Species Diversity 24: 229–246 Published online 25 November 2019 DOI: 10.12782/specdiv.24.229

Shallow-water Comatulids (Echinodermata: Crinoidea) from Ambon and Lombok Islands, Indonesia

Ichizo Kogo1,3, Toshihiko Fujita2, and Tsunemi Kubodera2 1 Osaka Museum of Natural History, 1-23 Nagai Park, Higashi-Sumiyoshi-ku, Osaka 546-0034, Japan E-mail: [email protected] 2 Department of Zoology, National Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba, Ibaraki 305-0005, Japan 3 Corresponding author (Received 28 March 2019; Accepted 11 September 2019)

Shallow-water comatulids were collected by scuba diving from Lombok and Ambon islands, Indonesia in 1992 and 1994. The collection consisted of a total of 35 species in 20 genera among Comatulidae, Himerometridae, Mariametridae, Colobometridae, and Antedonidae. Two species were newly recorded from Indonesian waters. Key Words: Comatulida, feather stars, Ambon Island, Lombok Island, Indonesia, shallow water.

shallower than 15 m, from Ambon Island on 25 Novem- Introduction ber–9 December 1992 and from Lombok Island on 3–15 February 1994. Sampling localities are shown in Fig. 1: Taxonomic studies of comatulids in Indonesian waters seven and three locations in Ambon and Lombok Island, were traditionally done by using the specimens collected respectively. The specimens were fixed in buffered 10% sea by expedition cruises, including the Challenger, the Sibo- water formalin and preserved in 70% ethanol. All specimens ga, and the Albatross Expeditions (Carpenter 1884, 1888; have been deposited in the National Museum of Nature and Hartlaub 1891; A. H. Clark 1912a, c, d, e, f, 1918, 1931, Science, Tsukuba (NSMT). 1936, 1941, 1947, 1950; Reichensperger 1913; A. H. Clark Morphological terms in the descriptions follow A. H. and A. M. Clark 1967). Recently, the French-Indonesian Clark (1915), Rowe et al. (1986), and Messing (1997, 2001). KARUBAR Expedition reported 15 species of 4 families of The following abbreviations and formulae are used. comatulids from deep waters (180–800 m) of the east Banda Cirri: the roman and arabic numerals mean the number Sea collected by dredging in 1991 (Messing et al. 2000). of cirri and the number of composed ossicles, respectively. The shallow-water (less than 50 m deep) comatulids of the Arms: Br means a division series. The preceding roman East Indian Archipelago were studied by Messing (1998b): numeral indicates the order from the base in ray (e.g., IBr: a total of about 100 comatulid species were recorded inclu- first division series in ray) and the following arabic number dung 43 species from the Banda Sea and 41 species from indicates the number of brachials, ossicles composing each Aru and Kai islands of Indonesia. Meanwhile in neighbor- division series (e.g., IBr 2: first division series composed of ing seas, Messing (1994) described 40 species of comatulids two brachials). Roman numbers hyphenated by plus sign at Madang, Papua New Guinea, Lane et al. (2000) listed 102 in parentheses indicating the position of syzygy (e.g., IIBr species from the South China Sea in the depth of 0–510 m, 4(3+4): second division series composed of 4 brachials Mekhova and Britayev (2012) revealed 32 species from where the third and fourth brachials joined by a syzygy). Southern Vietnam at 0–30 m of depth, and Messing and The lowercase br means a brachial of the division series Tay (2016) reported 39 species from Singaporean shallow- (with a preceding roman numeral) and free arm (without waters. roman numeral), and the following subscript arabic numer-

In this study, we report comatulid specimens from In- als indicate its position from the base (e.g., Ibr1: first brachi- donesian shallow waters (less than 15 m deep) of Ambon al of first division series, br1: first brachial of free arm). Plus Island in the Banda Sea and Lombok Island of the Lesser sign (+) indicates the position of syzygy (e.g., br1+2: first and Sunda Islands during a project study conducted by the Na- second arm brachials joined by syzygy). tional Museum of Nature and Science, Tokyo, in 1992 and Pinnules: P means a pinnule. The following subscript in- 1994. A synonymy, brief description of the specimens, and dicates the position from the arm base: arabic numerals distribution are given for each species. (e.g., P1, P2…) for the exterior pinnules, and alphabets (e.g., Pa, Pb…) for the interior pinnules on the inner side of arms. Pinnules on division series are labeled by Roman numerals Materials and Methods (e.g., PII , PIII…).

Specimens were collected by scuba diving at depths of

© 2019 The Japanese Society of Systematic Zoology 230 Ichizo Kogo et al.

Fig. 1. Sampling localities in Lombok and Ambon islands, Indonesia.

Material examined. 13 specimens. Ambon Is. NSMT Results E-8146 (1 specimen), Latuhalat, 10 m, December 1, 1992; NSMT E-8153 (1), Silale, 15 m, December 2, 1992. Lombok A total of 191 comatulid specimens, 110 and 81 speci- Is. NSMT E-8196 (1), Gili Air I., February 3, 1994; NSMT mens collected from Ambon and Lombok islands, Indone- E-8207 (1), E-8209 (1), E-8217 (1), E-8218 (1), E-8227 (1), sia, respectively, were referred to 35 species in 20 genera be- E-8228 (2), E-8231 (1), E-8237 (1), Gili Air I., February 5, longing to Comatulidae, Himerometridae, Mariametridae, 1994; NSMT E-8253 (1), Kodek, February 15, 1994. Colobometridae, and Antedonidae. Two species, Clarkco- Description. Centrodorsal large, thin discoidal, circu- manthus mirus and Clarkcomanthus exilis, are newly record- lar in aboral view, 6–9 mm across, 1–2 mm thick, with cir- ed from Indonesian waters. Comatulidae is the most diverse rus sockets in 1–2 irregular rows. Cirri XXIII–XLII, 20–29, family, represented by 20 species of 9 genera (excluding 15–28 mm long. Longest 2–3 cirrals (in 5th–10th) 1.2 times three unidentified species). longer than wide; cirrals beyond about 10th with a low distal transverse ridge; 6–7 distal ones with a blunt aboral spine. Order Comatulida A. H. Clark, 1908a Radials wholly concealed by centrodorsal or narrowly ex-

Family Comatulidae Fleming, 1828 posed. Division series very short. IBr 2; Ibr1 oblong, with Comatella nigra (Carpenter, 1888) proximal sides in contact laterally; Ibr2 (axil) triangular or rhombic, free laterally. II–IVBr all 2: IIIBr or IVBr twisted Actinometra nigra Carpenter, 1888: 304 (key). outward, consequently rays not in a plane. Arms 20–50

Comatella nigra—A. H. Clark 1908c: 208; 1909a: 395; 1911a: (usually 38–45), 80–115 mm long; first syzygy at br1+2 and 530; 1912a: 69, fig. 1; 1913: 3; 1918: 5, pl. 1; 1931: 92, pl. second syzygy at br3+4; middle brachials beyond br5 much 1 fig. 1, pl. 2 fig. 2, pl. 3 fig. 3; Reichensperger 1913: 83; shorter than wide with oblique articulation; distal brachial

H. L. Clark 1915: 101, 192; 1946: 25; A. M. Clark and margin everted and spiny. P1 excessively longer than other Rowe 1971: 15 (key); Gibbs et al. 1976: 110; Meyer and proximal pinnules, 21–25 mm long, of 47–60 segments; Macurda 1980: 83; Chen et al. 1988: 76, fig. 5; Fabricius combs of 13–22 teeth confluent with inner edge of pinnu-

1994: 1228 (table); Messing 1994: 239, 1998b: 189 (table); lars. Pinnule combs to P4–P6, rarely to P7–P10. First 2 pinnu- 2007: 99; Rowe and Gates 1995: 147; Lane et al. 2000: 476 lars of P2–P4 with a large fin-like expansion. (table); Kogo and Fujita 2005: 326; Mekhova and Britayev Distribution. Southern Vietnam (Mekhova and Britayev 2012: 917. 2012); Indonesia, northern Australia, Philippines (A. M. Comatella sp. aff. nigra—Kogo 2002: 4, fig. 3. Clark and Rowe 1971); Great Barrier Reef (Gibbs et al. 1976; Comatulids from Ambon and Lombok islands 231

Fabricius 1994); South China Sea (Lane et al. 2000); Papua twisted division series and stout arms, while this specimen New Guinea (Messing 1998b). Depth range: 0–120 m (Lane has plane division series and slender arms. et al. 2000). Distribution. Maldives, Sri Lanka, Bay of Bengal, Indo- nesia, northern Australia, Philippines, China (A. M. Clark Comatella stelligera (Carpenter, 1888) and Rowe 1971); Myanmar (A. M. Clark 1972); southern Vietnam (Mekhova and Britayev 2012); Papua New Guin- Actinometra stelligera Carpenter, 1888: 308, pl. 5 fig. 5a–d, ea, Micronesia (Meyer and Macurda 1980; Messing 1994); pl. 58 figs 1, 2; Hartlaub 1891: 104. Timor Sea (Rowe and Gates 1995); South Pacific Ocean Comatella stelligera—A. H. Clark 1909b: 136; 1912a: 68; (A. M. Clark and Rowe 1971); South China Sea (Lane et al. 1912b: 3; 1913: 3; 1918: 5, pl. 2; 1929: 636; 1931: 98, pl. 2000); Taiwan (Chen et al. 1988); Japan (A. M. Clark and 4 figs 6–9; 1936: 296; Reichensperger 1913: 84; H. L. Rowe 1971; Kogo 1998, 2006; Pilcher and Messing 2001; Clark 1915: 101; 1921: 13; 1946: 26; Gislén 1922: 18, figs Kogo and Fujita 2005). Depth range: 0–210 m (Lane et al. 7–9; 1940: 3; A. M. Clark and Rowe 1971: 15 (key); A. 2000). M. Clark 1972: 85; Gibbs et al. 1976: 110; Liao 1983: 263; Chen et al. 1988: 76, fig. 6; Liao and A. M. Clark 1995: 27, Alloeocomatella pectinifera (A. H. Clark, 1911c) fig. 12, pl. 1 fig. 2; Fabricius 1994: 1228 (table); Messing 1994: 239 (list); Rowe and Gates 1995: 147; Kogo 1998: Comissia pectinifer A. H. Clark, 1911c: 644; 1912a: 78; 1913: 14, fig. 9; 2002: 4; Lane et al. 2000: 476 (table); Pilcher and 6; 1929: 636; 1931: 255, pl. 25 figs 69, 70. Messing 2001: 16 (table); Kirkendale and Messing 2003: Comissia pectinifera—A. M. Clark and Spencer Davis 1966: 527; Kogo and Fujita 2005: 326; Mekhova and Britayev 598 (list); A. M. Clark and Rowe 1971: 14 (key); Messing 2012: 918. 1994: 239 (list). Actinometra maculata Carpenter, 1888: 307, pl. 5 fig. 1a–d, Comissia magnifica (not of Gislén, 1922)—Kogo 1998: 20, pl. 55 fig. 2; Hartlaub 1891: 105. fig. 15. Comatella maculata—A. H. Clark 1909b: 138; 1911b: 16; Alloeocomatella pectinifera—Messing 1995: 445, figs 3e–g, 1912a: 70; 1912b: 4; 1913: 3; 1918: 7; 1931: 112, pl. 20 fig. i, j, 5, 6; 1998b: 189 (table); 2001: 291; 2007: 99; Mar- 20; 1937: 98; Reichensperger 1913: 84; H. L. Clark 1915: shallCrossland and Price 1999: 25; Kogo 2002: 6; Kogo 101; 1921: 12; 1946: 26; Gislén 1922: 16, figs 4–6; A. M. and Fujita 2005: 328. Clark and Spencer Davis 1966: 598 (list); A. M. Clark and Rowe 1971: 15 (key); A. M. Clark 1972: 84; Meyer and Material examined. 4 specimens. Ambon Is. NSMT Macurda 1980: 83; Liao, 1983: 263, pl. 1; Chen et al. 1988: E-8151 (1 specimen), Silale, 15 m, December 2, 1992; NSMT 76, fig. 4; Fabricius 1994: 1228 (table); Messing 1994: 239 E-8167 (1), Silale, 15 m, December 3, 1992; NSMT E-8170 (list); Liao and A. M. Clark 1995: 27, pl. 1 fig. 1; Rowe and (1), Eri, 10 m, December 4, 1992. Lombok Is. NSMT E-8194 Gates 1995: 146; Lane et al. 2000: 476 (table); Kogo 2006: (1), Gili Air I., February 3, 1994. 225. Description. Centrodorsal thin discoidal, roundish or Actinometra notata Carpenter, 1889: 312, pl. 26 figs 6–12. pentagonal in aboral view, 4–7 mm across, 1–1.5 mm thick, Antedon basset-smithi Bell, 1894: 399, pl. 24. with cirrus sockets in 2 irregular rows. Cirri XXXII (or wholly lost), 18–19, about 15 mm long. Longest cirrals (4th– Material examined. 1 specimen. Lombok Is. NSMT 7th) about 2 times longer than wide; cirrals beyond 6th with E-8214 (1 specimen), Gili Air I., February 5, 1994. aboral spine. Radials wholly concealed by centrodorsal. IBr

Description. Centrodorsal discoidal, 4.2 mm across, 2: Ibr1 trapezoidal, free laterally, Ibr2 (axil) low pentagonal. 1 mm thick, with cirrus sockets in 1 partly 2 rows; aboral IIBr not present. Arms 10, 90–130 mm long; first syzygy at pole flat. Cirri XXII, 18–19, 13 mm long. Longest cirrals br3+4. P1 longest, 15–18 mm long, of 40–50 segments; combs (5th–7th) nearly 2 times longer than wide; cirrals beyond of 22–33 teeth confluent with outer edge of pinnulars, exces- 5th–6th with a distal transverse edge; 5–8 distal cirrals with sively long, nearly a half of pinnule length. Pinnule combs a middle transverse ridge. Radials wholly concealed by cen- to P4. trodorsal. IBr 2; Ibr1 oblong in contact laterally: Ibr2 (axil) Distribution. Indonesia, Philippines (A. M. Clark and low pentagonal. IIBr all 2. Rays in a plane. Ossicles of divi- Rowe 1971); Maldives, northern Australia, Papua New sion series and basal brachials smooth without distal spines. Guinea, Micronesia, Melanesia (Messing 1995, 1998b); Su-

Arms 20, 80 mm long. First syzygy at br1+2; middle brachials matra (MarshallCrossland and Price 1999); Japan (Kogo beyond br5 slightly shorter than wide, with oblique articu- 1998, 2002; Kogo and Fujita 2005). Depth range: 3–23 m lation and slightly everted distal margin. P1 longest, 16 mm (Messing 1995). long, of 45 segments; combs of 13–15 teeth confluent with inner edge of pinnulars. Pinnule combs to P4. First 2 pinnu- Comatula pectinata (Linnaeus, 1758) lars of P2–P4 each with a large fin-like expansion. Remarks. Among the 10 Comatella specimens from Gili Asterias pectinata Linnaeus, 1758: 663. Air of Lombok Island, 9 larger specimens (20–50 arms) Comatula pectinata—A. H. Clark 1909a: 394; 1909b: 148; were identified as C. nigra, and only this smallest specimen 1931: 339, pl. 20 fig. 49, pl. 33 fig. 100, pl. 34 fig. 101, pl. (20 arms) was as C. stelligera. The larger specimens have 35 fig. 102, pl. 36 fig. 103, pl. 37 fig. 104, pl. 38 figs 105– 232 Ichizo Kogo et al.

