in this study and sequences

of congeners K. hectori and K. rainfordi and various Amblygobius spp. and as outgroup sequences of Signigobius

biocellatus, Valenciennea puellaris, and Exyrias belissimus retrieved from GenBank was constructed with the same

software (see Table 2 for sequence details). A phylogenetic tree was then inferred by maximum likelihood (ML) in

PhyML 3.0 (Guindon et al. 2010; Guindon & Gascuel 2003) under the TN + G + I 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. Inference of the maximum

likelihood phylogeny was conducted via a software plugin for PhyML in Genious Pro.

Results

Koumansetta Whitley 1940

Koumansetta Whitley 1940: 425, type species Koumansetta rainfordi Whitley 1940, by original designation and monotypy.

Seychellea Smith 1957: 726, type species Seychellea hectori Smith 1957, by original designation and monotypy.

Diagnosis. (1) pelvic fins united to form a complete pointed pelvic disc; (2) cephalic suborbital row a absent; (3)

oculoscapular transverse rows trp and tra long, extending dorsally well above level of rows x

and x

; (4) snout

pointed, prominent, long, 0.7–0.8 in eye diameter, 2.8–3.0 in head length, with gently sloping dorsal profile,

KOVAČIĆ ET AL.

456

overhanging mouth; (5) mouth subterminal; (6) 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 (Fig. 1); (7) first two dorsal-fin spines longest

and elongate, remaining spines progressively shorter; (8) pelvic frenum absent; (9) color mostly brown to brown-

green with narrow yellow or orange longitudinal stripes on body and head and two black spots posteriorly, one

ocellated, partly or completely on the second dorsal fin, and another spot dorsoposteriorly on caudal peduncle; (10)

scales mostly ctenoid, predorsal area scaled at midline.

This diagnosis distinguishes members of the genus Koumansetta from 15 valid species presently assigned to

Amblygobius, and from two other genera of the “Va l e n ci e n n e a -lineage” (Signigobius and Valenciennea) sensu

Agorretta et al. (2013). Characters (1) and (2) distinguishes the genus from Valenciennea, character (2) from

Signigobius. The diagnosis was confirmed to be different compared to all comparative material examined of 13

valid species presently assigned to Amblygobius and compared to published data in the species descriptions of two

additional recently described Amblygobius species (Allen & Erdmann 2016) in characters (3)–(9) (Table 3).

Character (10) distinguishes Koumansetta only from "Amblygobius nocturnus species group” sensu Allen &

Erdmann (2016) and Amblygobius sewardii (Playfair 1867).

FIGURE 1. Gill rakers and structures of Koumansetta hoesei sp. nov.: A) gill rakers on lower limb, B) gill rakers on upper

limb, C) gill structures on upper limb. Non-type material, SMF 35872, 32.4 + 8.3 mm, Al Lith, Saudi Arabia.

Description. Body moderately elongate, laterally compressed. Depth at anal-fin origin 4.8 (4.5–5.2) in SL.

Head slightly laterally compressed. Snout pointed, prominent, long, 0.7–0.8 in eye diameter, 2.8–3.0 in head

length, with gently sloping dorsal profile, overhanging mouth. Eyes lateral, interorbital broad. Anterior nostril

short, tubular, erect, without process from rim; posterior nostril slit-like, with a low, raised ridge, closer to anterior

nostril than to anterior margin of eye. Mouth subterminal, moderately small, oblique. Posterior angle of jaws below

anterior edge of eye or slightly in front or behind the vertical. Teeth in upper and lower jaws unicuspid and pointed,

more or less curved inward. Upper jaw with caniniform teeth of similar size at the front, followed with inner row of

smaller slender sharp teeth laterally; lower jaw with an outer row of caniniform teeth of similar size at the front

ending laterally with single large curved canine and the inner row of smaller slender sharp teeth. Tongue reduced,

adnate, truncate in front. Chin without fold or barbels. Cranial roof covered by dorsal axial musculature. Gill

openings lateral, ending forward at pectoral-fin base. Branchiostegal membranes attached to entire lateral margin

of isthmus. Gill rakers poorly developed, rakers on lower limb of anterior arch slender, weak and soft, 6–7 in

457

NEW KOUMANSETTA FROM THE RED SEA

number, 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 (Fig. 1). Two dorsal fins, first fin with VI spines, first two dorsal-fin spines

elongate, remaining spines progressively shorter, second fin I/15–17; anal fin I/15–17. Pectoral-fin rays 15–18.

Pectoral girdle without flaps on anterior edge. Uppermost rays of pectoral fin within membrane. Pelvic fin united to

form a complete pelvic disc, pointed. Pelvic frenum absent. Caudal fin rounded, shorter than head length, with 13

branched and 16–17 segmented rays. Body covered with small, mainly ctenoid scales, 49–61 in lateral series;

transverse scale series 14–19. Predorsal area, including nape, breast and belly with cycloid scales, the area between

first dorsal-fin base and pectoral fin also with cycloid scales. No scales on cheek and opercle (a few small cycloid

scales occasionally present on upper opercle). Predorsal area scaled, scales not reaching the vertical through the

posterior edge of eye, predorsal scales 16–26. Head with anterior oculoscapular canal with pores B’, C (single), D

(single), E, F, G, and H’; posterior oculoscapular canal with K’ and L’; preopercular canal with M’, N, and O’.

Suborbital rows of sensory papillae: row a absent, row b longitudinal, short, beginning anteriorly below posterior

part of eye, not reaching above row d, row c as 10–16 transverse sub-orbital rows of sensory papillae, row d

longitudinal. Oculoscapular transverse rows trp and tra long, extending dorsally well above level of rows x. No

interorbital papillae. Head and body brown, reddish brown to greenish brown or brownish to greenish grey with

narrow yellow or orange longitudinal stripes, black ocellus entirely on or partially on the second dorsal fin, and

black spot dorsoposteriorly on caudal peduncle.

Key to the species of Koumansetta

1a First dorsal fin without ocellus; head and body with narrow orange longitudinal stripes; longitudinal scale series 55–61. . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. rainfordi

1b First dorsal fin with distinct black ocellus; head and body with narrow yellow longitudinal stripes; longitudinal scale series 49–

54. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2a Pectoral-fin rays 15; middorsal stripe and the dorsolateral pair of stripes going from snout in U shape backwards, each splitting

into double stripe at the level of eyes and converging again above posterior margin of opercle on predorsal area; lowermost

stripe on body ending below middle of second dorsal fin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. hoesei sp . nov.

2b Pectoral-fin rays 16–17; middorsal stripe and the lateral pair of stripes going from snout in U shape backwards retain single,

not diverging and doubling at the level of eyes; lowermost stripe on body extending to posterior caudal peduncle. . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . K. hectori.

Koumansetta hoesei Kovačić, Bogorodsky & Alpermann, sp. nov.

Hoese’s goby

Figs. 2–4, Table 1

Seychellea hectori (non Smith)—Clark et al. 1968: 22 (Dahlak Archipelago, listed); Goren 1979: 51 (Eilat, Wasset, Nocra;

description); Dor 1984: 250 (Red Sea, listed).

Amblygobius hectori (non Smith)—Randall 1983: 163 (Red Sea); Goren & Dor 1994: 62 (Red Sea, listed); Khalaf & Disi 1997:

185 (Jordan, description); Debelius 1998: 179 (Egypt and Sudan, photographs); Khalaf 2004: 46 (Jordan, listed).

Koumansetta hectori (non Smith)—Lieske & Myers 2004: 192 (Abu Galum, photograph); Golani & Bogorodsky 2010: 48

(Red Sea, listed).

Holotype. SMF 35723 [KAU14-796], female, 28.8 + 7.4 mm, Red Sea, Saudi Arabia, Al Lith, 15 November 2014,

coll. T.J. Alpermann & S.V. Bogorodsky.

Paratypes (all from the Red Sea, Saudi Arabia). KAUMM 442 [KAU11-224], male, 23.2 + 5.9 mm, Al Lith,

29 March 2011, coll. T.J. Alpermann & S.V. Bogorodsky; KAUMM 443 [KAU13-493], female, 38.5 + 10.5 mm,

Al Wajh, 16 June 2013, coll. T.J. Alpermann & S.V. Bogorodsky; KAUMM 444 [KAU11-516], female, 35.2 + 8.3

mm, Rabigh-Masturah, 7 April 2011, coll. S.V. Bogorodsky; PMR VP4048, male, 31.6 + 8.3 mm, Al Lith, 17

November 2014, coll. T.J. Alpermann & S.V. Bogorodsky; PMR VP4049, male, 40.8 + 10.3 mm, Maqna, 13 April

2011, coll. S.V. Bogorodsky; SMF 35707, female, 25.9 + 6.7 mm, Farasan Archipelago, 26 February 2012, coll.

T.J. Alpermann & S.V. Bogorodsky; SMF 35724 [KAU14-797], male, 24.4 + 6.7 mm, Al Lith, 15 November 2014,

coll. T.J. Alpermann & S.V. Bogorodsky; SMF 35867 [KAU12-845], female, 33.7 + 8.0 mm, Al Lith, 5 March

2012, coll. T.J. Alpermann & S.V. Bogorodsky.

KOVAČIĆ ET AL.

458

FIGURE 2. Koumansetta hoesei sp. nov. A: SMF 35723 [KAU14-796], holotype, female, 28.8 + 7.4 mm, freshly collected, Al

Lith, Saudi Arabia, Red Sea; B: SMF 35724 [KAU14-797], paratype, male, 24.4 + 6.7 mm, freshly collected, Al Lith, Saudi

Arabia, Red Sea; C: SMF 35723 [KAU14-796], holotype, preserved. D: SMF 35723, paratype, preserved. Photos by S.V.

Bogorodsky (A & B) and M. Kovačić (C & D).

Non-type material. MNHN 1977-0743 (2: 27.0–32.0 mm SL), Djibouti, Gulf of Tadjoura. All other material

from Saudi Arabia: KAUMM 445 (2: 20.8 & 27.5 mm SL), Jeddah, Obhur, 3 July 2013, coll. T.J. Alpermann &

S.V. Bogorodsky; KAUMM 446 [KAU14-846] (1: 34.5 mm SL), Al Lith, 16 November 2014, coll. T.J. Alpermann

& S.V. Bogorodsky; KAUMM 447 (5: 23.5–29.6 mm SL), Al Lith, 16 November 2014, coll. T.J. Alpermann &

S.V. Bogorodsky; KAUMM 448 (2: 24.6 & 30.5 mm SL), Al Lith, 19 November 2014, coll. T.J. Alpermann & S.V.

Bogorodsky; KAUMM 449 (1: 24.9 mm SL), Al Lith, 5 March 2012, coll. T.J. Alpermann & S.V. Bogorodsky;

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NEW KOUMANSETTA FROM THE RED SEA

KAUMM 450 (2: 25.5 & 30.0 mm SL), Al Lith, 7 March 2012, coll. T.J. Alpermann & S.V. Bogorodsky; PMR

VP4050 (4: 24.1–27.5 mm SL), Farasan Archipelago, 28 February 2012, coll. T.J. Alpermann & S.V. Bogorodsky;

SMF 35868 (1: 32.7 mm SL), Farasan Archipelago, 21 February 2012, coll. T.J. Alpermann & S.V. Bogorodsky;

SMF 35869 [KAU13-494] (1: 35.1 mm SL), Al Wajh, 16 June 2013, coll. T.J. Alpermann & S.V. Bogorodsky;

SMF 35870 [KAU13-495] (1: 19.9 mm SL), Al Wajh, 16 June 2013, coll. T.J. Alpermann & S.V. Bogorodsky;

SMF 35871 [KAU13-580] (1: 36.8 mm SL), Al Wajh, 20 June 2013, coll. T.J. Alpermann & S.V. Bogorodsky;

SMF 35872 (13: 18.6–33.0 mm SL), Al Lith, 18 November 2014, coll. T.J. Alpermann & S.V. Bogorodsky.

