Smithiana Bulletin The South African Institute for Aquatic Biodiversity ISSN: 1684-4130 Num. 13, 2011, pp. 51-73

Taxonomic review of Western Indian Ocean goatfishes of the genus Mulloidichthys (Family Mullidae), with description of a new species and remarks on colour and body form variation in Indo-West Pacifi c species

Franz Uiblein

Institute of Marine Research, Nordnesgaten 33, 5817 Bergen, Norway, and South African Institute for Aquatic Biodiversity, Grahamstown, South Africa email: franz@imr.no

Received 26 August; accepted 2 October 2010

Code Number: sm11004

ABSTRACT.

The taxonomy of the goatfish species of the genus Mulloidichthys from the Western Indian Ocean is reviewed. Four species are recognized: M. ayliffe sp. nov., M. flavolineatus, M. pfl uegeri and M. vanicolensis. As colour may vary considerably in live fish and fades in preserved fish, and meristic characters are rather conservative in this genus, a large set of 41 morphometric characters was also examined. All seven valid Mulloidichthys species are included in the comparisons. The Western Indian Ocean M. ayliffe sp. nov. differs from its Pacific sister species M. mimicus in number of lateral-line scales, several morphometric features, the position of the conspicuous bluish dorso-midlateral body stripe in relation to the lateral line, and the width of the yellow mid-lateral body stripe. The new species can also be distinguished from the other congenerics by colour, meristic and morphometric characters. A key is provided for the Western Indian Ocean species, which includes the South Pacifi c species, M. mimicus. The fi rst evidence for geographic variation in body form among Mulloidichthys species with marked differences between the Pacific and Indian Ocean is presented. These results complement earlier findings of interoceanic divergence in the number of gill rakers in M. vanicolensis. The implications of these fi ndings are discussed with respect to the need for DNA–based comparative studies on population differentiation and the ecological functions of colour patterns in Mulloidichthys species.

RÉSUMÉ.

La taxonomie de l’espèce de rouget et du genre Mulloidichthys de l’ouest de l’Océan Indien est passée en revue. Quatre espèces sont reconnues : M. ayliffe sp. nov., M. fl avolineatus, M. pfluegeri et M. vanicolensis. Étant donné que la couleur peut varier pour les poissons vivants et se détériorer pour les poissons conservés et que les caractéristiques meristiques de ce genre sont plutôt conservatives (minimales), un échantillon important de 41 caractéristiques morphometriques a été également analysé. Dans ces comparaisons on trouvera également toutes les 7 espèces vivantes de Mulloidichthys. L’espèce M. ayliffe de l’ouest de l’Océan Indien se distingue de sa sœur espèce du Pacifi que M. mimicus par rapport au nombre d’écailles en ligne latérale, plusieurs éléments morphometriques, la position de la ligne dorsale par rapport à la ligne latérale ainsi que la dimension du corps latéral du milieu. La nouvelle espèce peut également être identifiée par sa couleur et ses caractéristiques meristiques et morphometriques. Le moyen de comprendre les espèces de l’ouest de l’Océan Indien, y compris l’espèce Pacifi que M. mimicus, est présenté dans cette analyse. La première preuve de la variation géographique dans la morphologie des espèces Mulloidichthys avec des différences importantes entre le Pacifi que et l’Indien est présentée. Les résultats complètent les trouvailles antérieures sur les divergences interocéaniques spécialement en ce qui concerne la famille de M. vanicolensis.

KEY WORDS: morphology, colour patterns, new species, Mullidae, Mulloidichthys, Western Indian Ocean

Recent reviews of the goatfi sh genera Parupeneus and Upeneus (Mullidae) from the Western Indian Ocean region have resulted in descriptions of ten new species, resurrection of two species, and several other novel insights into their diversity and distribution (Randall & Heemstra 2009; Randall & King 2009; Uiblein & Heemstra 2010; Uiblein & Heemstra 2011; Uiblein & Heemstra in press). These studies have highlighted the obvious need for regional taxonomic reviews of goatfishes including the genus Mulloidichthys in which three species are known from the Western Indian Ocean, M. fl avolineatus (Lacepède 1801, M. pfl uegeri (Steindachner 1900), and M. vanicolensis (Valenciennes 1831) (Ben Tuvia, in Smith & Heemstra 1986; Myers 1989).

Earlier studies of Mulloidichthys found considerable variation in colour pattern, meristic or morphometric characters among species or populations, but found no evidence for the presence of a fourth species in the region. For example, in their description of M. mimicus from the South Pacific, Randall & Guézé (1980) noted that the species mimics the colour pattern of the blue-striped snapper Lutjanus kasmira (Forsskål 1775) that may reduce predation risk when the two species shoal together. In the Indian Ocean, off Tanzania, Kenya and Sri Lanka, a blue-striped colour variant of M. vanicolensis that appeared to resemble M. mimicus was observed (Randall & Guézé 1980). These authors further noted that M. vanicolensis forms shoals with Lutjanus kasmira and may be able to rapidly change between the ‘typical’ and the mimic colour pattern.

Stepien et al. (1994) found a divergence in the number of gill rakers between populations of Mulloidichthys vanicolensis in the Western Indian Ocean compared to those from the Pacifi c. Allozyme studies conducted in parallel to the morphological comparisons suggested, however, a close genetic relationship exists between the two populations.

Based on their findings these authors recognized three closely related species of the circumtropical ‘martinicus’ complex of Mulloidichthys: the Indo-West Pacifi c M. vanicolensis, the East Pacifi c M. dentatus (Gill 1862) and the West Atlantic M. martinicus (Cuvier 1829).

This study identifies a fourth, hitherto undescribed, species of Mulloidichthys from the Western Indian Ocean that can be clearly distinguished from the other six currently recognized species. A detailed comparison of colour patterns and 50 meristic and morphometric characters was made of a total of 118 specimens. Special emphasis was placed on the use of relatively quickly identifiable colour, meristic, and morphometric characters. The new species, M. ayliffe sp. nov. is described and a key to the Western Indian Ocean species that includes also the South Pacifi c sister species, M. mimicus, is provided, and can be used for both fresh and preserved fi sh. Comparisons of populations from different areas of the Indian Ocean and between the Indian Ocean and South Pacific are made to further understand the extent of geographical variation among the Western Indian Ocean Mulloidichthys species.

Abbreviations and descriptions of morphometric and meristic characters are provided in Table 1. Morphometric characters were measured with an electronic caliper and are expressed as % SL. For comparison with earlier studies — and in order to facilitate their application in the field — the diagnostically most important morphometric characters are provided as ratios of SL in the key and in Table 2c. Morphometric ratios less than 100 are given to two signifi cant digits.

Only meristic characters that vary among species are referred to in the diagnoses and comparisons: the number of pectoral-fin rays, rudimentary and developed gill rakers on lower and upper limb, and lateral-line scales. In order to verify the presence of the first minute dorsal-fin spine, characteristic for the genus, a stereomicroscope was used. In difficult cases the scales at the base of the second spine were moved or radiographs were examined. Gill rakers were identified as rudimentary if their length was less than their width. The gill raker in the angle between the upper and lower limbs of the first gill arch was included in the count for the lower limb. Lateral-line scale counts do not include scales on the caudal fi n.

