Chloeia pulchella Baird 1868

Chloeia pulchella Baird, 1868 reinstated Figs 45–47 Chloeia incerta de Quatrefages, 1866: 388; Hartman 1959: 131; Solís-Weiss et al. 2004: S2; Salazar-Vallejo et al. 2014: 11 (list) (partim). Chloeia pulchella Baird, 1868: 234–235; Hartman 1959: 132; Barroso & Paiva 2011: 422, Tab. 1. Chloeia pulchella var. pallida Baird, 1868: 235. Chloeia flava: Grube 1877: 509; Izuka 1912: 223–225, Pl. 2, Fig. 4, Pl. 22, Figs 3–5; Monro 1924: 71–72 (partim); Fauvel 1947: 24 (partim) (non (Pallas, 1766)). Chloeia flava var. pulchella: Horst 1912: 19, Pl. 7, Fig. 3; Fauvel 1917: 190–191; Fauvel 1947: 24–25, Fig. 20i; Fauvel 1953: 95, Fig. 46b; Hartman 1959: 131; Bleeker & van der Spoel 1992: 128. Type locality. Northeast coast of Australia (Baird 1868: 234). Type material. Northeastern Australia. Two syntypes of Chloeia pulchella (BMNH 1971.238), Raine Island reefs, H.M.S. Herald, F.M. Rayner, coll. Holotype of Chloeia pulchella var. pallida (BMNH 1846.10.14.60), Raine Island, J.B. Jukes, coll. (no further data). “ Mers de la Chine ”. One syntype of C. incerta de Quatrefages, 1866 (MNHN IA-TYPE 250), no further data (complete, pale, 73 mm long, 16 mm wide, 38 chaetigers). India. One syntype of C. incerta de Quatrefages, 1866 (MNHN IA-TYPE 248), Pondicherry, no further data (data in variation) Additional material. Australia. One specimen (AM W4453), Queensland, Deception Bay (27°09´S, 153°05´E), Aug. 1969, no further data (depressed, almost colorless; anterior and posterior ends bent dorsally; middorsal spots oval, longer than wide, barely visible along anterior and median segments; body 57 mm long, 18 mm wide, 35 chaetigers). One specimen (AM W 4455), New South Wales, Port Hacking (34°04´15″ S, 151°16´30″ E), 20 Sep. 1938, no further data (juvenile, bent ventrally; middorsal oval spots, longer than wide, visible along most chaetigers; body 9 mm long, 3 mm wide, 18 chaetigers). One specimen (AM W 30895), Queensland, Weipa, Lorim Point Wharf (12°40´S, 141°57´E), 1 Oct 1999, CRC Reef Research Centre, coll. (bent ventrally; pharynx partially exposed; middorsal spots oval, longer than wide, along body; median antenna purplish along posterior surface, 1/3 as long as caruncle; dorsal cirri dark purple; caruncular median ridge purplish; branchial stems pale, branches purplish; body 30 mm long, 10 mm wide, 36 chaetigers). One specimen (AM W 201650), Queensland, Cape Melville (14°10´S, 144°30´E), 8 Feb- 1979, no further data (bent ventrally; middorsal spots blackish, oval, longer than wide, visible along body; lateral bands on anterior notopodial surfaces paler; dorsal cirri dark purple; branchial stems pale, branches blackish; chaetae transparent, with yellowish band distally; right neuropodium of chaetiger 7 missing, duplicate in chaetiger 11; body 39 mm long, 10 mm wide, 32 chaetigers). One specimen (BMNH 1930.10.4.17), Cape Boileau, Northwestern Australia, B. Grey, coll., no further data (partially dried-out, yellowish; middorsal spots oval, longer than wide, dark purple, almost black; branchiae from chaetiger 4, stems pale, branches blackish; body twisted, anterior end compressed in small container, 61 mm long, 16 mm wide, 36 chaetigers). One specimen (MNHN A 53.4), no depth data, 1875, M. Casagneau, coll. (body grayish; middorsal spots dark purple, oval, longer than wide along body; dorsal cirri and branchiae pale, branchiae stiff, vertical; anterior eyes 3× larger than posterior ones; body bent laterally, 67 mm long, 14 mm wide, 41 chaetigers). One specimen (NTM W22040), NW Crocodile Island, MilingimbiAmhem Landing (11°41.162´S, 135°9.168´E), 2.7 m, Feb. 