Diagnosis. Dorsal-fin rays VI + I,15–16; anal-fin rays I,15; pectoral-fin rays 15; longitudinal scale series 50–

54; scales in transverse series 14–17; predorsal scales 19–23; cheek and opercle naked; scales cover the basal 1/4–

1/6 of the caudal fin, becoming rapidly smaller and cycloid; prepectoral area with about 3–6 vertical series of fine

cycloid scales covering the entire base of the pectoral fin; prepelvic area with cycloid scales, 9–11 in the midventral

row; interorbital 90–100% of eye diameter; width of body at anal-fin origin 10.8–12.4% of SL; caudal peduncle

depth 64.6–73.7% of its length; head depth 55.2–60.5% of head length; length of 11th second dorsal-fin ray 9.9–

11.5% of SL; length of 8th anal-fin ray 10.2–13.5% of SL; longitudinal yellow stripes present only on upper half of

body; mid-dorsal stripe and the pair of dorsolateral stripes diverging before eyes and then converging halfway

between eye and first dorsal fin, additional yellow stripe between mid-dorsal and dorsolateral stripes extending

from side of nape continuing along dorsal-fin bases and almost reaching, with interruptions, to ocellus on back;

head brown above and whitish below yellowish white demarcation stripe, starting anteriorly at angle of jaw and

going back to middle of pectoral-fin base; another yellowish white stripe beginning from the posterior part of upper

jaw across upper cheek, opercle, and upper pectoral-fin base becoming a demarcation stripe behind pectoral fin,

dividing the brown upper part of body from the grey lower part, and disappearing below middle of second dorsal

fin; ocellus in first dorsal fin dark red or red-brown with black center and edged with black; weak orange-red streak

is visible above ocellus.

Description (morphometric values in the text are presented first for the holotype followed by ranges for

paratypes; meristic values, if variable, the same). Body moderately elongate, compressed laterally, the depth at

pelvic-fin origin 4.8 (4.5–5.0) in SL, width at pelvic-fin origin 6.7 (6.7–7.5) in SL, depth at anal-fin origin 4.8 (4.5–

5.2) in SL, width at anal-fin origin 9.0 (8.1–9.3) in SL. Head length 3.3 (3.3–3.5) in SL, head slightly laterally

compressed (head width 1.9 (1.7–2.0) in head length, head depth 1.8 (1.7–1.8) in head length). Predorsal profile

gently convex. Snout with moderately sloping profile, overhanging mouth and somewhat pointed from dorsal view,

longer than eye diameter: snout length 0.7 (0.7–0.8) in eye diameter, 2.8 (2.8–3.0) in head length, 9.3 (9.3–10.7) in

SL. Anterior nostril tubular, short, lacking process from rim, halfway between anteriormost margin of snout and

anterior margin of eye. Posterior nostril slit- like, with a low, raised ridge, closer to anterior nostril than to anterior

margin of eye. Eyes lateral, with the upper edge at a level of dorsal profile of head, eye diameter 4.0 (3.6–4.4) in

head length, 13.1 (12.2–15.4) in SL. Interorbital wide, equal or narrower than eye diameter (width 1.1 (1.0–1.1) in

eye diameter, 14.4 (12.9–15.4) in SL). Mouth subterminal, moderately small, oblique. Rear margin of upper jaw

ending posteriorly slightly before or at vertical through anterior margin of eye, upper-jaw length 3.3 (3.1–3.3) in

head length, 11.1 (10.4–11.7) in SL. Each side of upper jaw with an outer row of 3–5 frontal, backwards and

inwards curved, caniniform teeth of similar size, and inner row of 7–12 smaller slender sharp teeth laterally. In one

large specimen (KAUMM 443), short intermediate row of five small teeth is visible laterally on each side. Each

side at front of lower jaw with an outer row of 3–4 caniniform teeth of similar size but less curved than those in

upper jaw, ending laterally with single large curved canine and the inner row of 10–15 smaller slender sharp teeth

laterally. Tongue reduced, adnate, truncate in front. Chin lacking barbels. Branchiostegals broadly fused with

isthmus, the gill openings lateral, ending forward at pectoral-fin base, little if at all wider than pectoral-fin base.

Gill rakers poorly developed, gill rakers on lower limb of first gill arch slender and soft, 6–7 in number, the upper

limb of first gill arch with 2 slender, weak and soft gill rakers anteriorly, followed by 4–5 short, also soft, broad

structures (Fig. 1). No spines on preopercle. No dermal crest anterior to first dorsal fin. Caudal-peduncle depth 8.0

(7.7–8.4) in SL, caudal-peduncle length 5.8 (5.4–6.1) in SL, caudal-peduncle depth in its length 1.4 (1.4–1.5).

Fins: Two separate dorsal fins, VI + I,15 (15: 5, 16: 4); anal-fin rays I,15; pectoral-fin rays 15; branched

caudal-fin rays 13, segmented 16–17 (16: 5, 17: 4); pelvic-fin rays I/5 + 5/I. The first dorsal-fin base 8.5 (7.9–9.9)

in SL; the second dorsal-fin base 3.8 (3.4–3.8) in SL; anal-fin base 4.2 (3.7–4.2) in SL. First two dorsal-fin spines

elongate (in holotype the second spine was broken) but not filamentous, remaining spines progressively shorter, the

sixth reaches the origin of the second dorsal fin. The first spine of the first dorsal fin 4.1 (3.2–4.1) in SL; second

KOVAČIĆ ET AL.

460

spine broken in holotype (3.5–4.5) in SL, third spine 6.9 (5.5–6.9) in SL; fourth spine 9.3 (7.3–9.3) in SL; fifth

spine 9.3 (8.2–9.6) in SL; sixth spine 12.0 (10.0–13.2) in SL. Dorsal fins connected by a low membrane. The first

dorsal-fin origin clearly behind a vertical at pectoral-fin base. The second dorsal-fin origin above anus. Soft rays of

the second dorsal fin progressively shorter to the

twelfth ray, or third to twelfth rays subequal in length, with last

three rays abruptly longer, and forming a lobe. In some specimens, the second soft ray is longer than the first ray in

the second dorsal fin. Origin of anal fin below the second dorsal-fin soft ray. Second anal-fin ray longer than first

ray, then rays progressively shorter to twelfth ray, the four last rays elongate with the last ray longer than the second

soft ray. Both soft anal and dorsal fins when depressed reaching only to the first caudal-fin rays. Pectoral fin

broadly rounded; extending posteriorly to below origin of the second dorsal fin, and only in one male reaching also

a vertical through origin of anal fin, the length 3.5 (3.5–3.8) in SL; uppermost and lowermost pectoral-fin rays not

branched; upper rays within membrane. Pelvic fins united, pointed, with fifth soft ray longer than other rays; 4.0

(3.3–4.0) in SL; not reaching anus in females, reaching anus in males or even anal fin in some of them; pelvic

frenum absent. Pectoral and pelvic fins longer than postorbital part of head. Caudal fin rounded, shorter than head,

1.2 (1.1–1.2) in head length, 3.9 (3.6–4.2) in SL.

Scales: body covered with small ctenoid scales, cycloid at axil of pectoral fin, nape, entire belly, and in the area

between first dorsal fin and pectoral fin; longitudinal scale series 50–54 (holotype 52, paratypes 50: 4, 51: 4, 52: 1,

54: 2; one side of 6 specimens too damaged for count due to tissue sampling for DNA) with 3–5 more scales on

caudal-fin origin, transverse scale series 14–17 (holotype 15, paratypes 14: 7, 15: 2, 16: 3, 17: 2; side of 3

specimens too damaged for count due to tissue sampling for DNA), circumpeduncular scales 17–20 (holotype 19,

paratypes 17: 2, 19: 5, 20: 1). Scales cover a basal 1/4–1/6 of caudal fin, becoming rapidly smaller and cycloid.

Other fins unscaled. Predorsal area scaled, scales not reaching to the vertical through the posterior edge of eye,

ending at the level of pore G, interorbital area naked, predorsal scales in median row 19–23 (holotype 19, paratypes

19: 2, 20: 3, 21: 1, 22: 2, 23: 3). Cheek and opercle naked. Prepectoral area with about 3–6 vertical series of fine

cycloid scales covering the entire base of pectoral fin (holotype 4, paratypes 3: 2, 4: 3, 5: 2, 6: 1). Prepelvic area

scaled with cycloid scales, 9–11 in the midventral row (holotype 10, paratypes 9: 2, 10: 5, 11: 1).

FIGURE 3. Cephalic sensory pores system and papillae of Koumansetta hoesei sp. nov. Paratype, PMR VP4048, male, 31.6 +

8.3 mm, Al Lith. Terminology in text. Drawing by M. Kovačić.

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NEW KOUMANSETTA FROM THE RED SEA

FIGURE 4. Koumansetta hoesei sp. nov., underwater photographs. A: Al Wajh, Saudi Arabia; B: Lahami Bay, southern Egypt;

C: Farasan Archipelago, Saudi Arabia. Photos by S.V. Bogorodsky.

KOVAČIĆ ET AL.

462

Cephalic sensory pores system and papillae (Fig. 3): Anterior oculoscapular canal with pores B’, C (single), D

(single), E, F, G, and H’; posterior oculoscapular canal with K’ and L’; preopercular canal with M’, N, and O’.

Rows of head sensory papillae were counted in all specimens, but some rows were visible only in part of specimens

despite staining and the lowest count in ranges could be underestimated due to miscount of non-visible papillae.

Preorbital: upper row r

longitudinal and long (9–16); upper row s

(3–8) transverse at posterior nostril, middle row

(3–8) transverse at anterior nostril, lower row s

(3) above upper lip and not visible in all specimens. Lateral

series c in four parts: superior c

2–9 papillae below posterior nostril; middle superior c

1–4 papillae below

anterior nostril; middle inferior row c

(3–5) and inferior row c

(4–5) above upper lip. Suborbital rows: row a

absent. Row b (6–8) longitudinal, beginning anteriorly below posterior part of eye i.e. between pupil and posterior

edge of eye. Row c as 10–12 transversal rows below eye, 8–10 in front of row b, 1–3 papillae above level of row b,

0–1 below (1: 4–7, 2: 2–7, 3: 1–6 , 4: 2–5, 5: 1–5, 6: 2–4, 7: 1–7, 8: 3–5, 9: 1–10, 10: 1–9, 11: 0–4, 12: 0–5). Row

d continuous (17–20) or divided in supralabial and posterior part (3 + 11 to 5 + 12), ending posteriorly below pupil.