Measurements showing high overall intraspecific variation, e.g. fin distances from the snout, were not included in the diagnoses and only rarely in comparisons. Body depth measurements were only considered when there was consistent co-variation with other closely correlated measurements.

No juvenile or subadult M. ayliffe sp. nov. were available for comparative examinations. Because goatfishes vary significantly during ontogeny due to allometric changes in body form (Uiblein & Heemstra 2010), morphological comparisons need to be restricted to distinct life-history stages and size classes. In the current study only fish > 12 mm SL were examined, except for a single 79 mm M. mimicus.

To explore the inter- and intraspecifi c distinction among species, forms or populations, various statistical methods were used. Chi2-Test was used for comparing meristic characters in widely overlapping populations; Principal Component Analysis (PCA) with size-adjustment based on the residuals gained from log-log regressions of the morphometric variables with standard length was used to obtain information on optimal distinction among species and/or deviating forms (e.g., Uiblein & Winkler 1992; Uiblein & Heemstra 2010). In order to facilitate correct identifi cation and avoid terminological misunderstandings regarding the comparison of body stripes, the following schema was developed (Fig. 1). Generally, two types of body stripes can be distinguished by colour and width.

The fi rst consists of bluish body stripes, with similar width to or narrower than the barbels, that vary in colour intensity between species (Table 2).

The stripes are blue to pale blue in Mulloidichthys ayliffe sp.nov. and M. mimicus, and pale blue to whitish in M. dentatus, M. fl avolineatus, M. martinicus and M. vanicolensis. The stripes may vary considerably in number, but the dorso-mid-lateral and the ventro-mid-lateral stripe are present in all six species, the latter stripe sometimes overlain by white-silvery body colour. In M. mimicus and M. ayliffe sp. nov. three additional bluish body stripes may occur, a ventro-lateral, a ventral, and a dorsolateral stripe, the latter usually consisting of a series of blue dots.

The second type of stripes consists of wider, yellow stripes (Fig. 1), their width varying from pupil diameter to more than orbit diameter (Table 2a). The yellow, mid-lateral stripe occurs in fresh specimens of all striped species and is bordered by the bluish dorso-mid-lateral stripe above and the bluish ventro-mid-lateral stripe below. Yellow stripes may also be present more ventrally or dorsally, mostly in association with additional bluish body stripes. Due to the yellow ‘background’ body coloration in M. ayliffe sp. nov. and M. mimicus the yellow stripes appear to be less contrasting compared to those of M. fl avolineatus and the ‘martinicus’ complex. The yellow body stripes fade in preserved fish and hence can be used only for fi eld identifi cation or freshly-caught fish. The bluish stripes and, in particular, the dorso-mid-lateral stripe are frequently retained in recently preserved M. ayliffe sp. nov. and M. mimicus.

Colour characters that fade in preserved fi sh were only included as supplementary information in the key and diagnoses along with comments on their restricted applicability. Colour photographs for each species were selected using primarily material that was also examined in the current study, or material that was identifiable (Plate 1).

Because M. ayliffe sp. nov. resembles M. mimicus considerably and misidentifi cations have been made to date, the latter is also included in the key and the species accounts.

Complementary information on species distributions was obtained from the literature.

Genus Mulloidichthys Whitley 1929

Mulloidichthys Whitley 1929: 122. Type species Mullus fl avolineatus Lacepède 1801. Type by being a replacement name for Mulloides Bleeker 1849.

DIAGNOSIS. Dorsal fins VIII + 9; anal fin I, 6; pelvic fins I, 5; pectoral-fin rays 15–18; principal caudalfin rays 7 + 8 (median 13 branched); gill rakers 6–10 + 19–27 = 26–35; lateral-line scales 33–39 (plus 3–4 on caudal base); lateral line complete; small conical teeth on both jaws; body oblong, slightly compressed; barbel length in adults (>125 mm SL) 4.2–5.7 times in SL, snout length 6.1–9.1 times in SL, larger than postorbital length (8.4–11 times in SL); fresh fish with either a rather uniformly redcoloured body or a silvery-white to yellow body with a yellow mid-lateral body stripe or band, bordered by two narrower bluish (blue to bluewhite) stripes.

DISTRIBUTION. In all major oceans, tropical to subtropical, one species in the Atlantic.

REMARKS. Seven species are recognized as valid, M. ayliffe sp. nov. from the Western Indian Ocean, M. fl avolineatus, M. pfl uegeri, and M. vanicolensis from the Indo-Pacifi c, M. mimicus and M. dentatus from the Pacifi c, and M. martinicus from the subtropical and tropical Atlantic. No recent revision of the genus exists.

This key is based exclusively on adult fish (> 125 mm SL), see also Tables 2, 3, 4, 5, 6, Figs. 2, 3, Plates 1, 2.

1a. Second dorsal-fin height 7.7–8.2 times in SL, caudal-peduncle depth 3.0–3.3 times or more in body depth at fi rst dorsal-fin origin, caudal-peduncle width 1.5–1.8 times in caudal-peduncle depth; body reddish in fresh fish, lateral body stripes absent (Indo-Pacifi c) ................................................................. M. pfl uegeri

1b. Second dorsal-fin height 5.5–7.5 times in SL, caudal-peduncle depth 2.4–2.9 times in body depth at fi rst dorsal-fin origin, caudal-peduncle width 1.9–3.3 times in caudal-peduncle depth; wide yellow and narrow bluish (blue, pale blue or whitish) body stripes present in fresh fi sh ............................................2

2a. Body and head depth shallow, body depth at anal-fin origin 4.7–5.6 times in SL, maximum head depth 4.5–5.2 times in SL, 19–22 gill rakers on lower limb, dark oval or rectangular blotch mid-laterally on body below first dorsal fin, sometimes weak or absent in fresh or preserved fi sh (Indo-Pacifi c) ............. ..........................................................................................................................................................M. fl avolineatus

2. Body and head moderately deep, body depth at anal-fin origin 3.7–4.8 times in SL, maximum head depth 3.9–4.8 times in SL, 19–26 gill rakers on lower limb, no dark blotch on body .................................3

3a. Gill rakers on lower limb 23–26, body depth at anal-fin origin 4.0–4.8 times in SL, second dorsal-fi n base 1.1–1.4 times in its height, yellow mid-lateral body stripe conspicuous, its width equal or less than pupil diameter, bluish (or whitish) body stripes weak, not retained in preserved fi sh (Indo-Pacifi c) ........... M. vanicolensis

3b. Gill rakers on lower limb 19–23, body depth at anal-fin origin 3.7–4.4 times in SL, second dorsal-fi n base 0.9–1.2 times in its height, yellow mid-lateral body stripe inconspicuous, its width larger than pupil diameter, bluish body stripes conspicuous, dorso-mid-lateral stripe frequently retained in freshly preserved fi sh ........................................................................................................................................................ 4

4a. Lateral-line scales 35–37, body depth at anal fin origin 3.9–4.4 times in SL, maximum head depth 4.0–4.5 times in SL, bluish dorso-mid-lateral body stripe crosses lateral line posterior to fi rst dorsal-fi n base (Indian Ocean) ..........................................................................................................................M. ayliffe sp. nov.