2004, R.S. Williams, coll. (bent ventrally; middorsal spots subrectangular, tips rounded; median antenna 1/3 as long as caruncle; dorsal cirri dark purple; branchial stems pale, branches purplish; anterior eyes 3–4× larger than posterior ones; body 9 mm long, 4.5 mm wide, 24 chaetigers). One specimen (ZMH P19546), complete, Dampier, Hampton Harbour, Western Australia, 7 Oct. 1972, donated by the Western Australian Museum (almost colorless, posterior end bent ventrally; middorsal spots oval, longer than wide, reddish along body; chaetae with yellowish tips; 88 mm long, 23 mm wide, 41 chaetigers). One specimen (ZMH V7424), Cape Moreton, Queensland, reef, no date or depth data, H. Schmidt, coll. (bent ventrally, pharynx exposed by fracture of ventral wall; dorsal oval spots, longer than wide, only visible along chaetigers 5–13, faded thereafter; body and chaetae brownish; body 86 mm long, 15 mm wide, 38 chaetigers). One specimen (ZMH V7703), Brisbane River, Queensland, no date or depth data, H. Schmidt, coll. (colorless, twisted, anterior end collapsed by pressing it in a small container; middorsal spots oval, longer than wide, visible along anterior 2/3 of body; body 107 mm long, 22 mm wide, 39 chaetigers). One specimen (ZMH V19545), Hampton Harbor, Dampier, Western Australia, intertidal flats, mud and boulders, 7 Oct. 1972, gift from Western Australia Museum (colorless, bent ventrally; middorsal spots oval, longer than wide throughout body; 70 mm long, 15 mm wide, 38 chaetigers). Indonesia. One specimen (ZMA VPol 153.1), Irian Jaya, RV Siboga Exped., Sta. 162 (between Loslos and Broken Isl., Salawatti, W coast), 18 m, dredge, sand, shells, 18 Aug. 1899 (anterior fragment, pale, lips pale; 18 mm long, 11 mm wide, 13 chaetigers). One specimen (ZMA VPol 153.2), Maluku, RV Siboga Exped., Sta. 258 (Kei Isl., Tual anchorage), 22 m, reef exploration, dredge, stones + coral, 12– 16 Dec. 1899 (middorsal spots oval throughout body; lips blackish; body brownish, bent ventrally, 46 mm long, 10 mm wide, 38 chaetigers; median chaetigers with harpoon-chaetae with well-defined spur). One specimen (ZMA VPol 155), Batavia (Jakarta), Java, no further data (bent ventrally, with oval spots; body 102 mm long, 23 mm wide, 40 chaetigers). One specimen (ZMB 752), SMS Gazelle Expedition, 1874–1876, West Papua, Sarawatti, coral reef, F. von Schleinitz, coll. (complete; middorsal spots oval, longer than wide, darker along anterior chaetigers; notochaetae with distal yellow band; body 80 mm long, 11 mm wide, 40 chaetigers). Thailand. One specimen (ZMH P19979), Mua Min, Gulf of Thailand, 1989, H. Thielen, coll. (data in variation). One specimen (ZMH P19980), Laem Ngon, Gulf of Thailand, 1989, H. Thielen, coll. (complete, colorless; middorsal spots oval, longer than wide, narrower in posterior chaetigers; lateral bands faded off; dorsal cirri and branchiae pale; chaetal tips wth yellowish-green pigmentation along anterior body third; body 106 mm long, 18 mm wide, 38 chaetigers). Singapore. One specimen (MNHN 398.1), Reef, no depth or date data, Dawydoff, coll. (complete, dorsal spots oval, longer than wide; diffuse spots along anterior notopodial surfaces; anterior eyes 2 larger than posterior ones; median antenna as long as caruncle; body 30 mm long, 7 mm wide, 29 chaetigers). Sri Lanka. One specimen (BMNH 1923.3.23.5), donation of the Colombo Museum, no further data (damaged, smashed in a small container, constricted by oppression in chaetigers 10–11; pale, middorsal spots oval, purple; median antenna 1/3 as long as caruncle, slightly longer than lateral antennae; lateral antennae slightly longer than