Callionymus omanensis, a new species of dragonet from Oman, north-western Indian Ocean (Teleostei: Callionymidae).

Journal of Fish Biology (2014) 85, 1303–1319 doi:10.1111/jfb.12460, available online at wileyonlinelibrary.com

Callionymus omanensis, a new species of dragonet from Oman, north-western Indian Ocean (Teleostei: Callionymidae) R. Fricke*†‡#, L. A. Jawad§ and J. M. Al-Mamry‖ *Im Ramstal 76, 97922 Lauda-Königshofen, Germany, †Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, 70191 Stuttgart, Germany, §4, Tin Turn Place, Flat Bush, Manukau, Auckland, New Zealand and ‖Marine Science and Fisheries Centre, Ministry of Agriculture and Fisheries Wealth, Muscat, Sultanate of Oman (Received 19 July 2013, Accepted 28 May 2014) A new species of deep-living dragonet Callionymus omanensis from Oman is described on the basis of a single male specimen collected in a trawl from 500 m depth off the coast of Oman. The new species is characterized within the subgenus Bathycallionymus by having a small branchial opening; head short (3⋅9 in proportion to standard length); eye large (2⋅4 in proportion to head length); preopercular spine with a long, upcurved main tip, with a small antrorse barb and a larger antrorse spine, and with a strong antrorse spine laterally at the preopercular-spine base, ventral margin smooth; first dorsal fin slightly higher than second dorsal fin (male); second dorsal fin distally straight; 17 pectoral fin rays; distal end of caudal fin slightly pointed, with two median unbranched rays bearing short filaments; first dorsal fin with basal black spot reaching from first to fourth membranes, third membrane with an ocellated distal black blotch; second dorsal fin with vertical dark grey bars; distal three-fourths of anal fin black; upper half of caudal fin with oblique dark grey bars; pelvic fin dark grey, second ray basally with a black blotch. The new species is compared with similar species. Revised keys to callionymid species of the western Indian Ocean and the Red Sea, as well as species of the subgenus Bathycallionymus, are presented. © 2014 The Fisheries Society of the British Isles

Key words: Arabian Sea; fishes; identification key; morphology; taxonomy.

INTRODUCTION Dragonets of the family Callionymidae are a group of benthic living fishes occurring in the upper 900 m of all temperate, subtropical and tropical oceans of the world, and a few species found in estuarine and freshwater habitats. They are characterized by a depressed body, a triangular head when seen from above, the eyes large, situated dorsally on the head, the presence of a preopercular spine that bears additional points and serrae, the gill opening reduced to a small pore, swimbladder absent, two dorsal fins, the first with thin, flexible spines, the second with soft rays and jugular pelvic fins which are separated from each other, but each connected with the pectoral fin base by a membrane. The Indo-Pacific species of the family were revised by Fricke ‡Author to whom correspondence should be addressed. Tel.: +49 9343 600801; email: [email protected] #urn:lsid:zoobank.org:pub:AA1A56B7-E04A-4F22-9EF7-04495BC7AD85

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R . F R I C K E E T A L.

(1983a), who distinguished 126 valid species from the area, including three species from the Mascarenes. Fricke (2002), in a checklist of the callionymids of the world, listed a total of 182 valid species in 10 genera. Subsequently, six additional species were described (Callionymus kanakorum Fricke 2006 and Protogrammus antipodus Fricke 2006 from New Caledonia, Tonlesapia tsukawakii Motomura & Mukai 2006 and Tonlesapia amnica Ng & Rainboth 2011 from Cambodia and Vietnam, respectively, Synchiropus tudorjonesi Allen & Erdmann 2012 from Papua, Indonesia and Callionymus profundus Fricke & Golani 2013 from the northern Red Sea), and Eleutherochir mccaddeni Fowler 1941 was removed from the synonymy of Eleutherochir opercularis (Valenciennes 1837) by Yoshigou et al. (2006), bringing the worldwide total to 189 species in the family. Fricke (1981) defined the kaianus species-group of the genus Callionymus as having a combination of large eyes, four flexible spines in the first dorsal fin, nine rays each in the second dorsal and anal fins, one or two median unbranched caudal fin rays that are often filamentous, a characteristic shape of the preopercular spine with a small antrorse barb in addition to one to three larger curved points on the dorsal margin and (usually) a characteristic black blotch on the third membrane of the first dorsal fin. Fricke (1981) distinguished 13 species in the group, most of which are endemic in small distribution ranges within the Indo-West Pacific. Nakabo (1982) described the genus Bathycallionymus with Callionymus kaianus Günther 1880 as the type species. Fricke (2002) treated Bathycallionymus (the former kaianus species-group) as a subgenus of the genus Callionymus, and distinguished 18 species; reasons for the treatment as a subgenus rather than a genus, including the monophyly of the Callionymus clade, the invalidity of Nakabo’s alternative classification and the preference for nomenclatural stability were discussed by Fricke (2002). Fricke (2006) described C. kanakorum, another member of the subgenus, from New Caledonia, and later (Fricke et al., 2011) recorded the species from Espiritu Santo, Vanuatu. Fricke & Golani (2013) described C. profundus from the northern Red Sea. Species of the subgenus Bathycallionymus live on deeper soft bottoms on the continental slope or on seamounts; they bury in the substratum, usually only leaving the eyes visible. Callionymids typically occur in harem groups, with one male controlling a larger home range living together with several females. Spawning usually takes place around dusk; the courting pair ascends and releases the eggs well above the seabed, following a complex courtship behaviour where the spreading of the first dorsal fin or flashing blue ‘lights’ (iridescent blue spots) are frequent motor patterns. The eggs and larvae are pelagic; during transformation into juveniles they shift to a benthic life style. A single specimen of an undescribed species of the subgenus, which was previously identified as Callionymus bentuviai (non Fricke, 1981), was trawled in deep water off the southern coast of Oman, north-western Indian Ocean. As the species is apparently extremely rare, and very difficult to collect, the new species is described here on the basis of a single specimen, bringing the total number of species known in the subgenus to 20.