4b. Lateral-line scales 38–39, body depth at anal fin origin 3.7–4.0 times in SL, maximum head depth 3.0–4.1 times in SL, bluish dorso-mid-lateral body stripe crosses lateral line below fi rst dorsal-fin base (Pacifi c) ................................................................................................................................................................ M. mimicus

Holotype. SAIAB 86367, 175 mm, KwaZulu-Natal, Sodwana Bay, Mellow Yellow Reef, South Africa, 27°31.863’S 32°42.48’E, 12–19 m, collected by Neville Ayliffe, Phillip C. Heemstra and Elaine Heemstra

Paratypes. Western Indian Ocean, South Africa: SAIAB 86368, 223–245 mm, KwaZulu-Natal, Coral Gardens, Sodwana Bay, 27°31.34’S, 32°41.15’E, 5–12 m; Tanzania: BPBM 17620, 155 mm, Mafi a Island, Chole Island, Chole Bay, reef, 6 m; Kenya: SAIAB 18057, 190-194 mm, Shimoni, 4°39’S, 39°23’E; SAIAB 13907, 197–230 mm, Shimoni, 4°39’S, 39°23’E; Seychelles: SAIAB 18055, 210 mm, Amirante Islands, Poivre Island, 05°46’S, 53°19’E; Oman: BPBM 36010, 183–187 mm, Southern Oman, Kuria Muria Islands, Sawda Island, E end, 4–8 m; BPBM 36024, 6, 171–217 mm, Southern Oman, Kuria Muria Islands, Sawda Island, SW side, 10 m; BPBM 41008, 218 mm.

Non-Types. Western Indian Ocean, Sri Lanka: BPBM 18770, 154 mm, Trincomalee, Dutch Point, 5 m; BPBM 27749, 202 mm, Trincomalee, fi sh market; BPBM 31285, 152–192 mm, Trincomalee, Koddiyar Patu, Foul Point, E side of lighthouse, 8°32’N, 81°19’E, 0–8 ft.

DIAGNOSIS. Pectoral fins 16 or 17; gill rakers 7–8 + 19–23 = 27–31; lateral-line scales 35–37; body depth at fi rst dorsal-fi n origin 26–29% SL; body depth at anus 23–25; caudal-peduncle depth 10–11; caudalpeduncle width 4.1–5.3; maximum head depth 22– 25; head depth through eye 17–20; head length 28– 31; snout length 11–13; orbit length 6.9–8.5; upper jaw length 9.0–11; barbel length 19–23; caudal-fi n length 28–31; anal-fin height 14–17; pelvic-fi n length 19–22; pectoral-fin length 19–22; pectoral-fi n width 4.2–5.2; fi rst dorsal-fi n height 21–24; second dorsal-fin height 14–17% SL; body, head and fi ns yellow in live fish, becoming orange dorsally and anteriorly in freshly collected fish; body and head with two to five straight bluish (blue or pale blue) lateral body stripes, with one to four yellow stripes between, bluish stripes approximately as wide as barbels; yellow mid-lateral stripe wider than pupil diameter; dorso-mid-lateral stripe most prominent, from behind upper orbit to behind end of second dorsal-fin base, crossing lateral line at 55–60% SL, well posterior to fi rst dorsal fin base; head and body of preserved fish pale-brown to brown, lateral body stripes sometimes retained.

DESCRIPTION. Measurements in % SL and counts are given in Table 4; morphometric data as ratios of SL for holotype, data for paratypes in brackets. Body elongate, its depth at fi rst dorsal-fin origin 3.6 [3.4–3.9], body depth at anal-fin origin 4.2 [4.0–4.4], maximum head depth 4.4 [4.0–4.5], head length 3.5 [3.2–3.6], snout length 8.5 [7.6–8.9], orbit length 13 [13–14], barbel length 4.8 [4.4–5.1], subequal to caudal-peduncle length (4.7 [4.7–5.3]), pelvic-fi n length (4.7 [4.7–5.1]), and pectoral-fin length (4.6 [4.8–5.2]), anal-fin height 6.9 [6.0–7.1], fi rst dorsalfin height 4.3 [4.3-4.8], second dorsal-fin height 6.7 [6.5–7.2].

Mouth small, maxilla not reaching a vertical at front of orbit, upper jaw length 9.8 [8.8–11] in SL; small conical teeth on both jaws, placed in one outer row and more irregularly behind in the front of jaws, and in a single row more posteriorly; no teeth on roof of mouth; anterior nostril a small vertically elliptical opening about ²/₃ orbit diameter in front of eye; posterior nostril a narrow slit covered by a membrane next to edge of upper orbit; longest gill filaments on first gill arch about ²/₃ orbit diameter; longest gill raker on first arch about ²/₃ in longest gill filament; a single flat spine at posterior edge of operculum at level of about mid of eye.

Scales very finely ctenoid; head fully scaled; fins naked except base of caudal fin; dorsal fi n of holotype behind 4th lateral line scale, origin of second dorsal above 18th scale, origin of anal fin below 19th scale origin; lateral line following contour of back; pored scales of lateral line with many branched tubules.

Fresh colour (based on holotype (HT), two paratypes, and underwater photographs) (Plates 1, 2). Body, head and fins mostly yellow in life, becoming slightly orange dorsally and anteriorly on in freshly collected fish; outer margins of jaws white to pale bluish; eyes with black pupils surrounded by red iris; iris at dorsal margin of orbit often bluish (see also description of upper-lateral stripe below).

Two to five straight, bluish body stripes (Fig. 2, Plate 1), their width about equal to barbel width, the dorso-mid-lateral stripe being the most prominent, reaching from behind the upper orbital margin to below rear end of soft dorsal-fin base, and continued anteriorly by a weak pigmentation of the iris along the dorsal orbital margin, and a tiny bluish patch (not always present) immediately in front of the upper edge of orbit; this stripe follows the lateral line for a distance from snout of 1.8 in SL (HT) [1.7– 1.8 in PTs], crossing the lateral line well posterior to the fi rst dorsal-fin base; the bluish ventro-midlateral stripe is the second-most conspicuous body stripe in this new species, consisting in a straight line from behind lower orbital margin to well behind anal-fin base, continuing below the eye as a wave-like line, which at its anterior end points towards the snout tip or upper jaw, but not reaching them; the ventro-lateral body stripe, i.e., the bluish stripe below the ventro-mid-lateral body stripe, forms a straight line from head to caudalfin base, passing ventral to pectoral fin base; this stripe is at a similar vertical distance from the midlateral stripe as the dorso-mid-lateral stripe, and sometimes continues anteriorly as a series of small bluish patches reaching from operculum to about the posterior margin of maxilla. Some specimens (e.g., the paratype in Plate 1) show an additional bluish, ventral stripe that extends close to the ventro-lateral body stripe, from behind operculum to anal-fin base, along the ventral body margin, and unites with the ventro-lateral stripe above the anal-fin base. Larger fish may also show a weak indication of a fifth bluish dorso-lateral body stripe as a series of close-set, small blue spots reaching from behind operculum to about end of fi rst dorsal fin base or to half body at maximum, passing closely beneath the fi rst dorsal-fi n base.

