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Smithiana Special Publication
The South African Institute for Aquatic Biodiversity
ISSN: 1684-4149
Vol. 1, Num. 2, 2003, pp. 1-10

Smithiana, Publications in Aquatic Biodiversity, Special Publication 2 October, 2003, pp. 1-10

A new flatfish of the Indo-Pacific genus Asterorhombus (Pleuronectiformes:Bothidae)

Dannie A. Hensley1 and John E. Randall2

1) Department of Marine Sciences, University of Puerto Rico, Box 9013, Mayagüez, PR 00681-9013, USA.
2) Bishop Museum, 1525 Bernice St., Honolulu, HI 96817-2704, USA.

Code Number: sp03001


Five species of Asterorhombus are currently recognized, A. bleekeri, A. osculus, A. annulatus, A. intermedius, and A. fijiensis. A new species, Asterorhombus filifer, is described from 19 specimens. It is easily separated from A. bleekeri, A. osculus, and A. annulatus by its palmate gill-rakers and its detached and longer first dorsal-fin ray. It is distinguished from A. fijiensis by less body depth, longer first dorsal-fin ray on the average, simpler membrane on this ray, narrower interorbital width, and both eyes usually with one tentacle (vs. 1-9 tentacles on upper eye only). Asterorhombus fijiensis and A. filifer are the only species of the genus with males having a wider interorbital than females. Asterorhombus filifer most closely resembles A. intermedius, which differs in having a longer first dorsal ray, smooth edge on the membrane of this ray, usually more lateral-line scales, and males with greater interorbital width. Asterorhombus filifer is the most wide-ranging species of the genus, extending from the western Indian Ocean to the Hawaiian and Society islands, and is the only species of Asterorhombus occurring on the Pacific Plate.


Tanaka (1915) described Asterorhombus stellifer in Japanese as a new genus and species of bothid fish from a single specimen 110 mm in SL obtained from the Nagasaki Market. Norman (1934: 60) acknowledged Tanaka’s help in translating the diagnosis to English but noted that the diagnosis was very brief; he was unable to treat the genus and species with certainty.

Amaoka (1969) recognized Asterorhombus as a valid genus and synonymized A. stellifer with Platophrys (Arnoglossus) intermedius Bleeker (1865). Thus, A. stellifer Tanaka (1915) [= A. intermedius (Bleeker, 1865)] is the type species of the genus, fide Amaoka, 1969. Norman (1934) considered A. intermedius a species of Arnoglossus. Chabanaud (1948) put the species into the genus Asterorhombus without comments. Amaoka provided a detailed description of Asterorhombus in which he classified only the single species A. intermedius. An important character used by Amaoka to define this genus was the presence of palmate gill-rakers, i.e. short, broad, with strong spines (see Amaoka, 1969: Fig. 112C or Amaoka et al., 1994: Fig. 2A).

Norman (1931) described Engyprosopon fijiensis from a specimen 83 mm in TL from Levuka, Fiji. Norman (1934) wrote, “In the peculiar form of the gill-rakers, as well as in other characters, this species resembles Arnoglossus intermedius (Bleeker) and should perhaps be included in that genus.”Hensley (1984, 1986) placed E. fijiensis in Asterorhombus.

Amaoka and Arai (1998) described a new species, Asterorhombus osculus, from Western Australia, redescribed Arnoglossus bleekeri Macleay (1881) from Queensland, Gulf of Carpentaria, and Western Australia, and classified it in Asterorhombus. Norman (1934) had placed Arnoglossus bleekeri in Engyprosopon.

Norman (1934) synonymized Anticitharus annulatus Weber (1913) from Indonesia with Arnoglossus intermedius (Bleeker). Amaoka and Mihara (2001) examined the three syntypes of A. annulatus and recently collected 29 specimens from New Caledonia. They concluded that A. annulatus is a valid species of Asterorhombus, thus bringing the total species of the genus to five.

Asterorhombus osculus, A. bleekeri, and A. annulatus do not have palmate gill-rakers (Amaoka and Arai, 1998; Amaoka and Mihara, 2001) as was used in the generic definition given by Amaoka (1969) and Amaoka et al. (1994).