MATERIALS AND METHODS Methods follow Fricke (1983a); fin-ray counts follow Fricke (1983b). The starting point for length measurements is the middle of the upper lip. Standard length (LS ) was measured from

© 2014 The Fisheries Society of the British Isles, Journal of Fish Biology 2014, 85, 1303–1319

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the tip of the upper lip to the middle of the urohyal and caudal fin base. The predorsal (1) length was measured from the middle of the upper lip to the base of the first spine of the first dorsal fin; the predorsal (2) length corresponded to the base of the first ray of the second dorsal fin. The last ray of the second dorsal and anal fins is always divided at its base; counts in the key include this divided ray as one. For the pectoral fins, two values for the fin-ray number are provided, as the numbers of the left and the right fin are counted. In the identification keys, males and females are keyed out separately only if there are significant differences between male and female morphology and colouration, and if females of two different species are more similar to each other than to the males of the same species. Species are classified based on Fricke (2002); the classification differs in some respect from Eschmeyer (2014) who inconsistently follows usage since 1990 and variably applies the classifications of Nakabo (1982) and Fricke (2002). Nomenclature follows Eschmeyer (2014). References and journals follow Fricke (2014) and Fricke & Eschmeyer (2014a); abbreviations of museum collections (see below) follow Fricke & Eschmeyer (2014b). The holotype was collected using a bottom trawl for 2⋅78 km (1⋅5 nautical miles) at a speed of c. 5⋅6 km h−1 (3 knots), in a tow of 30 min. A demersal trawl with a 40 mm cod-end was used.

RESULTS CALLIONYMUS OMANENSIS NEW SPECIES

Oman dragonet #urn:lsid:zoobank.org:act:BC67390A-3537-4D66-9E93-F418CCE71A48 Holotype [Figs 1, 2 and 4(a)]

SMF 34898, 111⋅3 mm LS , male, off Salalah City, southern Oman, Arabian Sea, north-western Indian Ocean, 16∘ 23′ 19⋅11′′ N; 54∘ 34′ 52⋅02′′ E, 500 m depth, bottom trawl, R.V. Al-Mustaquila, 2007. Diagnosis

The new species can be distinguished from all other members of the subgenus Bathycallionymus by the combination of the following characteristics: a small branchial opening; head short (3⋅9 in LS ); eye large (2⋅4 in head length); preopercular spine with a long, upcurved main tip, a small antrorse barb and a larger antrorse spine, and a strong antrorse spine at its base, ventral margin smooth; first dorsal fin slightly higher than second dorsal fin (male); second dorsal fin distally straight; 17 pectoral fin rays; caudal fin distally slightly pointed, with two median unbranched rays bearing short filaments; first dorsal fin with basal black spot reaching from first to fourth membranes, third membrane with an ocellated distal black blotch; second dorsal fin with vertical dark grey bars; distal three-fourths of anal fin black; upper half of caudal fin with oblique dark grey bars; pelvic fin dark grey, second ray basally with a black blotch. The new species differs from the similar C. profundus (Fig. 3) in a more slender preopercular spine with the distance of the two dorsal points more than the length of the longest point (stouter, distance less than the longest point in C. profundus; see Fig. 4), 17 pectoral fin rays (18–20 pectoral fin rays in C. profundus), the dark pelvic fin with a black blotch basally on second ray (pale in C. profundus, with a few dark spots), the first dorsal fin with a basal black blotch reaching from first to fourth membranes, and a distal ocellate black blotch on third membrane (basal black blotch only reaching from second to third membranes, and third membrane distally dark but not ocellate in C. profundus), the distal three-fourths of the anal fin dark grey (only distal one-fourth of the anal fin dark


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Fig. 1. Callionymus omanensis n. sp., SMF 34898, holotype, male, 111⋅3 mm standard length, caught off Salalah City, southern Oman, 500 m depth. Scale indicates 10 mm.

in C. profundus), the thorax in the male plain white (dark grey in C. profundus), the snout dorsally with an oblique dark band (without a band, but with a few dark spots in C. profundus), the dorsal part of the pectoral fin base pale but with a dark streak behind the pectoral fin (dorsal part black, no dark streak behind pectoral fin in C. profundus), and the dorsal half of the caudal fin with four oblique dark streaks (pale, with a few dark spots in C. profundus). Callionymus omanensis is distinguished from Callionymus bentuviai Fricke 1981 in having just a single dorsal point on the preopercular spine (additional to the small antrorse barb and the main tip; two points in C. bentuviai), 17 pectoral fin rays (19–22 in C. bentuviai), the short median filaments of the caudal fin (very long in C. bentuviai), the anal fin with the distal three-fourths dark grey (distal one-fourth dark in C. bentuviai), the dorsal half of the caudal fin with four oblique dark streaks (with one horizontal dark streak in C. bentuviai), and the second membrane of the first dorsal fin of the male not deeply incised (deeply incised in C. bentuviai). Callionymus omanensis differs from Callionymus africanus (Kotthaus 1977) in having just a single dorsal point on the preopercular spine [additional to the small antrorse barb and the main tip; two points in C. africanus; Fig. 4(c)], first dorsal fin with the basal dark blotch reaching from first to fourth membranes, and a distal black ocellus on third

Fig. 2. Callionymus omanensis n. sp., SMF 34898, holotype, male, 111⋅3 mm standard length, caught off Salalah City, southern Oman, 500 m depth. Fresh colouration.



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Fig. 3. Callionymus profundus Fricke & Golani 2013, HUJ 16989, holotype, male, 65⋅1 mm standard length, Eilat, Israel, Gulf of Aqaba, northern Red Sea, 410–480 m depth.

membrane (basal black blotch reaching from second to third membranes, no distal ocellus on third membrane in C. africanus), the colouration of the second dorsal fin (with vertical dark grey bars in C. omanensis n. sp.; with two horizontal rows of dark spots and three rows of white spots in C. africanus), the anal fin with the distal three-fourths dark grey (distal one-fourth dark in C. africanus), and the dorsal half of the caudal fin with four oblique dark streaks (pale, with a scattered dark spots in C. africanus). Description

D IV + viii, 1; A viii, 1; P1 ii–iii, 13–14, i (total 17); P2 I, 5; C (ii), i, 3, ii, 2, (iii). Proportions are given in Table I. Body elongated and depressed. Head depressed. Eyes large. Interorbital very narrow, 4⋅7 in eye diameter. Preopercular spine with a long, upcurved main tip, ventral margin smooth, base with a strong antrorse spine, one small antrorse barb and one large, curved

Fig. 4. Left preopercular spines of (a) Callionymus omanensis n. sp., SMF 34898, holotype, male, 111⋅3 mm standard length (LS ), Oman, (b) Callionymus profundus, HUJ 16989, holotype, male, 65⋅1 mm LS , Israel, Red Sea, (c) Callionymus africanus, RMNH 19984, female, 83⋅0 mm LS , Somalia Basin seamount and (d) Callionymus bentuviai, HUJ 6675, male, 84⋅9 mm LS , Eritrea, Red Sea. Scales indicate 2 mm.