One to several yellow bands are formed by the yellow ground colour of the body between the bluish body stripes; the width of the mid-lateral yellow stripe is between pupil and orbit diameter, the yellow ventro-lateral stripe below is about the same width; all other yellow stripes, if formed, are narrower.

Pectoral fins hyaline, pelvic and anal fi ns sometimes whitish-yellow at bases and distal-most areas; caudal fins with some light red patches from origin to lobes, inner margins of caudal-fi n lobes yellow-hyaline; dorsal fins slightly transparent in between posterior spines or rays; barbels white.

Preserved colour. Body and head of preserved fi sh pale-whitish to pale-brown, body of recently preserved fish dorsally darkerwith one to three dark body stripes in similar form and position to the bluish dorso-mid-, ventro-mid- and ventrolateral stripes in fresh fish; all three stripes retained in holotype, the dorso-mid-lateral stripe reaching from behind upper orbit to behind second dorsalfin base, the ventro-mid-lateral stripe from behind operculum to caudal peduncle, and the ventro-lateral stripe from below pectoral-fin base to mid anal-fin base; pectoral fins hyaline; all other fi ns whitish-pale.

DISTRIBUTION. Western Indian Ocean: Natal, South Africa, Tanzania, Kenya, Oman, Seychelles, Sri Lanka, Andaman Islands.

ETYMOLOGY. The name ‘ayliffe’ is used as a noun in apposition; it honours Mr Neville Ayliffe, a former dive operator at Sodwana, who has assisted the South African Institute of Aquatic Biodiversity in acquiring important fish collections during many years. He collected the holotype and two paratypes with a speargun from shallow reefs in Sodwana Bay, KwaZulu-Natal, South Africa.

COMPARISONS. Mulloidichthys ayliffe differs from the Western Indian Ocean species as follows: from M. fl avolineatus it differs in a deeper body and caudal peduncle, larger maximum head depth, longer dorsal-fi n bases, frequent presence of more than two bluish body stripes, a wider yellow midlateral stripe, and absence of a mid-lateral dark oval or rectangular blotch below fi rst dorsal-fi n base; it differs from M. pfl uegeri in a deeper caudal peduncle, shallower suborbital, shorter snout, larger eyes, shorter jaws, longer dorsal-fi n bases, higher anal fin, smaller pectoral-fin width, higher dorsal fins, and presence of lateral body stripes; and it differs from M. vanicolensis in fewer gill rakers, modally fewer lateral-line scales, lower second dorsal fin, bluish body stripes darker and often more than two present, and wider yellow mid-lateral body stripe.

Non-Western Indian Ocean species: Mulloidichthys ayliffe differs from the Eastern Pacific M. dentatus in longer dorsal- and anal-fi n bases, higher first dorsal fin, bluish body stripes darker and often more than two present, and wider yellow mid-lateral body stripe; it differs from the Western Atlantic M. martinicus in a longer caudal peduncle, longer second dorsal-fin base, shorter caudal fi n, bluish body stripes darker and often more than two present, and wider yellow mid-lateral body stripe; and it differs from the Central Pacifi c M. mimicus in fewer lateral-line scales, shallower body at anal-fin origin, lower maximum head depth, shorter snout, longer fi rst dorsal-fin base, bluish dorso-mid-lateral body stripe crossing lateral line further posteriorly, and yellow mid-lateral stripe narrower.

REMARKS. The use of colour patterns should allow easy fi eld identifi cation of Mulloidichthys ayliffe and distinction from co-occurring congenerics. The main colour difference to M. mimicus is the position of the bluish dorso-mid-lateral body stripe in relation to the lateral line. While the yellow mid-lateral stripe and the bluish dorso- and ventro-mid-lateral body stripes of M. ayliffe have a similar vertical placement as in M. dentatus, M. flavolineatus, M. martinicus, and M. vanicolensis, the yellow mid-lateral stripe differs in width and is less conspicuous due to the lack of overall yellow ground body colouration in the latter four species. Field observations suggest that the bluish body stripes may vary in intensity and contrast in M. ayliffe (e.g., Randall & Guézé, 1980). No information on night and resting colouration is currently available.

The combination of body depth at anal-fi n origin and length of fi rst dorsal-fin base allows clear separation between M. ayliffe from M. mimicus (Fig. 3). And, these two species can be separated from the closely related M. vanicolensis by combining second dorsal-fin height with total number of gill rakers (Fig. 2).

No significant differences in colour and morphological characters among populations from the southwestern Indian Ocean (East-Africa and Seychelles), Oman, and Sri Lanka were found (Table 4).

No juveniles or subadults were available for study.

Mulloidichthys ayliffe attains 25 cm SL; typical habitats of this species are shallow coral reef areas including submarine caves to 19 m depth.

Mullus fl avolineatus Lacépède 1801: no locality stated. No types known (Eschmeyer 2010). Mulloides fl avolineatus: Ben-Tuvia, in Smith & Heemstra 1986: 610, Plate 69, colour photo. Mulloidichthys fl avolineatus: Randall 1995: 239, Fig. 621 (colour photo), Fricke 1999: 307–309; Heemstra & Heemstra 2004: 261, colour painting; Heemstra et al. 2004: 3322; Randall 2005: 292, 2 colour photos; Taquet & Diringer 2007: 260, colour photo.

DIAGNOSIS. Pectoral fins 15–18; gill rakers 7–10 + 19–22 = 26–31; lateral-line scales 34–38; body depth at fi rst dorsal-fi n origin 21–26% SL; body depth at anus 16–21; caudal-peduncle depth 8.3–9.8; caudalpeduncle width 3.2–4.6; maximum head depth 19– 22; head depth through eye 16–19; head length 27– 31; snout length 12–15; orbit length 5.8–7.8; upper jaw length 8.1–9.5; barbel length 18–22; caudalfin length 29–33; anal-fin height 13–16; pelvic-fi n length 19–22; pectoral-fin length 19–21; pectoral-fi n width 3.5–4.9; fi rst dorsal-fi n height 19–23; second dorsal-fin height 14–16% SL; body silvery white, sometimes intermingled with yellow, darker above lateral line; head silvery white to yellowish, darker on dorsal part of snout and dorsally from midorbit; one straight yellow mid-lateral body stripe, its width subequal to pupil diameter, bordered by two pale bluish (sometimes whitish), narrow midlateral stripes; a dark oval to rectangular blotch at yellow mid-lateral body stripe below fi rst dorsalfin base, sometimes only faintly visible due to color changes, often retained in preserved fi sh; dorsal and caudal fins white to yellowish, partly hyaline, pectoral fins pale rose, partly transparent, pelvic and anal fins whitish and partly transparent; barbels white; in preserved fish body generally pale brown or ventrally pale and dorsally darkened, head pale brown; recently collected material with a broad dark band laterally.