In the present paper we describe a new species of Asterorhombus with palmate gill-rakers. It is the most wide-ranging of the genus, occurring from the Comoro Islands to the Hawaiian Islands. It is the only species known to occur on the Pacific Plate. A revision of the genus Asterorhombus, which will include a key to the species, is planned by the first author.


Methods for making counts and measurements on whole specimens followed those of Hubbs and Lagler (1949) with the following changes: because all dorsal- and anal-fin rays are unbranched, all ray elements were counted as individual rays; lengths of pelvic-fin bases were measured from the base of the first ray to the base of the last ray. Unless otherwise stated, measurements on whole specimens refer to those made on the ocular side. Standard length (SL) was used throughout. These measurements were made with dial calipers or an ocular micrometer to the nearest 0.1 mm. For regression analysis and analysis of covariance (ANCOVA), all variates were transformed to natural logarithms (ln) in all analyses. In all analyses SL was treated as the independent variable (X). Differences were considered significant at the P < 0.05 level. Regressions used in tests for sexual dimorphism were compared by ANCOVA according to procedures of Snedecor and Cochran (1967). Tests for allometry were performed with the geometric-mean-functional-regression model of Ricker (1973). In this model, confidence limits are determined for the slope (v). If unity is outside these limits, allometry is assumed (positive if below, negative if above); isometry is assumed if unity is within the limits.

The three species of Asterorhombus with palmate gill-rakers (A. intermedius, A. fijiensis, and the new species) also have the first dorsal-fin ray separated from the rest of the fin. Two important characters that differentiate these species are the length of this ray and the morphology of the membrane on the ray. Until recently, the ray’s membrane has gone unnoticed by those describing or illustrating these species. An example of this is Norman’s (1934:197-198, Fig. 146) account of A. intermedius. Here he mentioned and showed the first dorsal-fin ray as “somewhat prolonged and expanded distally.”In another important description and drawing of the same species, Amaoka (1969:111-113, Fig. 66) described the first-dorsal fin ray as “prolonged,”but made no mention of the membrane of this ray or showed a membrane in his drawing. Recently the morphology of this membrane has been described and illustrated for both A. intermedius and A. fijiensis(Amaoka et al, 1994; Senou et al., 1994; Lin et al., 1995; Hensley and Amaoka, 2001). It is important to note that the membrane of this ray is very fragile, tends to fold and curl easily, and in A. intermedius and the new species, where it runs nearly the entire length of the ray, it is very often completely furled around the ray. This is often true of freshy caught specimens and very frequent in preserved material. Fig. 2 shows a paratype of the new species where the membrane is furled around the ray except for near the tip. In the holotype (Fig. 1) the membrane is not furled around the ray, although it became furled once fixed and preserved. Thus, it is very important to gently pull the membrane away from the ray with a probe when trying to examine it.

Sex was determined by inspection of gonads through a small incision made in the right side of the abdominal cavity. Institutional abbreviations are as listed by Leviton et al. (1985).

Asterorhombus filifer, sp. nov.

Figs. 1, 2, 3A

Arnoglossus intermedius [non Bleeker, 1865]: Woods, 1966: 65-66, Pl.124, fig. C (Marshall Ids, Bikini Atoll).

HOLOTYPE: BPBM 34871, 98.3 mm SL, male, NW Hawaiian Ids, Midway Atoll, wreckage at shore between cargo pier and launching ramp, depth 3-5 m, rotenone, J.E. Randall & J.L. Earle, 18 September 1991.