107; H. L. Clark 1915: 101; A. M. Clark and Rowe 1971: M. Clark 1972: 76; Fishelson 1974: 184 (list); Gibbs et al. 14 (key); A. M. Clark 1972: 85; Gibbs et al. 1976: 110; Fa- 1976: 108; Tortonese 1977: 275 (list); Meyer and Macurda bricius 1994: 1228 (table); Liao and A. M. Clark 1995: 28, 1980: 75; Marshall and Rowe 1981: 382; Chen et al. 1988: fig. 13; Rowe and Gates 1995: 148; Kogo 1998: 23, fig. 17; 75, figs 3, 23D; Fabricius 1994: 1228 (table); Liao and A. 2002: 7; Lane et al. 2000: 476 (table). M. Clark 1995: 14, fig. 4; Messing 1994: 239 (list); 1998b: Comatula cf. pectinata—Messing and Tay 2016: 631, fig. 189 (table); Rowe and Gates 1995: 139; Kogo 1998: 17, fig. 3D–F. 12; 2002: 6; MarshallCrossland and Price 1999: 25; Kogo Actinometra pectinata—Carpenter 1888: 284, pl. 53 figs 15– and Fujita 2005: 327; Mekhova and Britayev 2012: 915; 22. Messing and Tay 2016: 633, fig. 4A–C. Alecto purpurea Müller, 1843: 132. Capillaster cf. multiradiatus—Messing 2007: 98. Comatula purpurea—A. H. Clark 1911a: 532; 1918: 271, pl. Capillaster clarki Reichensperger, 1913: 85. 14 fig. 16; 1931: 360, pl. 39 fig. 109, pl. 40 fig. 111, pl. 41 Comaster fimbriata—A. H. Clark 1908c: 201. fig. 114. Comaster coppingeri—A. H. Clark 1908c: 202.

Material examined. 3 specimens. Ambon Is. NSMT Material examined. 17 specimens. Ambon Is. NSMT E-8143 (1 specimen, juvenile), E-8147 (2), Latuhalat, 10 m, E-8115 (1 specimen), E-8118 (2), E-8133 (1), Pombo I., December 1, 1992. 10 m, November 28, 1992; NSMT E-8138 (1), Tial, 10 m, Description. Centrodorsal thin discoidal, 1–2.2 mm November 30, 1992; NSMT E-8150 (1), Latuhalat, 15 m, De- across, up to 0.6 mm thick, roundish in aboral view, with cember 1, 1992; NSMT E-8171 (1), E-8172 (1), E-8174 (1), cirrus sockets in 1 aligned row. Cirri VII–VIII, 11–14, E-8175 (1), E-8176 (1), E-8177 (1), E-8182 (1), Eri, 10 m, 5–8 mm long. Longest cirrals (4th–5th) 1.2 times longer than December 4, 1992. Lombok Is. NSMT E-8208 (1), E-8213 wide; cirrals smooth without aboral tubercle. Radials nar- (1), Gili Air I., February 5, 1994; NSMT E-8250 (1), E-8259 rowly exposed, in contact laterally. IBr 2(1+2): Ibr1 trape- (1), Kodek, February 15, 1994. zoidal, in contact laterally; Ibr2 (axil) triangular, free later- Description. Centrodorsal thin discoidal, circular or pen- ally. IIBr not present. Arms 10; anterior arms 85 mm, pos- tagonal in aboral view, 3–5 mm across, 0.7–1.2 mm thick, terior arms 50 mm long; first syzygy at br1+2, second usually with cirrus sockets in 1–2 irregular rows. Cirri XVI–XXVI, th th at br3+4. P1 longest, 7–8 mm long, of 30–32 segments; combs 12–24, 8–13 mm long. Longest cirrals (5 –7 ) 1.2–1.5 times of 4–7 teeth confluent with outer edge of pinnulars. Pinnule longer than wide; cirrals beyond 7th with aboral spine. Ra- combs to P2. dials almost wholly concealed by centrodorsal. IBr 2: Ibr1 Remarks. Comatula purpurea (Müller, 1843) was synony- trapezoidal, in contact laterally; Ibr2 (axil) triangular, free mized with Comatula pectinata by Rowe and Gates (1995: laterally. IIBr 4(3+4), rarely 2. IIIBr usually 4(3+4). Arms

148). These two species had been distinguished mainly by 12–22, 50–160 mm long. First syzygy at br2+3; middle bra- the arrangement of cirri (A. H. Clark 1931: 320, in key): chials beyond br5 wedge-shaped, excessively shortened to cirri are segregated in the interradial angles in the for- about 1/3 long of width, with spiny distal margin. PII lon- mer species, while not in the latter. The specimens (NSMT gest, 9–14 mm long, of 32–44 segments; combs of 10–17 E-8143 and E-8147) show both types of cirri arrangement. teeth confluent with outer edge of pinnulars; first comb

We agree the synonymization by Rowe and Gates (1995). tooth transverse. Pinnule combs to P4 or P5, rarely to middle Distribution. Singapore (Messing and Tay 2016); Indone- pinnules. sia, northern Australia, Philippines (A. M. Clark and Rowe Distribution. Western Indian Ocean, Red Sea, Anda- 1971); Great Barrier Reef (Gibbs et al. 1976; Fabricius 1994); man Is., Nicobar Is., Myanmar, Indonesia, China (A. M. South China Sea (Lane et al. 2000). Depth range: 0–120 m Clark and Rowe 1971; A. M. Clark 1972); southern Vietnam (Lane et al. 2000). (Mekhova and Britayev 2012); Sumatra (MarshallCrossland and Price 1999); Singapore (Messing and Tay 2016); Papua Capillaster multiradiatus (Linnaeus, 1758) New Guinea, Micronesia (Meyer and Macurda 1980; Mess- ing 1994, 2007); South China Sea (Lane et al. 2000), Tai- Asterias multiradiata Linnaeus, 1758: 663; Müller 1843: 133. wan (Chen et al. 1988); Japan (A. M. Clark and Rowe 1971; Comatula (Alecto) multiradiata—Müller 1849: 261. Kogo 1998, 2002; Kogo and Fujita 2005). Depth range: 0–ca. Actinometra multiradiata—Carpenter 1882: 521, 523; 1888: 300 m (Lane et al. 2000). 322, pl. 66 figs 1–3. Comaster multiradiata—A. H. Clark 1909a: 391. Phanogenia multibrachiata (Carpenter, 1888) Capillaster multiradiata—A. H. Clark 1909b: 134; 1911a: 530; 1911b: 16; 1912a: 76; 1912b: 4; 1913: 5; 1918: 14; Actinometra multibrachiata Carpenter, 1888: 299, pl. 56 figs 1931: 173, pl. 8 fig. 5, pl. 11 fig. 30, pl. 13 fig. 34, pl. 14 figs 3, 4. 35, 36, pl. 81 figs 222, 223; 1934: 10; 1936: 296; 1937: 99; Comaster multibrachiata—A. H. Clark 1912a: 86; 1931: 437, Reichensperger 1913: 84; H. L. Clark 1915: 101; 1921: 14; pl. 46 fig. 142. 1946: 27; Gislén 1938b: 9. Comaster multibrachiatus—A. M. Clark and Rowe 1971: 16 Capillaster multiradiatus—A. M. Clark and Spencer Davis (key); Messing 1994: 239 (list); 1998b: 189 (table); Liao 1966: 598 (list); A. M. Clark and Rowe 1971: 15 (key); A. and A. M. Clark 1995: 25, fig. 11; Rowe and Gates 1995: Comatulids from Ambon and Lombok islands 233

146; Kogo 1998: 26, fig. 20; Lane et al. 2000: 476 (table). NSMT E-8219 (1), E-8236 (1), Gili Air I., February 5, 1994. Phanogenia multibrachiata—A. H. Clark 1909a: 392; 1913: Description. Centrodorsal small, pentagonal or stellate 13; Messing 1998a: 206 (list), 2001: 290; Kogo 2002: 7; in aboral view, 1.5–2.5 mm across, up to 0.5 mm thick, lack- Kogo and Fujita 2005: 329; Mekhova and Britayev 2012: ing cirrus sockets and cirri. Radials trapezoidal, in contact

920. laterally. IBr 2: Ibr1 oblong or trapezoidal, in contact later- ally, Ibr2 (axil) low pentagonal, free laterally. IIBr 4(3+4) Material examined. 5 specimens. Ambon Is. NSMT or 2 (1+2). IIIBr–VBr 2(1+2), rarely 4(3+4). Division se- E-8155 (1 specimen), E-8157 (1), Silale, 15 m, December 2, ries narrow and widely separated from each other. Aboral 1992, E-8184 (1; juvenile of 14 arms), Leahari, December 8, interradial area of disk with many calcareous depositions.

1992. Lombok Is. NSMT E-8246 (2), Gili Air I., February 5, Arms 40–97, 50–200 mm long. First syzygy at br1+2. Bra- 1994. chials of middle to distal arm with spiny distal margins. PII Description. Centrodorsal thin discoidal, circular longest, 9–25 mm long, of 35–50 segments with distal comb or roundish pentagonal in aboral view, 5–6 mm across, of 10–16 teeth confluent with outer edge of pinnulars; first

1–2 mm thick, with cirrus sockets in 2–3 irregular rows. comb tooth transverse. Pinnule combs to P4, rarely to mid- Cirri short and stout, XXV–XLVI, 10–13, 10–14 mm long. dle and distal pinnules. Pinnulars of middle to distal pin- Longest cirrals (3rd–5th) 2 times longer than wide; succeed- nules with spiny distal margin. A dark mid-aboral line usu- ing cirrals gradually shortened, with distal transverse ridge ally present on division series and proximal arms. or aboral spine. Radials narrowly exposed, in contact lat- Distribution. Maldives, Bay of Bengal, Singapore, Indo- erally. IBr 2: Ibr1 oblong in contact laterally; Ibr2 (axil) low nesia, China (A. M. Clark and Rowe 1971); southern Viet- pentagonal, free laterally. IIBr–VBr 2(2+1) or 4(3+4). nam (Mekhova and Britayev 2012); Malaysia (Owen et al.

Arms 55–124, 77–158 mm long; first syzygy at br1+2. PII lon- 2009); Micronesia (Zmarzly 1984; Messing 1998b; Kirkend- gest, 13–17 mm long, of 40–62 segments; combs of 7–12 ale and Messing 2003; Owen et al. 2009); South China Sea teeth confluent with outer edge of piulars; first comb tooth (Lane et al. 2000); South Pacific Ocean (A. M. Clark and transverse. Pinnule combs to P4, rarely to middle pinnules. Rowe 1971); Japan (A. M. Clark and Rowe 1971; Pilcher and Distribution. Bay of Bengal, Indonesia, Philippines, Messing 2001; Kogo and Fujita 2005). Depth range: 4–50 m China (A. M. Clark and Rowe 1971); southern Vietnam (Lane et al. 2000). (Mekhova and Britayev 2012); western, northern, and east- ern Australia (Rowe and Gates 1995); Sulu Sea, Papua New Comaster nobilis (Carpenter, 1884) Guinea (Messing 1994, 1998b); South China Sea (Lane et al. 2000); Taiwan (Liao and A. M. Clark 1995); Japan (A. M. Actinometra nobilis Carpenter, 1884: 55; 1888: 336, pl. 65. Clark and Rowe 1971; Kogo 2002; Kogo and Fujita 2005). Comanthus nobilis—A. H. Clark 1909a: 395. Depth range: 20–83 m (Lane et al. 2000). Comanthus (Comanthina) nobilis—A. H. Clark 1909b: 143. Comanthina nobilis—Hoggett and Rowe 1986: 126; Rowe Phanogenia gracilis (Hartlaub, 1890) et al. 1986: 243, fig. 8b; Rowe and Gates 1995: 142; Kogo 1998: 31, fig. 26; Messing 1998b: 189 (table); Mar- Actinometra gracilis Hartlaub, 1890: 187; 1891: 111, pl. 5 fig. shallCrossland and Price 1999: 25; Lane et al. 2000: 475 55. (table). Comaster gracilis—A. H. Clark 1909b: 139; 1912a: 84; 1913: Comaster nobilis—Messing 1998a: 206 (list); Pilcher and 12; 1918: 35 (key); 1931: 430, pl. 47 figs 143, 144; 1936: Messing 2001: 16 (table); Kogo 2002: 9; Kogo and Fujita 296; 1937: 99; 1952: 266; A. M. Clark and Spencer Davis 2005: 331; Mekhova and Britayev 2012: 917, fig. 7. 1966: 598 (list); A. M. Clark and Rowe 1971: 16 (key); A. Comanthina schlegelii (not of Carpenter, 1881)—Reichens- M. Clark 1972: 84; Meyer and Macurda 1980: 81; Zmarzly perger 1913: 88; A. H. Clark 1931: 466 (part), pl. 54 fig. 1984; 108, fig. 3d; Fabricius 1994: 1228 (table); Messing 159, pl. 55 figs 160, 161; Gislén 1938b: 9. 1994: 239 (list); 1998b: 189 (table); Liao and A. M. Clark Comanthina schlegeli (not of Carpenter, 1881)—Utino- 1995: 24 (key); Kogo 1998: 25, fig. 19; Lane et al. 2000: mi and Kogo 1965: 265; Liao and A. M. Clark 1995: 18 476 (table). (part), fig. 8. Phanogenia gracilis—Messing 1998a: 206 (list); 2001: 290; 2007: 98; Pilcher and Messing 2001: 16 (table); Kirken- Material examined. 6 specimens. Ambon Is. NSMT dale and Messing 2003: 524; Kogo and Fujita 2005: 329; E-8117 (1 specimen), Pombo I., 10 m, November 28, 1992; Owen et al. 2009: 1517; Mekhova and Britayev 2012: 920. NSMT E-8148 (1), Latuhalat, 10 m, December 1, 1992; NSMT E-8161 (1), Silale, 15 m, December 2, 1992. Lombok Material examined. 17 specimens. Ambon Is. NSMT Is. NSMT E-8205 (1), Gili Air I., February 3, 1994; NSMT E-8108 (1 specimen), Pombo I., lagoon, November 27, 1992; E-8220 (1), E-8244 (1), Gili Air I., February 5, 1994. NSMT E-8131 (1), E-8134 (1), E-8137 (1), Pombo I., 10 m, Description. Centrodorsal small, pentagonal or stellate in November 28, 1992; NSMT E-8156 (1), E-8158 (1), E-8159 aboral view, 4–5 mm across, up to 0.5 mm thick, with 0–16 (1), E-8164 (1), E-8165 (2), Silale, 15 m, December 2, 1992; small cirrus sockets in 1 aligned row. Cirri weak or absent, NSMT E-8192 (2), Lilibooi, 15 m, December 9, 1992. Lom- 0–XI, 14–16, up to 15 mm long; cirrals smooth without ab- bok Is. NSMT E-8198 (3), Gili Air I., February 3, 1994; oral spine. Radials trapezoidal, in contact laterally. IBr 2: 234 Ichizo Kogo et al.

th th Ibr1 oblong, IBr2 (axil) low pentagonal, both in contact later- 11–13 mm long. Longest cirrals (4 –6 ) slightly longer ally. IIBr 4(3+4), rarely 2: IIbr1 in contact laterally, IIbr2 free than wide; cirrals smooth without aboral spine. Radials laterally. IIIBr usually inner side 4(3+4) and outer 2 (‘2-4- narrowly exposed, in contact laterally. IBr 2: Ibr1 oblong, 4-2 order’). IVBr and VBr all 4(3+4). Interradial area with in contact laterally; Ibr2 (axil) low pentagonal, free later- thin irregular polygonal perisomic plates. Arms 66–109, ally. IIBr 4(3+4). IIIBr usually 4(3+4) on inner side and 2