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Table I. Proportions of the holotype of Callionymus omanensis n. sp. Catalogue number proportions Head length in LS Body depth in LS Body width in LS Caudal peduncle length in LS Caudal peduncle depth in LS Eye diameter in LH Preorbital length in LH Interorbital distance in LH Upper jaw length in LH Urogenital papilla length in LH Predorsal (1) length in LS Predorsal (2) length in LS Preanal length in LS Prepectoral length in LS Prepelvic length in LS Length of first spine of first dorsal fin in LS Length of second spine of first dorsal fin in LS Length of third spine of first dorsal fin in LS Length of fourth spine of first dorsal fin in LS Length of first ray of second dorsal fin in LS Length of fifth ray of second dorsal fin in LS Length of last ray of second dorsal fin in LS Length of first ray of anal fin in LS Length of fifth ray of anal fin in LS Length of last ray of anal fin in LS Pectoral fin length in LS Pelvic fin spine length in LS Pelvic fin length in LS Caudal fin length in LS

SMF 34898 3⋅9 8⋅7 5⋅8 4⋅5 21⋅8 2⋅4 4⋅2 19⋅8 3⋅1 9⋅0 3⋅4 2⋅2 1⋅9 2⋅5 3⋅6 4⋅9 5⋅0 5⋅4 7⋅8 5⋅1 5⋅1 4⋅7 8⋅9 7⋅5 6⋅4 5⋅7 15⋅4 3⋅9 2⋅9

LS , standard length; LH , head length.

point on dorsal margin, formula 1 −2 1. Cephalic lateral-line system with a preorbital branch, a short suborbital branch, a disconnected preopercular branch and a supraoccipital commissure connecting lines of opposite sides. Occipital region smooth, with two low bony protuberances. Body lateral-line system without dorsal branches, but with one commissure dorsally on the caudal peduncle connecting the lateral lines of the opposite sides; lateral line reaching along about half the length of the upper median unbranched caudal fin ray. Urogenital papilla relatively long. First dorsal fin slightly higher than second dorsal fin in the male, without filaments, the membranes only slightly incised (sexual dimorphism of the first dorsal fin presumed, judging from the situation in the other species of the subgenus). Second dorsal fin distally straight, rays unbranched, the last divided at its base, posterior branch of last ray slightly elongated in male. Anal fin beginning on vertical through second membrane of second dorsal fin. Anal fin distally mostly straight; anal fin rays unbranched, the last divided at its base. Pectoral fin reaching to base of second anal fin membrane



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when adpressed. Pelvic fin reaching to base of second anal fin ray when adpressed. Membrane connecting fifth pelvic fin ray with pectoral fin base ending opposite eighth pectoral fin ray (starting count from most dorsal pectoral fin rays). Caudal fin in male distally slightly pointed, with the two median unbranched rays elongated, bearing short filaments. Colour immediately after collection (Fig. 2)

Head and body yellowish brown, cheeks yellow, thorax yellowish white. Dorsum with irregularly arranged grey and white spots. Eye dorsally yellowish brown, laterally silvery yellow. Sides of head and opercle with irregularly arranged dark grey blotches. A semicircular dark band behind the pectoral fin base (usually covered by the pectoral fin). Sides of body with one row of large, irregularly shaped dark grey blotches below the lateral line. Dorsal part of caudal peduncle with small white spots. First dorsal fin in the male yellow, with a large black blotch surrounded by white lines basally on first to fourth membranes. A wavy white line distally on first to second membranes. Third membrane with a distal ocellated black blotch. Second dorsal fin yellow, with nine vertically orientated oblique black streaks that are surrounded by yellow pigment and interspersed by white pigment; membranes otherwise yellow, with a distal, horizontal white line. Anal fin basally light grey, distal three-fourths black. Caudal fin white, upper half with four oblique dark grey streaks that are narrowly margined by yellow, lower half with a black streak. Pelvic fin grey. Pectoral fin yellowish, basally white. Colour in preservative

Head and body beige, ventrally white; thorax in male white, without dark pigment. Dorsum with irregularly arranged white spots, and with a short oblique dark streak below the last membrane of second dorsal fin. Snout with a transverse dark grey band. Eye with dark grey pigment. Sides of head and opercle with irregularly arranged grey blotches. A semicircular dark band behind the pectoral fin base (usually covered by the pectoral fin). Sides of body with one row of elongated dark grey blotches below the lateral line. Dorsal part of caudal peduncle with small white spots. First dorsal fin in the male pale, with a large black blotch basally on first to fourth membranes. Third membrane with a distal ocellated black blotch. Second dorsal fin white, with seven vertically orientated oblique dark grey streaks; membranes otherwise pale. Anal fin basally light grey, distal three-fourths black. Caudal fin pale, upper half with four oblique dark grey streaks, lower half with a narrow dark grey streak. Pelvic fin plain dark grey, basally with an elongated black blotch on second ray. Pectoral fin translucent, without markings. Sexual dimorphism

Unknown (only the male holotype is known). Distribution

The species is only known from the type locality, off Salalah, southern Oman, Arabian Sea, north-western Indian Ocean. The holotype was collected at a depth of 500 m. The species is probably restricted to deep water of the southern coast of Oman, judging from the narrow distribution ranges of other species of the subgenus, and the


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presence of similar species in other areas, which are not expected to co-occur with the new species. Etymology

The name of the new species refers to its type locality, Oman. Comparisons

The new species is a member of the subgenus Callionymus (Bathycallionymus) as defined by Fricke (1981, 2002) and Nakabo (1982). It is similar to C. profundus (Fricke & Golani, 2013) from the Gulf of Aqaba, northern Red Sea, and C. bentuviai (Fricke, 1981) from the southern Red Sea in having the second dorsal and anal fins with a straight distal margin, the first dorsal fin without filaments and the caudal fin bearing two filaments. Other species of the subgenus have the second dorsal and anal fins with convex distal margins, one or more filaments in the first dorsal fin of the male, and one or no filaments of the caudal fin. The new species differs from C. profundus, C. bentuviai and C. africanus according to the information given in the Diagnosis. A revised key to callionymid species of the Red Sea and western Indian Ocean is presented below to distinguish C. omanensis n. sp. from potentially co-occurring species. Also, a key to species of the subgenus Bathycallionymus is provided to characterize the new species among other related species (updated, based on a key presented by Fricke & Golani, 2013).