PARATYPES: INDIAN OCEAN: South Africa; SAIAB (RUSI) 64708, 28.0 mm SL, sex undetermined, KwaZulu-Natal, Sodwana Bay, north of Sponge Reef, 27°31' 04" S, 32°41’39" E, 26-28 m, P.C. Heemstra, 17 May 2001. Comoro Ids; ANSP 145284, 95.6 mm, female, Grande Comore, west side of fringing reef, 11°41’33" S, 43°14’27" E, 24.4 m or less, R/V Anton Bruun cr. 9, 27 November 1964. SAIAB (RUSI) 29059, 101 mm, female, Grande Comore, west coast north of Moroni, coral reef, 100-150 m offshore, 20-28 m, P.C. Heemstra, M.J. Smale, and R. Sanders, 20 October 1986. Seychelles; ANSP 107779, 59.6 mm, male, Amirante Ids, vicinity St. Joseph Id,southwest of Resource Id, 15.2-27.4 m, D. Dockins et al., 10 March 1964. ANSP 107789, 50.6 mm, male, Amirante Ids, D’Arros Id, off east side of Id, 29-33.5 m, D. Dockins et al., 9 March 1964. ANSP 108722, 76.1 mm, male, Amirante Ids, D’Arros Id, of east side of Id, 15.2-27.4 m, J.E. Böhlke et al., 5 March 1964. Mauritius; BPBM 24776, 60.5 mm, male, west coast of Flic en Flac, reef and somesand, 57 m, rotenone, J.E. Randall, D. Pelicier, J.S. Dench, and R.M. Bray, 7 April 1987. WESTERN PACIFIC OCEAN AND ADJACENT SEAS: Australia; AMS I.22578-038, 47.0 mm SL, sex undetermined, northern Queensland, Escape Reef, back reef slope, sand and coral in cave and gutter, 150°49' S, 145°50' E, D.J. Blake, G.R. Allen, and W.A. Starck, 28 January 1981. Chesterfield Bank; BPBM 33653, 42.6 mm, female, lagoon, southwest side, off Ile Longue, patch reef, 19°52' S, 158°19.8' E, 9 m, rotenone, J.E. Randall, M.L. Kulbicki, P.J. Doherty, et al., 24 August 1988. New Caledonia; BPBM 34263, 85.6 mm, male, lagoon near southeast end of St. Vincent Pass, rubble and sand near reef, 21°2.1' S, 165°57.8' E, 4 m, rotenone, M. Kulbicki, J.E. Randall, et al., 21 March 1990.Philippines; SAIAB (RUSI) 52650, 63.2 mm, male, Luzon, Bolinao, 16°23' N, 119°53' E, 30-35 m, P.C. Heemstra, 8 October 1995. USNM 260363, 47.5 mm, female, Negros, east of Bais, 36.6 m or less, V.G. Springer et al., 17 June 1978. Marshall Ids; USNM 141789, 24.1 mm, sex undetermined, Bikini Atoll, Arji Id, lagoon, 91 m offshore, 6-12 m, rotenone, E.S. Herald and V.E. Brock, 7 August 1946. Hawaiian Ids; BPBM 10028, 35 mm, sex undetermined, Oahu, off rocky islet at SW end of Waimea Bay, cave entrance, 10.7 m, Chemfish, J.E. Randall, L.A. Randall, and P.M. Allen, 27 July 1970. BPBM 22623, 60.8 mm, male, Oahu, Kaneohe Bay, W.J. Baldwin. MNHN 2001-1130, 79.2 mm, female, collected with holotype; NSMT-P 60932, 104.5 mm, female, Oahu, Kahe Pt., sand near edge of reef, 14 m, hand net, R. Holcom, early March 1997. Society Ids; ROM 61627, 31.0 mm, sex undetermined, Moorea, 300 m west of pass out of Cook Bay, 17°28’37" S, 149°49’45" W, dead coral, rock, coarse sand in broad (15 m wide) gully with gently sloping sides, 21-26 m, rotenone, R. Winterbottom, 13 December 1989.

DIAGNOSIS: A species of Asterorhombus having the following combination of characters: palmate gill-rakers; uniserial dentition in both jaws; first dorsal-fin ray separated from remainder of fin, elongate, 0.8-1.7 in head length, posterior membrane on ray broad, smooth-margined, and running nearly entire length of ray (Fig. 3A); both eyes usually with one tentacle, frequently branched; sexual dimorphism in interorbital width in specimens larger than ca. 60 mm SL, 8.9-18 in head length (1.6-3.1 % SL) in males, 15-21.2 (0.9-2.3 % SL) in females (Fig. 4); body depth 2.0-2.5 in SL (40.4-50.1 SL); lateral-line scales 56-67 (Fig. 5).