80–125 mm long. First syzygy at br3+4. PII longest, 20 mm on outer side of ray. IVBr and VBr 4(3+4). Interradial area long, of about 55 segments with distal comb of 7–10 teeth with thin irregular polygonal perisomic plates. Arms 48–83, confluent with inner edge of pinnulars; 3–5 pinnulars with 62–85 mm long; first syzygy at br3+4. PII longest, 13–17 mm paired teeth of same size at both edges; proximal 2–3 teeth long, of 34–50 segments; combs of 6–10 teeth confluent transverse. Pinnule combs to P5, rarely to P6–P10. with inner edge of pinnulars; 3–4 pinnulars with paired Remarks. Hoggett and Rowe (1986) distinguished this teeth of same size at both edges; first comb tooth transverse. species from C. schlegelii. On the other hand, Messing and Pinnule combs to P3 or P4, rarely to P5. Tay (2016) restored this species to synonymy under C. Distribution. Maldives, Sri Lanka, Bay of Bengal, Indo- schlegelii, following A. H. Clark (1931). These two species nesia, northern and western Australia, Philippines, China, are distinguishable by number of arms (Carpenter 1884: 55), South Pacific Ocean (A. M. Clark and Rowe 1971; Rowe and by cirrus number, comb distribution, and color pattern and Gates 1995); southern Vietnam (Mekhova and Britayev (Rowe et al. 1986: 244). We treated here C. nobilis and C. 2012); Sumatra (MarshallCrossland and Price, 1999); Sin- schlegelii as separate species. gapore (Messing and Tay 2016); Borneo, South China Distribution. Southern Vietnam (Mekhova and Britayev Sea (Lane et al. 2000); Papua New Guinea (Messing 1994, 2012); Sumatra (MarshallCrossland and Price 1999); west- 1998a); Micronesia (Meyer and Macurda 1980; Zmarzly ern, northern, and eastern Australia, Coral Sea (Rowe et al. 1984; Messing 1998b, 2007; Kirkendale and Messing 2003); 1986); Sulu Sea, Papua New Guinea (Messing 1998b); Bor- Taiwan (Chen et al. 1988; Liao and A. M. Clark 1995); Japan neo, South China Sea (Lane et al. 2000); Japan (Kogo 1998, (A. M. Clark and Rowe 1971; Pilcher and Messing 2001; 2002; Pilcher and Messing 2001). Depth range: 8–92 m Kogo and Fujita 2005). Depth range: 3–30 m (Lane et al. (Rowe et al. 1986). 2000).

Comaster schlegelii (Carpenter, 1881) Clarkcomanthus luteofuscum (H. L. Clark, 1915)

Actinometra schlegelii Carpenter, 1881: 210. Comanthus luteofuscum H. L. Clark, 1915: 102. Actinometra schlegeli—Carpenter 1884: 55; 1888: 331 (key). Comanthus luteofusea—H. L. Clark 1921: 18. Comanthus (Comanthina) schlegelii—A. H. Clark 1911a: 536. Clarkcomanthus luteofuscum—Rowe et al. 1986: 233, figs 2c, Comanthus schlegelii—H. L. Clark 1915: 101; 1921: 20; A. H. d, 7a, b; Fabricius 1994: 1228 (table); Messing 1994: 239 Clark 1937: 99. (list); 1998b: 189 (table); 2007: 97; Rowe and Gates 1995: Comanthina schlegelii—A. H. Clark 1912a: 91; 1912b: 8; 142; Kogo 1998: 50, fig. 39; MarshallCrossland and Price 1913: 14; 1918: 46; 1931: 466 (part), pl. 53 fig. 158, pl. 64 1999: 25; Kirkendale and Messing 2003: 530; Kogo and fig. 180; H. L. Clark 1946: 33; A. M. Clark and Spencer Fujita 2005: 335; Summers et al. 2014: 336. Davis 1966: 598 (list); Gibbs et al. 1976: 109; Rowe et al. Comanthus samoana (not of A. H. Clark, 1909f)—A. H. 1986: 244, figs 2e, 8c, 8d; Messing 1994: 239 (list); 1998b: Clark 1931: 593 (part); 1936: 298 (part). 189 (table); Rowe and Gates 1995: 142; Kogo 1998: 30, Comanthus samoanus (not of A. H. Clark, 1909f)—Meyer fig. 24; MarshallCrossland and Price 1999: 25; Laneet al. and Macurda 1980: 81. 2000: 476 (table). Cenolia samoana (not of A. H. Clark, 1909f)—H. L. Clark Comanthina schlegeli—A. M. Clark and Rowe 1971: 16 1946: 36 (part). (key); Meyer and Macurda 1980: 75; Zmarzly 1984: 108, fig. 3c; Chen et al. 1988: 76, figs 9, 23G; Fabricius 1994: Material examined. 3 specimens. Ambon Is. NSMT 1228 (table); Liao and A. M. Clark 1995: 18 (part). E-8141 (1 specimen), E-8142 (1), Tial, 10 m, November 30, Comaster schlegelii—Messing 1998a: 206 (list); 2007: 96; 1992; NSMT E-8190 (1), Lilibooi, 15–20 m, December 9, Pilcher and Messing 2001: 16 (table); Kirkendale and 1992. Messing 2003: 528; Kogo and Fujita 2005: 331; Mekhova Description. Centrodorsal small, discoidal, 3.1–3.3 mm and Britayev 2012: 917; Messing and Tay 2016: 638. across, up to 1.2 mm thick, with cirrus sockets aligned in 1, Actinometra duplex Carpenter, 1888: 335. partly 2 rows. Cirr XVII–XX, 13–15, 8–10 mm long. Longest Comanthus duplex—A. H. Clark 1908c: 205. cirrals (5th–7th) 1.2 times longer than wide; cirrals beyond 7th with transverse ridge; 2–4 distal cirrals usually with ab- Material examined. 3 specimens. Ambon Is. NSMT oral spine. Radials mostly concealed by centrodorsal. IBr 2:

E-8127 (1 specimen), E-8132 (1), Pombo I., 10 m, November Ibr1 trapezoidal, in contact laterally, Ibr2 (axil) triangular, 28, 1992; NSMT E-8163 (1), Silale, 15 m, December 2, 1992. free laterally. IIBr 2 or 4(3+4). III–IVBr 4(3+4), rarely 2. Description. Centrodorsal thin discoidal, 4–5 mm across, Arms stout, 33–46, 60–75 mm long; anterior arms not much

1 mm thick, with cirrus sockets aligned in 1, partly 2 rows. longer than posterior arms; first syzygy at br3+4. PII longest, Cirri XVIII–XXV including 2–6 rudimentary ones, 15–17, 11–14 mm long, of 30–35 segments; combs of 10–12 dis- Comatulids from Ambon and Lombok islands 235

crete teeth confluent with outer edge of pinnulars. IIP , PIII, Kogo 1998: 51, fig. 40; MarshallCrossland and Price 1999: and P1 much longer than succeeding pinnules. Pinnule 25; Pilcher and Messing 2001: 16 (table); Kirkendale and combs to P2. Messing 2003: 529; Kogo and Fujita 2005: 335; Summers Distribution. Western, northern, and eastern Australia, et al. 2014: 336. Indonesia, Melanesia (Rowe et al. 1986); Sumatra (Mar- shallCrossland and Price 1999); Great Barrier Reef (Fabri- Material examined. 19 specimens. Ambon Is. NSMT cius 1994); Sulu Sea, Papua New Guinea, Micronesia (Mess- E-8169 (1 specimen), E-8180 (1), Eri, 10 m, December 4, ing 1994, 1998b, 2007; Kirkendale and Messing 2003); Japan 1992. Lombok Is. NSMT E-8199 (3), Gili Air I., February 3, (Kogo and Fujita 2005). Depth range: 0–18 m (Rowe et al. 1994; NSMT E-8206 (1), E-8212 (1), E-8216 (1), E-8222 (1), 1986). E-8223 (1), E-8225 (2), E-8229 (1), E-8230 (1), E-8234 (1), E-8238 (1), Gili Air I., February 5, 1994; NSMT E-8251 (2), Clarkcomanthus alternans (Carpenter, 1881) E-8254 (1), Kodek, February 15, 1994. Description. Centrodorsal small, discoidal, circular, Actinometra alternans Carpenter, 1881: 208; 1888: 330 (key). pentagonal or stellate in aboral view, 2–3 mm across, up Comantheria alternans—A. H. Clark 1912a: 90; 1913: 13; to 0.5 mm thick, with 0–12 cirrus sockets in 1 aligned row. 1931: 488; Reichensperger 1913: 87. Cirri absent or weak, if present, up to X, 11–13, 6–8 mm Comanthus alternans—A. H. Clark 1908c: 206; H. L. Clark long. Longest cirrals (4th–6th) about 1.5 times longer than 1915: 101; 1921: 16; Rowe et al. 1986: 224, fig. 6a; Fa- wide; 2–3 most distal cirrals with a small aboral spine or bricius 1994: 1228; Messing 1994: 239 (list); 1998b: 189 tubercle. Radials narrowly exposed, in contact laterally. IBr

(table); 2007: 97; Rowe and Gates 1995: 143; Kogo 1998: 2: Ibr1 oblong, in contact laterally; Ibr2 (axil) triangular or 33, fig. 26; 2002: 9; Lane et al. 2000: 476 (table); Kirken- low pentagonal, free laterally. IIBr and IIIBr 4(3+4) or 2. dale and Messing 2003: 530; Kogo and Fujita 2005: 332; Arms 14–c. 50; anterior arms 85–130 mm, posterior arms

Mekhova and Britayev 2012: 915. 35–80 mm long; first syzygy at br3+4. PII longest, 14–26 mm Clarkcomanthus alternans—Summers et al. 2014: 336. long, of 36–48 segments; combs of 6–11 teeth confluent with outer edge of pinnulars; distal pinnulars often with sec-

Material examined. 8 specimens. Ambon Is. NSMT ondary tooth. Pinnule combs to P2. E-8110 (1 specimen), Pombo I., lagoon, November 27, 1992; Distribution. Western, northern, and eastern Australia, NSMT E-8114 (1), Pombo I., 10 m, November 28, 1992. Indonesia, Melanesia (Rowe et al. 1986); Sulu Sea (Messing Lombok Is. NSMT E-8197 (1), Gili Air I., February 3, 1994; 1998b, 2007); Great Barrier Reef (Fabricius 1994); Papua NSMT E-8242 (1), Gili Air I., February 5, 1994; NSMT New Guinea (Messing 1994); Micronesia (Messing 1998b, E-8255 (2), E-8256 (1), E-8260 (1), Kodek, February 15, 2007; Kirkendale and Messing 2003; Japan (Kogo 1998; 1994. Pilcher and Messing 2001; Kogo and Fujita 2005). Depth Description. Centrodorsal thin, small, pentagonal or range: 1–144 m (Rowe et al. 1986). stellate in aboral view, 2.5–3.5 mm across, up to 0.4 mm thick, lacking cirrus sockets and cirri. Radials trapezoidal, Clarkcomanthus mirabilis in contact laterally. IBr 2: Ibr1 oblong, in contact laterally; (Rowe, Hoggett, Birtles, and Vail, 1986) Ibr2 (axil) low pentagonal, free laterally. IIBr 4(3+4). IIIBr 2, rarely 4(3+4); outer brachials twisted outward. IVBr and Comanthus mirabilis Rowe et al., 1986: 226, fig. 6B, C. VBr 4(3+4). Arms 48–122, 60–100 mm long; first syzygy at Clarkcomanthus mirabilis—Summers et al. 2014: 336. br3+4. PII longest, 9–16 mm long, of 32–44 segments; combs of 6–10 teeth confluent with outer edge of pinnulars; 2–3 Material examined. 1 specimen. Lombok Is. NSMT distal pinnulars with small secondary tooth. Pinnule combs E-8232 (1 specimen), Gili Air I., February 5, 1994. to P4, rarely to P5–P7. Description. Centrodorsal small, stellate in aboral view, Distribution. Southern Vietnam (Mekhova and Britayev 3 mm across, reduced to level with radial-circlet, lacking 2012); western, northern and eastern Australia, Indone- cirrus sockets and cirri. Radials trapezoidal, 2 mm long, in sia, New Guinea, New Caledonia, Philippines (Rowe et al. contact laterally. IBr 2: Ibr1 trapezoidal, in contact laterally; 1986); Sulu Sea, Papua New Guinea, Micronesia (Messing Ibr2 (axil) pentagonal, free laterally. IIBr 4(3+4). IIIBr 2, 1994, 1998b, 2007; Kirkendale and Messing 2003); South rarely 4(3+4). Arms 39; anterior arms 110 mm, posterior

China Sea (Lane et al. 2000); Japan (Rowe et al. 1986; Kogo arms 85 mm long; first syzygy at br3+4. PII longest, 18 mm 1998, 2002; Kogo and Fujita 2005). Depth range: 0–90 m long, of 49 segments; combs of 12–14 teeth confluent with (Lane et al. 2000). outer edge of pinnulars, some pinnulars with secondary

tooth; most proximal tooth transverse. Pinnule combs to P2, Clarkcomanthus littoralis (Carpenter, 1888) rarely to P3–P9. Disc covered with many nodules. Distribution. Philippines, Papua New Guinea (Messing Actinometra littoralis Carpenter, 1888: 346, pl. 67 figs 1, 2. 1994); northern and eastern Australia, Indonesia, New Cale- Clarkcomanthus littoralis—Rowe et al. 1986: 236, fig. 7c–f; donia. Depth range: 3–18 m (Rowe et al. 1986). Fabricius 1994: 1228 (table); Messing 1994: 239 (list); 1998b: 189 (table); 2007: 97; Rowe and Gates 1995: 141; 236 Ichizo Kogo et al.