DISTRIBUTION. Indo-West Pacific: Red Sea, East and South Africa, Madagascar and Mascarenes east to Hawaiian Islands, Line Islands and Pitcairn Group, north to southern Japan, south to Joseph Bonaparte Gulf (northern in front of Western Australia), New South Wales (Australia) at 36°S, Lord Howe Island, New Caledonia and Rapa Island.

COMPARISONS. Mulloidichthys fl avolineatus differs from the Western Indian Ocean species as follows: from M. ayliffe it differs in a shallower body and caudal-peduncle, smaller maximum head depth, shorter dorsal-fin bases, narrower yellow midlateral body stripe, and frequent presence of one mid-lateral dark oval or rectangular blotch below fi rst dorsal-fin base; it differs from M. pfl uegeri in a shallower body and head, thinner caudal peduncle, shorter jaws, smaller snout width, shorter fi rst dorsal and anal-fin bases, smaller pectoral-fi n width, higher second dorsal fin, presence of body stripes, and frequent presence of one mid-lateral dark oval or rectangular blotch below fi rst dorsalfin base; it differs from M. vanicolensis in fewer gill rakers, modally more pectoral-fin rays and fewer lateral-line scales, shallower body, lower maximum head depth, shorter jaws, lower second dorsal fi n, and frequent presence of one mid-lateral dark oval or rectangular blotch below fi rst dorsal-fi n base.

Non-Western Indian Ocean species: Mulloidichthys fl avolineatus differs from the Pacific M. dentatus in a shallower body and caudalpeduncle, and frequent presence of one mid-lateral dark oval or rectangular blotch below fi rst dorsalfin base; it differs from the W-Atlantic M. martinicus in a shallower body and caudal-peduncle, lower maximum head depth, shorter fi rst dorsal-fi n base, and frequent presence of one mid-lateral dark oval or rectangular blotch below fi rst dorsal-fi n base; and it differs from the Pacifi c M. mimicus in fewer lateral line scales, shallower body, caudal peduncle and head, shorter fi rst dorsal-fi n base, narrower yellow mid-lateral body stripe, and frequent presence of one mid-lateral dark oval or rectangular blotch below fi rst dorsal-fi n base.

REMARKS. This species varies considerably in body colouration with the yellow mid-lateral body stripe always present in diurnally active fi sh, but the dark mid-lateral oval or rectangular blotch being sometimes only faintly visible and the bluish mid-lateral body stripes sometimes changing to white and fading into the whitish-silvery or pale yellowish body colour in fresh fi sh.

In interspecific associations with Mulloidichthys vanicolensis the blotch may be completely ‘switched off’, thus making it considerably more diffi cult for an observer to distinguish between the two species. During a dive off Oahu, Hawaii, I observed a dense aggregation of both species close to a rocky wall above a sand bottom with both species showing identical coloration (the dark blotch in M. fl avolineatus being absent or only very diffi cult to detect), and the species being only identifi able on body proportions (Plate 2b). Both species were mixed in a large shoal, M. fl avolineatus appearing to be positioned slightly underneath M. vanicolensis. Resting or night colouration with dark-brown patches on pale body and the yellow stripe not visible (Plate 1F)

The Eastern Indian Ocean population of M. vanicolensis has a shorter caudal peduncle than populations in the Western Indian Ocean and Pacific. The Pacific Ocean population differs from Indian Ocean population in a higher pectoral-fi n ray number (Table 3), a shorter suborbital distance, and shorter barbels (Fig. 4).

A comparison of juvenile and subadult (<125 mm SL) morphological characteristics between M. fl avolineatus and M. vanicolensis is currently in preparation (Uiblein & Randall unpublished data).

Mulloidichthys fl avolineatus attains 29 cm SL; it inhabits reef areas to 35 m depth.

Mulloides pfl uegeri Steindachner 1900: 485–486, Table III, Fig. 4; type locality Honolulu, Oahu, Hawaii.

Mulloidichthys:pfl ugeri (non Steindachner): Myers 1989, 159, plate 74 I, colour photo.

Mulloidichthys:pfl ugeri Fricke 1999: 309; Heemstra et al. 2004: 3322; Randall 2005: 293, 2 colour photos; Taquet & Diringer 2007: 261, colour photo.

DIAGNOSIS. Pectoral fins 17-18; gill rakers 6–7 + 19–22 = 26–29; lateral-line scales 35–37; body depth at fi rst dorsal-fin origin 26–28% SL; body depth at anus 21–24; caudal-peduncle depth 8.4– 9.1; caudal-peduncle width 4.6–6.0; maximum head depth 24–26; head depth through eye 19–21; head length 28–31; snout length 13–16; orbit length 5.2–6.9; upper jaw length 11–12; barbel length 19– 21; caudal-fin length 29–33; anal-fin height 12–13; pelvic-fin length 20–24; pectoral-fin length 20–23; pectoral-fin width 5.4–5.7; fi rst dorsal-fi n height 18–21; second dorsal-fin height 12–13% SL; body ventrally white with rose flanks, ventral half of caudal peduncle white, dorsal half of body red becoming more intense further dorsally; head from eye dorsally and anteriorly to jaws red, inner jaw margins pale rose, operculum and posterior part of head from below eye and behind jaws whitish rose; dorsal fins rose with red base, caudal fi n rose, white at inner margin of lobes; pectoral fi ns rose, slightly transparent; pelvic and anal fi ns rose; barbels whitish-rose.

DISTRIBUTION. Indo-West Pacifi c: Mascarenes, eastern Indonesia east to Hawaiian and Marquesas islands, north to Ryukyu Islands, south to Tonga.

COMPARISONS. Mulloidichthys pfl uegeri differs from the Western Indian Ocean species as follows: from M. ayliffe in a shallower caudal peduncle, deeper suborbital, longer snout, smaller eyes, longer jaws, shorter dorsal-fin bases, lower anal fi n, larger pectoral-fin width, lower dorsal fins, and absence of lateral body stripes; from M. fl avolineatus it differs in a deeper body and head, thicker caudal peduncle, longer jaws, larger snout width, longer first dorsal- and anal-fin bases, larger pectoral-fi n width, lower second dorsal fin, absence of body stripes and absence of mid-lateral dark oval or rectangular blotch; from M. vanicolensis it differs in fewer gill rakers, shallower and thicker caudal peduncle, deeper suborbital, longer snout and jaws, lower anal and dorsal fins, larger pectoral-fi n width, and absence body stripes.