DESCRIPTION: Data for the holotype in parentheses are followed by those for the paratypes.

Morphometric data expressed as percent SL: head length (27) 27-31; body depth (46) 40-50; snout length (6.6) 6.6-9.7; upper-jaw length ocular side (11) 10-13, blindside (11) 10-13; lower-jaw length ocular side (15) 14-17, blind side (16) 14-18; lower-eye length (5) 5-8, upper-eye length (6) 6-7; interorbital width (2) 2-3 in males > 6 cm SL, 1-2 in females (Fig. 4); pectoral-fin length ocular side (13) 13-17, blind side (13) 11-14; length of third pelvic-finray ocular side (11) 9-11, blind side (not measurable) 810; length of pelvic-fin base ocular side (9) 8-11, blind side (6) 4-6; length of first dorsal-fin ray in specimens < 63 mm SL 27-39, in specimens > 76 mm SL (20) 17-28; length of dorsal-fin ray near middle of fin (12) 10-13; length of first anal-fin ray (not measurable) 5-8; length of anal-fin ray near middle of fin (11) 11-13.

Morphometrics expressed as ratios of larger measurements: head length (4) 3-4, body depth (2) 2-3, both in SL. Measurements in head length: snout length (4) 3.0-5; upper-jaw length ocular side (3) 2-3, blind side (3) 2-3; lower-jaw length ocular side (2) 2, blind side (2) 2;lower-eye length (5) 4-5, upper-eye length (5) 4-5; interorbital width (12) 9-18 in males > ca. 60 mm SL, 1333 in females (Fig. 4); pectoral-fin length ocular side (2) 2, blind side (2) 2-3; length of third pelvic-fin ray ocular side (3) 3, blind side (not measurable) 3-4; length of pelvic-fin base ocular side (3) 3-4, blind side (5) 5-8; length of first dorsal-fin ray in specimens < 63 mm SL 1, in specimens > 76 mm SL (1) 1-2; length of dorsal-fin ray near middle of fin (2) 2-3; length of first anal-fin ray (not measurable) 3-6; length of anal-fin ray near middle of fin (2) 2-3.

Dorsal-fin rays (88) 81-87; anal-fin rays (69) 63-68; pectoral-fin rays ocular side (12) 11-12, blind side (11) 910; pelvic-fin rays ocular side (6) 5-6 (usually 6), blind side (6) 5-6 (usually 6): lateral-line scales (67) 56-66 (Fig. 5): upper gill-rakers ocular side (1) 0-2, lower (10) 6-10, total (11) 7-11; vertebrae (counts not available) 10 + 26-28.

Anterior profile of head nearly straight, becoming curved above nuchal area; slight to moderate indentation anterior to ventral half of upper eye. Anterior margin of upper eye slightly posterior to anterior margin of lower eye. Both eyes usually with one monolobed or multilobed tentacle. Rear end of maxilla below or slightly posterior to anterior margin of lower eye. Interorbital region scaled on posterior 1/4-1/2 area in specimens > ca. 60 mm SL, often scaleless in smaller specimens.

Nostrils of ocular side at same horizontal level as dorsal margin of lower eye, about midway between snout tip and anterior margin of lower eye, anterior nostril tubular, posterior nostril slit-like; nostrils of blind side similar to those of ocular side, anterior nostril at vertical from base of second dorsal fin ray. Tip of isthmus at vertical from rear margin of lower eye. Teeth in single series in both jaws, equally developed on both sides, anterior teeth in both jaws not much enlarged.

Base of first pelvic-fin ray on ocular side at tip of isthmus; base of first pelvic-fin ray on blind side at vertical between third and fourth ray or base of fourth ray of pelvic-fin on ocular side. First dorsal-fin ray with very narrow membrane anteriorly; posterior membrane broad with smooth margin; both membranes extending from near base to tip of ray (Fig. 3A).