Clarkcomanthus exilis excessively longer than succeeding pinnules. Pinnule combs

(Rowe, Hoggett, Birtles, and Vail, 1986) to P2. Distribution. First record in Indonesia; previously Oxycomanthus exilis Rowe et al., 1986: 251, fig. 9a; Vail known from eastern Australia, New Caledonia (Rowe et al. 1987: 554 (table), 557; Fabricius 1994: 1228 (table); Mess- 1986; Fabricius 1994); Philippines (Messing 1998b); Micro- ing 1994: 239 (list); 1998b: 189 (table); 2007: 98; Rowe nesia (Kirkendale and Messing 2003). Depth range: 3–10 m and Gates 1995: 149; Kirkendale and Messing 2003: 533; (Rowe et al. 1986). Kogo 2002: 11; Kogo and Fujita 2005: 335. ?Oxycomanthus exilis—Kogo 1998: 49, fig. 38. Clarkcomanthus sp. 1 Clarkcomanthus exilis—Summers et al. 2014: 336. Material examined. 1 specimen. Ambon Is. NSMT Material examined. 1 specimen. Ambon Is. NSMT E-8116 (1 specimen), Pombo I., 10 m, November 28, 1992. E-8128 (1 specimen), Pombo I., 10 m, November 28, 1992. Description. Centrodorsal thin discoidal, pentagonal Description. Centrodorsal small, pentagonal in aboral in aboral view, 3.2 mm across, 0.7 mm thick, with 16 cirrus view, 2 mm across, 0.2 mm thick, with about 10 minute cir- sockets in 1 aligned row, only 3 rudimentary cirri retained. rus sockets in 1 aligned row. Cirri V including 3 rudimen- Radials trapezoidal, in contact laterally. IBr 2: Ibr1 oblong, in tary ones, 12, up to 6 mm long. Cirrals smooth without ab- contact laterally at base; Ibr2 (axil) low pentagonal, free lat- oral spine. Radials trapezoidal, in contact laterally. Rays in a erally. IIBr 4(3+4), rarely 2. IIIBr 2, rarely 4(3+4). IVBr 2. plane. IBr 2: Ibr1 trapezoidal, in contact laterally; Ibr2 (axil) Arms 31 or 32, 30–35 mm long; first syzygy at br3+4; middle low pentagonal, free laterally. IIBr and IIIBr all 4(3+4). brachials beyond br4 with everted and spiny distal edge. PII Arms presumably 34, 64 mm long, first syzygy at br3+4. PII longest, 9 mm long, of 41 segments; combs of 8 teeth non- longest, 14 mm long, of 37 segments; combs of 8–9 non- confluent, with pointed tip; distal 2 or 3 pinnulars with sec- confluent teeth. Pinnule combs to P1. ondary tooth; first comb tooth transverse. Pinnule combs to Distribution. First record in Indonesia; previously P3, rarely to P4. known from western, northern, and eastern Australia, Mela- Remarks. This specimen closely resembles to the speci- nesia (Rowe et al. 1986; Rowe and Gates 1995); Great Bar- mens of Clarkcomanthus mirus (NSMT E-8188 and 8247). rier Reef (Vail 1987); Papua New Guinea (Messing 1994); Though the components of the division series and the distri- Micronesia (Kirkendale and Messing 2003; Messing 2007); bution of pinnule combs are different: IIIBr series is usually

Japan (Kogo 2002; Kogo and Fujita 2005). Depth range: 4(3+4) and combs are limited to P2 in C. mirus, while IIIBr 3–25 m (Rowe et al. 1986). mainly of 2 and combs extend to P4 in this specimen. From these points, this specimen is regarded as a young individual Clarkcomanthus mirus of some species of Clarkcomanthus, other than C. mirus. (Rowe, Hoggett, Birtles, and Vail, 1986) Clarkcomanthus sp. 2 Oxycomanthus mirus Rowe et al., 1986: 255, figs 3c, 9c–d; Fabricius 1994: 1228 (table); Rowe and Gates 1995: 150; Material examined. 1 specimen. Ambon Is. NSMT Messing 1998b: 189 (table). E-8140 (1 specimen), Tial, 10 m, November 30, 1992. Oxycomanthus cf. mirus—Kirkendale and Messing 2003: Description. Centrodorsal discoidal, roundish in ab- 533. oral view, 1 mm across, 0.4 mm thick, with cirrus sockets Clarkcomanthus mirus—Summers et al. 2014: 336. in 1 aligned row. Cirri VI, 9, 5 mm long. Cirrals beyond 1st longer than wide, smooth, no aboral spine; opposing spine Material examined. 2 specimens. Ambon Is. NSMT prominent. Radials trapezoidal, in contact laterally. IBr

E-8188 (1 specimen), Lilibooi, 15–20 m, December 9, 1992. 2: Ibr1 oblong; Ibr2 (axil) pentagonal, free laterally; IIBr Lombok Is. NSMT E-8247 (1), Gili Trawangan I., February 4(3+4). Arms 11, 38 mm long; first syzygy at br3+4; brachi- 13, 1994. als of middle arm with spiny distal margin. P1 9 mm long, Description. Centrodorsal thin discoidal, roundish or of 26–28 segments; combs of 4–7 teeth non-confluent, with stellate in aboral view, 2.5–3 mm across, up to 0.5 mm thick, pointed tip; no secondary tooth. P2 and P3 absent. P4 3.5 mm with cirrus sockets in 1 aligned row. Cirri I–XII, 11–12, long, of 11 segments; combs of 0–4 teeth. P5 4 mm long, of 6 mm long. Longest cirrals (4th–7th) 2 times longer than 12 segments, no comb. Middle pinnules slender, not swol- wide; all cirrals smooth without aboral spine. Radials trap- len without recognizable gonad; pinnulars with spiny distal ezoidal, in contact laterally. IBr 2: Ibr1 oblong, in contact margin. laterally; Ibr2 (axil) low pentagonal, free laterally. IIBr and Remarks. This specimen is supposedly a young indi- IIIBr all 4(3+4). Arms 27–32; anterior arms 95–130 mm, vidual of a species of Clarkcomanthus, for having a small posterior arms 65–80 mm long; first syzygy at br3+4. Middle centrodorsal, non-confluent teeth of combs, and distalmost brachials beyond br5 wedge-shaped with everted distal mar- pinnule comb restricted to on proximal pinnules. The speci- gin. PII longest, 11–15 mm long, of 40 segments; combs of men resembles to C. exilis and C. mirus for holding non- 10–12 discrete teeth non-confluent, with pointed tip; 3–4 confluent teeth. Though the distalmost comb is on1 P or P2 pinnulars often with small secondary tooth. 1P similar to PII, in these two species, while on P4 in this specimen. It also Comatulids from Ambon and Lombok islands 237 resembles to Comanthus sp. (NSMT E-8195) in absence of E-8235 (1), E-8239 (1), E-8240 (1), E-8245 (1), Gili Air I.,

P2 and P3, but this specimen has non-confluent teeth, while February 3, 1994. Comanthus sp. has confluent teeth. Description. Centrodorsal small, thin discoidal, round, pentagonal or stellate in aboral view, 2.5–3.5 mm across, up Comanthus parvicirrus (Müller, 1841) to 0.5 mm thick, with cirrus sockets in 1 aligned row. Cirri short and weak, VI–X, 8–15, 7–12 mm long. Longest cirrals Alecto parvicirra Müller, 1841: 145. (5th–7th) 1.3 times longer than wide; 2–3 distal cirrals with a Actinometra parvicirra—Carpenter 1881: 204; 1882: 519; low transverse ridge. Radials trapezoidal, in contact laterally.

1888: 338, pl. 61, pl. 67 figs 3, 4; Hartlaub 1891: 96. IBr 2: Ibr1 oblong, in contact laterally; Ibr2 (axil) triangular, Comanthus (Comanthus) parvicirra—A. H. Clark 1911a: free or in contact laterally. IIBr and IIIBr 4(3+4), rarely 2. 536; Utinomi and Kogo 1965: 265. IVBr all 4(3+4). Arms 19–42; anterior arms 70–180 mm,

Comanthus (Validia) parvicirra—A. H. Clark 1911b: 18. posterior arms 40–80 mm long; first syzygy at br3+4. PII lon- Comanthus (Vania) parvicirra—A. H. Clark 1918: 54; Gislén gest, 10–13 mm long, of 34–48 segments; combs of 6–12 1922: 50, figs 41–43. discrete teeth confluent with outer edge of pinnulars; first Comanthus parvicirum—H. L. Clark 1915: 103. comb tooth transverse. Combed pinnule tips usually coiled. Comanthus parvicirra—A. H. Clark 1911b: 8 (list); 1912a: Pinnule combs to middle, often to distal pinnules. Middle 97; 1912b: 10; 1913: 17; 1929: 637; 1931: 631, pl. 29 fig. pinnulars of distal pinnule with smooth distal edge. 88, pl. 65 fig. 184, pl. 73 fig. 200, pl. 78 figs 209, 210, pl. 79 Distribution. South Africa (A. M. Clark 1972); western figs 211–214, pl. 80 figs 215–218, pl. 81 fig. 221; 1936: 299; Indian Ocean, Bay of Bengal, Indonesia, Philippines, China 1937: 99; Reichensperger 1913: 88; H. L. Clark 1915: 101, (A. M. Clark and Rowe 1971); southern Vietnam (Mekhova 103; 1921: 19; 1946: 39; Gislén 1927: 18; 1940: 8. and Britayev 2012); Sumatra (MarshallCrossland and Price Comanthus parvicirrus—A. M. Clark and Rowe 1971: 16 1999); Singapore (Messing and Tay 2016); Hong Kong (A. (key); A. M. Clark 1972: 77; 1982: 486 (table); Gibbs et M. Clark 1982); northern Australia (A. M. Clark and Rowe al. 1976: 109; Meyer and Macurda 1980: 80; Marshall 1971; Rowe et al. 1986; Rowe and Gates 1995); Great Barrier and Rowe 1981: 383; Liao 1983: 264; Zmarzly 1984: 108, Reef (Gibbs et al. 1976; Vail 1987; Fabricius 1994; Rowe and fig. 4; Rowe et al. 1986: 211, fig. 5a, b; Vail 1987: 553, Gates 1995); eastern Australia (Rowe and Gates 1995); Coral 554 (table); Chen et al. 1988: 76, fig. 10; Fabricius 1994: Sea (Rowe et al. 1986); Papua New Guinea (Messing 1994); 1228 (table); Messing 1994: 239 (list); 1998b: 189 (table); Micronesia (Meyer and Macurda 1980; Zmarzly 1984; Mess- 2007: 97; Liao and A. M. Clark 1995: 21, fig. 10, pl. 2 ing 1998b, 2007; Kirkendale and Messing 2003); South Pa- fig. 1; Rowe and Gates 1995: 144; Kogo 1998: 33, fig. 26; cific Ocean (A. M. Clark and Rowe 1971); South China Sea 2002: 10; MarshallCrossland and Price 1999: 26; Lane et (Lane et al. 2000); Japan (A. M. Clark and Rowe 1971; Kogo al. 2000: 476 (table); Pilcher and Messing 2001: 16 (table); 1998, 2002; Pilcher and Messing 2001; Kogo and Fujita Kirkendale and Messing 2003: 531; Kogo and Fujita 2005: 2005). Depth range: 1–110 m (Lane et al. 2000). 331; Mekhova and Britayev 2012: 916; Summers et al. 2014: 337; Messing and Tay 2016: 638. Comanthus wahlbergii (Müller, 1843) Alecto timorensis Müller, 1841: 145. Comatula timorensis—Müller 1849: 263. Alecto wahlbergii Müller, 1843: 131. Comanthus timorensis—A. H. Clark 1931: 603, pl. 64 fig. Comatula (Actinometra) wahlbergii—Müller 1849: 256. 181, pl. 75 fig. 204, pl. 76 fig. 205, pl. 77 figs 206, 207; Comanthus (Bennettia) wahlbergii—A. H. Clark 1911b: 17. 1936: 299; Gislén 1940: 7; H. L. Clark 1946: 38. Comanthus wahlbergii—A. H. Clark 1912a: 95; 1913: 16; Actinometra annulata Bell, 1882: 535, pl. 35. 1918: 49 (key); 1931: 588, pl. 65 fig. 183; 1937: 99; Gislén Comanthus (Comanthus) annulata—A. H. Clark 1911a: 530. 1938a: 8, fig. 7; Rowe et al. 1986: 228, figs 2a, 6d–h; Fa- Comanthus annulata—A. H. Clark 1912a: 98; 1912b: 9; bricius 1994: 1228 (table); Messing 1994: 239 (list); 1998b: 1913: 17; H. L. Clark 1921: 16. 189 (table); Rowe and Gates 1995: 145; Kogo 1998: 37, Comanthus annulatum—H. L. Clark 1915: 101. fig. 29; MarshallCrossland and Price 1999: 26; Lane et Comanthus (Vania) annulata—A. H. Clark 1918: 53; Gislén al. 2000: 476 (table); Kirkendale and Messing 2003: 531; 1922: 49, figs 37–40. Kogo and Fujita 2005: 332; Summers et al. 2014: 337. Comanthus intricata A. H. Clark, 1908b: 220. Comanthus wahlbergi—Vail 1987: 555. Comanthus (Comanthus) samoana A. H. Clark, 1909f: 30. Material examined. 16 specimens. Ambon Is. NSMT Comanthus samoana—A. H. Clark 1931: 593 (part); 1936: E-8104 (1 specimen, juvenile), Pombo I., November 26, 298 (part). 1992, ; NSMT E-8125 (1, 10 arms), E-8129 (1), Pombo I., Comanthus samoanus—Gibbs et al. 1976: 110. 10 m, November 28, 1992; NSMT E-8149 (1), Latuhalat, 15 m, December 1, 1992; NSMT E-8154 (1), E-8168 (1), Si- Material examined. 6 specimens. Ambon Is. NSMT lale, 15 m, December 2, 1992; NSMT E-8178 (1), Eri, 10 m, E-8106 (1 specimen), Pombo I., lagoon, November 27, December 4, 1992; NSMT E-8183 (1), Leahari, December 1992; NSMT E-8124 (1), Pombo I., 10 m, November 28, 8, 1992; NSMT E-8191 (1), Lilibooi, 15 m, December 9, 1992; NSMT E-8145 (1), Latuhalat, 10 m, December 1, 1992; 1992. Lombok Is. NSMT E-8193 (1), E-8211 (1), E-8221 (1), NSMT E-8160 (1), Silale, 15 m, December 2, 1992; NSMT 238 Ichizo Kogo et al.

E-8189 (1), Lilibooi, 15–20 m, December 9, 1992. Lombok New Guinea (Messing 1994, 1998b); Great Barrier Reef (Vail Is. NSMT E-8215 (1), Gili Air I., February 5, 1994. 1987; Fabricius 1994); Japan (Rowe et al. 1986; Kogo 2002; Description. Centrodorsal discoidal, circular in aboral Kogo and Fujita 2005). Depth range: 0–32 m (Rowe et al. view, 2.5–4 mm across, 0.3–1 mm thick, with flat aboral pole 1986). and cirrus sockets in 1 aligned, partly 2 rows. Cirri XV– XXIII, 14–16, 7–11 mm long. Longest cirrals (usually 4th– Comanthus suavia Rowe, Hoggett, Birtles, and Vail, 1986 5th) 1.3 times longer than wide; 4–7 distal cirrals with aboral spine or transverse ridge. Radials almost wholly concealed Comanthus suavia Rowe et al., 1986: 222, fig. 5e–g; Messing by centrodorsal. Rays in a plane. IBr 2: Ibr1 oblong, in con- 1994: 239 (list); 1998b: 189 (table); 2007: 97; Rowe and tact laterally, Ibr2 (axil) low pentagonal, free laterally. IIBr Gates 1995: 145; MarshallCrossland and Price 1999: 16; 4(3+4), rarely 2. IIIBr 4(3+4). Arms 20–42; anterior arms Obuchi 2013: 16, fig. 1, pl. 10a; Summers et al. 2014: 337. 60–105 mm, posterior arms 35–85 mm long; first syzygy at br3+4. PII (sometimes absent) longest, 10–12 mm long, of Material examined. 1 specimen. Ambon Is. NSMT 42–48 segments; combs of 7–10 teeth confluent with outer E-8226 (1 specimen), Gili Air I., February 5, 1994. edge of pinnulars; first comb tooth transverse; 3–4 distal Description. Centrodorsal thin discoidal, pentagonal in pinnulars with transverse ridge or paired teeth. Pinnule aboral view, 3 mm across, 0.3 mm thick, with cirrus sock- combs to P5, often to middle pinnules. ets in 1 aligned row. Cirri XI including 4 rudimentary ones, Distribution. Maldives, Sri Lanka, Sumatra (Mar- 13–15, 8–9 mm long. Longest cirrals (4th–7th) 1.3 times lon- shallCrossland and Price 1999); western, northern, and ger than wide; cirrals beyond 5th with transverse ridge. Ra- eastern Australia, Tasman Sea, New Zealand, Coral Sea, dials trapezoidal, in contact laterally. IBr 2: Ibr1 trapezoidal, Melanesia (Rowe et al. 1986; Rowe and Gates 1995); Great in contact laterally; Ibr2 (axil) low pentagonal, free laterally. Barrier Reef (Vail 1987; Fabricius 1994); Papua New Guinea IIBr 4(3+4) or 2. IIIBr all 4(3+4). Arms 39; anterior arms (Messing 1994, 1998b); Micronesia (Kirkendale and Mess- 144 mm long, posterior arms much shorter; first syzygy at ing 2003); South China Sea (Lane et al. 2000); Japan (Kogo br3+4. PII longest, 16 mm long, of about 48 segments; combs 1998; Kogo and Fujita 2005). Depth range: 1–103 m (Lane of 7–10 teeth confluent with outer edge of pinnulars; first et al. 2000). comb tooth transverse. Pinnule combs to P2, rarely to P3– P10. Middle pinnulars of distal pinnule with smooth distal Comanthus gisleni Rowe, Hoggett, Birtles, and Vail, 1986 margin. Distribution. Sri Lanka, Sumatra (MarshallCrossland Comanthus gisleni Rowe et al., 1986: 219, figs 4b, 5d; Vail and Price 1999); western, northern, and eastern Austra- 1987: 555; Fabricius 1994: 1228 (table); Messing 1994: 239 lia (Rowe et al. 1986; Rowe and Gates 1995); Papua New (list), 1998b: 189 (table); Rowe and Gates 1995: 144; Kogo Guinea, Micronesia (Rowe et al. 1986; Messing 1994, 1998b, 1998: 35, fig. 28; 2002: 10; MarshallCrossland and Price 2007); Japan (Obuchi 2013). Depth range: 3–20 m (Rowe et 1999: 26; Lane et al. 2000: 476 (table); Kogo and Fujita al. 1986). 2005: 332; Mekhova and Britayev 2012: 915, fig. 6; Sum- mers et al. 2014: 337. Comanthus sp.