Preoperculo-mandibular rows: external row e (8 + 11 to 23 + 23); and internal row i (10 + 13 to 16 + 15) divided

into anterior and posterior sections; mental row f longitudinal (4–6), not visible in some material. Oculoscapular

rows: anterior longitudinal row x

(8 + 5 to 12 + 8) divided into part above pores F to H’ and part behind above H’;

posterior longitudinal row x

(5–9) above pores K’ to L’; row z (4–10) transverse behind pore M’; row tra (8–22)

transverse, dividing row x

above H’; row trp (7–19) transverse, dividing rows x

and x

; row q transverse row (3–8)

behind pore H’ and below posterior part of row x

; transverse row y (4–8) behind pore L’; transverse axillary rows

(5–17), as

(5–16), as

(6–18) visible in some specimens; axillary rows la

(2–5) and la

(2–5) visible in some

specimens. Opercular rows: transverse row ot (15–24); superior longitudinal row os (3–10); and inferior

longitudinal row oi (7–12). Anterior dorsal rows: anterior transverse row n (10–23) long and curved, from the level

of center of eye to dorsal midline; transverse row o (6–9); longitudinal row g (6–15) anteriorly reaching near row o;

longitudinal row m (4–5) below and behind row g; longitudinal row h (5 + 3 to 8 + 4) divided; rows o, g, m, h

visible only in some specimens.

Color in life (Fig. 4). Upper two-thirds of head brown, with reddish hue anteriorly, above demarcation stripe.

Demarcation stripe whitish or yellowish, black along upper edge, extending from angle of mouth to the middle of

pectoral-fin base. Head whitish below the stripe. Another yellowish white stripe, black along lower edge,

beginning from the posterior part of upper jaw across upper cheek, opercle, and upper pectoral-fin base becoming a

demarcation stripe behind pectoral-fin and disappearing below middle of the second dorsal fin. Upper half of body

greenish brown to dark brown above this stripe; lower half of body below this demarcation stripe grey to dark grey,

shading to silvery whitish ventrally, sometimes with bluish hue on abdomen. A large black ocellus, rimmed by a

semicircular white mark below, on back below middle of the second dorsal fin, the spot extending into fin where

rimmed above in yellow; the black spot on back appears an open eye, sometimes with dark red anteriorly and

posteriorly; another black spot larger than pupil dorsally at posterior end of caudal peduncle, rimmed below in pale

yellow. Head with mid-dorsal stripe extending from the tip of snout diverging and then converging on interorbital

area and nearly reaching origin of the first dorsal fin. Three narrow longitudinal, black-edged, yellow dorsolateral

stripes on each side of head and body between mid-dorsal and demarcation stripes. First stripe, beginning from side

of nape above posterior margin of preopercle, running posteriorly along dorsal fin bases and ending close to ocellus

at the second dorsal fin. The second stripe on left and right body sides connected in an U-shape at the front of snout

and splitting into two stripes at top of snout, one through upper part of the eye and another across interorbital area,

stripes converging at vertical through posterior margin of opercle and continuing backwards to black ocellus, and

behind it continuing, more or less visible, along the top of caudal peduncle, where it ends at another black spot. The

third stripe extending from the upper lip through the lower part of eye and the upper part of the opercle, running

above lateral midline to caudal peduncle where it is usually broken into a short segment at middle of caudal-fin

base. First dorsal fin with a reddish brown band, rimmed above in yellow, along base; a black-edged dark red or

red-brown, with black center and broadly rimmed with yellow, ocellus over first three spines and adjacent third

membrane at midlevel of fin; an indistinct narrow reddish brown stripe on membranes between fourth and sixth

spines; and an orange-red streak in outer half of first membrane above ocellus. The second dorsal fin with a reddish

brown band rimmed above in yellow, circumflexing the black ocellus, along base; narrow white margin and an

indistinct reddish brown submarginal band, more obvious on posterior half; an indistinct narrow reddish brown

stripe in the middle of anterior half of fin; and yellow rear edge. Anal fin with transparent membranes anteriorly

becoming pale grey posteriorly, with narrow pale bluish white margin on posterior half and rear edge. Caudal fin

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NEW KOUMANSETTA FROM THE RED SEA

with curved white bar at base, rest of fin with transparent membranes. Pectoral and pelvic fins with translucent

membranes.

Color when fresh (Fig. 2A & B). Similar to color of alive gobies but ground color more reddish, posterior end

of caudal peduncle red-brown, reddish brown stripes in dorsal fins more obvious, and melanophores visible on

caudal and anal fins, the rest of these fins transparent.

Color preserved (Fig. 2C & D). Upper two-thirds of head brown, head whitish ventrally; upper half of body

brown; lower half of body below demarcation stripe buff (light brownish yellow). A large black spot (ocellus)

spread on back and on fin at middle of the second dorsal fin rimmed by a semicircular buff mark below and less

conspicuous pale edge above on fin; another black spot larger than pupil dorsally at posterior end of caudal

peduncle, pale rimmed below. Head with mid-dorsal whitish to buff stripe from the tip of snout diverging and then

converging on interorbital area nearly reaching origin of the first dorsal fin. The three dorsolateral stripes and the

demarcation line characteristic for alive fish also present in preserved material, only whitish to buff and dark

edged. First dorsal fin with a brown band along base; a black-edged ocellus with black center, rimmed whitish,

over first three spines and adjacent third membrane at midlevel of fin. The second dorsal fin with a dark band

passing along base, extending above and circumflexing whitish rimmed black ocellus; pigmented submarginal

band more obvious on posterior half. Anal fin also dark pigmented. Caudal fin poorly pigmented. Pectoral fins not

pigmented. Pelvic fin with small scattered melanophores, less dense in females.

Etymology. The specific epithet is in honor of Douglass F. Hoese, in recognition of his important contributions

to the systematics of gobioid fishes.

Distribution and habitat. Based on phylogenetic analysis, examined specimens and numerous photographs it

is revealed that K. hoesei sp. nov. is restricted to the Red Sea and the inner Gulf of Aden (Gulf of Tadjoura). A

coral-reef species usually seen hovering a short distance above the substratum at depths of 3–26 m in lagoons and

seaward reefs, usually solitary, but sometimes seen in a small group, each individual in a moderate distance from

others. Winterbottom & Emery (1986) found filamentous algae, harpacticoid copepods, amphipods, ostracods, and

nematodes in the gut of one specimen of Koumansetta hectori, and we expect a similar benthic, omnivorous diet in

K. hoesei sp. nov.

Remarks. Hitherto, Koumansetta hoesei sp. nov. has been misidentified as K. hectori. However, the new

species clearly differs from the latter in details of coloration and morphology: mid-dorsal stripe and the pair of

dorsolateral stripes diverging before eyes and then converging halfway distance between eye and first dorsal fin in

K. hoesei sp. nov. versus mid-dorsal stripe and both dorsolateral stripes single, not diverging in K. hectori.

Koumansetta hoesei sp. nov. has an additional yellow stripe between mid-dorsal and two dorsolateral stripes

extending from side of nape, continuing along dorsal-fin bases and disappearing before ocellus on back; in

contrast, K. hectori lacks such stripe. Lower head with two yellowish white stripes, lower stripe beginning from the

upper jaw and ending at the middle of pectoral-fin base, another stripe beginning from the lower edge of eye

running across opercle and upper pectoral-fin axil and continuing as demarcation stripe on body disappearing

below middle of the second dorsal fin in K. hoesei sp. nov. whereas only single demarcation stripe starts anteriorly

at angle of jaws going back over lower cheek, lower opercle and lower pectoral-fin base backwards to caudal

peduncle and caudal-fin base in K. hectori. Color of ocellus in first dorsal fin also differs, the ocellus is dark red or

red-brown with black center and edged with black in K. hoesei sp. nov., but ocellus is a black round spot in K.

hectori. In K. hoesei sp. nov. a weak orange-red streak is visible above ocellus versus up to three well defined red

marks above ocellus, one above the other, in K. hectori. Membranes of at least anterior half of anal fin semi-

translucent in K. hoesei sp. nov. versus most anal-fin membranes grey to dark grey in K. hectori. Koumansetta

hoesei sp. nov. can be distinguished from K. hectori also by morphological characters: pectoral-fin rays 15 versus

pectoral-fin rays 16–17; interorbital equal to eye diameter or to 1/10 smaller than eye diameter versus interorbital

2/3 to 9/10 of eye diameter; scales in transverse series 14–17 versus scales in transverse series 17–18; width at

anal-fin origin 10.8–12.4% of SL versus 9.9–10.7%; caudal-peduncle depth 64.6–73.7% of its length versus 75.4–

88.9%; head depth 55.2–60.5% of head length versus 47.3–54.4%; length of 11th second dorsal-fin ray 9.9–11.5%

of SL versus 11.8–14.2%; length of 8th anal-fin ray 10.2–13.5% of SL versus 13.7–14.9%. In addition to non-

overlapping morphometrics, some morphometric characters (anal-fin base, caudal peduncle length, pectoral-fin

length, second dorsal-fin base, snout to origin of anus, total dorsal bases) are rarely overlapping between two

species and can also be used as distinguishing character for most specimens (Table 1).

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TABLE 1 . Morphometric characters (as proportional measurements in %) of Koumansetta hoesei sp. nov. and K.

hectori. Characters are sorted in alphabetic order.

Species K. hoesei sp. nov. K. hectori

Specimen Holotype Paratypes Paratypes Specimens in

comparative

material

Specimens in

comparative

material

Sex Female Females Males Females Males

Number of specimens 1 3 5 2 5

Standard length (SL) in mm 28.8 33.7–38.5 23.2–40.8 26.5–32.9 32.3–39.3

% of standard length:

Anal-fin base 23.6 23.7–26.8 24.2–26.7 26.4–28.3 26.6–27.2

Anal-fin spine length 9.4 7.1–8.6 7.3–9.6 8.3–8.5 8.2–9.7

Anal-fin 1st ray length 13.5 11.4–13.4 12.7–13.9 12.8–13.1 13.0–14.0

Anal-fin 2nd ray length 14.9 14.5–14.8 14.8–17.2 15.5–18.5 15.0–15.9

Anal-fin 8th ray length 12.5 10.2–12.2 11.6–13.5 13.7–14.0 13.7–14.9

Anal-fin last ray length 16.0 15.4–17.1 13.9–18.6 14.0–14.7 14.3–17.0

Body depth at anal-fin origin 20.8 20.8–21.6 19.4–22.3 21.1–21.6 20.1–21.6

Body depth at pelvic-fin origin 20.8 20.2–21.6 20.3–22.3 21.5–23.1 20.9–22.6

Body width at anal-fin origin 11.1 10.9–11.3 10.8–12.4 10.0–10.2 9.9–10.7

Body width at pelvic-fin origin 14.9 13.4–13.6 13.5–15.0 12.8–14.0 11.4–12.7

Caudal fin length 25.7 23.6–27.0 25.2–27.5 24.9–25.2 24.5–26.3

Caudal peduncle depth 12.5 11.9–12.7 11.9–13.0 12.1–12.2 12.1–12.7

Caudal peduncle length 17.4 17.9–18.5 16.4–18.6 13.7–15.8 14.8–16.5

Eye diameter 7.6 6.5–7.4 6.9–8.2 7.0–7.5 6.9–7.4

First dorsal-fin base 11.8 10.1–10.4 10.8–12.7 10.9–13.4 12.6–14.5

First dorsal-fin1st spine length 24.7 26.7–31.2 24.2–30.9 24.9–29.8 27.5–38.7

First dorsal-fin2nd spine length damaged 22.3–24.9 22.0–28.2 22.6–25.2 23.6–31.9

First dorsal-fin3rd spine length 14.6 14.5–18.2 15.4–17.9 13.6–15.8 14.3–23.3

First dorsal-fin4th spine length 10.8 11.7–13.1 12.3–13.8 11.2–11.3 10.4–16.1

First dorsal-fin5th spine length 10.8 10.4–12.2 10.4–12.0 9.4–9.7 8.8–12.2

First dorsal-fin 6th spine length 8.3 7.7–9.6 7.6–10.0 6.7–7.2 8.0–10.4

Head length 30.2 28.2–28.6 28.2–30.6 28.9–30.2 27.5–29.1

Head width 16.3 14.8–16.9 15.9–17.0 15.5–17.0 13.7–16.6

Interorbital width 6.9 6.5–7.4 6.6–7.8 6.4–7.0 6.0–6.6

Pectoral-fin length 28.8 26.7–28.7 26.6–28.9 25.5–26.4 23.8–26.9

Pelvic-fin length 25.0 25.0–26.8 25.1–30.4 21.5–22.5 24.5–29.5

Pelvic to anus 28.1 25.0–28.8 24.7–28.0 24.5–25.2 22.3–26.0

Postorbital length 14.2 13.0–14.2 13.0–14.7 14.6–14.7 12.9–13.6

Second dorsal-fin base 26.4 26.4–28.6 26.6–29.4 28.3–33.4 29.5–31.4

Second dorsal-fin spine length 11.1 9.7–11.2 9.8–12.1 12.1–12.8 11.5–13.3

Second dorsal-fin 1st ray length 14.9 13.8–14.5 14.9–17.2 14.3–15.5 13.7–16.1

Second dorsal-fin 2nd ray length 13.9 13.6–15.6 13.9–16.2 16.1–16.2 14.6–15.8

Second dorsal-fin 3rd ray length 13.5 13.4–15.1 12.0–15.4 15.5–16.1 14.0–15.3

Second dorsal-fin 6th ray length 11.5 11.1–14.5 10.8–13.9 14.3–14.6 12.1–14.0

Second dorsal-fin 11th ray length 11.1 9.9–10.9 10.0–11.5 12.8–13.2 11.8–14.2

Second dorsal-fin last ray length 14.9 14.8–18.4 15.4–18.6 15.1–15.2 14.0–18.1

Snout length 10.8 9.4–9.8 9.5–10.7 9.1–9.4 8.5–9.6

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Koumansetta hoesei sp. nov. can easily be distinguished from another congener K. rainfordi by coloration;

most distinctive characters are: stripes on head and body yellow versus stripes orange to reddish in K. rainfordi;

longitudinal stripes present only on upper half of body versus longitudinal stripes present also on ventral side of

body in K. rainfordi; dorsolateral stripes anteriorly diverging and converging again versus dorsolateral stripes not

converging and diverging again in K. rainfordi; color of head reddish-brown to brown versus color of head light

green in K. rainfordi; no white spots on back just below both dorsal fins versus several white spots on back just

below both dorsal fins in K. rainfordi; first dorsal fin with distinct black ocellus versus no black ocellus on the first

dorsal fin in K. rainfordi; black ocellus on back and on the second dorsal fin at its middle versus black spot on the

outer half of the second dorsal fin in K. rainfordi. Koumansetta hoesei sp. nov. can be distinguished from K.

rainfordi also by morphological characters: pectoral-fin rays 15 versus pectoral-fin rays 16–18 in K. rainfordi;

longitudinal scale series 50–54 versus longitudinal scale series 55–61 in K. rainfordi; scales in transverse series

14–17 versus transverse series 17–19 in K. rainfordi; predorsal scales 19–23 versus predorsal scales 24–26 in K.

rainfordi.

The distribution of Koumansetta hoesei sp. nov. is sympatric with four species of Amblygobius (e.g., A.

albimaculatus (Rüppell 1830), A. esakiae Herre 1939, A. nocturnus (Herre 1945), and A. sewardii). It is easily

distinguished from all these species in its general appearance and in morphological characters as provided in the

present study. Allen & Erdmann (2012) included the Red Sea in the range of distribution of Amblygobius sphynx,

however, material from the southern Red Sea (USNM 339029) was re-examined herein and the specimen was

identified as A. albimaculatus.

TABLE 1. (Continued)

Species K. hoesei sp. nov. K. hectori

Specimen Holotype Paratypes Paratypes Specimens in

comparative

material

Specimens in

comparative

material

Snout to origin of first dorsal fin 37.8 35.3–37.2 35.3–39.8 38.9–39.2 35.7–37.5

Snout to origin of second dorsal fin 55.6 54.3–56.0 52.7–59.2 56.2–59.0 54.4–56.0

Snout to origin of anal fin 65.6 62.3–65.3 59.8–65.2 61.4–62.3 57.7–60.4

Snout to origin of anus 60.4 57.9–60.8 56.0–59.1 57.1–57.7 53.3–55.7

Snout to origin of pelvic fin 31.9 31.0–31.2 30.4–33.6 31.9–32.8 30.2–33.1

Total dorsal-fin bases 45.1 45.4–46.8 45.1–49.5 47.2–53.5 47.4–53.6

Upper-jaw length 9.0 8.6–8.8 8.6–9.7 8.2–8.3 8.0–9.3

% of caudal peduncle:

Caudal peduncle depth 72.0 64.6–71.0 66.7–73.7 76.2–88.9 75.4–83.7

% of head length:

Caudal-fin length 85.1 83.0–94.5 80.8–93.1 82.5–87.4 87.5–95.1

Cheek depth 14.9 14.7–15.5 14.0–15.6 14.7–16.3 13.0–17.0

Eye diameter 25.3 22.7–26.0 23.7–27.8 24.2–25.0 24.3–25.5

Head depth 55.2 56.8–60.0 57.1–60.5 47.4–50.0 47.3–54.4

Head width 54.0 52.6–59.1 50.7–59.6 53.7–56.3 49.5–57.7

Postorbital length 47.1 45.5–50.5 45.5–49.5 48.8–50.5 46.4–47.0

Snout length 35.6 33.0–34.7 32.9–36.1 30.0–32.6 31.0–33.3

Upper jaw length 29.9 30.0–31.0 30.1–32.5 28.4–28.8 29.0–31.9

% of eye diameter:

Interorbital width 90.9 95.7–100 94.7–100 85.0–100 85.2–96.0

Snout length 140.9 126.9–148.0 123.8–146.4 120.0–134.8 123.1–137.0

% of pelvic to anus:

Pelvic-fin length 88.9 92.8–100 100–114.8 87.7–89.2 106.0–114.9

KOVAČIĆ ET AL.

466

Koumansetta hectori (Smith 1957)

Hector’s goby

Fig. 5, Table 1

Seychellea hectori Smith 1957: 726 (type locality: western Indian Ocean, Seychelles, Mahé; holotype: SAIAB 214)—Smith

1959: 204 (western Indian Ocean); Smith & Smith 1963: 47 (listed).

Amblygobius hectori—Masuda et al. 1980: 274 (Japan, description); Masuda et al. 1984: 256 (Japan, description);

Winterbottom & Emery 1986: 5 (Chagos Archipelago, description); Randall & Goren 1993: 3 (Maldives, description);

Myers 1999: 242 (Palau, Truk, Pohnpei, description); Larson & Murdy 2001: 3595 (western Pacific, listed); Senou et al.

2007: 66 (Ryukyu Islands, listed); Randall & Lim 2000: 635 (South China Sea, listed); Adrim et al. 2004: 126 (Anambas

Island, listed); Kuiter & Tonozuka 2004: 338 (Bali, photo); Senou et al. 2004: 373 (Japan, photo); Anderson 2005: 96

(Maldives, photo); Shibukawa 2009: 261 (Andaman Sea, description); Seeto & Baldwin 2010: 52 (Fiji, listed);

Satapoomin 2011: 65 (Andaman Sea, listed); Chabanet et al. 2012 (Maldives, listed); Yoshigou 2013: 110 (Ryukyu

Islands, listed).

Koumansetta hectori—Allen & Adrim 2003: 59 (Indonesia, listed); Shao et al. 2008: 262 (southern Taiwan, listed); Fricke et al.

2011 (New Caledonia, listed); Allen & Erdmann 2012: 981 (East Indies, description, photo from the Philippines); Fricke et

al. 2013: 270 (Europa Island, listed); Motomura & Harazaki 2017: 133 (Yaku-shima Island, photo); Fricke et al. 2018: 334

(Madagascar, listed).

Diagnosis. Dorsal-fin rays VI + I,15–17; anal-fin rays I,15–16; pectoral-fin rays 16–17; longitudinal scale series

49–54; scales in transverse series 17–18; interorbital wide, 67–90 % of eye diameter*; rear margin of upper jaw

ending posteriorly slightly behind or at vertical through anterior margin of eye; first two dorsal-fin spines filiform,

elongate, remaining progressively shorter, the sixth dorsal-fin spine almost reaches the origin of the second dorsal;

body covered with small ctenoid scales, cycloid at axil of pectoral fin, nape and entire belly, also in the area

between first dorsal fin and pectoral fin; predorsal scales 16–22, scales not reaching to the vertical through the

posterior edge of eye, ending at the level of pore G or H'*; cheek and opercle naked; scales cover the basal 1/3–1/4

of caudal fin, becoming rapidly smaller and cycloid; prepectoral area with about 4–9 vertical series of fine cycloid

scales covering the entire base of the pectoral fin; prepelvic area with cycloid scales, 9–14 in the midventral row;

width at anal-fin origin 9.9–10.7% of SL; caudal peduncle depth 75.4–88.9% of its length; head depth 47.3–54.4%

of head length; length of 11th second dorsal-fin ray 11.8–14.2% of SL; length of 8th anal-fin ray 13.7–14.9% of

SL.

Color in life (Fig. 5). Upper two-thirds of head brown with reddish hue anteriorly, greyish ventrally; upper half

of body dark brown above pale yellowish white stripe extending from posterior part of upper jaw across lower

cheek, lower opercle, and middle of pectoral-fin base and ending at caudal-fin base; lower half of head and body

above this demarcation stripe grey to dark grey, usually with bluish hue on abdomen. A large black spot (ocellus),

rimmed by a semicircular white mark below, on back at middle of the second dorsal fin, the spot extending into fin,

where rimmed above in yellow; the black spot on back appears an open eye, sometimes with dark red anteriorly

and posteriorly; another black spot larger than pupil dorsally at posterior end of caudal peduncle, rimmed below in

yellow. Head with straight middorsal stripe from the tip of snout reaching origin of the first dorsal fin. Two narrow

longitudinal yellow dorsolateral stripes on each side of head and body between middorsal and demarcation stripes.

Upper stripe with U-shaped continuation at the front of snout from one side of body to the other side and

continuing across upper part of eye and inner surface of orbit (only upper edge of orbit is visible through stripe)

backwards to black ocellus, the stripe then continuing behind ocellus, more or less visible, along the top of caudal

peduncle, where it ends at another black spot. Lower stripe extending from the upper lip through the lower part of

eye, upper part of opercle, and upper pectoral-fin base running along lateral midline to the caudal peduncle where it

is bended or separate, continuing as slightly lower positioned pale yellow mark on caudal-fin base. First dorsal fin

with a reddish brown band, rimmed above in yellow, along base; a black ocellus, a little larger than pupil, broadly

rimmed with yellow over first three spines and adjacent third membrane at midlevel of fin; an indistinct narrow

reddish brown stripe on membranes between fourth and sixth spines; and an obvious orange-red streak or series of

2 or 3 elongate spots in outer half of first membrane. The second dorsal fin with a broad reddish brown band

rimmed above in yellow, circumflexing black ocellus, along base; narrow yellow margin and an indistinct reddish

brown submarginal band, more obvious on posterior half; an indistinct narrow reddish brown stripe in the middle

of anterior half of fin; and yellow rear edge. Anal fin with pale grey to dark grey membranes and with narrow pale

bluish white margin on posterior half and rear edge. Caudal fin with curved white bar at base, rest of fin with

transparent membranes. Pectoral and pelvic fins with translucent membranes.