Colour of fixed and preserved specimens: ground colour on ocular side tan. Some specimens with few or no markings. Specimens retaining markings with many scattered dark spots of variable sizes and degrees of darkness. Approximately five rows of spots discernible: 1) row of relatively small spots immediately proximal to bases of dorsal and anal fins, 2) another row of darker, larger spots immediately proximal to these, and 3) row of usually four distinct spots on straight section of lateral line, posteriormost usually darkest and most distinct. Median fins and ocular-side ventral fin with small, scattered dark spots. First ray of dorsal fin usually with 3-6 dark areas or bands. Interorbital region frequently with three dark bars, anterior- and posteriormost usually broader and darker than middle bar. Blind side light tan with no dark markings.

Colour in life: Based on colour photographs of BPBM 34871, 37832, 34263 and RUSI 52650. Spots more distinct than in preserved specimens. Median fins and ocular-side ventral fin also with whitish spots. Dark bands on first dorsal-fin ray very distinct, separated by whitish areas.

DISTRIBUTION: Indian Ocean: South Africa; Seychelles; Comoro Ids; Mauritius. Pacific Ocean: Coral Sea (Chesterfield Bank, New Caledonia, Great Barrier Reef); Society Ids (Moorea); Hawaiian Ids (Oahu, Midway Id); Marshall Ids (Bikini Atoll); Philippines (Luzon, Negros).

SIZE, HABITAT, SEXUAL DIMORPHISM, AND DEVELOPMENTAL VARIATION: The largest specimen examined was a female 104.5 mm in SL. The species is found near reefs, in gullies, or cave entrances on sand or coral- or rock-rubble bottoms at depths of 3-57 m.

Regressions of interorbital widths on SL showed males with significantly higher elevations (i.e. wider interorbital widths) than females when compared by ANCOVA (Fig. 4). Length of the first dorsal-fin ray shows negative allometry (v = 0.6538 ± 0.1321), i.e. it becomes proportionately shorter with increased SL (compare Figs. 1 & 2).

ETYMOLOGY: This species is named Asterorhombus filifer from the combination of the Latin filum, filament, and fero, carry, in reference to the long first dorsal-fin ray, especially as it appears in small specimens and when the membrane is folded around the ray (Fig. 2).

REMARKS: Asterorhombus filifer differs from A. bleekeri , A. osculus , and A.annulatus in having 1) palmate gill-rakers, 2) the first dorsal-fin ray detached from remainder of fin and elongate (0.8-1.7 in head length; Fig. 6), and 3) sexual dimorphism in interorbital width (Fig. 4). In addition, it has uniserial dentition in both jaws vs. teeth in 2-3 rows in upper jaws of A. bleekeri and A. osculus (uniserial in A. annulatus).

Asterorhombus filifer can be distinguished from A. fijiensis by the following characters: (information on A. fijiensis from Amaoka et al., 1994 and Hensley and Amaoka, 2001): body depth (1.7-1.9 in SL in A. fijiensis vs. 2.0-2.3 in Asterorhombus filifer; interorbital width in specimens larger than ca. 6 cm SL (3.7-5.8 in head length in males, 5.8-9.0 in females vs. 9-18 in males, 13-33 in females; Fig. 4); length of first dorsal-fin ray (1.5-2.8 vs. 0.8-1.7 in head length); morphology of membrane on first dorsal-fin ray (confined to tip of ray, with many folds, projections, and filaments vs. present on nearly entire length of ray, with smooth margins; Fig. 3A); ocular tentacles (1-9, often branched tentacles on upper eye, but absent on lower eye vs. both eyes usually with one tentacle, which is often branched).