palps; branchiae from chaetiger 4, stems pale, branches purple; body 76 mm long, 16 mm wide, 38 chaetigers). Japan. Four specimens (MNHN A53.1), Sta. 17, 1909, M. de Guerne, coll. (complete; middorsal spots oval, longer than wide, to narrow medially wider bands, along median and posterior chaetigers, one specimen with anterior chaetigers with round spots; dorsal cirri purplish; branchiae pale; anterior eyes 2×larger than posterior ones; body 68–88 mm long, 16–18 mm wide, 34–36 chaetigers). Four specimens (MNHN A53.2), Sta. 17, 1909, M. de Guerne, coll. (complete; middorsal spots oval, longer than wide, fading off; dorsal cirri and branchiae pale; anterior eyes 2×larger than posterior ones, 4× larger in one specimen; body 70–75 mm long, 16–19 mm wide, 34–36 chaetigers). Five specimens (MNHN A54 17), Miyajima Island, Hiroshima Bay. Inland Sea, Jun. 1908 (data in variation). One specimen (ZMH V646), Nagasaki, no depth or date data (identified by Michaelsen 1893), H. B̂ddinghaus, coll. (partially dehydrated, pharynx exposed, distorted, anterior end bent dorsally; middorsal spots oval, longer than wide, lateral bands fused anteriorly, continued laterally along notopodial anterior surfaces; median antenna and dorsal cirri dark purple; branchiae pale; body 92 mm long, 18 mm wide, 35 chaetigers). Two specimens (BMNH 1925.1.28.134 – 135), Inland Sea, no further data (middorsal spots oval; bipinnate branchiae from chaetiger 4; body bent ventrally, 73–84 mm long, 19–23 mm wide, 35–36 chaetigers). One specimen (ZMH V4457), Shikoku, Sanuki, no depth or date data, H. Lenz, coll. (partially dried out; middorsal spots oval, longer than wide, visible along body; lateral brownish bands along anterior parapodial surfaces very wide, brownish; dorsal cirr dark purple; branchiae pale; chaetae golden; venter pale; 72 mm long, 10 mm wide, 36 chaetigers). China. Three specimens (BMNH 1926.4.27.28 – 30), Amoy, Prof. Ping, coll., no further data (pale, middorsal spots oval, dark purple, longer than wide; lateral bands along notopododial anterior surfaces visible throughout body; dorsal cirri dark purple; branchiae from chaetiger 4, pinkish; largest specimen with pharynx exposed, rings smooth; body 34–55 mm long, 8– 8 mm wide, 31–32 chaetigers). Two specimens (BMNH 1926.4.27.31 – 32), Joachow, Prof. Ping, coll., no further data (whitish; middorsal spots oval, longer than wide, purple; one tapered with branchiae contracted, another one markedly contracted with branchiae not contracted, both with branchiae from chaetiger 4; median antennae 2× longer than lateral antennae; lateral antennae 3× longer than palps; body 45–52 mm long, 10–16 mm wide, 30–32 chaetigers). One specimen (ZMH V7733), Fockien, no depth or date data, G. Siemssen, coll. (bent ventrally; middorsal spots oval, longer than wide, along body; lateral bands barely visible, extensions along notopodial surfaces dark purple, wide; median antenna and dorsal cirri dark purple; branchiae pale; body 65 mm long, 18 mm wide, 34 chaetigers). One specimen (ZMH V8215), Fockien, no depth or date data, G. Siemssen, coll. (complete; anterior end distorted, pharynx exposed through fracture on dorsal body wall, probably after tearing out a fishing hook; middorsal spots oval, longer than wide, along anterior and median chaetigers, blunt rectangular in posterior chaetigers; lateral bands along notopodia dark purple, wide; roughly circular purple spots in neurochaetal bundle; body 100 mm long, 21 mm wide, 37 chaetigers). China Sea. One specimen (ZMH V645), no further data, H. Vollbarth, coll. (almost colorless; middorsal spots reddish, better defined along