467

NEW KOUMANSETTA FROM THE RED SEA

FIGURE 5. Koumansetta hectori, underwater photographs. A: Reunion; B: Maldives; C: Philippines. Photos by A. Diringer

(A), J. Greenfield (B), P. Poppe (C).

KOVAČIĆ ET AL.

468

Distribution and habitat. Widespread from islands of the western Indian Ocean (Seychelles, Reunion,

Mayotte Island, Europa Island, and Madagascar), Chagos Archipelago, Maldives, and Andaman Sea east to the

Indonesia, Malaysia (Tioman), Papua New Guinea, the Philippines, New Caledonia, Solomon Islands, Micronesia,

and Fiji, north to Taiwan and southern Japan (Ryukyu and Yaeyama Islands), no positive records from Australia.

The record from Fiji (easternmost record of the species) is based on material from USNM collected by Springer &

team (e.g. USNM 236679, USNM 235950). Usually seen solitary hovering a short distance above bottom near base

of coral reefs at depth of 3–30 m.

Remarks. Diagnosis of K. hectori is the combination of original description (Smith 1957) and the data from

the material examined. Some data therefore represent extended ranges based on both sources: dorsal-fin rays VI +

I,15–17 (original description: 16–17, studied material: 15–17); pectoral-fin rays 16–17 (original description: 16,

studied material: 16–17); longitudinal scale series 49–54 (original description: 50–54, studied material: 49–54);

vertical rows of scales in prepectoral area 4–9 (original description: 9, studied material: 4–7). Despite the

geographically distant origins of the comparative material of K. hectori (Comoros, Seychelles, Maldives,

Philippines), the found data match well the original description from Seychelles. Where the data observed for the

examined material differed from the original description, only the observed characters where included in the

diagnosis and marked with *. Two generic characters were obvious in examined material that were not mentioned

in original description: anterior nostril tubular, short, lacking process from rim; posterior nostril slit-like, closer to

anterior nostril than to anterior margin of eye; and gill rakers on lower limb of anterior arch slender, weak and soft,

6–7 in number. The differences could be caused by the less precise measuring or by a less precise account in the

original description, by lower quality of techniques (stereomicroscope or caliper) and by lack of staining method in

the original description. However, even excluding these characters, there are enough characters matching both the

original description and results for the studied material, so that this species be can positively identified among

congeners. Contrary to this, the published meristic data for this species that are clearly distant from both, the counts

in the original description and in this study, should be taken with caution; e.g. Allen & Erdmann (2012) gave a

range of longitudinal scale series of 65–70 and the count of first dorsal and anal fins I,14. Some ranges among

morphometric characters in Table 1 from the presently studied material from Comoros, Seychelles, Maldives, and

the Philippines are non-overlapping with the data in the original description based on material from the Seychelles.

Unfortunately, the morphometric methods were not described in Smith (1957), contrary to the precise description

of measurement methods in the present description. Considering that specimens from type locality (Seychelles)

were examined in the present study, noticed differences are probably results of different measurement methods.

Koumansetta rainfordi Whitley 1940

Old Glory

Fig. 6

Koumansetta rainfordi Whitley 1940: 426 (type locality: Australia, Queensland, Whitsunday Group, Hayman Island; holotype:

AMS IA.2029)—Allen & Adrim 2003: 59 (Indonesia, listed); Randall et al. 2004: 28 (Tonga, listed); Randall 2005: 541

(South Pacific, description, photograph from Papua New Guinea); Shao et al. 2008: 262 (southern Taiwan, listed); Fricke

et al. 2011 (New Caledonia, listed); Allen & Erdmann 2012: 982 (East Indies, description, photo from West Papua); Fricke

et al. 2014: 168 (Papua New Guinea, listed).

Amblygobius rainfordi—Randall et al. 1997: 394 (Great Barrier Reef, description); Myers 1999: 242 (Palau, Marshall Islands,

description); Randall & Lim 2000: 635 (South China Sea, listed); Hutchins 2001: 42 (Western Australia; listed); Larson &

Murdy 2001: 3595 (western Pacific, listed); Kuiter & Tonozuka 2004: 338 (Bali and Flores, photos); Seeto & Baldwin

2010: 52 (Fiji, listed).

Diagnosis. Dorsal-fin rays VI + I,15–17; anal-fin rays I,15–17; pectoral-fin rays 16–18; longitudinal scale series

55–61; scales in transverse series 17–19; interorbital wide, 75–100 % of eye diameter*; rear margin of upper jaw

ending at vertical through anterior margin of eye or slightly behind*; first two dorsal-fin spines filiform, elongate*,

remaining progressively shorter; scales ctenoid posteriorly, becoming cycloid anteriorly between first dorsal fin

and pectoral fin, at axil of pectoral fin, on nape and entire belly; predorsal scales 24–26 (published data: 24, studied

material: 24–26); scales not reaching to the level of the posterior edge of the eye on predorsal area, ending at the

level of pore G; cheek naked*, opercle naked, in some specimens with a few cycloid scales in upper part; scales

cover the basal 1/4–1/6 of the caudal fin, becoming rapidly smaller and cycloid; prepectoral area with about 5–9

469

NEW KOUMANSETTA FROM THE RED SEA

vertical series of fine cycloid scales covering the entire base of the pectoral fin; prepelvic area with cycloid scales,

11–16 in the midventral row.

Color in life (Fig. 6). Body grey to brownish or greenish grey, upper half of head and predorsal area pale

yellowish green, lower head olive. Head with a short, middorsal, orange to reddish stripe on interorbital area

broken into 2–3 segments, other six narrow orange stripes on each side of head and body, the second to fourth and

those along anal-fin base are dark-edged and the edge is again bordered by a thin bright green margin; first stripe

extending from top of snout along upper side of nape and nearly reaching the vertical through origin of first dorsal

fin; second stripe beginning from posterior nostril across inner surface of orbit and ending below origin of first

dorsal fin where continuing as longitudinal row of spots to below sixth-eights soft ray; third stripe on left and right

side of body with U-shape connection at the front of snout and continuing across eye backwards to below rear base

of dorsal fin; fourth stripe extending from upper lip across cheek, opercle and upper third of pectoral-fin base and

running along midlateral of body to posterior part of caudal peduncle; fifth stripe extending from chin across lower

opercle and lower third of pectoral-fin base and running along ventral side of body to lower part of caudal-fin base

where it meets with curved orange bar on caudal-fin base; sixth stripe beginning from chin and continuing along

ventral side of abdomen and along anal-fin base. Back with series of 5–7 white spots of variable size. Black spot

larger than pupil dorsally at posterior end of caudal peduncle, rimmed below in white. Bases of both dorsal fins

with black-edged orange stripe, another indistinct red-orange stripe above basal stripe. Second dorsal fin with

white margin and indistinct red-orange submarginal stripe, and with yellow-edged black spot on outer half in the

middle of fin, the spot varying individually in size. Anal-fin membranes pale grey anteriorly shading darker

posteriorly, margin of fin narrowly bluish white. Caudal fin with curved white bar at base, rest of fin with

transparent membranes and pale grey rays. Pectoral and pelvic fins with translucent membranes.

Distribution and habitat. Reported from the Western and eastern Australia (Queensland) throughout

Indonesia and Papua New Guinea east to Marshall Islands, Solomon Islands, Fiji, and Tonga, north to the Taiwan

and the Philippines. Occurs on rich-growth coral reefs at depths of 2–30 m; darts into corals when alarmed.

Remarks. Diagnosis represents the sum of published data (Whitley 1940; Randall 2005, Allen & Erdmann

2012 and the data from the examined comparative material). Some data therefore represent extended ranges based

on the published data and the material examined: dorsal-fin rays VI + I,15–17 (published data: 15–16, studied

material: 15–17); anal-fin rays I,15–17 (published data: 15–16, studied material: 15–17); longitudinal scale series

55–61 (published data: 55–60, studied material: 57–61); scales in transverse series 17–19 (published data: 19,

studied material: 17–18). Where the data observed for the studied material obviously differed from the original

description in Whitley (1940), only the observed characters where included in the diagnosis and marked with *.

The differences could be caused by differences in the methods applied or by the lower quality of stereomicroscope

used for original description compared to modern techniques that were helped by staining. Despite these

differences, there is no need for urgent redescription of the species considering the small number of described

species in the genus and enough differential characters that are present in both, the original description and studied

material, to positively identify this species among congeners.

Kuiter & Tonozuka (2004) reproduced two underwater photographs taken from Menjangan Island, Bali, of

individuals similar to K. rainfordi but differ in detail of coloration. Probably gobies from this locality represent an

undescribed species but specimens are needed for study.

Molecular phylogenetics

The maximum likelihood analysis resulted in a monophyletic clade consisting of the three species of Koumansetta

with K. rainfordi being sister to the lineage with K. hectori and K. hoesei sp. nov. (Fig. 7). Bootstrap support of this

clade was 100 percent and the clade with the latter two species also received very high support with 98 percent.

Divergence within species of Koumansetta was considerably low when compared with among species divergence

and the evolutionary divergence of K. hoesei sp. nov. from its closest relative K. hectori became evident. Besides

the high supported monophyly of the genus Koumansetta, only the clade consisting of Amblygobius phalaena, A.

semicinctus, A. cf. phalaena and A. albimaculatus received high bootstrap support (99 percent) above the species

level. The pattern of divergence of this clade and of other Amblygobius spp. included in the present analysis was

not resolved, or at least did not receive high support from bootstrapped analyses.

KOVAČIĆ ET AL.

470

FIGURE 6. Koumansetta rainfordi, underwater photographs. A: West Papua; B: Tulamben, Bali, Indonesia; C: Kimbe Bay,

Papua New Guinea; D: Sulawesi, Indonesia. Photos by A. Ryanskiy.

471

NEW KOUMANSETTA FROM THE RED SEA

FIGURE 7. Maximum likelihood phylogenetic tree inferred from mitochondrial COI sequences (652 bp) of species of

Koumansetta and Amblygobius, and as outgroup Signigobius biocellatus, Valenciennea puellaris, and Exyrias belissimus.

Sequences are denoted by GenBank accession if available or by Barcode of Life Database (BOLD) ID (marked with *).

Additionally, locality from where specimens were collected is given. Sequence from holotype of Koumansetta hoesei sp. nov.

SMF 35723 is shown in bold. Numbers on braches correspond to bootstrap proportions in percent as retrieved from 1.000

replicates higher than 50 percent. Scale bar represents average number of nucleotide substitutions in relation to branch length.

KOVAČIĆ ET AL.

472

TABLE 2. Information on sequences used for inference of a Maximum Likelihood phylogeny of Koumansetta and Amblygobius species based on partial mitochondrial COI sequences. Holotype of

Koumansetta hoesei sp. nov. marked with *.