Non-Western Indian Ocean species: Mulloidichthys pfl uegeri differs from the Pacifi c M. dentatus in a shallower and thicker caudal peduncle, deeper head and suborbital, longer jaws, longer second dorsal- and anal-fin bases, shorter anal and second dorsal fins, larger pectoral-fin width, and absence body stripes; it differs from the Atlantic

M. martinicus in a shallower and thicker caudal peduncle, deeper head, deeper suborbital, smaller eyes, longer jaws, longer interdorsal distance, longer second dorsal-fin base, shorter anal and second dorsal fins, and absence body stripes; and it differs from the Pacifi c M. mimicus in fewer lateralline scales, shallower body, shallower and thicker caudal peduncle, deeper suborbital, smaller eyes, longer jaws, larger interdorsal distance, lower anal fin, larger pectoral-fin width, lower second dorsal fin, and absence of body stripes.

REMARKS. The red body colour in Mulloidichthys pfl uegeri may vary considerably in intensity with appearance of four broad vertical dark-red bars on pale-red ground colour as shown in photographs of fresh fish in Randall (2005: 293) and Taqet & Diringer (2007: 261). This pattern possibly represents the resting and/or night colour.

The Pacific Ocean specimens of M. pfl uegeri have a deeper body and head, a longer caudal fi n and a lower first dorsal fin than those from the Indian Ocean; the sample size from both areas would have to be larger to investigate these differences in more detail.

No information on juvenile morphological characteristics is currently available.

The distribution of Mulloidichthys pfl uegeri appears to be restricted to oceanic islands, similar to Upeneus taeniopterus Cuvier, 1829 (Uiblein & Heemstra 2010); M. pfl uegeri attains 40 cm SL; largest species of the genus; it occurs on shallow bottoms to 110 m depth.

Upeneus vanicolensis Valenciennes, in Cuvier & Valenciennes 1831: 521; type locality: Vanicolo Island, Santa Cruz Islands, Solomon Islands, Southwestern Pacific, 11°37’S, 166°58’E.

Mulloides vanicolensis Ben-Tuvia, in Smith & Heemstra 1986: 611, Plate 69, colour painting.

Mulloidichthys vanicolensis Randall 1995 (in part): 239, Fig. 621 (colour photo showing a mixed school of M. ayliffe and M. vanicolensis), Fricke 1999: 309–310; Heemstra & Heemstra 2004: 261; Heemstra et al. 2004: 3322; Randall 2005: 293, 2 colour photos; Taquet & Diringer 2007: 261, colour photo.

DIAGNOSIS. Pectoral fins 15–17; gill rakers 7–10 + 23–26 = 31–35; lateral-line scales 36–38; body depth at fi rst dorsal-fi n origin 25–30% SL; body depth at anus 21–25; caudal-peduncle depth 9.2–11; caudalpeduncle width 3.0–4.9; maximum head depth 21– 25; head depth through eye 16–20; head length 28– 32; snout length 11–14; orbit length 6.2–9.4; upper jaw length 9.4–11; barbel length 19–24; caudal-fi n length 29–34; anal-fin height 14–18; pelvic-fi n length 20–24; pectoral-fin length 20–24; pectoral-fi n width 3.9–5.4; fi rst dorsal-fi n height 20–25; second dorsal-fin height 15–18 %SL; body silvery white ventrally, yellow above lateral line; head silvery white to pale rose below eye and behind jaws, with some yellowish patches, darker rose on snout and yellowish above mid eye; one straight yellow midlateral body stripe from eye to caudal-fin base, its width subequal to pupil diameter, becoming wider on posterior part of caudal peduncle, bordered by two narrow pale bluish (sometimes whitish) midlateral stripes from eye to behind anal fi n-base; dorsal caudal, pelvic and anal fins yellow, pectoral fin hyaline; barbels white; in preserved fi sh body pale brown or ventrally pale and dorsally darkened, head pale brown; recently collected material with a broad dark band mid-laterally.

DISTRIBUTION. Indo-West Pacific: Red Sea, East Africa, Comoro Islands and Mascarenes, east to the Hawaiian Islands, Line Islands and Pitcairn Group, north to southern Japan, south to Joseph Bonaparte Gulf (northern in front of Western Australia), New South Wales (Australia), Lord Howe, Norfolk, Kermadec Islands, New Caledonia, Tonga and Gambier Islands.

COMPARISONS. Mulloidichthys vanicolensis differs from the Western Indian Ocean species as follows: from M. ayliffe it differs in more gill rakers, more lateral-line scales, higher second dorsal fi n, bluish body stripes weaker (sometimes whitish), and yellow mid-lateral body stripe narrower; it differs from M. fl avolineatus in more gill rakers, modally fewer pectoral-fin rays and more lateral-line scales, a deeper body, larger maximum head depth, longer jaws, higher second dorsal fin, and absence of one mid-lateral dark oval or rectangular blotch below fi rst dorsal-fin base; and it differs from M. pfl uegeri in more gill rakers, deeper and thinner caudal peduncle, shallower suborbital, shorter snout and jaws, higher anal and dorsal fi ns, larger pectoral-fin width, and presence of a yellow midlateral body stripe.

Non-Western Indian Ocean species: Mulloidichthys vanicolensis differs from the Pacific M. dentatus in more gill rakers, longer anal-fi n base, and lower dorsal fins; it differs from the Atlantic M. martinicus in more gill rakers and shorter snout; and from the Pacifi c M. mimicus it differs in more gill rakers, modally fewer lateral-line scales, shallower body at anal-fin origin, higher second dorsal fi n, bluish body stripes weaker (sometimes whitish), and yellow mid-lateral body stripe narrower.

REMARKS. Mulloidichthys vanicolensis varies considerably in body coloration with the yellow mid-lateral body stripe always present in active fish, but the bluish mid-lateral body stripes sometimes changing to white due to blending with the general whitish-silvery or pale yellowish body colour in fresh fish. Resting or night colouration with patches of red on pale body and yellow stripes only faintly visible (Randall 2005: 293).

The southwesten Indian Ocean specimens of M. vanicolensis have a thinner caudal peduncle than the populations form Oman and the Eastern Indian Ocean and the latter differs from the Western Indian Ocean specimens in having a longer caudal peduncle. The main differences between the Indian Ocean and Pacific populations are a higher anal fi n and a longer pectoral fin in the latter (Fig. 5).

Mulloidichthys vanicolensis attains 31 cm SL; it occurs on shallow bottoms to 113 m depth.

Mulloidichthys mimicus Randall & Guézé 1980: 603–609, 2 figs. (colour photos); type locality: west side of Sentinelle de l’Est, Nuku Hiva, Marquesas Islands, South Pacific; Randall 2005: 292, 2 colour photos.