KEY TO SPECIES OF CALLIONYMIDAE OF THE RED SEA AND WESTERN INDIAN OCEAN 1a. Opercle with a free flap of skin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1b. Opercle without a free flap of skin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 2a. Body with lateral fold of skin below lateral line; lower lip without fleshy papillae; A rays unbranched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Diplogrammus) 3 2b. Body without lateral fold of skin below lateral line; dorsal margin of lower lip with a row of erect fleshy papillae; A rays branched . . . . . . . . . . . . . . (Draculo) 6 3a. Second spine of first dorsal fin longer than first spine of second dorsal fin . . . . 4 3b. Second spine of first dorsal fin as long as or shorter than first spine of second dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4a. Body depth 12–13% LS ; pectoral fin rays 16–17 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplogrammus gruveli Smith 1963 4b. Body depth 17–18% LS ; pectoral fin rays 20 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplogrammus pygmaeus Fricke 1981 5a. First spine of first dorsal fin with long filament in male; main tip of preopercular spine upcurved; anal fin without distal black margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplogrammus infulatus Smith 1963 5b. First spine of first dorsal fin without filament in male; main tip of preopercular spine straight; anal fin with narrow distal black margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diplogrammus randalli Fricke 1983



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6a. First dorsal fin with a single spine; second dorsal fin with 11 rays (the last divided at its base); anal fin with 12 branched rays (the last divided at its base) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Draculo celetus (Smith 1963) 6b. First dorsal fin with four spines; second dorsal fin with 9–10 rays (the last divided at its base); anal fin with 10 branched rays (the last divided at its base) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Draculo maugei (Smith 1966) 7a. Lateral line with numerous long branches above and below; preopercular spine simple, without accessory spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracallionymus costatus (Boulenger 1898) 7b. Lateral line mostly unbranched; preopercular spine with one or more accessory spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 8a. Snout shorter than eye diameter; soft D rays branched (in specimens >3 cm); gill opening sublateral; no antrorse spine present at preopercular spine base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Synchiropus) 9 8b. Snout equal to, or longer than, eye diameter; soft D rays unbranched; gill opening dorsal; antrorse spine at base of preopercular spine . . . . . . . . . . (Callionymus) 14 9a. Preopercular spine with one dorsal point additional to main tip . . . . . . . . . . . . . 10 9b. Preopercular spine with two to five dorsal points additional to main tip . . . . . 11 10a. Caudal fin pale or with one distal dark bar; caudal fin length 33–40% LS ; second dorsal fin pale or with distal dark bar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Synchiropus monacanthus Smith 1935 10b. Caudal fin with two dark bars; caudal fin length 27–32% LS ; second dorsal fin spotted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Synchiropus stellatus Smith 1963 11a. Preopercular spine with two dorsal points additional to main tip . . . . . . . . . . . . 12 11b. Preopercular spine with three to five dorsal points additional to main tip . . . . 13 12a. Main tip of preopercular spine straight; preopercular spine always without small antrorse spines at base; first dorsal fin dark, striped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Synchiropus marmoratus (Peters 1855) 12b. Main tip of preopercular spine upcurved; preopercular spine usually with one or two small antrorse points at base; first dorsal fin light, with dark blotches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Synchiropus sechellensis Regan 1908 13a. Dorsal spines filamentous in male; lower pectoral fin base without black spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Synchiropus postulus Smith 1963 13b. Dorsal spines not filamentous in male; lower pectoral fin base with black spot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Synchiropus minutulus Fricke 1981 14a. Both dorsal and ventral margins of preopercular spine with antrorse spines or serrae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 14b. Only dorsal margin of preopercular spine with curved points or antrorse serrae (one or two antrorse spines may be present at base of preopercular spine) . . . . 17 15a. Snout densely covered with numerous spines Callionymus spiniceps Regan 1908 15b. Snout without spines (occasionally except for a small preorbital spine) . . . . . . 16 16a. Over eye tentacle present; lower margin of preopercular spine with one to three curved points additional to antrorse spine at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus muscatensis Regan 1905 16b. Over eye tentacle absent; lower margin of preopercular spine with four to nine curved points additional to antrorse spine at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus oxycephalus Fricke 1980 17a. Upper margin of preopercular spine with small antrorse serrae . . . . . . . . . . . . . 18


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17b. Upper margin of preopercular spine with large curved points (occasionally with one additional antrorse serra) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 18a. Second dorsal fin with eight rays (the last divided at its base) . . . . . . . . . . . . . . 19 18b. Second dorsal fin with nine rays (the last divided at its base) . . . . . . . . . . . . . . . 22 19a. Anal fin with seven rays (the last divided at base) . . . . . . . . . . . . . . . . . . . . . . . . . 20 19b. Anal fin with eight rays (the last divided at base) . . . . . . . . . . . . . . . . . . . . . . . . . 21 20a. Main tip of preopercular spine upcurved; dorsal margin of preopercular spine with five to six small antrorse serrae . . . . . . . . . . . . . . . Callionymus flavus Fricke 1983 20b. Main tip of preopercular spine straight; dorsal margin of preopercular spine with 8–15 small antrorse serrae . . . . . . . . . . . . . . Callionymus delicatulus Smith 1963 21a. First spine of first dorsal fin filamentous; anal fin distally black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus margaretae Regan 1905 21b. First dorsal fin without filaments; anal fin pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus tenuis Fricke 1981 22a. Anal fin with nine rays (the last divided at its base); first spine of first dorsal fin in male separate from second spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus filamentosus Valenciennes in Cuvier & Valenciennes 1837 22b. Anal fin with eight rays (the last divided at its base); first spine of first dorsal fin in male connected with second spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 23a. First spine of first dorsal fin with a long filament . . . . . . . . . . . . . . . . . . . . . . . . . . 24 23b. First spine of first dorsal fin without a filament . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 24a. Lower margin of caudal fin pale . . . . . . . . . . . . Callionymus aagilis Fricke 1999 24b. Lower margin of caudal fin black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 25a. Third spine of first dorsal fin distally with black blotch; dorsal margin of preopercular spine with 6–12 small antrorse serrae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus gardineri Regan 1908, male 25b. Third spine of first dorsal fin without distal black blotch; dorsal margin of preopercular spine with three to six small antrorse serrae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus margaretae Regan 1905 26a. Anal fin with a distal black bar . . . . . . . . . . . . . Callionymus persicus Regan 1905 26b. Anal fin pale . . . . . . . . . . . . . . . . . . . . . Callionymus gardineri Regan 1908, female 27a. First dorsal fin with three spines; jaws extremely protractile, forming a broad tube when protracted . . . . . . . . . . . . . . . . . . . . . . Callionymus hindsii Richardson 1844 27b. First dorsal fin with four spines; jaws moderately protractile, narrow when protracted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 28a. Anal fin with seven rays (the last divided at base) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus stigmatopareius Fricke 1981 28b. Anal fin with eight or nine rays (the last divided at base) . . . . . . . . . . . . . . . . . . 29 29a. Dorsal margin of preopercular spine with a small antrorse barb near main tip, and with one or two large curved points . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 29b. Dorsal margin of preopercular spine without a small antrorse barb near main tip, but with one or more large curved points . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 30a. Caudal fin distally convex or slightly pointed, with one or two filaments . . . . 31 30b. Caudal fin distally convex, without filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 31a. Preopercular spine with two curved points (additional to small antrorse barb) on dorsal side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus bentuviai Fricke 1981 31b. Preopercular spine with one curved point (additional to small antrorse barb) on dorsal side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32