Material examined. 1 specimen. Ambon Is. NSMT Material examined. 1 specimen. Lombok Is. NSMT E-8179 (1 specimen), Eri, 10 m, December 4, 1992. E-8195 (1 specimen), Gili Air I., February 3, 1994. Description. Centrodorsal thin discoidal, roundish in ab- Description. Centrodorsal thin discoidal, circular in ab- oral view, 3 mm across, 0.3 mm thick, with cirrus sockets in oral view, 0.8 mm across, with 10 cirrus sockets in 1 aligned 1 aligned row. Cirri XIV, including 2 rudimentary ones, 13– row. Cirri IV retained, 8, 4 mm long, with knobby articula- 14, 7–8 mm long. Longest cirrals (5th–6th) 1.3 times longer tion. Longest cirrals (2nd–3rd) 2.5 times longer than wide; than wide; cirrals beyond 7th shorter than wide with blunt middle and distal cirrals slightly longer than wide, without aboral crest. Radials trapezoidal, in contact laterally. IBr 2: aboral spine; opposing spine prominent. Rays in a plane. Di-

Ibr1 trapezoidal, in contact laterally at base; Ibr2 (axil) low vision series widely separated each other. Radials oblong, in triangular, free laterally. IIBr 2, rarely 4(3+4). IIIBr 4(3+4), contact laterally at base. IBr 2; Ibr1 rectangle, slightly wider rarely 2. Arms 28, 65–85 mm long; first syzygy at br3+4. PII than long, Ibr2 (axil) pentagonal. Arms 10, 45 mm long; first longest, 14 mm long, of about 40 segments; combs of about syzygy at br3+4. P1 longest, 5 mm long, of 21 segments, with 6 teeth confluent with outer edge of pinnulars; first comb 8 teeth confluent teeth with outer edge of pinnulars; first tooth transverse. Pinnule combs to middle, often to distal comb tooth transverse; combed pinnulars with transverse pinnules. Middle pinnulars of distal pinnule fringed with bar; some pinnulars longer than wide. P2 and P3 absent; P4 small distal spines. 4 mm long, of 10 segments, with 0–4 confluent teeth. 5P Distribution. Sri Lanka, Sumatra (MarshallCrossland 4 mm long, of 10 segments, no comb. Pinnules slender; no and Price 1999); southern Vietnam (Mekhova and Britayev recognizable gonad; middle and distal pinnulars spiny at

2012); Thailand, western, northern, and eastern Australia, distal margin. Pa and Pd present; Pb and Pc absent. Coral Sea, Melanesia (Rowe et al. 1986); Timor Sea (Rowe Remarks. This specimen belongs to the genusComanthus and Gates 1995); Borneo (Lane et al. 2000); Sulu Sea, Papua for having a small centrodorsal, exposed radials and a few Comatulids from Ambon and Lombok islands 239 cirri, though could not be identified to a particular species. A. M. Clark 1995); Japan (A. M. Clark and Rowe 1971; Kogo Some proximal pinnules are not developed probably be- and Fujita 2005). Depth range: 0–50 m (Lane et al. 2000). cause this specimen is a juvenile. Family Himerometridae A. H. Clark, 1908b Anneissia bennetti (Müller, 1841) Himerometra robustipinna (Carpenter, 1881)

Alecto bennetti Müller, 1841: 146. Actinometra robustipinna Carpenter, 1881: 201; 1888: 330 Comatula (Alecto) bennetti—Müller 1849: 264. (key). Actinometra bennetti—Carpenter 1881: 212; 1882: 526; Himerometra robustipinna—A. H. Clark 1908c: 213; 1909c: 1888: 381 (table); Hartlaub 1891: 95. 7 (list); 1913: 25; 1918: 74 (key); 1941: 193, pl. 16 fig. 60, Comanthus (Comanthus) bennetti—A. H. Clark 1909b: 147. pl. 17 fig. 63, pl. 18 figs 68, 69; H. L. Clark 1946: 42; A. Comanthus (Cenolia) bennetti—A. H. Clark 1918: 50. M. Clark and Rowe 1971: 21 (key); Gibbs et al. 1976: 111; Cenolia bennetti—H. L. Clark 1946: 36. Meyer et al. 1984: 180; Fabricius 1994: 1228 (table); Mess- Comanthus bennetti—A. H. Clark 1912a: 93; 1913: 15; 1931: ing 1994: 239 (list); 1998b: 189 (table); Rowe and Gates 531, pl. 66 figs 186, 187. pl. 67 fig. 188, pl. 68 fig. 189, pl. 1995: 155; Kogo 1998: 59, fig. 47; MarshallCrossland and 82 fig. 225; 1936: 298; 1952: 266; A. M. Clark and Rowe Price 1999: 27; Lane et al. 2000: 476 (table); Pilcher and 1971: 16 (key); Gibbs et al. 1976: 109; Meyer and Mac- Messing 2001: 16 (table); Kogo and Fujita 2005: 337; Me- urda 1980: 78; Zmarzly 1984: 108, fig. 3a, b; Meyer et al. khova and Britayev 2012: 912; Messing and Tay 2016: 644. 1984: 180; Liao and A. M. Clark 1995: 20. Antedon kraepelini Hartlaub, 1890: 183. Oxycomanthus bennetti—Rowe et al. 1986: 259, figs 3b, 9g; Himerometra kraepelini—Reichensperger 1913: 96. Chen et al. 1988: 77, figs 11, 23 A, B; Fabricius 1994: 1228 Antedon crassipinna Hartlaub, 1891: 32, pl. 1 figs 1, 3, 5. (table): Messing 1994: 239 (list); 1998b:189 (table); 2007: Himerometra crassipinna—Reichensperger 1913: 98. 97; Rowe and Gates 1995: 149; Kogo 1998: 42, fig. 33; Antedon inopinata Bell, 1894: 398. MarshallCrossland and Price 1999: 26; Lane et al. 2000: 476 (table); Pilcher and Messing 2001: 16 (table); Kirken- Material examined. 3 specimens. Ambon Is. NSMT dale and Messing 2003: 532; Kogo and Fujita 2005: 333; E-8103 (1 specimen), Pombo I., 10 m, November 25, 1992. Mekhova and Britayev 2012: 918. Lombok Is. NSMT E-8204 (1), Kodek, February 15, 1994; Anneissia bennetti—Summers et al. 2014: 337; Messing and NSMT E-8257 (1), Gili Air I., February 3, 1994. Tay 2016: 639. Description. Centrodorsal large, thick discoidal or low hemispherical, 7–10 mm across, 3–4 mm thick, with cirrus Material examined. 8 specimens. Ambon Is. NSMT sockets in 2–3 crowded rows; aboral pole deeply concave. E-8109 (1 specimen), Pombo I., lagoon, November 27, Cirri long and stout, XXXII–LVIII, 26–35, 25–32 mm long. 1992; NSMT E-8126 (2), E-8130 (1), E-8135 (1), E-8136 (1), Proximal cirrals mostly as long as wide; 7–10 distal cirrals Pombo I., 10 m, November 28, 1992; NSMT E-8162 (1), Si- slightly shorter than wide with blunt aboral tubercle. Ra- lale, 15 m, December 2, 1992. Lombok Is. NSMT E-8224 (1), dials wholly concealed by centrodorsal. IBr 2: Ibr1 oblong, Gili Air I. February 5, 1994. proximal angle in contact laterally; Ibr2 (axil) low pentago- Description. Centrodorsal large, hemispherical or globu- nal, free laterally. IIBr 4(3+4), rarely 2. IIIBr 2 or 4(3+4). lar, 8–14 mm across, 3–4.5 mm thick with cirrus sockets IVBr 4(3+4), usually arising from outer sides of IIIBr. Arms in 2–3 crowded rows. Cirri very stout, XXX–LVII, 25–33, 42–50, 85–125 mm long; first syzygy at br3+4; middle-distal th th 22–40 mm long. Longest cirrals (6 –14 ) as long as wide; brachials about 3 times wider than long. PII longest and succeeding cirrals shorter than wide; distal cirrals with blunt stoutest, 18 mm long, of 23–28 segments; proximal pinnu- aboral tubercle. Radials wholly concealed by centrodorsal. lars much larger than distal ones. PII–PIV similar in shape, IBr 2: Ibr1 excessively short; Ibr2 (axil) low pentagonal, in excessively stouter and longer than succeeding pinnules. close to laterally. IIBr–VBr 4(3+4), rarely 2, free laterally. Distribution. Sri Lanka, Bay of Bengal, Indonesia, north-

Arms 40–80, 80–150 mm long; first syzygy at br3+4. PII lon- ern Australia, Philippines, China, South Pacific Ocean (A. gest, 22–30 mm long, of 56–62 segments; combs of 10–14 M. Clark and Rowe 1971); southern Vietnam (Mekhova teeth non-confluent, with pointed tip. Pinnule combs to P3– and Britayev 2012); Sumatra (MarshallCrossland and Price P4. 1999); Singapore (Messing and Tay 2016); western, north- Distribution. Bay of Bengal, Indonesia, northern Aus- ern, and eastern Australia (Rowe and Gates 1995); Papua tralia, Philippines, China (A. M. Clark and Rowe 1971); New Guinea (Messing 1994, 1998b); South China Sea (Lane southern Vietnam (Mekhova and Britayev 2012); Sumatra et al. 2000); Japan (A. M. Clark and Rowe 1971; Kogo 1998; (MarshallCrossland and Price, 1999); Singapore (Messing Kogo and Fujita 2005). Depth range: 0–57 m (Lane et al. and Tay 2016); western Australia (Rowe and Gates 1995); 2000). Papua New Guinea (Messing 1994, 1998b); Micronesia (A. H. Clark 1931, 1952; Meyer and Macurda 1980; Zmarzly Heterometra amboinae (A. H. Clark, 1912a) 1984; Kirkendale and Messing 2003; Messing 2007); South Pacific Ocean (A. M. Clark and Rowe 1971); South China Craspedometra amboinae A. H. Clark, 1912a: 35 (list); 1918: Sea (Lane et al. 2000); Taiwan (Chen et al. 1988; Liao and 78 (footnote); Reichensperger 1913: 99. 240 Ichizo Kogo et al.