Species GenBank accession Voucher BOLD ID Museum catalog

number

Collection locality

Amblygobius albimaculatus GU805111 ADC09_240.4#1 DSFSF535-09 n.a. Mozambique

JF492825 ADC09_240.4 #2 DSFSF592-09 n.a. Mozambique

MH514921 KAU12-226 n.a. SMF 35873 Saudi Arabia

MH514922 KAU14-896 n.a. n.a. Saudi Arabia

Amblygobius buanensis MH514923 ROM-T00683 n.a. ROM 81256 Palau

Amblygobius cf. phalaena AF391378 AMBLYPHAELII n.a. n.a. French Polynesia, Society

Islands

n.a. SCIL-268 SCIL-268 USNM 435157 French Polynesia, Society

Islands

n.a. AUST-019 AUST-019 USNM 424012 French Polynesia, Austral

Islands

n.a. AUST-013 AUST-013 USNM 424009 French Polynesia, Austral

Islands

Amblygobius decussatus KP194244 HBL-LI-2001-67 LIFS087-08 n.a. Queensland, Australia

KP194955 HBL-LI-2001-135 LIFS124-08 n.a. Queensland, Australia

KP194952 2676 LIFS461-08 n.a. Queensland, Australia

FJ582722 HLC-11158 TZAIC450-05 n.a. Philippines

FJ582719 HLC-13096 TZAIB473-06 n.a. Philippines

FJ582720 HLC-13198 TZAIB575-06 n.a. Philippines

FJ582721 HLC-13199 TZAIB576-06 n.a. Philippines

FJ582723 HLC-10978 TZAIC278-05 n.a. Philippines

Amblygobius nocturnus n.a. VERDE-2015-18_GAL-57 PHILV304-15 USNM 436170 Philippines

Amblygobius phalaena AF391369 AMBLYPHAEL n.a. n.a. French Polynesia, Society

Islands

FJ582724 HLC-13170 TZAIB547-06 n.a. Philippines

KP194751 2469 LIFS254-08 n.a. Queensland, Australia

FJ582725 HLC-13171 TZAIB548-06 n.a. Philippines

KP194499 HBL-LI-2000-96 LIFS052-08 n.a. Queensland, Australia

KP194021 HBL-LI-2000-151 LIFS066-08 n.a. Queensland, Australia

……continued on the next page

473

NEW KOUMANSETTA FROM THE RED SEA

ABLE 2

. (Continued)

Species GenBank accession Voucher BOLD ID Museum catalog

number

Collection locality

KP194353 2684 LIFS469-08 n.a. Queensland, Australia

KP194327 3081 LIFS866-08 n.a. Queensland, Australia

KP194685 GT 2846 LIFS631-08 CSIRO H 7595-07 Queensland, Australia

KP194153 GT 2847 LIFS632-08 CSIRO H 7595-07 Queensland, Australia

KP194792 2683 LIFS468-08 n.a. Queensland, Australia

KP194896 HBL-LI-2001-154 LIFS137-08 n.a. Queensland, Australia

JQ431409 MBIO1188.4 MBFA698-07 MNHN_2008-907 French Polynesia

Amblygobius semicinctus n.a. n.a. SAIAD524-12 n.a. Kenya

Amblygobius sphynx n.a. VERDE-2015-16_GAL-55 PHILV281-15 USNM 436147 Philippines

Koumansetta hectori JQ349683 NBE0117 SBF055-11 n.a. Madagascar

JQ349687 NBE0118 SBF056-11 n.a. Madagascar

JQ349685 NBE0507 SBF223-11 n.a. Madagascar

JQ349684 NBE0547 SBF252-11 n.a. Madagascar

JQ349686 NBE0119 SBF057-11 n.a. Madagascar

Koumansetta hoesei sp. nov. MH514912 KAU11-224 n.a. KAUMM 442 Saudi Arabia (Red Sea)

MH514913 KAU11-516 n.a. KAUMM 444 Saudi Arabia (Red Sea)

MH514914 KAU14-122 n.a. n.a. Saudi Arabia (Red Sea)

MH514915 KAU14-123 n.a. n.a. Saudi Arabia (Red Sea)

MH514916 KAU14-796 * n.a. SMF 35724 * Saudi Arabia (Red Sea)

MH514917 KAU14-797 n.a. SMF 35723 Saudi Arabia (Red Sea)

MH514918 KAU14-846 n.a. KAUMM 446 Saudi Arabia (Red Sea)

MH514919 KAU12-845 n.a. SMF 35867 Saudi Arabia (Red Sea)

MH514920 KAU14-121 n.a. n.a. Saudi Arabia (Red Sea)

Koumansetta rainfordi KP194772 2771 LIFS556-08 n.a. Queensland, Australia

KP194249 2825 LIFS610-08 n.a. Queensland, Australia

Valenciennea puellaris KU176381 ADC2013 240.104A #1 DSFSG963-13 n.a. South Africa

Signigobius biocellatus HQ536673 C219 (voucher LACM:T-001079)

GBGCA2078-13 n.a. n.a.

Exyrias belissimus HQ909463 C249 (voucher LACM:T-000169)

GBGCA1945-13 n.a. n.a.

KOVAČIĆ ET AL.

474

TABLE 3. Diagnostic characters of Koumansetta compared to Amblygobius species.

number of

character in

diagnosis

Koumansetta:

K. hectori

K. hoesei sp. nov.

K. rainfordi

Group 1) in text:

A. albimaculatus

A. phalaena

A. semicinctus

A. sphynx

Group 2) in text:

A. buanensis

A. bynoensis

A. decussatus

A. stethophthalmus

Group 3) in text:

A. linki

Group 4) in text:

A. calvatus

A. cheraphilus

A. esakiae

A. nocturnus

A. tekomaji

Group 5) in text:

A. sewardii

3 oculoscapular transverse

rows trp and tra long,

extending dorsally well

above level of rows x1 and

oculoscapular transverse

rows trp and tra short,

below level of rows x1 and

x2 or shortly passing above

oculoscapular transverse

rows trp and tra short,

below level of rows x1 and

x2 or shortly passing above

oculoscapular transverse

rows trp and tra short,

shortly passing rows x1 and

oculoscapular transverse

rows trp and tra short,

below level of rows x1 and

x2 or shortly passing above

oculoscapular transverse

rows trp and tra short,

below level of rows x1 and

4 snout pointed, prominent,

long, 0.7–0.8 in eye

diameter, 2.8–3.0 in head

length, with gently sloping

dorsal profile, overhanging

mouth

snout blunt, rounded,

ending anteriorly at level or

behind the mouth, 0.8–0.9

in eye diameter, 3.1–3.8 in

head length

snout blunt, rounded,

ending in the level of

mouth with dermal flap

falling on upper lip, 0.7–1.0

in eye diameter, 3.0–3.5 in

head length

snout blunt, rounded,

moderately oblique, ending

behind vertical of mouth

anterior edge, 1.0 in eye

diameter, 3.8-3.9 in head

length

snout blunt, rounded,

ending anteriorly at level or

behind the mouth, 1.0-1.3

in eye diameter, 4.3-5.8 in

head length

snout blunt, rounded,

moderately oblique, ending

behind vertical level of

mouth anterior edge, 0.7 in

eye diameter, 4.3-4.4 in

head length

5 mouth subterminal mouth terminal mouth terminal mouth terminal mouth terminal mouth terminal

6 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

the upper limb of first gill

arch with 4-6 long

filaments on joint branch

the upper limb of first gill

arch with 4-6 long

filaments on joint branch

the upper limb of first gill

arch with 4 long filaments

on joint branch

the upper limb of first gill

arch with 5 long filaments

on joint branch

the upper limb of first gill

arch with 5 long filaments

on joint branch

......continued on the next page

475

NEW KOUMANSETTA FROM THE RED SEA

TABLE 3.

(Continued)

number of

character in

diagnosis

Koumansetta:

K. hectori

K. hoesei sp. nov.

K. rainfordi

Group 1) in text:

A. albimaculatus

A. phalaena

A. semicinctus

A. sphynx

Group 2) in text:

A. buanensis

A. bynoensis

A. decussatus

A. stethophthalmus

Group 3) in text:

A. linki

Group 4) in text:

A. calvatus

A. cheraphilus

A. esakiae

A. nocturnus

A. tekomaji

Group 5) in text:

A. sewardii

7 first two dorsal-fin spines

longest and elongate,

remaining spines

progressively shorter

first dorsal fin with third to

fourth or fifth spines the

longest, elongate or not

first dorsal fin with third to

fifth spines longest, fifth

slightly longer, slightly

elongate or not

first dorsal fin with third to

fifth spines longest, fifth

slightly longer, not

elongate

first dorsal fin with third to

fifth spines longest, not

elongate

first dorsal fin with third to

fourth spines longest, not

elongate

8 pelvic frenum absent pelvic frenum weakly to

well developed (1/5 to 1/3

of spine length in midline

height)

pelvic frenum weakly to

well developed (1/8 to 1/3

of spine length in midline

height)

pelvic frenum weakly

developed (1/5 of spine

length in midline height)

pelvic frenum well

developed (1/3 to 1/2 of

spine length in midline

height)

pelvic frenum well

developed (1/2 to 2/3 of

spine length in midline

height)

9 color pattern mostly brown

to brown-green with

narrow yellow or orange

longitudinal stripes on body

and head and two black

spots posteriorly, one

ocellated, partly or

completely on the second

dorsal fin, and another spot

dorsoposteriorly on caudal

peduncle

body with variable other

pattern details but always

with five vertical bars on

body and black spot present

at upper part of operculum

body with variable other

color pattern but always

with two to four

longitudinal stripes, which

are either restricted to head

and anterior body (A.

bynoensis and A.

stethophthalmus) or

completely along side of

body reaching caudal fin

(A. buanensis and A.

decussatus)

body with three long

longitudinal brown stripes

body with 1-3 long

longitudinal brownish or

reddish stripes on each side

or just head with

longitudinal rows of

elongated spots

body grey to brown with

pale blue vertical bands

KOVAČIĆ ET AL.

476

Discussion

In this study, we provide arguments for the validity of the genus Koumansetta, and in addition to the two described

species K. rainfordi and K. hectori, we describe a new species Koumansetta hoesei sp. nov., which is endemic to

the Red Sea and the Gulf of Aden.

In the past, Koumansetta hoesei sp. nov. has repeatedly been misidentified as K. hectori; the results of our

detailed morphological examination allow to clearly distinguish the two species. Furthermore, details of coloration,

most evident in the pattern of stripes along head and body, are distinctive for K. hoesei sp. nov. and its Indo-West

Pacific congener K. hectori. The phylogenetic analysis of partial mitochondrial COI showed that K. hoesei sp. nov.

is evolutionary markedly divergent from its closest relative K. hectori. Assuming rates of molecular evolution

similar to those that consistently are found in distinct groups of tropical reef fishes (see e.g., the compilation in

Lessios (2008) for mitochondrial COI) the observed differences in sequence composition between the two

congeners indicate longstanding evolutionary divergence in the range of some few millions of years. This gross

estimation of the timing of divergence of K. hectori and K. hoesei sp. nov. from their common ancestor hints at an

onset of speciation in the early Pleistocene, when the Red Sea repeatedly became effectively isolated from the

Indian Ocean due to lowered sea levels during periodic glacial periods (see Klausewitz (1989) and references

therein). Most likely at the onset or very early in this period, the two sister species evolved from their common

ancestor in allopatry as a consequence of reproductive isolation of Red Sea and Indian Ocean/Indo-Pacific

populations. As in other Red Sea endemic species with Indian Ocean (or Indo-Pacific) sibling species that—

deduced from their evolutionary age—must have persisted during later glacial sea level low stands that followed

their divergence, it must be concluded that glacial refugia have existed for K. hoesei sp. nov. to survive these

periods. Such refugia may either have been located inside the Red Sea in certain areas with permissive ecological

conditions and/or outside the Red Sea in the Gulf of Aden region (see e.g. discussion of this topic in Klausewitz

(1989), and more recently reviewed in DiBattista et al. (2017)).