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32a. Pectoral fin rays 17; preopercular spine slender, distance of the two dorsal points more than the length of the longest point [Fig. 4(a)]; pelvic fin dark, with a black blotch basally on second ray; first dorsal fin (male) with a basal black blotch reaching from first to fourth membranes; dorsal half of caudal fin with four oblique dark streaks . . . . . . . . . . . . . . . . . . . . . . Callionymus omanensis n. sp. 32b. Pectoral fin rays 18–20; preopercular spine stout, distance of the two dorsal points less than the length of the longest point [Fig. 4(b)]; pelvic fin pale, with a few dark spots; first dorsal fin (male) with a basal black blotch reaching from second to third membranes; dorsal half of caudal fin pale, with a few dark spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus profundus Fricke & Golani 2013 33a. Second dorsal fin with vertical black streaks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 33b. Second dorsal fin with horizontal rows with white and dark blotches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus africanus (Kotthaus 1977), male 34a. Black blotch on first dorsal fin on first or second to third membranes; cheeks pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus kotthausi Fricke 1981 34b. Black blotch on first dorsal fin only on third membrane; cheeks with spots or suborbital dark streak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 35a. Suborbital dark streak present; second half of anal fin black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus africanus (Kotthaus 1977), female 35b. No suborbital dark streak, but many brown blotches encircled with darker brown on cheeks; anal fin pale, distal margin dusky . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus regani Nakabo 1979 36a. Dorsal margin of preopercular spine with one or two curved points . . . . . . . . . 37 36b. Dorsal margin of preopercular spine with three to seven curved points . . . . . . 38 37a. Main tip of preopercular spine long, curved; preopercular spine not upcurved at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus carebares Alcock 1890 37b. Main tip of preopercular spine short, straight; preopercular spine upcurved at base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus mascarenus Fricke 1983 38a. Caudal fin elongated, with two long median filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus cooperi Regan 1908 38b. Caudal fin distally convex, without filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 39a. Second spine of first dorsal fin longer than first (may be both filamentous in males) Callionymus erythraeus Ninni 1934 39b. Second spine of first dorsal fin shorter than first . . . . . . . . . . . . . . . . . . . . . . . . . . 40 40a. Preopercular spine strongly upcurved at base; male with black blotch surrounded by white posteriorly on first dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus mascarenus Fricke 1983 40b. Preopercular spine only slightly upcurved at base; male without black blotch posteriorly on first dorsal fin. . . . . . . . . . . . . . . . . Callionymus marleyi Regan 1919

KEY TO THE SPECIES OF SUBGENUS BATHYCALLIONYMUS OF THE GENUS CALLIONYMUS 1a. Dorsal margin of preopercular spine with a small antrorse barb additional to large, curved points; head in LS more than 3⋅2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2


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1b. Dorsal margin of preopercular spine usually without a small antrorse barb additional to large, curved points; head in LS 2⋅7–3⋅1; northern Indian Ocean . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus carebares Alcock 1890 2a. Second dorsal and anal fins very high, males with convex distal margins . . . . . 3 2b. Second dorsal and anal fins relatively low, with straight distal margins . . . . . . . 8 3a. First spine of first dorsal fin filamentous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3b. First spine of first dorsal fin not filamentous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 4a. Second dorsal fin relatively low, first ray about 1⋅0, fifth ray about 0⋅9 in head length; preopercular spine with two curved points (additional to one small antrorse barb) on its dorsal side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4b. Second dorsal fin high, first ray c. 1⋅1, fifth ray c. 0⋅8 in head length; preopercular spine with one large curved point (additional to one small antrorse barb) on its dorsal side. South China Sea . . . . . . . . . . . . . . Callionymus altipinnis Fricke 1981 5a. Caudal fin distally convex, with one long filament; filament on first spine of first dorsal fin long, nearly twice as long as second spine; Taiwan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus formosanus Fricke 1981 5b. Caudal fin distally convex, with two filaments; filament on first spine of first dorsal fin relatively short, not twice as long as second spine; Indonesia or Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6a. First dorsal fin with a large black blotch covering most of third membrane, but without a black blotch on first membrane; second dorsal fin pale, without markings; upper half of pectoral fin base pale; north-western Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus bifilum Fricke 2000 6b. First dorsal fin with a large black blotch distally on second and third membranes, and a small black blotch distally on first membrane; second dorsal fin with vertical dark streaks; upper half of pectoral fin base with a dark blotch; western Indonesia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus semeiophor Fricke 1983 7a. Main tip of preopercular spine long and slender; distal half of anal fin black, tips of fin rays white; distal margin of caudal fin regularly shaped; black blotch on third membrane of first dorsal fin relatively large, central in position (not reaching distal margin); Philippines . . . . . . . . . . . . . . . . Callionymus guentheri Fricke 1981 7b. Main tip of preopercular spine short; anal fin with a distal black streak on each membrane, distal two-thirds of membrane and tips of fin rays dark brown; distal margin of caudal fin irregularly shaped; black blotch on third membrane of first dorsal fin very small, extremely distal in position; Japan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus sokonumeri Kamohara 1936 8a. Caudal fin convex, without filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 8b. Caudal fin convex or slightly pointed, with one or two filaments . . . . . . . . . . . . 16 9a. Anal fin with a narrow distal yellow margin or colourless; sides of body with a row of indistinct brownish blotches or without blotches . . . . . . . . . . . . . . . . . . . 10 9b. Anal fin with a dark brown or black margin; sides of body with a row of small distinct black blotches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 10a. First spine of first dorsal fin with a long filament, more than twice as long as second spine; western Indonesia . . . . . . . . . . . . Callionymus bleekeri Fricke 1983 10b. First spine of first dorsal fin not filamentous, about as long as second spine; south-western Indian Ocean . . . . . . . . . . . . . . . . Callionymus regani Nakabo 1979 11a. Main tip of preopercular spine long and slender; India . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus kotthausi Fricke 1981