anterior and posterior chaetigers, lateral bands wide, barely visible along anterior region; few dorsal cirri on site, reddish; body 36 mm long, 9 mm wide, 29 chaetigers). Vietnam. One specimen (MNHN 54.8), RV Lanessan, Sta. unnumb., Chult Beach, 20 May 1930, M.A. Krempf, coll. (partially dehydrated; middorsal spots reddish, fading out, oval, longer than wide; dorsal cirri and branchiae pale; chaetae with distal yellowish banding; anterior eyes 2–3 larger than posterior ones; body 72 mm long, 17 mm wide, 38 chaetigers). New Guinea. One specimen (BMNH 1925.1.28.139), Dune Island, no further data (pale; middorsal spots oval, longer than wide, purple; median antenna ¾ as long as caruncle, about 2× longer than lateral antennae; lateral antennae slightly longer than palps; branchiae brownish from chaetiger 4; chaetae golden; body slightly bent ventrally, 77 mm long, 20 mm wide, 37 chaetigers). Papua New Guinea. One specimen (AM A2248), Fairfax Harbour (09°26´34″ S, 147°05´45″ E), 1877–1878, no further data (stiff, partially dehydrated, slightly bent laterally; middorsal spots dark purple, oval, longer than wide, visible along body; dorsal cirri pale; branchial stems colorless, branches purplish; neurochaetae with yellowish band distally; body 60 mm long, 16 mm wide, 39 chaetigers). New Caledonia. One specimen (MNHN A53.3), north off New Caledonia, 10 m, Mar. 1826 (complete, posterior end bent ventrally; middorsal spots barely visible, oval, longer than wide, reddish; dorsal cirri and branchiae pale orange; eyes faded off; body 61 mm long, 18 mm wide, 36 chaetigers). Three specimens (MNHN A54.13), 1880, no further data (complete, larger specimens bent ventrally; all with middorsal spots oval, longer than wide; median specimen with dorsal cirri purplish, branchiae brownish; others with dorsal cirri and branchiae colorless; anterior eyes barely larger than posterior ones in smallest specimen, 2–3× larger in larger specimens; body 16–53 mm long, 4–10 mm wide, 26–33 chaetigers). One specimen (MNHN Musorstom NC987), Sta. 987 (20°16.3´S, 164°00.3´E), 18–19 m, 30 Apr. 1988 (bent ventrally; anterior end slightly compressed; middorsal spots oval, longer than wide, blackish; dorsal cirri purplish; branchiae pale; body 34 mm long, 8 mm wide, 31 chaetigers). Diagnosis. Chloeia with bipinnate branchiae from chaetiger 4, progressively smaller posteriorly; middorsal spots almond-shaped or oval, longer than wide, central in each segment; harpoon notochaetae without spurs; neurochaetae spurred, furcates and acicular. Description. Syntypes (BMNH 1971.238) complete, shortest with anterior end bent dorsally, longest flaccid (Fig. 45A); body fusiform, 44–45 mm long, 8 mm wide, 33 chaetigers. Syntypes pale, almost whitish. Dorsum with almond-shaped or oval, longer than wide, brownish spots, wider medially, often with well-defined tips, especially anteriorly; lateral brownish bands continued towards anterior parapodial surfaces visible only along a few anterior chaetigers (Fig. 45B). Dorsal cirri brownish, darker along anterior surfaces, and in cirrophores. Bipinnate branchiae with pale stems, lateral branches brownish. Venter pale. Prostomium anteriorly entire. Eyes blackish, anterior eyes slightly larger than posterior ones. Median antenna inserted at anterior caruncular margin, brownish or pale, 1/3 as long as caruncle, size proportion to lateral antennae unknown because lateral antennae broken. Lateral antennae bases close to each other. Palps broken. Mouth ventral on chaetiger 2–3. Pharynx not exposed. Caruncle pale, twisted in one syntype (Fig. 45C), sigmoid in the other, trilobed, tapered, reaching