The 32 diagnostic nucleotide substitutions that allow distinction between K. hoesei sp. nov. 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). In agreement with expectations from this relatively low degree of molecular

divergence, and in contrast to the more divergent sibling species in Koumansetta and Cheilinus, no prominent

morphological divergence was found in C. crenidens from Red Sea and Indian Ocean populations.

The three species of Koumansetta share morphological synapomorphies that presumably have evolved as the

result of adaptive processes. In Koumansetta, adaptation shaped the typical “Amblygobius” body form for a

hyperbenthic lifestyle, further departing from the usual goby-shaped body to an almost wrasse-shaped body. For

example, K. hectori and K. hoesei sp. nov. in appearance resemble juveniles of the wrasse Halichoeres marginatus

Rüppell 1835 living in the same habitat (SB, unpublished data). The evolutionary significance of this gross

morphological adaptation with regards to niche evolution is evident as all three species of Koumansetta occur in a

different habitat than the 15 species of Amblygobius. They are typical inhabitants of coral reef areas hovering a

short distance above or within corals or coral rock whereas species of Amblygobius live in areas close to coral reefs

above soft substrata where they build their burrow.

The Koumansetta diagnosis, however, is not just the mix of characters where each character distinguishes

Koumansetta from just one or a part of Amblygobius species. In fact, the characters (3)-(9) in the diagnosis of

Koumansetta work against all remaining Amblygobius species (Table 3). These synapomorphies confirm a quite

distinguished evolutionary history and substantial adaptations of a monophyletic Koumansetta, providing no

justification for further keeping these species in Amblygobius.

As expected from the marked morphological distinction of the genus Koumansetta from Amblygobius species,

the molecular phylogeny of partial mitochondrial COI confirmed that the genus Koumansetta is a monophyletic

group of species, which is well divergent from all Amblygobius species included in the phylogeny (Fig. 7). The

477

NEW KOUMANSETTA FROM THE RED SEA

information content of the partial COI sequence used in the present ML analysis, however, was not sufficient to

resolve deeper phylogenetic relationships among clades of species in residual Amblygobius species. Although it

appears from the present phylogeny that Koumansetta is the closest sister to a clade consisting of only a part of

residual Amblygobius (i.e. the clade with A. albimaculatus, A. phalaena, A. semicinctus, A. cf. phalaena and A.

sphynx), this phylogenetic grouping did not receive support from bootstrapped analyses (support value for the

branch joining these two clades was 40 percent (not shown in Fig. 7)). This means, there is no clear evidence for a

paraphyletic “Amblygobius” and additional analyses based on more than just the barcoding portion of the

mitochondrial COI gene is required to resolve the phylogenetic placement of Koumansetta in relation to

Amblygobius spp.

However, independent of the question on a possible paraphyly of Amblygobius, the species of the genus where

divided into several groups based on some of the morphological characters that we studied for comparison with

Koumansetta (all but two species of Amblygobius were studied herein and data from the literature were used for the

two residual species). Our observations are preliminary and we suggest that no taxonomical decisions be made

before they are confirmed by examination of a larger number of specimens and before a more detailed analysis of

phylogenetic relationships among species of Amblygobius and Koumansetta reveals there evolutionary

relationships. However, our examination of our data suggest the following five groups.

Group 1) four species: A. albimaculatus (Rüppell 1830), A. phalaena (Valenciennes 1837), A. semicinctus

(Bennett 1833), and A. sphynx (Valenciennes 1837) (type species) which differ from all other Amblygobius in

pelvic frenum size (character 8 in Table 3) (overlapping with group 2); body coloration (character 9 in Table 3);

head lateral line system with 14–21 suborbital rows c (slight overlap with group 2); snout length 3.1–3.8 in SL

(overlapping with group 2); interorbital broad, 1.0–1.2 in eye diameter, 3.9–4.3 in head length; body high, depth at

pelvic-fin origin 3.7–3.9 in SL, at anal-fin origin 3.8–4.1 in SL; caudal peduncle high, 6.5–6.9 in SL; and predorsal

scaled to the interorbital with middorsal count of 26–32 scales (overlapping with group 2).

Group 2) four species: A. buanensis Herre 1927, A. bynoensis (Richardson 1844) [type species of

Odontogobius Bleeker 1874], A. decussatus (Bleeker 1855) and A. stethophthalmus (Bleeker 1851) which differ

from all other Amblygobius in snout shape and size (character 4 in Table 3); pelvic frenum size (character 8 in

Table 3) (overlapping with group 1); body coloration (character 9 in Table 3); head lateral line system with 14–21

suborbital rows c (slightly overlapping with group 1); caudal peduncle depth 7.0–8.1 in SL; and predorsal scaled to

the interorbital with middorsal count of 23–30 scales (overlapping with group 1).

Group 3) A. linki Herre 1927, single species differs from all other Amblygobius species in body coloration

(character 9 in Table 3); head lateral line system with 13–14 suborbital rows c, 4 above and 1 below row b; shallow

caudal peduncle depth 8.0–8.8 in SL (overlapping with group 4); predorsal scaled to the level of posterior

preopercular edge with middorsal count of 21 scales; anal-fin count low, I/12, pectoral-fin count low, 16 rays.

Group 4): “A. nocturnus species group”, also proposed by Allen & Erdmann (2016) contains five species:

Amblygobius calvatus Allen & Erdmann 2016; A. cheraphilus Allen & Erdmann 2016; A. esakiae Herre 1939;

Yabotichthys nocturnus Herre 1945; and Seychellea tekomaji Smith 1945 which differ from all other Amblygobius

species in pelvic frenum size (character 8 in Table 3); body coloration (character 9 in Table 3); head lateral line

system with 8–9 suborbital rows c, 2–3 above and 1 below row b (overlapping with group 5); snout length 4.3–5.8

in head length (overlapping with group 5), 1.0–1.3 in eye diameter; shallow caudal peduncle depth, 8.0–8.8 in SL

(overlapping with A. linki); cycloid scales (overlapping with group 5); and opercle naked (overlapping with group

4).

Group 5) Amblygobius sewardii (Playfair 1867) with its Red Sea synonym Biat magnusi Klausewitz 1968

(synonymy following H. Larson’s pers. comm.), single species differs from all other Amblygobius species in

characters pelvic frenum size (character 8 in Table 3); body coloration (character 9 in Table 3); head lateral line

system with 9 suborbital rows c, 3 above and 1 below row b (overlapping with group 4); dentition in upper and

lower jaws; snout length 4.3–4.4 in head length; body slender, depth at pelvic-fin origin 5.4 in SL, at anal-fin origin

6.0-6.1 (overlapping with group 4); shallow caudal peduncle depth, 10.9–11.0 in SL; cycloid scales (shared with

group 4); opercle naked (shared with group 4), anal-fin count high, I/18; and high count of scales in the

longitudinal and transverse series, 83 + 5 to 85 + 5 and 26–27, respectively.

In conclusion, the combined morphological synapomorphies and genetic evidence on monophyly of

Koumansetta presented herein justifies the maintenance of the genus following Randall’s (2005) opinion, despite

later synonymization (e.g., Kottelat 2013) or partial acceptation of the genus (Eschmeyer et al. 2017). However,

KOVAČIĆ ET AL.

478

taxonomical decisions in Kottelat (2013) and Eschmeyer et al. (2017), even being without elaborated explanations,

questioned Randall (2005) and therefore asked for counteraction regarding the status of the genus Koumansetta,

and caused the need for including a detailed re-description of the genus in this study. Regarding the seemingly

unresolved situation of the remaining species currently assigned to Amblygobius, additional study is underway to

clarify the distribution, evolutionary relationships and the taxonomy of these species.

Acknowledgements

Susanne Dorow and Jennifer Steppler are gratefully thanked for technical assistance at SMF and we are also very

grateful to Don Stacey from ROM, Sandra Raredon and Jeff Williams from USNM and Arnold Suzamoto and

Loreen O’Hara from BPBM for the kind loan of the material used in this study. We also thank Roger Bills and

Nkosinathi Mazungula for providing images and X-rays of the holotype and of paratypes of Seychellea hectori,

Gavin Gouws (SAIAB) who kindly provided a photograph and sequences of A. semicinctus, and Eric Holm (ROM)

who provided tissue subsamples of specimens of A. buanensis. Alain Diringer, Jean-Marie Gradot, Jim Greenfield,

Philippe Poppe, and Andrey Ryanskiy are acknowledged with thanks for contributing underwater photographs and

Amy Driskell and Jeffrey Williams (USNM) are thanked for photographs of fresh specimens and sequence data for

A. nocturnus and A. sphynx. We heartily thank Ewald Lieske for organizing and for acquiring funds for a field trip

to Lahami Bay, southern Egypt. We also thank Christine Thacker (LACM) for providing information on collection

localities of specimens of A. phalaena and A. cf. phalaena as well as tissue samples of various species for

continuation of this study. Matthias Juhas and Stephanie Simon (SMF) are thanked for assisting in molecular

genetic analyses. We further 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, Mohamed Gabr (KAU), Fareed Krupp (SRI) for their help in the

realization of the present study. This work has been supported in part by grant to MK of the Croatian Science

Foundation under the project IP-2016-06-5251.

Comparative material examined

Amblygobius albimaculatus (Rüppell 1830): BPBM 20410, 1 female, 70.2 + 18.3 mm, Sudan, Port Sudan harbour;

SMF 35873, 1 male 74.1 + 24.5 mm, Saudi Arabia, Farasan Archipelago; USNM 339029, 1 male, 63.5 + 20.1

mm, Eritrea, Sciumma Island, ½ mile of the south west shore.

Amblygobius buanensis Herre 1927: BPBM 31365, 1 female, 47.4 + 13.3 mm & 1 male, 48.3 + 13.5 mm, Palau,

Koror, east side, Ebadul's Pier.

Amblygobius bynoensis (Richardson 1844): USNM 244181, 1 female, 74.7 + 20.1 mm & 1 male, 79.3 + 20.7 mm,

Australia, Queensland, Gulf of Carpenteria, Duyfken Point near Weipa.

Amblygobius decussatus (Bleeker 1855): ROM 75033, 1 male, 49.6 + 15.0 mm, Palau, western coast of Babeldaob

Amblygobius esakiae Herre 1939: ROM 49497, 1 female, 48.8 + 18.8 mm & 1 male, 46.6 + 19.2 mm, Philippines,

Negros I., Dumaguate.