Heterometra amboinae—A. H. Clark 1918: 78 (key); 1941: 1998: 62, fig. 49; 2002: 18; Kogo and Fujita 2005: 339. 297, pl. 29 fig. 123, pl. 31 figs 131–136; A. M. Clark and Rowe 1971: 22 (key). Material examined. 3 specimens. Ambon Is. NSMT E-8112 (1 specimen), E-8113 (1), E-8122 (1), Pombo I., Material examined. 2 specimens. Lombok Is. NSMT 10 m, November 28, 1992. E-8249 (2 specimens), Kodek, February15, 1994. Description. Centrodorsal low hemispherical, 4–6 mm Description (larger specimen). Centrodorsal discoidal, across, 2 mm thick, with cirrus sockets in 2–3 alternating 7 mm across, 3 mm thick, with cirrus sockets in 1–2 ir- rows and concave aboral pole. Cirri XXVII–XXXV, 19–25, regular rows and flat aboral pole. Cirri stout, XXIII, 33–35, 17–27 mm long. Longest cirrals (6th –12th) about 1.2 times 25–30 mm long. Cirrals wholly shorter than wide; beyond longer than wide; cirrals beyond 11th with low aboral crest. th about 11 with prominent aboral spine. Radials wholly con- Radials wholly concealed by centrodorsal. IBr 2: Ibr1 ob- cealed by centrodorsal. IBr 2: Ibr1 oblong, in contact later- long, free laterally; Ibr2 (axil) low pentagonal. IIBr and IIIBr ally; Ibr2 (axil) low triangular, free laterally. IIBr 4(3+4). all 2. Division series with rounded adambulacral lateral pro- Arms stout, 16, 135 mm long; first syzygy at br3+4; br4–br10 cesses. Arms 20–30, 80–140 mm long; first syzygy at br3+4. about 3 times wider than long with parallel articulation, be- P2 and P3 similar, stiff and erect, 14–16 mm long, of 13–16 yond br10 wedge-shaped. P3 longest, stiffened, 20 mm long, segments, much longer and stouter than P1 and P4. of 30 segments; proximal pinnulars shorter than wide with a Distribution. Western Indian Ocean, Red Sea, Maldives, keel toward tip. P2 similar to P3, but slightly shorter. P2 and Sri Lanka, Bay of Bengal, Indonesia, western, northern, and P3 much longer and stouter than PII and P1. eastern Australia, Philippines, China (A. M. Clark and Rowe Remarks. Four Heterometra species, namely H. varii- 1971; Rowe and Gates 1995); southern Vietnam (Mekhova pinna (Carpenter, 1882), H. affinis (Hartlaub, 1890), H. am- and Britayev 2012); Sumatra (MarshallCrossland and Price boinae (A. H. Clark, 1912a), and H. sarae A. H. Clark, 1941, 1999); Singapore (A. M. Clark 1972; Messing and Tay 2016); were recorded from Ambon (A. H. Clark 1941). In these 4 Sulu Sea, Papua New Guinea, Micronesia (Meyer and Ma- species, H. amboinae is the most closely related to H. sarae curda 1980; Messing 1994, 1998b, 2007; Kirkendale and in sharing excessively short middle-distal brachials. Accord- Messing 2003); Great Barrier Reef (Fabricius 1994); South ing to A. H. Clark (1941: 230–232), brachials beyond br4 are Pacific Ocean (A. M. Clark and Rowe 1971); South China 3–4 times wider than long in H. amboinae, while 6–8 times Sea (Lane et al. 2000); Japan (A. M. Clark and Rowe 1971; wider than long in H. sarae. Based on these characteristics, Kogo 1998, 2002; Pilcher and Messing 2001; Kogo and Fu- present specimens were identified with H. amboinae. jita 2005). Depth range: 0–245 m (Lane et al. 2000). Distribution. Sri Lanka, Singapore, Indonesia (A. H. Clark 1941; A. M. Clark and Rowe 1971). Depth range: Liparometra articulata (Müller, 1849) 0–50 m (A. H. Clark 1941). Comatula (Alecto) articulata Müller, 1849: 263. Family Mariametridae A. H. Clark, 1909e Antedon articulata—Carpenter 1888: 226 (key), 379 (table). Stephanometra indica (Smith, 1876) Dichrometra articulata—A. H. Clark 1912a: 152. Liparometra articulata—A. H. Clark 1913: 31; 1918: 103: Comatula indica Smith, 1876: 406. 1941: 461, pl. 53 figs 240–242; H. L. Clark 1946: 46; A. M. Stephanometra indica—A. H. Clark 1909c: 10 (list); 1911b: Clark and Rowe 1971: 24 (key); Meyer et al. 1984: 182; 26; 1912a: 135; 1913: 29; 1918: 97; 1937: 88; 1952: 267; H. Fabricius 1994: 1228 (table); Rowe and Gates 1995: 159; L. Clark 1946: 45; A. M. Clark and Spencer Davis 1966: Lane et al. 2000: 477 (table); Kogo 2002: 19; Kogo and Fu- 598 (list); A. M. Clark and Rowe 1971: 24 (key); A. M. jita 2005: 337. Clark 1972: 107, fig. 10f–h; Meyer and Macurda 1980: ?Liparometra articulata—Kogo 1998: 64, fig. 52. 86; Marshall and Rowe 1981: 394; Fabricius 1994: 1228 (table); Messing 1994: 239 (list); 1998b: 189 (table); 2007: Material examined. 12 specimens. Ambon Is. NSMT 100; Rowe and Gates 1995: 159; Kogo 1998: 63, fig. 50; E-8144 (1 specimen), Latuhalat, 10 m, December 1, 1992; MarshallCrossland and Price 1999: 27; Lane et al. 2000: NSMT E-8173 (1), E-8181 (1), Eri, 10 m, December 4, 1992; 477 (table); Pilcher and Messing 2001: 16 (table); Kirken- NSMT E-8185 (1), E-8186 (2), E-8187 (1), Lilibooi, 15 m, dale and Messing 2003: 534; Kogo and Fujita 2005: 338; December 9, 1992. Lombok Is. NSMT E-8233 (1), E-8241 Rankin and Messing 2008: 11, figs 1–3a–c, 4a–c, 10–14; (1), E-8243 (1), Gili Air I., February 5, 1994; NSMT E-8252 Mekhova and Britayev 2012: 915; Messing and Tay 2016: (2), Kodek, February 15, 1994. 653. Description. Centrodorsal low hemispherical, 4–7 mm Antedon monacantha Hartlaub, 1890: 179. across, 2–3 mm thick, with cirrus sockets in 2–3 alternating Stephanometra monacantha—Reichensperger 1913: 102. rows and small flat aboral pole. Cirri XXVIII–XLII, 21–33, Antedon spicata Carpenter, 1881: 190; 1888: 225 (key), 380; 14–31 mm long. Longest cirrals (7th–13th) 1.2 times longer 1889: 310, pl. 27 figs 3–5. than wide; cirrals beyond 6th–12th with a sharp aboral crest. Stephanometra spicata—A. H. Clark 1909c: 10 (list); 1912a: Radials almost wholly concealed by centrodorsal. Division 132; 1918: 95 (key); 1934: 11; 1937: 88, 100; 1941: 424, pl. series short, with low aboral synarthrial tubercle and thick

49 figs 223, 224, pl. 91 fig. 447; Gislén 1938b: 13; Kogo adambulacral lateral processes. IBr 2: Ibr1 oblong; Ibr2 (axil) Comatulids from Ambon and Lombok islands 241 low pentagonal, in contact laterally. IIBr and IIIBr all 2. Distribution. Western India, Pakistan, Maldives, Bay of

Arms 14–41, 53–139 mm long; first syzygy at br3+4. P2 and Bengal, Indonesia, northern Australia, Philippines, China P3 similar, longest, 10–14 mm long, of 22–28 segments, lon- (A. M. Clark and Rowe 1971); Sri Lanka (A. M. Clark and ger and stouter than P1, P4 and succeeding pinnules. Rowe 1971; MarshallCrossland and Price 1999); southern Distribution. Northern Australia, Indonesia, China (A. Vietnam (Mekhova and Britayev 2012); Hong Kong (A. M. M. Clark and Rowe 1971); Great Barrier Reef (Fabricius Clark 1982; Liao and A. M. Clark 1995); Papua New Guinea 1994; Rowe and Gates 1995); South China Sea (Lane et al. (Messing 1994, 1998b); Micronesia (Messing 1998b; Kirken- 2000); Japan (A. M. Clark and Rowe 1971; Kogo 2002; Kogo dale and Messing 2003); Hawaiian Is., South Pacific Ocean and Fujita 2005). Depth range: 0–69 m (Lane et al. 2000). (A. M. Clark and Rowe 1971); South China Sea (Lane et al. 2000); Japan (A. M. Clark and Rowe 1971; Kogo 1998; Kogo Lamprometra palmata (Müller, 1841) and Fujita 2005). Depth range: 0–50 m (Lane et al. 2000).

Alecto palmata Müller, 1841: 144. Family Colobometridae A. H. Clark, 1909d Comatula (Alecto) palmata—Müller 1849: 261. Cenometra bella (Hartlaub, 1890) Antedon palmata—Carpenter 1888: 226 (key); Hartlaub 1891: 49, pl. 3 fig. 27. Antedon bella Hartlaub, 1890: 174; 1891: 43, pl. 2 figs 23, 26. Himerometra palmata—A. H. Clark 1907a: 356 (list). Cenometra bella—A. H. Clark 1909c: 8 (list); 1912a: 153; Dichrometra palmata—A. H. Clark 1909c: 12 (list); 1911b: 1936: 307; 1947: 27, pl. 3 figs 9, 10; A. M. Clark and Rowe 27; 1912a: 148; Reichensperger 1913 : 103. 1971: 19 (key); Meyer and Macurda 1980: 88; Chen et al. Lamprometra palmata palmata—A. H. Clark 1941: 474, pl. 1988: 78, figs 16, 23E; Fabricius 1994: 1228 (table); Mess- 53 figs 243–246, pl. 54 figs 248–252, pl. 55 fig. 257; A. M. ing 1994: 239 (list); 1998b: 189 (table); 2007: 96; Rowe Clark 1972: 104, fig. 10a–e; Liao and A. M. Clark 1995: and Gates 1995: 134; MarshallCrossland and Price 1999: 41, fig. 21; Kogo 1998: 65, fig. 53. 27; Lane et al. 2000: 477 (table); Pilcher and Messing Lamprometra palmata—A. H. Clark 1913: 33; 1918: 100 2001: 16 (table); Kirkendale and Messing 2003: 535; Kogo (key); 1929: 641; 1934: 11; 1936: 303; 1937: 100; H. L. and Fujita 2005: 341; Mekhova and Britayev 2012: 910; Clark 1915: 101; 1946: 47; Gislén 1938b: 14; 1940: 10; A. Messing and Tay 2016: 641, fig. 7B. M. Clark and Spencer Davis 1966: 598 (list); A. M. Clark Cenometra bella var. magnifica Gislén, 1940: 12, pl. 3 figs 12, and Rowe 1971: 24 (key); Meyer and Macurda 1980: 84; 13. A. M. Clark 1982: 486 (table); Chen et al. 1988: 78, fig. Antedon abbotti A. H. Clark, 1907b: 148. 21; Messing 1994: 239; 1998b: 189 (table); 2007: 100; Himerometra unicornis A. H. Clark, 1908c: 216; 1909a: 399. Rowe and Gates 1995: 158; MarshallCrossland and Price Cenometra delicata A. H. Clark, 1909a: 398. 1999: 27; Lane et al. 2000: 477 (table); Pilcher and Mess- ing 2001: 16 (table); Kogo 2002: 20; 2006: 231; Kirkendale Material examined. 5 specimens. Ambon Is. NSMT and Messing 2003: 535; Kogo and Fujita 2005: 340; Mek- E-8119 (1 specimen), Pombo I., 10 m, November 28, 1992. hova and Britayev 2012: 914; Rankin and Messing 2008: Lombok Is. NSMT E-8210 (1), Gili Air I., February 5, 1994; 25, figs 2e–f, 5, 16–19; Messing and Tay 2016: 650. NSMT E-8248 (1), E-8258 (2). Kodek, February 15, 1994. Antedon moorei Bell, 1894: 401. Description. Centrodorsal thick discoidal or low hemi- Lamprometra moorei—Gislén 1938b: 14, fig. 3. spherical, 3–6 mm across, 1–3 mm thick, with cirrus sock- Antedon conjungens Carpenter, 1889: 311, pl. 27 figs 1, 2. ets in 1–2 irregular rows. Cirri stout, XVI–XXX, 30–38, 15–24 mm long; cirrals wholly wider than long; middle Material examined. 6 specimens. Ambon Is. NSMT cirrals with transverse ridge and several distal cirrals with E-8111 (4 specimens), Pombo I., 10 m, November 28, 1992; paired aboral spines. Radials almost wholly concealed by NSMT E-8139 (1), Tial, 10 m, November 30, 1992. Lombok centrodorsal. Division series with rounded or broad adam-

Is. NSMT E-8200 (1), Gili Air I., February 3, 1994. bulacral lateral processes. IBr 2: Ibr1 square, in contact lat- Description. Centrodorsal thin discoidal to low hemi- erally at base; Ibr2 (axil) pentagonal, free laterally. IIBr and spherical, 2–6 mm across, up to 1.3 mm thick, with cir- IIIBr all 2. Arms 17–30, 75–120 mm long. First syzygy at rus sockets in 2–3 irregular rows and flat aboral pole. Cirri br3+4. P2 longest, abruptly stout and horn-like, 10–13 mm XXII–XXXVI, 20–24, 13–18 mm long. Longest cirrals (3–4 long, of 18–22 segments, much longer than P3 and succeed- in 5th–12th) about 1.2 times longer than wide; cirrals beyond ing pinnules; middle to distal pinnulars with spiny distal th 12 gradually shortened, with a minute aboral spine or tu- margin. P1 slender, intermediate in length between P2 and bercle. Radials almost wholly concealed by centrodorsal,. P3. IBr 2: Ibr1 trapezoidal, proximal angle in contact laterally; Distribution. Bay of Bengal, Indonesia, Philippines, Ibr2 (axil) low pentagonal, free laterally. IIBr and IIIBr all 2. China (A. M. Clark and Rowe 1971); southern Vietnam Division series with low aboral synarthrial tubercle. Arms (Mekhova and Britayev 2012); Sumatra (MarshallCrossland

16–39, 42–72 mm long; first syzygy at br3+4. P2 longest, and Price 1999); western, northern, and eastern Australia 7–11 mm long, of 16–21 segments, much longer and stouter (Rowe and Gates 1995); Great Barrier Reef (Fabricius 1994); than P1 and P3; proximal pinnulars stout, gradually attenu- Sulu Sea, Papua New Guinea (Messing 1998b); Micronesia ated with flagellate tip. (Meyer and Macurda 1980; Messing 1998b; Kirkendale and 242 Ichizo Kogo et al.

Messing 2003); South Pacific Ocean (A. M. Clark and Rowe 1–2 lows and depressed aboral pole. Cirri long, reaching 1/3 1971); South China Sea (Lane et al. 2000); Taiwan (Chen et of arm length, X–XVII, 41–52, 26–33 mm long. Longest cir- al. 1988); Japan (A. M. Clark and Rowe 1971; Pilcher and rals (6th–12th) 1.5–2 times longer than wide; cirrals beyond Messing 2001; Kogo and Fujita 2005). Depth range: 0–55 m 4th with flared and spiny distal margin; about 20 distal cir- (Lane et al. 2000). rals with prominent paired aboral spines. Radials oblong, in

contact laterally. IBr 2: Ibr1 oblong, free laterally; Ibr2 (axil) Petasometra clarae (Hartlaub, 1890) low pentagonal. Division series and first brachials with ab- oral synarthrial tubercle. Arms 10, 70–90 mm long; first syz-

Antedon clarae Hartlaub, 1890: 174; 1891: 41, pl. 2 fig. 19. ygy at br3+4; proximal brachials with lateral expansion and Cyllometra clarae—A. H. Clark 1912a: 158. tuft of spines on distal edge. Oral pinnules mostly composed Petasometra clarae—A. H. Clark 1918: 113; 1947: 109, pl. 13 of long segments (2–3 times longer than wide) with tuft of

figs 62–64; A. M. Clark and Rowe 1971: 20 (key); Fabri- sharp distal spines. P2–P4 longest, 13–15 mm long, of 13–15 cius 1994: 1228 (table); Messing 1994: 239 (list); 1998b: segments. Pa always (also Pb and Pc in juvenile individual) 189 (table); Rowe and Gates 1995: 137. absent. Distribution. Indonesia, northern Australia, Philippines, Material examined. 2 specimens. Lombok Is. NSMT South Pacific Ocean (A. M. Clark and Rowe 1971); south- E-8201 (2 specimens), Gili Air I., February 3, 1994. ern Vietnam (Mekhova and Britayev 2012); Sumatra (Mar- Description. Centrodorsal discoidal, about 3 mm across, shallCrossland and Price 1999); Singapore (Messing and 1 mm thick, with cirrus sockets compactly in 1 aligned row Tay 2016); western, northern, and eastern Australia (Rowe and flat aboral pole. Cirri XV–XVI, 24, 11–13 mm long; cir- and Gates 1995); Sumatra, Borneo (MarshallCrossland and rals wholly shorter than long; beyond 4th with transverse Price, 1999; Lane et al. 2000); Sulu Sea, Micronesia (Messing ridge. Radials narrowly exposed, in contact laterally at base. 1998b); South China Sea (Lane et al. 2000); Taiwan (Chen