chaetiger 4 or 6. Median ridge with upper margin blackish, with about 24–30 vertical folds, partially concealing lateral lobes. Lateral lobes narrow, with about 32 vertical folds. Bipinnate branchiae from chaetiger 4, continued throughout body, parallel along most body segments; progressively larger to chaetigers 12–13, diminishing in size in posterior chaetigers. Median segments with 8–9 lateral branches. Parapodia biramous, notopodia with cirriform branchiae along chaetigers 1–4 (chaetiger 4 with both types of branchiae), 1/3–1/5 as long as dorsal cirri. Dorsal cirri longer than bipinnate branchiae along most chaetigers, 3–4× longer in posterior chaetigers. Second ventral cirri with cirrophores 2× longer and slightly wider, and cirrostyle 2× longer than adjacent ones, directed dorsally. Other ventral cirri directed ventrolaterally, as long as one and a half the length of subsequent segments. Chaetae most complete. Complete chaetae with distal fragile hoods, rarely eroded. Notochaetae in anterior chaetigers furcates (Fig. 45D), major tines 5—6× longer than minor ones. Median chaetigers with harpoon notochaetae (Fig. 45F), spurs barely visible. Neurochaetae all furcates, major tines 3—4× longer than minor ones in anterior chaetigers (Fig. 45E), up to 5—6× longer in median chaetigers, including aciculars (Fig. 45G). Posterior region tapered (Fig. 45H); pygidium with anus terminal; anal cirri pale, sausage-shaped, 6–7× longer than wide. Live pigmentation (after photo from a Queensland specimen by Nyoni (2020)). Body salmon pink with blackish dorsal cirri and golden chaetae. Dorsal pigmentation with blackish middorsal oval to fusiform spots, each longer than wide, in a whitish halo, wider laterally, barely developed along anterior and posterior spots margins; lateral bands continued towards anterior parapodial surfaces. Branchiae with white stems and dark red lateral branches. Caruncle with median ridge blackish. There are two videos available on the internet (Iromongara 2009, Senja 2021, 1:27). Variation. The holotype of C. p. var. pallida (BMNH 1846.10.14.60) is complete, slightly twisted to the left, almost colorless; body 49 mm long, 12 mm wide, 35 chaetigers. Eyes blackish, small, anterior eyes 2× larger than posterior ones. Antennae mucronate; median antenna 1/3 as long as caruncle (Fig. 46A), slightly longer than lateral antennae. Lateral antennae with bases separate from each other, 2× longer than palps. Caruncle sinuous, reaching chaetiger 5, vertical folds in median ridge and lateral lobes difficult to count. Middorsal spots drop-shaped to fusiform along chaetigers 4–11; following segments with thinner, teardrop to oval spots (Fig. 46B) continued to a few segments before pygidium. Branchiae from chaetiger 4, progressively larger to chaetiger 9, shorter in segments of the last body third. Anal cirri subcylindrical (Fig. 46C), whitish, 3× longer than wide. The syntype of C. incerta de Quatrefages, 1866 (MNHN IA-TYPE 248) is brownish, beheaded (Fig. 46D), with a ventral dissection along chaetigers 14–19; body 84 mm long, 15 mm wide, 39 chaetigers. Median segments with oval middorsal spots (Fig. 46E). Posterior region tapered (Fig. 46F), with thinner middorsal bands. Three large specimens from Japan (MNHN A54 17) have middorsal spots oval (Fig. 47A, C, E); bipinnate branchiae from chaetiger 4, some removed from median chaetigers for observing middorsal spots; body 70–79 mm long, 13–16 mm wide, 32–35 chaetigers. Despite being of similar size, the middorsal spots have slightly different width, but this is not size-dependent, because they are rather oval in