DIAGNOSIS. Pectoral fins 15–17; gill rakers 6–8 + 21–23 = 28–31; lateral-line scales 38–39; body depth at fi rst dorsal-fi n origin 28–30% SL; body depth at anus 25–27; caudal-peduncle depth 10–11; caudalpeduncle width 4.0–4.7; maximum head depth 24– 26; head depth through eye 18–20; head length 28– 31; snout length 12–14; orbit length 7.2–8.5; upper jaw length 9.5–11; barbel length 20–22; caudal-fi n length 27–31; anal-fin height 13–17; pelvic-fi n length 20–23; pectoral-fin length 20–22; pectoral-fi n width 4.5–5.2; fi rst dorsal-fi n height 20–23; second dorsal-fin height 14–16% SL; head, body and fi ns yellow in live fish, blending dorsally and anteriorly into orange after collection; body and head covered by two to five straight bluish (blue or pale blue) body stripes, with one to four yellow stripes in between; bluish stripes approximately as wide as barbel, yellow mid-lateral body stripe wider than orbit diameter; bluish dorso-mid-lateral stripe most prominent, from above orbit to behind end of second dorsal-fin base, crossing lateral line at 44-48% SL, below posterior end of fi rst dorsal-fi n base; head and body of preserved fi sh pale-brown to brown, bluish lateral body stripes sometimes retained.

DISTRIBUTION. South Pacific Islands, a single record from Kauai, Hawaiian Islands.

COMPARISONS. The Pacifi c Mulloidichthys mimicus differs from the Indian Ocean M. ayliffe sp. nov. in more lateral-line scales, a deeper body at analfin origin, higher maximum head depth, longer snout, shorter fi rst dorsal-fin base, bluish dorsomid-lateral body stripe crossing lateral line farther anterior, and wider yellow mid-lateral body stripe; it differs from M. fl avolineatus in more lateral-line scales, a deeper body, caudal peduncle and head, longer fi rst dorsal-fin base, wider yellow midlateral body stripe, and absence of one mid-lateral dark oval or rectangular blotch below fi rst dorsal fin; it differs from M. pfl uegeri in more lateral-line scales, a deeper body, deeper and thinner caudal peduncle, shallower suborbital, larger eyes, shorter jaws, smaller interdorsal distance, higher anal fi n, smaller pectoral-fin width, higher second dorsal fin, and presence of body stripes; and it differs from M. vanicolensis in fewer gill rakers, modally more lateral-line scales, a deeper body at anal-fi n origin, lower second dorsal fin, bluish body stripes more conspicuous and wider yellow mid-lateral body stripe.

Non-Western Indian Ocean species: Mulloidichthys mimicus differs from the Pacific M. dentatus in more lateral-line scales, a deeper body, higher maximum head depth, longer second dorsal- and anal-fin bases, bluish body stripes more conspicuous and wider yellow mid-lateral body stripe; and it differs from the Atlantic M. martinicus in more lateral-line scales, a deeper body at anal-fi n origin, higher maximum head depth, longer analfin base, shorter caudal fin, bluish body stripes more conspicuous, and wider yellow mid-lateral body stripe.

REMARKS. The use of colour patterns should allow easy fi eld identifi cation of Mulloidichthys mimicus and distinction from co-occurring congenerics, as has been noted for M. ayliffe.

No information on night and resting colour, and on population differences is currently available.

The single juvenile measured clearly differs in a shallower body, caudal peduncle and head, shorter snout and jaws, longer orbit, shorter barbels and longer caudal peduncle compared with the adult specimens.

A single record of this species from Kauai, Hawaiian Islands, is based on a photograph from a local fisherman. It is unclear, however, if this record represents a naturally occurring population in the area or is derived from accidental introduction.

Mulloidichthys mimicus attains 26 cm SL; it occurs in shallow reef habitats to 15 m depth

The present account is the fi rst comprehensive review of the goatfi sh genus Mulloidichthys in the Western Indian Ocean. Apart from the description of a new species for this region, detailed comparisons between all known species of this genus and an identification key, data on morphological differentiation among populations from the Indian Ocean and between the Indian and Pacific Oceans are also presented.

A diagnosis of Mulloidichthys species has to be based on a combination of colour, meristic and morphometric characters with special emphasis on the latter. Bluish and yellow body stripes and overall body coloration are to some extent useful for the distinction between species. But colour patterns do change considerably, e.g. during resting, at night, or in interspecifi c aggregations. Preserved fishes are often monochromatic (i.e., pale-brown or brownish). Meristic characters behave rather conservatively in this genus and show relatively little variation. By including a large dataset of morphometric variables in the description, diagnosis and comparisons it was possible to clearly distinguish all species.

This study is the first to provide clear evidence for geographic variation in body form among Mulloidichthys species with marked differences between the Pacific and Indian Oceans. These results complement earlier findings of interoceanic divergence in the number of gill rakers reported for M. vanicolensis by Stepien et al. (1994). A re-examination of their allozyme-based fi ndings by using modern DNA techniques might prove a valuable approach towards a better understanding of the extent of geographic differentiation and isolation among Mulloidichthys populations.

As also observed in other goatfi sh genera (e.g., Upeneus, Uiblein & Heemstra 2010), considerable allometric changes in body form occur in Mulloidichthys species. An example is the comparison of a single juvenile with adult M. mimicus. A detailed comparison among lifehistory stages and species with more information on phenotypic diversity among Pacifi c populations and forms of M. fl avolineatus and M. vanicolensis is currently in preparation (Uiblein & Randall unpublished data).

Another rewarding future area of research would be to study the relationship between colour patterns in Mulloidichthys species and their possible functions with regard to mimicry and inter- or intraspecific communication. Our observation of the close colour resemblance in M. fl avolineatus and M. vanicolensis (Plate 2) indicates that colour mimicry may also occur among those species. The advantages for either species are however unclear. Possibly the relatively smaller and more elongated M. fl avolineatus hides among M. vanicolensis to reduce predation risk. To go any further with such an assumption it will be necessary to understand both the costs and benefits of colour changes in M. fl avolineatus, as ‘switching off’ the species-specifi c dark lateral blotch may increase the confusion among the two species, which — at least during the reproductive period — may entail an increased risk of hybridization.

I thank Jack Randall, Phil Heemstra, Elaine Heemstra, Neville Ayliffe, Gerry Allen, Romain Causse, John Earle, Richard Field, Jerry Finan, Lori O’Hara, Dennis Polack, and Arnold Suzumoto for photographs of fresh or preserved fi sh, assistance with collection material, and/or providing additional information or comparative data. Thanks to John Earle for organizing a dive trip for in situ studies of goatfishes off Oahu, Hawaii for me. The hospitality and assistance of Lori O’Hara and Arnold Suzumoto, Bishop Museum, Honolulu, Hawaii, and Ronald de Ruiter and Martin van Ojien, Naturalis Museum, Leiden, The Netherlands, during collection visits is gratefully acknowledged. Thanks to the South African Institute of Aquatic Biodiversity and the Nansen Programme of the Center for Developmental Fisheries at the Institute of Marine Research (IMR), Bergen, for travel support. The comments of Gerry Allen, Phil Heemstra, Wouter Holleman, Jack Randall and Elaine Heemstra on earlier versions of the manuscript are gratefully acknowledged

Mulloidichthys dentatus: East Pacifi c, Mexico: USNM 65581, 165 mm, Acapulco, Albatross Expedition, 1904-1905; RMNH 5145, 226 mm, Las Tres Marias; USNM 43241, 156 mm, Sonora, Bay of Guaymas.