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11b. Main tip of preopercular spine short . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 12a. First spine of first dorsal fin filamentous; pelvic fin distally spotted . . . . . . . . . 13 12b. First spine of first dorsal fin not filamentous; pelvic fin distally with a dark area, but not spotted . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 13a. First spine of first dorsal fin with a relatively long filament, at least 1⋅5 times as long as second spine; upper half of caudal fin spotted, but without a distal dark bar; Wallis and Futuna . . . . . . . . . . . . . . . . . . . . . . Callionymus futuna Fricke 1998 13b. First spine of first dorsal fin with a relatively short filament, at most 1⋅2 times as long as second spine; upper half of caudal fin not spotted, but with a small distal dark bar; New Caledonia . . . . . . . . . . . . . . . Callionymus kanakorum Fricke 2006 14a. First dorsal fin much higher than second dorsal fin, first spine >1⋅5 times longer than first ray of second dorsal fin; northern Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus kailolae Fricke 2000 14b. First dorsal fin only slightly higher than second dorsal fin, first spine not more than 1⋅2 times longer than first ray of second dorsal fin . . . . . . . . . . . . . . . . . . . 15 15a. Preopercular spine with one dorsal point (additional to a small antrorse barb); pectoral fin base with a large dark brown area; back marbled with olive green in fresh specimens; opercle with large white spots; lower part of caudal fin distally black; second dorsal fin with a basal row of dark blotches; western Indonesia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus whiteheadi Fricke 1981 15b. Preopercular spine with two dorsal points (additional to a small antrorse barb); pectoral fin base with two black streaks; back yellowish brown, with white blotches surrounded by semicircular dark bands; opercle with small dark spots; lower part of caudal fin with a broad curved dark band, but distally colourless; second dorsal fin without a basal row of blotches or spots; East Africa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus africanus (Kotthaus 1977) (female) 16a. First spine of first dorsal fin with a long filament . . . . . . . . . . . . . . . . . . . . . . . . . . 17 16b. First spine of first dorsal fin without a filament . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 17a. Main tip of preopercular spine short, largest point on its dorsal margin with a basal hook; second dorsal fin colourless, with transverse white lines; sides of body with a row of distinct black blotches; Indonesia . . Callionymus kaianus Günther 1880 17b. Main tip of preopercular spine long and slender, largest point on its dorsal margin without a basal hook; second dorsal fin with rows of blotches; sides of body with a row of indistinct brownish blotches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 18a. Anal fin with a dark margin; second dorsal fin with two rows of white blotches and a distal dark margin; main tip of preopercular spine c. 2⋅5–3⋅0 times longer than largest point on the dorsal side; body with light blotches edged with semicircular dark lines; north-eastern Australia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus moretonensis Johnson 1971 18b. Anal fin pale, without a dark margin; second dorsal fin with a basal and two to three more distal rows of dark spots; main tip of preopercular spine c. 1⋅0–1⋅5 times longer than largest point on the dorsal side; body with minute blackish spots forming rings and blotches; Japan . . . . . . . . . . . Callionymus ochiaii Fricke 1981 19a. Caudal fin with two long filaments which are nearly twice as long as remainder of fin; southern Red Sea . . . . . . . . . . . . . . . . . . Callionymus bentuviai Fricke 1981 19b. Caudal fin with one or two relatively short filaments which are not longer than the remainder of the fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20


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20a. Preopercular spine with two curved points (additional to small antrorse barb) on dorsal side; Indian Ocean or western Pacific . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 20b. Preopercular spine with one curved point (additional to small antrorse barb) on dorsal side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 21a. First dorsal fin with a large dark area, with a branch reaching anteriorly to first spine; second dorsal fin with vertical dark streaks; Indonesia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus semeiophor Fricke 1983 (female) 21b. First dorsal fin with a black blotch on third membrane (rarely with an additional blotch on the same membrane), occasionally reaching from second to fourth spines; second dorsal fin with rows of dark and white spots (but without vertical dark streaks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 22a. Cephalic lateral line in postorbital region with a long branch running downwards; first dorsal fin with a large black blotch basally on the third membrane, with one or two branches reaching to second membrane; pectoral fin base with two dark streaks; anal fin with a distal black margin (usually including tips of fin rays); caudal fin without two median transverse black lines; second dorsal fin with a basal, a median and a distal row of light spots and two median rows of black spots; East Africa . . . . . . . . . . . . . . . . . . . . Callionymus africanus (Kotthaus 1977) 22b. Cephalic lateral line in postorbital region without a long branch; first dorsal fin with a relatively small distal black blotch on the third membrane; pectoral fin base with a dark area; distal half of anal fin black, tips of fin rays white; caudal fin with two median transverse black lines; second dorsal fin with a basal and a median row of black spots; Philippines . . . . Callionymus guentheri Fricke 1981 (female) 23a. Pectoral fin rays 17; preopercular spine slender, distance of the two dorsal points more than the length of the longest point [Fig. 4(a)]; pelvic fin dark, with a black blotch basally on second ray; first dorsal fin (male) with a basal black blotch reaching from first to fourth membranes; dorsal half of caudal fin with four oblique dark streaks; southern Oman, north-western Indian Ocean . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callionymus omanensis n. sp. 23b. Pectoral fin rays 18–20; preopercular spine stout, distance of the two dorsal points less than the length of the longest point [Fig. 4(b)]; pelvic fin pale, with a few dark spots; first dorsal fin (male) with a basal black blotch reaching from second to third membranes; dorsal half of caudal fin pale, with a few dark spots; northern Gulf of Aqaba, Red Sea . . . . . . . . . Callionymus profundus Fricke & Golani 2013