specimens 70 or 79 mm long, whereas they are thinner in a 75 mm long (Fig. 47B, D, F). These specimens also have some remnants of a dark transverse band over the middorsal oval spot, and paler lateral bands. Another large specimen from Thailand (ZMH P19979) is complete, 111 mm long, 17 mm wide, 38 chaetigers, and has a better defined pigmentation pattern (Fig. 47G); the middorsal spots are oval, longer than wide, visible along body, rectangular in posterior chaetigers; lateral bands surrounding middorsal spots anteriorly, fused to middorsal spots along anterior body third, directed laterally along notopodial anterior surfaces (Fig. 47H); some anterior neuropodia with circular purple spots; dorsal cirri and branchial stems pale, branchial branches dark purple; chaetae with a yellowish band along their distal half in anterior region, becoming distal third or fourth in median and posterior chaetigers; anterior eyes 2–3× larger than posterior ones. Specimens from Indonesia (ZMA VPol 153.2) have blackish lips and harpoon-chaeetae with better defined spurs. Further, the middorsal oval spots are long lasting when most other pigments fade off, but there are some videos available in internet made with specimens from Malaysia (Iromongara 2009, Senja 2021, 1:27), showing specimens with a pale pink or whitish dorsal background and having only the middorsal spots, and lateral bands visible along most segments. Remarks. Chloeia pulchella Baird, 1868, reinstated, was described from Northeastern Australia; because of the presence of bipinnate branchiae from chaetiger 4, becoming progressively smaller posteriorly, and by having middorsal spots oval or round, it belongs in the group flava. However, in C. pulchella dorsal spots are oval, and it resembles C. maculata Potts, 1909, described from the Western Indian Ocean. These two species differ, however, especially after the type of dorsal spots, and the type of harpoon notochaetae, and neurochaetae. In C. pulchella the middorsal spots are oval, longer than wide, and central in each segment, the harpoon notochaetae do not have spurs or smooth tines, and the neurochaetae are spurred or furcates, with short minor tines, whereas in C. maculata the middorsal spots are half-oval, displaced towards posterior segmental half, the harpoon notochaetae have short smooth tines, and the neurochaetae have long minor tines. Baird (1868) did not explain the etymology; the specific epithet is the diminutive of the Latin adjective pulcher: beautiful. The dorsal pigmentation pattern included “a distinct rather broad line of a very dark colour—not squareshaped, as in C. flava, but extending the length to more than half of the breadth of the segment” (Baird 1868: 235). After a slightly larger, paler specimen collected close to the type locality of the syntypes, he proposed the variety Chloeia pulchella var. pallida because it had less intense pigmentation in chaetae and body, and colorless branchiae, although chaetae were similar. He indicated that the specimen was “nearly colourless in all parts except the peculiar marks on the dorsal surface of the segments.” No further details were given, and the specific epithet is based upon the paler body pigmentation. These differences are herein disregarded as for sustaining a subspecific taxon, and consequently the variety is included as a junior synonym. On the other hand, Horst (1912:19) followed M’Intosh (1885: 9) by regarding C. pulchella as a junior synonym of C. flava, but retained the name because his specimens, of similar size than the syntypes “probably represent young individuals” of C. flava, such that they “c