IBr 2: Ibr1 oblong, free laterally; Ibr2 (axil) pentagonal. IIBr et al. 1988); Japan (Kogo 1998; Pilcher and Messing 2001; 4(3+4). Division series with aboral synarthrial tubercle and Kogo and Fujita 2005). Depth range: 0–122 m (Lane et al. weak adambulacral processes. Arms 10–12, 35–65 mm long; 2000). first syzygy at br3+4. P2 longest, 7–8 mm long, of 18–21 seg- ments, much longer than succeeding pinnules; middle pin- Oligometra serripinna (Carpenter, 1881) nulars about 1.5 times longer than wide with smooth distal margin. P1 similar to P2, slightly shorter. Pa always, Pb some- Antedon serripinna Carpenter, 1881: 182. times absent. Oligometra serripinna—A. H. Clark 1908a: 126 (list); 1918: Distribution. Indonesia (A. M. Clark and Rowe 1971); 130; A. M. Clark and Rowe 1971: 19 (key); Vail 1987: 555; western, northern, and eastern Australia (Rowe and Gates Liao and A. M. Clark 1995: 47, fig. 26; MarshallCrossland 1995); Papua New Guinea (Messing 1994, 1998b). Depth and Price 1999: 28; Messing and Tay 2016: 642, fig. 7C, D, range: 0–40 m (Rowe and Gates 1995). F. Oligometra serripinna serripinna—A. H. Clark 1947: 217. Colobometra perspinosa (Carpenter, 1881) Material examined. 10 specimens. Ambon Is. NSMT Antedon perspinosa Carpenter, 1881: 178; 1888: 193 (key); E-8120 (1 specimen), E-8123 (8), Pombo I., 10 m, November Hartlaub 1891: 85, pl. 5 fig. 54. 28, 1992; NSMT E-8166 (1), Silale, 10 m, December 3, 1992. Cyllometra perspinosa—A. H. Clark 1907a: 357 (list). Description. Centrodorsal discoidal, 1.5–2 mm across, Colobometra perspinosa—A. H. Clark 1912a: 164; 1913: 1–1.5 mm thick, with cirrus sockets in 1 aligned row and 37; 1918: 125; 1936: 308; 1947: 117, pl. 14 figs 67–69; flat aboral pole. Cirri X–XIV, 15–20, 6–9 mm long. Cirrals Gislén 1940: 14, pl. 3 fig. 14; H. L. Clark 1946: 52; A. M. wholly wider than long; beyond 3rd with prominent trans- Clark and Rowe 1971: 19 (key); Chen et al. 1988: 78, verse ridge; 3–4 most distal cirrals rarely with paired aboral figs 18, 23F; Messing 1994: 239 (list); 1998b: 189 (table); spines. Radials narrowly exposed, in contact laterally. IBr 2:

Rowe and Gates 1995: 134; Kogo 1998: 76, fig. 61; Mar- Ibr1 oblong, free laterally; Ibr2 (axil) pentagonal. Arms 10, shallCrossland and Price 1999: 27; Lane et al. 2000: 477 35–60 mm long; first syzygy at br3+4. P2 longest and stout- (table); Pilcher and Messing 2001: 16 (table); Kogo and est, 5–6 mm long, of 12–15 segments, much longer than Fujita 2005: 342; Mekhova and Britayev 2012: 910; Mess- succeeding pinnules; middle pinnulars about 2 times lon-

ing and Tay 2016: 639. ger than wide with spiny distal margin. P1 intermediate in length between P2 and P3. Pa usually absent. Material examined. 4 specimens. Ambon Is. NSMT Distribution. Western Indian Ocean, Red Sea, Maldives, E-8105 (1 specimen), Pombo I., lagoon, November 22, 1992; Sri Lanka, Bay of Bengal, Philippines, Indonesia, Japan (A. NSMT E-8121 (1), Pombo I., 10 m, November 28, 1992; M. Clark and Rowe 1971); Sumatra (MarshallCrossland and NSMT E-8152 (1), Silale, 15 m, December 2, 1992. Lombok Price 1999); Singapore (Messing and Tay 2016); Great Bar- Is. NSMT E-8202 (1, juvenile), Gili Air I., February 3, 1994. rier Reef (Vail 1987); South China Sea (Lane et al. 2000); Description. Centrodorsal hemispherical, 4–5 mm Taiwan (Liao and A. M. Clark 1995). Depth range: 0–90 m across, 2 mm thick, with cirrus sockets sparsely arranged in (Lane et al. 2000). Comatulids from Ambon and Lombok islands 243

Family Antedonidae Norman, 1865 times longer than wide with smooth distal margin. P2 inter- Dorometra nana (Hartlaub, 1890) mediate in length between P1 and P3. Distribution. Indonesia (A. M. Clark and Rowe 1971); Antedon nana Hartlaub, 1890: 170; 1891: 89, pl. 5 figs 57, 58; Great Barrier Reef (Fabricius 1994). Depth range: 2–15 m A. H. Clark 1907a: 353 (list). (A. H. Clark and A. M. Clark 1967). Iridometra nana—A. H. Clark 1908a: 131 (list); 1912a: 231; 1912b: 9; 1913: 54; 1937: 102 (list); H. L. Clark 1915: 106. Dorometra nana—A. H. Clark 1918: 216; H. L. Clark 1921: Discussion 25; Gislén 1922: 133, figs 123, 124; 1940: 15, pl. 2 figs 9–11; A. H. Clark and A. M. Clark 1967: 71, fig. 3d; A. Although our surveys are very limited to show the com- M. Clark and Rowe 1971: 24 (key); A. M. Clark 1972: 142; plete comatulid fauna of Ambon and Lombok islands, but Meyer and Macurda 1980: 91; Zmarzly 1984: 108; Mess- the fauna may be characterized by widely-distributed spe- ing 1994: 239 (list); 1998b: 189 (table); Fabricius 1994: cies. Of 31 identified species reported in this study, 16 spe- 1228 (table); Rowe and Gates 1995: 122; Kirkendale and cies (52%) were collected at both islands. The most fre- Messing 2003: 537; Kogo and Fujita 2005: 350. quently occurring species in this study were Comanthus par- Dorometra cf. nana—Messing and Tay 2016: 655. vicirrus (collected at 7 sites), Capillaster multiradiatus (at 6 sites), Comanthus wahlbergii and Liparometra articulata (at Material examined. 1 specimen. Lombok Is. NSMT 5 sites). Succeeding common species were Comatella nigra, E-8203 (1 specimen), Gili Air I., February 3, 1994. Alloeocomatella pectiniferra, Phanogenia multibrachiata, P. Description. Centrodorsal hemispherical, 2 mm across, gracils, Comaster nobilis, Clarkcomanthus alternans, C. lit- 1.3 mm thick, with cirrus sockets in 2–3 alternating rows toralis, Anneissia bennetti, Himerometra robustipinna, Lam- and papillose aboral pole. Cirri all lost, presumably XL–L. prometra palmata, Cenometra bella, and Colobometra per- Radials wholly concealed by centrodorsal. IBr 2 with aboral spinosa (collected at 3–4 sites). Among these 16 species, 11 synarthrial tubercle: Ibr1 oblong, free laterally; Ibr2 (axil) species were occupied by Comatulidae. rhombic. Arms 10, tips broken, presumably 30–40 mm long; The 31 species collected in this study are also occurring in first syzygy at br3+4. Proximal pinnules slender. P3 longest, the neighboring seas. Out of 40 species collected at Madang, 6 mm long, of 17 segments, about 1.5 times longer than Papua New Guinea (Messing 1994), 25 species (63%) are

P1; middle pinnulars nearly 3 times longer than wide with also found in this study. Similarly, 28 of 46 species (61%) at everted distal margin. P2 resembles to P1, slightly shorter. northern Papua New Guinea, 27 of 40 species (68%) in Sulu P3–P5 with gonad. Sea, and 15 of 19 species (79%) at Chuuk Atoll, Caroline Distribution. Bay of Bengal, Indonesia, Philippines, Islands (Messing 1998b), 20 of 47 species (43%) at western northern Australia, China (A. M. Clark and Rowe 1971); Philippines and 8 of 20 species (40%) at northern Borneo Maldives (A. M. Clark 1972); Singapore (Messing and Tay (Lane et al. 2000, ‘zone 3’ and ‘zone 4’, respectively), 20 of 2016); western, and eastern Australia (Rowe and Gates 32 species (63%) in the Bay of Nhatrang, southern Vietnam 1995); Papua New Guinea (Messing 1994, 1998b); Micro- (Mekhova and Britayev 2012), and 13 of 39 species (33%) at nesia (Meyer and Macurda 1980; Zmarzly 1984; Kirkend- Singapore (Messing and Tay 2016), are found in this study. ale and Messing 2003); South China Sea (Lane et al. 2000); Furthermore, 25 of 31 species collected in this study extend Japan (A. M. Clark and Rowe 1971; Kogo and Fujita 2005). their distribution northwardly to subtropical southwestern Depth range: 0–60 m (Lane et al. 2000). Japan (Kogo and Fujita 2005). The majority of the species from Ambon and Lombok islands are widely distributed to Euantedon polytes A. H. Clark, 1936 the western Pacific Ocean. In this study, the family Comatulidae occupy 67.6% (23 Euantedon polytes A. H. Clark, 1936: 315, pl. 7 fig. 5, pl. 8 species) of the total number of species (34 species) collected fig. 6; A. H. Clark and A. M. Clark 1967: 106; A. M. Clark at Ambon and Lombok islands, which supports that this and Rowe 1971: 26 (key); Fabricius 1994: 1228 (table). family is the most diverse in the shallow-water comatulid fauna in the tropical west Pacific. In the neighboring tropi- Material examined. 5 specimens. Ambon Is. NSMT E- cal seas mentioned above, Comatulidae is the most domi- 8107 (5 specimens), Pombo I., lagoon, November 27, 1992. nant at the average ratio of 55.8% (40.9 to 72.2%) in 7 out Description. Centrodorsal low hemispherical, 3 mm of 8 areas (Messing 1994, 1998b; Lane et al. 2000; Mekhova across, 1 mm thick with cirrus sockets in 2–3 alternat- and Britayev 2012). In Singapore waters, on the contrary, ing rows and flat and smooth aboral pole. Cirri about XL, Comatulidae is the second family occupied 28.2% (11 spe- 13–15, 8–12 mm long. Longest cirrals (5th–7th) about 2 times cies) follows with Himerometridae 35.9% (14 species) longer than wide; distal cirrals shorter than wide, no aboral (Messing and Tay 2016). The ratio of Comatulidae in this spine. Radials narrowly exposed, in contact laterally. IBr 2, study is relatively higher than those reported from these with aboral synarthrial tubercle: Ibr1 oblong, free laterally; neighboring seas in the tropical western Pacific. This may Ibr2 (axil) rhombic. Arms 10, about 80 mm long; first syzy- be due to the difference in surveyed depths. The depths are gy at br3+4. P1 longest, 10–14 mm long, of 22–26 segments, 0–15 m in this study, while from 0 to 30–50 m in the neigh- nearly 3 times longer than P3–P5; middle pinnulars about 2 boring seas. 244 Ichizo Kogo et al.

pine Islands. Proceedings of the United States National Museum Acknowledgments 36: 391–410. Clark, A. H. 1909b. On a collection of the crinoids from Copenhagen Museum. Videnskabelige Meddelelser fra den Naturhistoriske We are deeply grateful to Kashim Moosa, Woro W. Kas- Forening i Kjøbenhavn 1909: 115–194. toro, Mudjiono, and Aznam Aziz (Indonesian Institute of Clark, A. H. 1909c. A revision of the crinoid families Thalassometridae Sciences, Research Center for Oceanography), for their as- and Himerometridae. Proceedings of the Biological Society of sistance in sampling. We also thank to Charles G. Messing Washington 22: 1–22. (Nova Southeastern University) and Marc Eléaume (Mu- Clark, A. H. 1909d. New recent Indian crinoids. Proceedings of the Bio- séum National d’Histoire Naturelle, Paris), and an anony- logical Society of Washington 22: 143–152. mous reviewer for their critical advice on the manuscript. Clark, A. H. 1909e. New genera and higher groups of unstalked cri- noids. Proceedings of the Biological Society of Washington 22: The manuscript was greatly improved by the editor Dr. 173–178. Hironori Komatsu. This work was supported by MEXT Clark, A. H. 1909f. Five new species of recent unstalked Crinoids. Pro- KAKENHI Grant Number 04041103 to TK and grants ceedings of the United States National Museum 37: 29–34. from the Director General of National Museum of Nature Clark, A. H. 1911a. On a collection of unstalked crinoids made by Unit- and Science, Japan to TF. This is also a contribution of JSPS ed State Fisheries Steamer Albatross in the vicinity of the Philip- Asian CORE Comsea Program and of Core-to-Core Pro- pine Islands. Proceedings of the United States National Museum gram, B. Asia-Africa Science Platforms. 39: 529–563. Clark, A. H. 1911b. The recent crinoids of the coasts of Africa. Proceed- ings of the United States National Museum 40: 1–51. References Clark, A. H. 1911c. A new unstalked crinoid from Christmas Island. Annals and Magazine of Natural History, Series 8 7: 644–645. Clark, A. H. 1912a. The crinoids of the Indian Ocean. The Trustees of the Bell, F. J. 1882. An attempt to apply a method for formulation to the Indian Museum, Calcutta, 325 pp., 61 figs. species of the Comatulidae; with a description of a new species. Clark, A. H. 1912b. The crinoids of the Natural History Museum at Proceedings of the Zoological Society of London 6: 530–536, pl. Hamburg. Smithsonian Miscellaneous Collections 60: 1–33. 35. Clark, A. H. 1912c. Seventeen new East Indian crinoids belonging to Bell, F. J. 1894. On the echinoderms collected during the voyage of the families Comasteridae and Zygometridae. Proceedings of the H.M.S.‘Penguin’ and by H.M.S.‘Egerina’ when surveying Maccles- Biological Society of Washington 25: 17–28. field Bank. Proceedings of the Zoological Society of London 1884: Clark, A. H. 1912d. Preliminary description of eleven new crinoids be- 392–413. longing to the families Himerometridae, Mariametridae, and Col- Carpenter, P. H. 1881. The Comatulae of the Leyden Museum. Notes obometridae, discovered by the ‘Siboga’ in the Dutch East Indies. from the Leyden Museum 3: 173–217. Annals and Magazine of Natural History, Series 8 10: 31–41. Carpenter, P. H. 1882. Descriptions of new or little-known Comatulae, I Clark, A. H. 1912e. Descriptions of eleven new crinoids belonging to & II. Journal of the Linnean Society 16: 487–526. the families Calometridae and Thalassometridae discovered by Carpenter, P. H. 1884. Report upon the Crinoidea during the voyage of the Siboga in the Dutch East Indies. Zoologischer Anzeiger 39: H.M.S. Challenger during the years 1873–76. Part I. General mor- 420–428. phology, with descriptions of the stalked crinoids. Report on the Clark, A. H. 1912f. Descriptions of twenty new recent unstalked cri- Scientific Results of the voyage of H.M.S. Challenger, Zoology 11: noids, belonging to the families Antedonidae and Atelecrinidae, i–x, 1–442, pls 1–62. from the Dutch East Indies. Notes Leyden Museum 34: 129–155. Carpenter, P. H. 1888. Report upon the Crinoidea collected during the Clark, A. H. 1913. Noted on the recent crinoids in the British Museum. voyage of H.M.S. Challenger, during the years 1873–1876. Part II. Smithsonian Miscellaneous Collections 61: 1–89. The Comatulae. Report on the Scientific Results of the voyage of Clark, A. H. 1915. A monograph of the existing crinoids. Vol. 1, The H.M.S. Challenger, Zoology 26: i–x, 1–402, pls 1–70. Comatulids. Pt. 1. Bulletin of the United States National Museum Carpenter, P. H. 1889. Report on the Comatulae of the Mergui Archi- 82: i–iii, 1–406, pls 1–17. pelago, collected from the trustees of the Indian Museum, Calcut- Clark, A. H. 1918. The unstalked crinoids of the Siboga Expedition. Si- ta, by Dr. J. Anderson. Journal of the Linnean Society 21: 304–316, boga Expeditie 42b: i–ix, 1–300, pls 1–28. pls 26–27. Clark, A. H. 1929. On some recent crinoids in the collection of the Chen, J. C., Chang, K. H., and Chen, C. P. 1988. Shallow water crinoids British Museum. Journal of the Linnean Society 36: 635–664, pls of Kenting National Park, Taiwan. Bulletin of the Institute of Zool- 40–44. ogy, Academia Sinica 27: 73–90. Clark, A. H. 1931. A monograph of the existing crinoids. Vol. 1, The Clark, A. H. 1907a. New genera of recent free crinoids. Smithsonian Comatulids. Pt. 3. Superfamily Comasterida. Bulletin of the Unit- Miscellaneous Collections 50: 343–365. ed States National Museum 82: i–vii, 1–816, pls 1–82. Clark, A. H. 1907b. Descriptions of new species of recent unstalked Clark, A. H. 1934. On a collection of crinoids from the Raffles Museum, crinoids from the coasts of northeastern Asia. Proceedings of the Singapore. Proceedings of the Biological Society of Washington United States National Museum 33: 127–156. 47: 9–14. Clark, A. H. 1908a. New genera of unstalked crinoids. Proceedings of Clark, A. H. 1936. Biological results of the Snellius Expedition. 1. The the Biological Society of Washington 21: 125–136. unstalked crinoids of the Snellius Expedition. Temminkia 1: 295– Clark, A. H. 1908b. New genera and species of crinoids. Proceedings of 317, pls 7–9. the Biological Society of Washington 21: 219–232. Clark, A. H. 1937. Crinoidea. The John Murray Expedition, 1933–34, Clark, A. H. 1908c. Preliminary notice of a collection of recent crinoids Scientific Reports 4(4): 88–108, pl. 1. from the Philippine Islands. Smithsonian Miscellaneous Collec- Clark, A. H. 1941. A monograph of the existing crinoids. Vol. 1, The tions 52: 199–234. Comatulids. Pt. 4a. Superfamily Mariametrida. Bulletin of the Clark, A. H. 1909a. On a collection of recent crinoids from the Philip- Comatulids from Ambon and Lombok islands 245