Asterorhombus intermedius most closely resembles A. filifer. It was not uncommon to find A. filifer misidentified as A. intermedius in various collections, and there has been at least one published misidentification (i.e. Woods, 1966). They resemble each other in their lesser body depth and having a narrow interorbital region compared to A. fijiensis. In addition, both species have the membrane of the first dorsal-fin ray along most or the entire length of the ray (Fig. 3), i.e. not the complicated membranous structure limited to the tip of the ray as found in A. fijiensis (Amaoka et al. 1994, Figs. 1 & 3). However, A. filifer and A. intermedius differ in several characters. Comparing male and female interorbital widths regressed on SL for A. intermedius shows no significant difference in elevations, i.e. no sexual dimorphism in this character (Fig. 7) . This is in contrast to A. filifer where elevations of regression lines are significantly different (Fig. 4). Although both species are similar in having the first dorsal-fin ray membrane on most of the length of the ray, they show differences in length of the ray and morphology of the membrane. This ray is much longer in A. filifer compared to A. intermedius (Fig. 6; ray length 12 mm in 104.7 mm SL A. intermedius holotype ) and A. fijiensis (Hensley and Amaoka, 2001). Edges of the membrane in A. filifer are smooth (Fig. 3A), while in A. intermedius they show sharp indentations, branching, and filamentous extensions (Fig. 3B; also see Lin et al., 1995: Fig. 4 and Hensley and Amaoka, 2001; membrane morphology in holotype of A. intermedius is very similar to that in Fig. 3B). Although both species usually have tentacles on both eyes, those in A. intermedius are only rarely branched (Hensley and Amaoka, 2001); and unbranched in holotype of A. intermedius, while most specimens of A. filifer have branched tentacles. Although there is a slight overlap in lateral-line scale counts, A. filifer usually has a higher count than A. intermedius (Fig. 5); (most lateral-line scales in holotype of A. intermedius were missing).

Assuming collecting of Asterorhombus species has been adequate to discern at least some rough patterns of distribution, three patterns can be seen. Three species of Asterorhombus appear to have restricted distributions: 1) A. osculus - Western Australia; 2) A. bleekeri - north and east coast of Queensland, including Gulf of Carpentaria, and Western Australia (Amaoka and Arai, 1998); and 3) A. annulatus - Flores, Timor, and Coral Sea (Amaoka and Mihara, 2001). Two species are much more broadly distributed: 1) A. fijiensis - western Indian Ocean, Western Australia, Ryukyu, South China Sea, Indo-Australian Archipelago, northern Great Barrier Reef, and Fiji; and A. intermedius - western Indian Ocean, including the Red Sea, to southern Japan, Taiwan, Indo-Australian Archipelago, and Tonga Islands (Lin et al., 1995; Hensley and Amaoka, 2001). The third pattern is shown by A. filifer. It extends from the western Indian Ocean to the Marshall, Hawaiian and Society islands; and it is the only species of Asterorhombus known to occur on the Pacific Plate.

Amaoka et al. (1994) were able to observe and photograph a live A. fijiensis. They found that the complicated membranous structure at the tip of the first dorsal-fin ray when in its natural state of folding, bore a remarkable resemblance to a small crustacean or fish. The live A. fijiensis would extend the first ray, bring the membranous structure near its mouth, and vibrate it. Although they never observed a prey species attracted to the membranous structure, they called it the esca and the ray the illicium. Senou et al. (1994) reported that the first dorsal-fin ray of A. intermedius and its wide membrane with complicated edging resembled a sandworm moving on the bottom. Lin et al. (1995) observed A. intermedius and saw that the first dorsal-fin ray was held up in the water and moved, resembling floating leaf-like algae. Shirai and Kitazawa (1998) observed a live A. intermedius and placed frozen krill in the tank. The fish approached the krill and rotated the first dorsal-fin at about a 45° angle above the vertical plane. It kept approaching the krill and finally darted at it and attacked it. The first dorsal-fin ray and membrane and associated behavior of these fishes have been interpreted as aggressive mimicry. If this is corroborated, they are the first known pleuronectiform fishes to exhibit this. An obvious need here is for more observations on live species of Asterorhombus. There have been no observations of living A. filifer. Although the first dorsal-fin ray of this species lacks the esca-type structure at the tip of the ray as in A. fijiensis, and the highly elaborate sculpturing of the membrane of this ray as in A. intermedius, it may possibly serve as a lure mimicking a worm or small eel.