DISCUSSION Most species of the subgenus Bathycallionymus have narrow, restricted distribution ranges. The typical habitat includes soft bottom (sand or mud) on the continental slope; this habitat is extremely patchy, and it is difficult to collect here. Furthermore, similar to other callionymids, members of the subgenus Bathycallionymus live in small harem groups, and also in a continuous stretch of soft bottom habitat in a very patchy distribution. The steep slope habitat, together with the patchy distribution type, and extremely restricted distribution ranges, suggests that most species are very rare, and due to difficulty in collecting they are even more rare in collections. Several species of the group are known by a single specimen only (C. bleekeri, C. formosanus, C. kailolae, C. omanensis n. sp. and C. profundus). Larger quantities of specimens of the subgenus are only



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collected if they occur on seamounts which have a flat top covered with soft bottom habitats in the appropriate depth range. The new species is most similar to species from the northern and southern Red Sea (C. profundus and C. bentuviai) and East Africa (C. africanus). A barrier separating the distribution ranges of C. omanensis n. sp. from those of the neighbouring C. bentuviai and C. africanus is unknown, although an upwelling zone in southern Arabia (Wyrtki, 1971) may play a role as a dispersal barrier. In an alternative classification by Nakabo (1982), C. omanensis n. sp. would be a member of the genus Bathycallionymus. Here, Bathycallionymus is treated as a subgenus of Callionymus. C O M PA R AT I V E M AT E R I A L

Callionymus africanus: ZIM 5533 (holotype), seamount north-east of Mombasa, Kenya; BMNH 1939.5.24.1422 (one specimen), near Zanzibar, Tanzania; RMNH 19984 (six specimens), seamount on Chain Ridge, off Somalia; ZIM 5534 (30 paratypes), seamount north-east of Mombasa, Kenya. Callionymus altipinnis (all China): MSL 0001 (holotype), Hong Kong; CAS 46967 (10 paratypes), Hainan; CAS 46968 (one paratype), Hainan; CAS 46969 (one paratype), Hainan. Callionymus bentuviai (all Eritrea, southern Red Sea): HUJ 9935 (holotype); HUJ 6674 (two specimens); HUJ 6675 (one specimen); HUJ 8068 (two paratypes); HUJ 10463 (two specimens); ZIM 5532 (two paratypes). Callionymus bifilum: CSIRO CA.3532 (holotype), Joseph Bonaparte Gulf, Western Australia; BMNH 1892.1.14.26-27 (two paratypes), Holothuria Bank, Western Australia. Callionymus bleekeri: RMNH 4872 (holotype), Java, Indonesia. Callionymus carebares: BMNH 1890.11.28.18–24 (seven syntypes), Ganjam, India; MNHN 1890-0335-0340 (six syntypes), Ganjam, India; ZSI F.12740-F.12741 (two syntypes), Ganjam, India; ZSI F.12742 and F.12858 (15 syntypes), Ganjam, India; AMS I.28730-001 (one specimen), Maputo, Mozambique; BMNH 1903.5.14.34 (one specimen), Karachi, Pakistan; BMNH 1904.5.25.218-220 (three specimens), Sea of Oman, Oman; BMNH 1939.5.24.1384 (one specimen), Gulf of Oman, Oman; BMNH 1939.5.24.1385-1409 (24 specimens), Arabian Sea; BMNH 1939.5.24.1410-1421 (15 specimens), Gulf of Aden, Yemen; FMNH 5740 (one specimen), Sea of Oman, Oman; IRSN 1797 (two specimens), Bay of Bengal, India; SMNS 23508 (one specimen), Thailand, Andaman Sea (new record). Callionymus formosanus: CAS 46972 (holotype), Pescadores Strait, Taiwan. Callionymus futuna (all Wallis and Futuna, Futuna Island shelf): MNHN 1995-0521 (holotype); MNHN 1995-0522 (two paratypes); SMNS 18823 (one paratype). Callionymus guentheri (all Philippines): BMNH 1879.5.14.567 (holotype), Zamboanga; CAS 32668 (two paratypes), Batangas, Luzon; CAS 32801 (one paratype), Marinduque; CAS 32897 (11 paratypes), Batangas, Luzon; CAS 32905 (seven paratypes), Camarines Sur; CAS 32916 (five paratypes), Quezon, Luzon; CAS 32997 (one paratype), Balayan Bay; CAS 33067 (one paratype), Luzon; CAS 33703 (one paratype), Balayan Bay; CAS 33864 (three paratypes), Camarines Sur; CAS 33879 (one paratype), Marinduque; CAS 34074 (one paratype), Marinduque; CAS 34154 (seven paratypes), Marinduque; CAS 34190 (two paratypes), Ragay Gulf; CAS 34197