Amblygobius linki Herre 1927: ROM 74900, 1 female, 24.9 + 7.3 mm & 1 male, 30.0 + 10.1 mm, Palau, Babeldaob

Amblygobius nocturnus (Herre 1945): BPBM 35217, 1 female, 37.0 + 11.3 mm, Japan, Ogasawara archipelago,

Chichijima; BPBM 37640, 1 female, 39.4 + 11.1 mm, Indonesia, Mentawai Islands, Siberut Island, Sarabua

Bay.

Amblygobius phalaena (Valenciennes 1837): BPBM 38657, 1 female, 28.5 + 8.1 mm, Indonesia, Bali, northeast

coast at Tulamben.

Amblygobius semicinctus (Bennett 1833): BPBM 32952, 1 female, 70.5 + 23.9 mm, Maldives, South Malé Atoll,

Embudu Island, south side.

479

NEW KOUMANSETTA FROM THE RED SEA

Amblygobius sewardii (Playfair 1867): SMF 13823, 1 female, 60.1 + 20.7 mm & 1 male, 57.1 + 20.0 mm, Gulf of

Suez, Egypt, Al Tur.

Amblygobius sphynx (Valenciennes 1837): ROM 82005, 1 female, 57.1 mm, caudal fin damaged, Palau, Koror.

Amblygobius stethophthalmus (Bleeker 1851): USNM 424325, 1 female, 66.8 + 16.3 mm & 1 male, 62.8 + 15.9

mm, Singapore, Raffles Light House.

Amblygobius tekomaji (Smith 1959): ROM 36789, 1 female, 25.2 + 7.2 mm, Chagos Archipelago, Great Chagos

Bank.

Koumansetta hectori (Smith 1957): USNM 339658, 1 female, 32.9 + 8.3 mm & 1 male, 36.4 + 8.9 mm, Seychelles,

Mahe; ROM 54351, male, 37.3 + 10.1 mm, Philippines, Negros Oriental, Siquijor Island; ROM 58884, 2

males, 39.3 + 10.6 and 38.6 + 10.5 mm, Comoros, Moheli; ROM 54952, 1 female, 26.5 + 6.6 mm, 1 male, 39.3

+ 10.6 mm, Maldives, North Male Atoll.

Koumansetta rainfordi Whitley 1940: USNM 361749, 1 female, 42.1 mm, caudal fin damaged, 1 male, 35.0 + 10.1

mm, Pacific, Vanuatu, Efate Island; ROM 85171, 1 male, 31.4 + 8.5 mm, Indonesia, Raja Ampat, Waigeo

Island; ROM 38758, 2 females, 49.9 + 11.5 and 51.0 + 13.0 mm, 1 male, 44.8 + 12.4 mm, Australia,

Queensland, Great Barrier Reef.

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An updated checklist of the Red Sea gobioid species (Teleostei: Gobiiformes), with four new records

Article

Full-text available

Apr 2023

An updated checklist and status assessment of the gobioid species (Gobiiformes) in the Red Sea is provided. Of the 162 species, 141 belong to the Gobiidae, nine to Microdesmidae, five to Xenisthmidae, six to Schindleriidae and one to Kraemeriidae. The Shrimpgoby (Cryptocentrus steinhardti) and the Sand Goby (Hazeus ingressus) are reported from Eilat, Israel and Abu Dabab, Marsa Alam, Egypt. Both species, which belong to Indo-West Pacific genera, were originally described from the eastern Mediterranean Sea and were unknown from the Red Sea so far. Records of these species are based on underwater photographs. Eviota pseudostigma, a species known from islands of the Western Indian Ocean, was photographed and collected from Mangrove Bay, El Quseir, Egypt, and represents a new record for the Red Sea. Eviota oculopiperita, described from the north-eastern Red Sea was found on the western side of the Red Sea. A new record of the microdesmid fish Gunnellichthys irideus, based on underwater photographs taken from the southern Egypt, is reported. Previous records of Paragobiodon echinocephalus from the Gulf of Aqaba are regarded as misidentification of P. modestus.

New Distributional Records of Twelve Reef Fishes from Lakshadweep Waters, India

Article

Full-text available

Apr 2022

Previous fish faunal studies have indicated that 144 families, 432 genera, and 856 species are extant in the Lakshadweep archipelago. In the present study, 12 new records are added to the Lakshadweep fish diversity. The new records include Harlequin sweetlips (Plectorhinchus chaetodonoides), Three-stripe fusilier (Pterocaesio trilineata), Hector’s goby (Koumansetta hectori), Large toothed cardinalfish (Cheilodipterus macrodon), Coral hawkfish (Cirrhitichthys oxycephalus), One-spot wormfish (Gunnellichthys monostigma), Imposter fangblenny (Plagiotremus phenax), Disco blenny (Meiacanthus smithi), Maldives damselfish (Amblyglyphidodon indicus), Orange spotted spinefoot (Siganus guttatus), Spotfin burrfish (Chilomycterus reticulatus), and filament finned parrotfish (Scarus altipinnis). Plectorhinchus chaetodonoides, S. altipinnis and G. monostigma are new records to Indian fish fauna. The current study enhances the fish faunal records of Lakshadweep archipelago to being represented by 435 genera and 868 extant species, with Koumensetta, Gunnellichthys and Chilomycterus genera being recorded here for the first time. The observations were made through underwater visual census by SCUBA and snorkeling in the lagoon and fore-reef area of a single island, Kavaratti, one of the 12 atolls of Lakshadweep archipelago, from January to March 2021. Identifications of the new records were confirmed through photography and videography.

Endemic Fishes of the Red Sea

Chapter

Full-text available

Jan 2019

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.

Beyond Oman’s shores: records of Oxyurichthys omanensis (Teleostei: Gobiidae) from the central and southern regions of the Western Indian Ocean

Article

May 2025
ZOOTAXA

The goby genus Oxyurichthys (Gobionellinae) currently comprises 21 recognized species distributed across the tropical Indo-West Pacific region, with 12 species inhabiting the Western Indian Ocean (WIO). A recent examination of specimens from a mudflat habitat in northern Oman, located in the northwestern Indian Ocean, led to the description of a new species, Oxyurichthys omanensis. In this study, we re-examined collections of Oxyurichthys specimens from the central and southern WIO, previously identified as O. ophthalmonema. Our results provide robust evidence for the presence of O. omanensis in these regions, specifically from Kenya, Mayotte, and Madagascar. Here, we present a detailed morphological assessment of genotyped voucher specimens, enhanced by comprehensive descriptive information to further clarify the identification and distribution of Oxyurichthys species in the central and southern WIO.

Description of a new snake eel Ophichthus olivaceus (Teleostei: Anguilliformes, Ophichthidae) from the Red Sea

Article

Full-text available

Mar 2020

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.

A new species of Lioteres (Pisces, Gobiidae) from Kwazulu, with a revised checklist of South African gobies and comments on the generic relationships and endemism of western Indian Ocean gobioids. Royal Ontario Museum, Life Sciences Occasional Papers Series

Article

Full-text available

Jan 1979

Review of the gobioid fishes of the Chagos Archipelago, Central Indian Ocean

Article

Full-text available

Jan 1986

Annotated checklist of the fishes of Madagascar, southwestern Indian Ocean, with 158 new records

Article

Full-text available

Feb 2018

An annotated checklist of the fish species of the Madagascar EEZ (southwestern Indian Ocean) comprises a total of 1,798 species in 247 families. 158 species are recorded from Madagascar for the first time. The majority of the species is autochthonous; 28 species have been introduced, mainly in freshwater habitats. The fish fauna is mostly marine (95.4% of the total number of native fish species), with the Gobiidae, Labridae, Serranidae, Pomacentridae and Apogonidae being the families with most representatives; among the 90 native freshwater fish species (adults mainly occurring in freshwater), the Cichlidae are the dominating family, but there are also two endemic families, the Bedotiidae (16 species) and Anchariidae (6 species). The fish fauna at Madagascar is typical for offshore, high islands in the southwestern Indian Ocean. Zoogeographically, the main element of the marine fish fauna of Madagascar consists of widespread tropical Indo-Pacific species (978 species, 58.3% of the total native marine species). A total of 13 species (3.3%) are found worldwide, either circumtropical or circumtropical including warm temperate zones. A total of 215 species (12.8%) are widespread in the Indo-West Pacific. An additional 453 species (27.0%) are Indian Ocean endemics, including 233 western Indian Ocean endemics (13.9%), 73 southwestern Indian Ocean endemics (4.4%), 16 species endemic to Madagascar and Mascarenes (1.0%), 4 species endemic to Madagascar and Comoros (0.2%), 3 species endemic to Madagascar and Madagascar Ridge (0.2%), and 37 marine species endemic to Madagascar (2.2%). Most of the autochthonous freshwater fishes are endemic to Madagascar (87 species, 96.7% of the native freshwater species).

Morphological and molecular divergence between Crenidens crenidens (Forsskål, 1775) and C. indicus Day, 1873 (Perciformes: Sparidae) and notes on a Red Sea endemic lineage of C. crenidens

Article

Full-text available

Jul 2017

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.

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

Article

Full-text available

Aug 2016

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.

Marine fishes recorded from the Anambas and Natuna islands, South China Sea

Article

Full-text available

Mar 2004
RAFFLES B ZOOL

Four hundred and thirty species of fish from marine localities in the Anambas and Natuna groups of islands in the South China Sea were recorded between 11(th) and 22(nd) Mar 2002. Two hundred and seventy three species are represented by collections, while 157 species were recorded only by visual census. Among these, seven species are recorded for the first time in the South China Sea.

Descriptions Of Two New Gobies (Gobiidae: Amblygobius) From The Tropical Western Pacific Ocean

Article

Jan 2016

Two new species belonging to the Indo-Pacific gobiid genus Amblygobius are described from mud-bottom habitats. Amblygobius calvatus n. sp. is described on the basis of 9 specimens, 23.7–48.0 mm SL, from the El Nido area of northern Palawan in the Philippines. Diagnostic features for the new species include usual counts of 15 segmented dorsal and anal-fin rays, scales entirely cycloid, no scales on the head (including the side of the nape and upper opercle), 80–86 longitudinal body scales, 24–26 transverse body scales, a strongly lanceolate caudal fin, a grayish-brown color in life with two orange-brown stripes on the head and body, 8–11 small black spots or saddles on the upper back, a blackish moustache-like marking above the upper lip, a horizontally oval orange-brown spot on the opercle, and a white pectoral-fin base with a central, horizontally-elongate, reddish-brown marking. Amblygobius cheraphilus n. sp. is described from 11 specimens, 14.6–32.9 mm SL, collected near the town of Alotau in Milne Bay Province of Papua New Guinea. It differs from congeners on the basis of a combination of features, including usual counts of 13 segmented dorsal and anal-fin rays, scales entirely cycloid, no scales on the head except for the side of the nape, 56–60 longitudinal scales, 14–18 transverse scales, a moderately lanceolate caudal fin, a grayish color in life with two reddish-brown stripes on the head and body with the lower stripe containing a prominent oval dark-brown spot on the opercle and ending in a dark-brown spot on the caudal-fin base, a series of small brown saddles on the back and predorsal region, and a faint ocellus on the upper caudal-fin rays.

Evolutionary history and zoogeography of the Red Sea ichtyofauna

Article

Jan 1989

W. Klausewitz

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

Checklist of the fishes of Western Australia

Article

Jan 2001

J. Barry Hutchins

Illustrations of some Australian fishes

Article

G.P. Whitley

Redescription of the genus Koumansetta (Teleostei: Gobiidae), with description of a new species from the Red Sea

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