COMPARATIVE MATERIAL EXAMINED: Asterorhombus intermedius: INDIAN OCEAN INCLUDING RED SEA: Red Sea; BMNH 1935.9.30.66-68, 3 specimens, 52.3-59.1 mm SL, males and females. BPBM 19871, 84.3 mm, female. MNHN 1967-593, 3, 77.0-122.3 mm, male and females. MNHN 1966-131, 2, 48.2-56.1 mm, male and female. MNHN 1966-132, 88.8 mm, female. MNHN 1966134, 105.3 mm, female. MNHN 1966-133, 95.5 mm, male. USNM 260387, 2, 43.2-66.8 mm, females. USNM 257040, 8, 39.1-82.9, males and females. USNM 260388, 49.3 mm, female. USNM 257041, 107.7 mm, female. Madagascar; SAIAB (RUSI) 52648, 10, 74.5-100.1 mm, males and females; SAIAB (RUSI) 52651, 26.2 mm, sex unknown. Zanzibar; MNHN 1964-244, 2, 66.0-80.4 mm, females. USNM 260391, 2, 76.8-89.5 mm, male and female. Seychelles; ANSP 107788, 2, 56.7-87.2 mm, females. ANSP 107783, 34.3 mm, female. BMNH 1908.3.23:131-2, 2, 96.6-107.9 mm, females. MNHN 1982-22, 102.2 mm, male.

MNHN 1987-1870, 99.1 mm, male. Maldive Ids; BMNH 1901.12.31:114-115, 2, 32.2-69.0 mm, male and female. Sri Lanka; USNM 257039, 3, 33.9-60.2 mm, males. USNM 260390, 102.4 mm, female. Western Australia (Dampier Archipelago); NTM S.11924.008, 81.7 mm, male. WESTERN PACIFIC OCEAN INCLUDING TIMOR AND ARAFURA SEAS; NTM S.13321-005, 2, 79.0-79.3 mm, males. NTM S.1330-002, 57.6-64.2 mm, male and female. NTM S.12982-001, 80.8 mm, male. NTM S.11899-008, 73.0 mm, male. NTM S.10011-019, 27.8 mm, female. Australia (Queensland); AMS I.20555-010, 97.2 mm, female. AMS IA .3002, 67.0-72.8 mm, males. AMS E.6689-90, 4, 72.6-75.9 mm, males and females. ANSP 113517, 20.1 mm, sex unknown. ANSP 113512, 47.1 mm, male. ANSP 113499, 3, 27.1-52.6 mm, females and sex unknown. ANSP 113515, 2, 38.8-45.0 mm, male and female. ANSP 113518, 43.8 mm, female. BMNH 1925.7.22.49-52, 4, 47.0-89.2 mm, females. SIO 69-222, 70.5 mm, male. Solomon Ids; BPBM 15974, 112.6 mm, female. Tonga; ANSP 153436, 7, 48.2-78.7 mm, males and females. Sulawesi; RMNH 6746 (Holotype), 104.7 mm, female. USNM 260377, 2, 72.7-98.3 mm, females. USNM 260362, 65.3 mm, female. Hong Kong; NTUM 02485, 84.4 mm, male. CAS-SU 60949, 114.6 mm, male. Japan (Shikoku); BSKU 8376, 120.3 mm, female.


For the loan of and information regarding specimens we are very grateful to K. Amaoka, M.E. Anderson, M.L. Bauchot, A.C. Bentley, J. Chambers, M. Desoutter, W.N. Eschmeyer, P.C. Heemstra, M.A Hine, M. Holloway, T. Iwamoto, S.L. Jewett, M. McGrouther, V. Mthombeni, J. G. Ness, P. Pruvost, D. Rennis, B.C. Russell, W.G. Saul, S.-C. Shen, G. Shinohara, W.F. Smith-Vaniz, A.Y. Suzumoto, M.J.P. van Oijen, H.J. Walker, A. Wheeler, and R.Winterbottom. D.L. Ballantine’s help with the photography and Loreen R. O’Hara’s help with the drawing are greatly appreciated.


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