Mulloidichthys fl avolineatus: Western Indian Ocean, South Africa: SAIAB 86370, 254 mm, KwaZulu-Natal, Ribbon Reef, Sodwana Bay, 27°29.37’S, 32°41.38’E, 12–18 m; Chagos: SAIAB 15361, 170–179 mm, NW corner of Isle Boddam on ocean side; Mascarenes: SAIAB 68799, 5: 130–132 mm, Rodrigues, off Port Mathurin, Ile Hollandaise; SAIAB 70580, 180 mm, Rodrigues, northofGrandBay; Eastern Indian Ocean, Indonesia: RMNH 13299, 2: 177 mm, Sumatra, Sabang Bay, Pulu Weh; West Pacifi c, Guam: BPBM 77, 235 mm, Mariana Islands; Vanuatu: BPBM 962, 178 mm, Efate Island; Hawaii: BPBM 1749, 185 mm, Oahu, Honolulu; BPBM 1750, 172 mm, Oahu, Honolulu; BPBM 25457, 126 mm, Oahu, Waianae coast; BPBM 25674, 174 mm, Oahu, Honolulu market; BPBM 4087, 287 mm, Laysan; BPBM 4088, 138–226 mm, Lisianski; Marcos Island: BPBM 7087, 210 mm, N end; reef flat, 3 ft; BPBM 7088, 197 mm, reef flat, 3 ft; Rapa Island: BPBM 12937, 164 mm, E side of Akatamiro Bay, 8 ft; Phoenix Islands: BPBM 15299, 3: 146-154 mm, Hull Island, Orona Atoll; Midway Atoll: BPBM 15308, 152 mm, reef; Caroline Islands: BPBM 24628, 163 mm, Puluwat Atoll, lagoon side, 07°20’N, 149°11’E, at surface; Marquesas Islands: BPBM 2140, 200 mm, Nukuhiva; New Zealand: RMNH 11308, 210 mm; Indonesia: RMNH 29720, 135–151 mm, Java Sea, Selat Linta, E of Komodo, Indonesian-Dutch Snellis II Exp., 1984, 8º30’S, 119º34.6’E; RMNH 29994, 218 mm, Bay of Sanggar, N of Sumbawa, near edge of coastel reefflat, Indonesian-Dutch Snellius II exp. 1984; RMNH 13300, 145 mm, Jakarta, Bay of Batavia.

Mulloidichthys martinicus: West Atlantic, Ascension Island: USNM 267497, 270 mm, Northeast Bay; Netherlands Antilles: RMNH 22268, 165–179 mm, Curacao, Bullenbaai; Antigua: USNM 170110, 181 mm, Atlantic, Barbados.

Mulloidichthys mimicus: Holotype: BPBM 12638, 204 mm, Pacific, Marquesas Islands, Nukuhiva, Taiohae Bay; W side of Sentinelle de l’Est, 15 m; Paratypes, West Pacifi c, Line Islands: BPBM 4079, 183 mm, Palmyra Atoll (head broken); BPBM 31897, 164 mm, Kiritimati Atoll, Bay of Wrecks, N end; reef, coral and rubble drop-off, 11 m; BPBM 7738, 6: 213–255 mm, Teraina Island, W end; wreck of the ‘Southbank’, 6–7,5 m; Marquesas Islands: BPBM 12135, 169 mm, Uapou, Hakahetau; S side of bay, 6–11 m; BPBM 11901, 79 mm, Tahuata Island, off point at S end of Vaitahu Bay, 18 m. Mulloidichthys pfl uegeri: Western Indian Ocean, Mascarenes: MNHN 1965-23, 273 mm, Réunion, 21°7‘1“S, 55°34‘59“E; MNHN 1965-29, 188 mm, Réunion, 21°7‘1“S, 55°34‘59“E; SAIAB 70557, 333 mm, Rodrigues, north of Grand Bay; West Pacifi c, Hawaii: BPBM 8479, 258 mm, Oahu, Honolulu fish market; USNM 55516, 208 mm, Maui, Lahui; Marquesas Islands: BPBM 11057, 355 mm, Fatuhiva, off point at N end of Hanauu Bay, 60–75 ft; USNM 267494, 247 mm, Tahiti, Papeete market.

Mulloidichthys vanicolensis: Western Indian Ocean, South Africa: SAIAB 46235, 185 mm, Aliwal Shoal southern ledges; SAIAB 86369, 215 mm, KwaZulu-Natal, Coral Gardens, Sodwana Bay, 27°31.34’S, 34°41.15’E, 5–12 m; Mozambique: SAIAB 51025, 152 mm, Pinda Island, 14°13’S, 40°46’E; SAIAB 60425, 172 mm, Baixo Sao Joan, 26°24‘S, 32°55‘E; Tanzania: SAIAB 18056, 143-152 mm, Pemba Island, 05°08‘S, 39°40‘E; Oman: BPBM 36011, 208– 220 mm, Southern Oman, Kuria Muria Islands, Sawda Island, E end, 4–8 m; BPBM 36023, 6: 195– 224 mm, Kuria Muria Islands, Southern Oman, Sawda Island, SW side, 10 m; BPBM 39473, 4: 173–256 mm; Mauritius: SAIAB 1338, 3: 132–272 mm; SAIAB 5672, 135 mm; Eastern Indian Ocean, Indonesia: RMNH 25449, 153–199 mm, Sumatra, Sabang Bay, Pulu Weh; West Pacifi c, Hawaii: BPBM 1715, 179 mm, Oahu, Honolulu; BPBM 4082, 225 mm, Oahu, Honolulu; BPBM 4083, 194 mm, Oahu, Honolulu; BPBM 8804, 196-205 mm, Oahu, Honolulu fish market; BPBM 25675, 162 mm, Oahu, Honolulu market; Marcus Island: BPBM 2412, 287 mm; Pitcairn Group: BPBM 13266, 166 mm, Pitcairn Island, off Bounty Bay, reef, 80 ft; BPBM 16576, 227–237 mm, Oeno Atoll, N side of atoll; off small boat passage, 40–60 ft; Johnston Atoll: BPBM 29587, 151 mm, Johnston Island; under concrete ramp, 1–3 m; Samoa: RMNH, 11245, 183 mm, Savaii; French Polynesia: RMNH, 11297, 148 mm, Tahiti; Netherlands New Guinea: RMNH 25446, 144 mm, Hollandia, Landingstage, Viss. Onderz. Holl. Exp. Brongersma, cs. 1954/55; Indonesia: RMNH 31737, 130 mm, Java Sea, Selat Linta, E of Komodo, Indonesian-Dutch Snellis II Exp. 1984, 8º30‘S, 119º34.6‘E; RMNH, 25451, 130 mm, Jakarta, Bay of Batavia.

LITERATURE CITED

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