United States National Museum 82: i–vii, 1–603, pls 1–61. Hartlaub, C. 1890. Beitrag zur Kenntniss der Comatuliden-Fauna des Clark, A. H. 1947. A monograph of the existing crinoids. Vol. 1, The Indischen Archipels. Nachrichten von der Königlichen Gesell- Comatulids. Pt. 4b. Superfamily Mariametrida (concluded) and schaft der Wissenschaften und der Georgs-August-Universität zu Superfamily Tropiometrida. Bulletin of the United States National Göttingeni 5: 168–187. Museum 82: i–vii, 1–473, pls 1–43. Hartlaub, C. 1891. Beitrag zur Kenntniss der Comatulidenfauna des Clark, A. H. 1950. A monograph of the existing crinoids. Vol. 1, The Indischen Archipels. Nova Acta der Kaiserlische Leopoldinisch- Comatulids. Pt. 4c. Superfamily Tropiometrida. Bulletin of the Carolonisch. Deutschen Akademie der Naturforscher 58: 5–120, United States National Museum 82: i–vii, 1–383, pls 1–32. pls 1–5. Clark, A. H. 1952. Echinoderms from the Marshall Islands. Proceedings Hoggett, A. K. and Rowe, F. W. E. 1986. A reappraisal of the family Co- of the United States National Museum 102: 265–303. masteridae A. H. Clark, 1908 (Echinodermata: Crinoidea) with Clark, A. H. and Clark, A. M. 1967. A monograph of the existing cri- the description of a new subfamily and a new genus. Journal of the noids. Vol. 1, The Comatulids. Pt. 5. Suborders Oligophreata (con- Linnean Society (Zoology) 88: 103–142. cluded) and Macrophreata. Bulletin of the United States National Kirkendale, L. and Messing, C. G. 2003. An annotated checklist and key Museum 82: i–x, 1–860. to the Crinoidea of Guam and the Commonwealth of the North- Clark, A. M. 1972. Some crinoids from the Indian Ocean. Bulletin of ern Mariana Islands. Marine Biology 152: 523–546. the British Museum (Natural History) Zoology 24: 75–156, table Kogo, I. 1998. Crinoids from Japan and its adjacent waters. Osaka Mu- 15. seum of Natural History, Special Publication 30: 1–148. Clark, A. M. 1982. Echinoderms of Hong Kong. Pp. 485–501. In: Mor- Kogo, I. 2002. Report on the crinoids collected from the Nansei Islands, ton, B. S. and Tseng, C. K. (Eds) The Marine Flora and Fauna of Japan, during a cruise of the training vessel Toyoshio Maru in Hong Kong and Southern China, Hong Kong, 1980. Hong Kong 1999. Bulletin of the Osaka Museum of Natural History 56: 1–44. University Press, Hong Kong. Kogo, I. 2006. Comatulid fauna (Echinodermata: Crinoidea) of the Sa- Clark, A. M. and Rowe, F. W. E. 1971. Monograph of Shallow-water gami Sea and a part of Izu Islands, central Japan. Memoirs of the Indo-West Pacific Echinoderms. Trustees of the British Museum National Science Museum, Tokyo 41: 223–246. (Natural History), London, x+238 pp., 31 pls. Kogo, I. and Fujita, T. 2005. Geographical distribution of crinoids Clark, A. M. and Spencer Davis, P. 1966. Echinoderms of the Maldive (Echinodermata) in southwestern Japan. National Science Muse- Islands. Annals and Magazine of Natural History, Series 13 8: um Monographs 29: 297–355. 597–621, pl. 18. Lane, D. J. W., Marsh, L. M., VandenSpiegel, D., and Rowe, F. W. E. Clark, H. L. 1915. The comatulids of Torres Strait: with special reference 2000. Echinoderm fauna of the South China Sea: an inventory to their habits and reactions. Papers from the Department of Ma- and analysis of distribution patterns. Raffles Bulletin of Zoology, rine Biology of the Carnegie Institution of Washington 8: 99–125. Supplement 8: 459–493. Clark, H. L. 1921. The echinoderm fauna of Torres Strait: its composi- Liao, Y. 1983. The echinoderms of the Xisha Islands, Guangdong Prov- tion and its origin. Papers from Department of Marine Biology of ince, China. V. Crinoidea. Studia Marina Sinica 20: 263–277, pl. 1. the Carnegie Institution of Washington 10: 1–223, pls 1–38. Liao, Y. and Clark, A. M. 1995. The Echinoderms of Southern China. Sci- Clark, H. L. 1946. The echinoderm fauna of Australia, its composition ence Press, Beijing, 614 pp., 23 pls. and its origin. Publication of the Carnegie Institution of Washing- Linnaeus, C. 1758. Systema naturae per regna tria naturae, secundum ton 566: 1–567. classes, ordines, genera, species, cum characteribus, differentiis, syn- Fabricius, E. K. 1994. Spatial patterns in shallow-water crinoid com- onymis, locis. Tomus I. Editio decima, reformata. Holmiae, Salvius, munities on the central Barrier Reef. Australian Journal of Marine viii+824 pp. and Freshwater Research 45: 1225–1236. Marshall, J. I. and Rowe, F. W. E. 1981. The crinoids of Madagascar. Bul- Fishelson, L. 1974. Ecology of the northern Red Sea crinoids and their letin du Muséum Nationale d’Histoire Naturelle, Paris, 4ème Série epi- and endozoic fauna. Marine Biology 26: 183–192. 3: 379–413. Fleming, J. 1828. A History of British Animals, Exhibiting the Descriptive MarshallCrossland, J. I. and Price, A. R. G. 1999. Indian Ocean echi- Characters and Systematical Arrangement of the Genera and Spe- noderms collected during the Sindbad Voyage (1980–81): 4. Cri- cies of Quadrupeds, Birds, Reptiles, Fishes, Mollusca, and Radiata of noidea. Bulletin of the Natural History Museum of London (Zool- the United Kingdom; Including the Indigenous, Extirpated, and Ex- ogy) 65: 23–29. tinct Kinds, Together with Periodical and Occasional Visitants. Bell Mekhova, E. S. and Britayev, T. A. 2012. Fauna of unstalked crinoids and Bradfute, Edinburgh, xxiii+565 pp. (Crinoidea: Comatulida) of the Bay of Nhatrang, Southern Viet- Gibbs, P. E., Clark, A. M., and Clark, C. M. 1976. Echinoderms from nam. Paleontological Journal 46: 909–926. the northern region of the Great Barrier Reef, Australia. Bulletin Messing, C. G. 1994. Comatulid crinoids (Echinodermata) of Madang, of the British Museum (Natural History) 30: 103–144. Papua New Guinea, and environs: Diversity and ecology. Pp. Gislén, T. 1922. The crinoids from Dr. S. Bock’s expedition to Japan 237–243. In: David, B., Guille, A., Féral, J.-P., and Roux, M. (Eds): 1914. Nova Acta Regiae Societatis Scientiarum Upsaliensis 5: Echinoderms through Time. Proceedings of the Eighth International 1–183, pls 1–2. Echinoderm Conference, 1993. A. A. Balkema, Rotterdam. Gislén, T. 1927. Papers from Dr. Th. Mortensen’s Pacific expedition Messing, C. G. 1995. Alloeocomatella, a new genus of reef-dwelling 1914–16. 37. Japanese crinoids. Videnskabelige Meddelelser frå feather star from the tropical Indo-West Pacific (Echinodermata: Dansk Naturhistorisk Forening i Kjøbenhavn 83: 1–69, pls 1–2. Crinoidea: Comatulidae). Proceedings of the Biological Society of Gislén, T. 1938a. Crinoids of S. Africa. Kunglica Svenska Vetenskap- Washington 108: 436–450. sakademiens Handlingar, Series 3 17: 1–22, pls 1–2. Messing, C. G. 1997. Living comatulids. Pp. 3–30. In: Waters, J. A. and Gislén, T. 1938b. Crinoids from French Indo China and Macclesfield Maples, C. G. (Eds) Geology of Echinoderms, The Paleontological Bank. Kunglica Fysiographiska Sällskapets i Lund Forhandlingar Society Papers, 3. Carnegie Museum of Natural History, Pitts- 7: 3–21. burgh. Gislén, T. 1940. A collection of crinoids from the South Sea Islands. Messing, C. G. 1998a. Revision of the Recent Indo-West Pacific comat- Kunglica Svenska Vetenskapsakademiens Handlingar, Series 3 18: ulid genus Comaster Agassiz. Part 1. The type species of Comaster 3–16, pls 1–3. and Phanogenia Lovén. Invertebrate Taxonomy 12: 191–209. 246 Ichizo Kogo et al.

Messing, C. G. 1998b. An initial re-assessment of the distribution and a combined approach to explain the morphological and ecological richness of the East Indian shallow-water crinoid fauna. Pp. 187– diversity in Phanogenia gracilis Hartlaub, 1893 (Echinodermata: 192. In: Mooi, R. and Telford, M. (Eds) Echinoderms: San Francis- Crinoidea) sensu lato: two species or intraspecific variation? Ma- co. Proceedings of the Ninth International Echinoderm Conference, rine Biology 156: 1517–1529. San Francisco, 1996. Balkema, Rotterdam. Pilcher, N. J. and Messing, C. G. 2001. A preliminary checklist of the Messing, C. G. 2001. A key to the genera of Comasteridae (Echinoder- shallow-water crinoids of the Kerama Islands, Okinawa, Japan. mata: Crinoidea) with the description of a new genus. Bulletin of Report of Akajima Marine Science Laboratory 12: 15–17. [In Japa- the Biological Society of Washington 10: 277–300. nese] Messing, C. G. 2007. The crinoid fauna (Echinodermata: Crinoidea) of Rankin, D. L. and Messing, C. G. 2008. A revision of the comatulid Palau. Pacific Science 61: 91–111. genus Stephanometra A. H. Clark with a diagnosis of the genus Messing, C. G., Améziane, N., and Eléaume, M. 2000. Echinodermata Lamprometra A. H. Clark (Echinodermata: Crinoidea). Zootaxa Crinoidea: Comatulids of the KARUBAR Expedition to Indone- 1888: 1–35. sia: the families Comasteridae, Asterometridae, Calometridae and Reichensperger, A. 1913. Ungestielte Crinoiden der Aru- und Kei-In- Thalassometridae. Mémoires de le Museum National d’Histoire seln. Abhandlungen der Senckenbergischen Naturforschenden Naturelle, Paris 184: 627–702. Gesellschaft 35: 79–108. Messing, C. G. and Tay, T. S. 2016. Extant Crinoidea (Echinodermata) Rowe, F. W. E., Hoggett, A. K., Birtles, R. A., and Vail, L. L. 1986. Revi- of Singapore. Raffles Bulletin of Zoology, Supplement 34: 627–658. sion of some comasterid genera from Australia (Echinodermata, Meyer, D. L. and Macurda, D. B. Jr. 1980. Ecology and distribution of Crinoidea), with descriptions of two new genera and nine new the shallow-water crinoids of Palau and Guam. Micronesica 16: species. Journal of the Linnean Society 86: 197–277. 59–99. Rowe, F. W. E. and Gates, J. 1995. Zoological Catalogue of Australia, Vol. Meyer, D. L., LaHaye, C. A., Holland, N. D., Arneson, A. C., and Strick- 33, Echinodermata. CSIRO, Melbourne, 510 pp. ler, J. R. 1984. Time-lapse cinematography of feather stars on the Smith, E. A. 1876. Diagnosis of new species of Mollusca and Echino- Great Barrier Reef, Australia: demonstrations of posture changes, dermata from the island of Rodriguez. Annals and Magazine of locomotion, spawning and possible predation by fish. Marine Bi- Natural History, Series 4 17: 404–406. ology 78: 179–184. Summers, M. M., Messing, C. G., and Rouse, G. W. 2014. Phylogeny of Müller, J. 1841. Über die Gattungen und Arten der Comatulen. Archiv Comatulidae (Echinodermata: Comatulida): A new classification für Naturgeschichte 7: 139–148. and assessment of morphological characters for crinoid taxonomy. Müller, J. 1843. Neue Beiträge zur Kenntnis der Arten von Comatulen. Molecular Phylogenetics and Evolution 80: 319–339. Archiv für Naturgeschichte 9: 113–136. Tortonese, E. 1977. Report on Echinoderms from the Gulf of Aqaba Müller, J. 1849. Über die Gattung Comatula Lam. und ihre Arten. Ab- (Red Sea). Italian Journal of Zoology 12: 273–290. handlungen der Koniglichen Akademie der Wissenschaften zu Utinomi, H. and Kogo, I. 1965. On some crinoids from the coastal sea Berlin 1847: 237–265. of Kii Peninsula. Publications of the Seto Marine Biological Labo- Norman, A. M. 1865. On the genera and species of British Echinoder- ratory 13: 263–286, 1 pl. mata. 1. Crinoidea, Ophiuroidea, Asteroidea. Annals and Maga- Vail, L. 1987. Dial patterns of emergence of crinoids from within a reef zine of Natural History, Series 3 15: 98–129. at Lizard Island, Great Barrier Reef, Australia. Marine Biology 93: Obuchi, M. 2013. Two new tropical comatulids (Echinodermata: Cri- 551–560. noidea: Comasteridae) from Okinoshima Island, Kochi, new re- Zmarzly, D. L. 1984. Distribution and ecology of shallow-water crinoids cords for Japan. Kuroshio Biosphere 9: 15–26, pl. 1. at Enewetak Atoll, Marshall Island, with an annotated checklist of Owen, C. L., Messing, C. G., Rouse, G. W., and Shivji, M. S. 2009. Using their symbionts. Pacific Science 38: 105–122.