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(nine paratypes), Camarines Sur; CAS 34205 (one paratype), Marinduque; CAS 34272 (one paratype), Ragay Gulf; CAS 34278 (four paratypes), Marinduque; CAS 34286 (three paratypes), Marinduque; CAS 34401 (one paratype), Balayan Bay; CAS 34426 (one paratype), Batangas, Luzon; CAS 34468 (one paratype), Balayan Bay; CAS 46966 (four paratypes), Quezon; SMNS 8510 (two specimens), Occidental Mindoro; SMNS 9097 (one specimen), Occidental Mindoro; USNM 150916 (one specimen), Cabuyan Grande Island. Callionymus kaianus (all Kai Islands, Maluku, Indonesia): BMNH 1879.5.14.565 (holotype); ZMUC P.6498 (one specimen); ZMUC P.6499 (one specimen). Callionymus kailolae: AMS I.22807-021 (holotype), north of Port Hedland, Western Australia. Callionymus kanakorum: MNHN 2000-5519 (holotype), Grande Terre, New Caledonia; SMNS 8550 (one paratype), Grande Terre, New Caledonia; SMNS 12047 (one paratype), Grande Terre, New Caledonia. Callionymus kotthausi: ZIM 5535 (holotype), Cochin, India; ZIM 5536 (11 paratypes), Cochin, India. Callionymus moretonensis (all Australia): AMS I.15608-001 (holotype), Cape Moreton, Queensland; CAS 24764-24767 (four paratypes), Cape Moreton, Queensland; QM I.9156–I.9157 (two paratypes), Cape Moreton, Queensland; SMNS 12172 (five specimens), Arafura Sea, Northern Territory; SMNS 12173 (eight specimens), Arafura Sea, Northern Territory; SMNS 14791 (13 specimens), off Newcastle, New South Wales; WAM P.25739-003 (one specimen), Cape Moreton, Queensland. Callionymus ochiaii: FAKU 23261 (holotype), Kagoshima Prefecture, Japan; FAKU 23257-23260 and 23275 (five paratypes), Kagoshima Prefecture, Japan; USNM 160485 (one specimen), Goto Island, Japan. Callionymus profundus: HUJ 16989 (holotype), Eilat, Israel, Gulf of Aqaba, Red Sea. Callionymus regani (all Saya de Malha Bank, western Indian Ocean): HUMZ 72408 (holotype); BMNH 1908.3.23.263 (one specimen); HUMZ 72325 (one paratype); HUMZ 72359 (one paratype); HUMZ 42405 and 42407 (two paratypes); HUMZ 73395 (one paratype); HUMZ 73448–73449 (two paratypes); HUMZ 73681 (one paratype). Callionymus semeiophor: NTM S.10764-001 (holotype), Sumbawa, Indonesia; NTM S.10760-002 (one paratype), Lombok, Indonesia. Callionymus sokonumeri: NSMT P.21026-21027 (2), Mimase, Japan; NSMT 23873 (one specimen), Kagoshima, Japan. Callionymus whiteheadi: BMNH 1980.6.20.1 (holotype), Bali, Indonesia; BMNH 1980.11.25.2 (one paratype), Java, Indonesia. We would like to thank the following persons for information, loan of specimens or permission to examine specimens under their care: D. F. Hoese, J. Leis, M. McGrouther, J. Paxton (AMS), O. Crimmen (BMNH), W. N. Eschmeyer, T. Iwamoto (CAS), P. Last (CSIRO), T. Nakabo (FAKU), D. Golani (HUJ), K. Amaoka (HUMZ), J.-P. Gosse (IRSN), P. Pruvost (MNHN), K.-C. Au (MSL), K. Matsuura (NSMT), H. K. Larson (NTM), R. J. McKay (QM), M. J. P. vam Oijen (RMNH), S. Smith, J. T. Williams (USNM), G. R. Allen, J. B. Hutchins, S. Morrison (WAM), H. Wilkens (ZIM), J. G. Nielsen (ZMUC) and P. K. Talwar (ZSI). We are grateful to T. Alpermann (SMF) for providing a catalogue number for the holotype of the new species. We would also like to thank the Ministry of Fisheries Wealth, Marine Science and Fisheries Centre, the Ministry of Fisheries Wealth and the Directorate of Agriculture and Fisheries Developmental Fund for giving us the opportunity to work on the fish samples to examine the qualitative and quantitative distribution



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of marine organisms in the Sultanate of Oman, and for providing the appropriate financial support.

References Fricke, R. (1981). The kaianus-group of the genus Callionymus (Pisces: Callionymidae), with descriptions of six new species. Proceedings of the California Academy of Sciences(Series 4) 42, 349–377. Fricke, R. (1983a). Revision of the Indo-Pacific Genera and Species of the Dragonet Family Callionymidae (Teleostei). Braunschweig: J. Cramer. Fricke, R. (1983b). A method of counting caudal fin rays of actinopterygian fishes. Braunschweiger Naturkundliche Schriften 1, 729–733. Fricke, R. (2002). Annotated checklist of the dragonet families Callionymidae and Draconettidae (Teleostei: Callionymoidei), with comments on callionymid fish classification. Stuttgarter Beiträge zur Naturkunde, Serie A (Biologie) 645, 1–103. Fricke, R. (2006). Two new species and a new record of dragonets from New Caledonia (Teleostei: Callionymidae). Stuttgarter Beiträge zur Naturkunde, Serie A (Biologie) 696, 1–14. Fricke, R. & Golani, D. (2013). Callionymus profundus n. sp., a new species of dragonet from the Gulf of Aqaba (Gulf of Eilat), Red Sea (Teleostei: Callionymidae). Stuttgarter Beiträge zur Naturkunde A, Neue Serie 6, 277–285. Fricke, R., Earle, J. L., Pyle, R. L. & Séret, B. (2011). Checklist of the fishes. In The Natural History of Santo (Bouchet, P., Le Guyader, H. & Pascal, O., eds), pp. 343–409. Paris: MNHN and Marseille: PNI. Nakabo, T. (1982). Revision of the genera of dragonets (Pisces, Callionymidae). Publications of the Seto Marine Biological Laboratory 27, 77–131. Wyrtki, K. (1971). Oceanographic Atlas of the International Indian Ocean Expedition. Washington, DC: National Science Foundation. Yoshigou, H., Ohta, I. & Yoshino, T. (2006). First record of a callionymid fish, Eleutherochir mccaddeni, from Japan. Japanese Journal of Ichthyology 52, 189–193.

Electronic References Eschmeyer, W. N. (Ed.) (2014). Catalog of Fishes. San Francisco, CA: California Academy of Sciences. Available at http://research.calacademy.org/research/Ichthyology/Catalog/ fishcatmain.asp/ (last accessed 6 April 2014). Fricke, R. (Ed.) (2014). Literature in the Catalog of Fishes. San Francisco, CA: California Academy of Sciences. Available at http://research.calacademy.org/research/Ichthyology/ Catalog/fishcatmain.asp/ (last accessed 6 April 2014). Fricke, R. & Eschmeyer, W. N. (2014a). Journals in the Catalog of Fishes. San Francisco, CA: California Academy of Sciences. Available at http://research.calacademy.org/research/ Ichthyology/Catalog/journals.asp/ (last accessed 6 April 2014). Fricke, R. & Eschmeyer, W. N. (2014b). A Guide to Fish Collections in the Catalog of Fishes. San Francisco, CA: California Academy of Sciences. Available at http://research. calacademy.org/research/Ichthyology/Catalog/collections.asp/ (last accessed 6 April 2014).


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