Chloeia violacea Horst, 1910

Salazar-Vallejo, Sergio I., 2023, Revision of Chloeia Savigny in Lamarck, 1818 (Annelida, Amphinomidae), Zootaxa 5238 (1), pp. 1-134 : 116-118

Chloeia violacea Horst, 1910

Figs 1E , 55 , 56

Chloeia violacea Horst, 1910: 174, 1912: 22 , Pl. 7, Fig. 8 , Pl. 8, Figs 8–11 ; Monro 1937: 253–254; Fauvel 1953: 95, Fig. 46e ; Hartman 1959: 132; Gallardo 1968: 56–57, Pl. 10, Figs 13 , 14 ; Bleeker & van der Spoel 1992: 128 (design. lectotype); Barroso & Paiva 2011: 422, Tab. 1; Salazar-Vallejo et al. 2014: 11 (list).

Type material. Indonesia. Lectotype ( ZMA V.Pol 161.3 , designated by Bleeker & van der Spoel 1992: 128), Maluku, RV Siboga Exped ., Sta. 174 (Ceram (Seram), N coast, Waru Bay or Bima Bay ), 18 m, townet, dredge, reef expl., muddy bottom, 28–29 Aug. 1899 . Paralectotype ( ZMA V.Pol 161.1 ), Lesser Sunda Islands, RV Siboga Exped ., Sta. 47 (Bay of Bima, near South Fort ), 55 m, trawl, dredge and shore expl., muddy bottom with coral sand, 8–12 Apr. 1899 (these two lots were included in the original description) .

Additional material. Indonesia. One specimen ( ZMA V.Pol 161.2 ), Irian Jaya, RV Siboga Exped ., Sta. 164 (01°42.5´S, 130°47.5´E), 32m, dredge, sand, stones, shells, 20 Aug. 1899 (many branchiae and chaetae lost; body pinkisk, without pigmentation, fusiform, anterior prostomial area dark purple; anterior eyes 3–4× larger than posterior ones; bipinnate branchiae from chaetiger 4; 13 mm long, 3 mm wide, 23 chaetigers). One specimen ( ZMA V.Pol 161.4 ), Maluku, Kei Island, off Nuhu Jaan, RV Siboga Exped ., Sta. 260 (05°36.5´S, 132°55.2´E), 90m, dredge, sand + coral + shells, 16–18 Dec. 1899 (many branchiae and chaetae lost; body without pigmentation, fusiform, anterior eyes 3× larger than posterior ones; bipinnate branchiae from chaetiger 4; 11 mm long, 3 mm wide, 22 chaetigers). One specimen ( ZMA V.Pol 2056 ), Lesser Sunda Islands, RV Siboga Exped ., Sta. 313 (E of Dangar Besar, anchorage), 36 m, dredge, trawl + reef expl., sand, coral, muddy bottom, 14–16 Feb. 1900 (damaged). One specimen ( NTM W18632 ), West Natuna , Sta. NB18B (03°40.28´N, 107°56.00´E), 56 m, 3 Aug. 2001, I. Al-Hakim, coll. (early juvenile, anterior end bent laterally; body wall grayish; dorsal cirri dark purple; branchial stems purplish; body depressed, 3.5 mm long, 1.5 mm wide, 14 chaetigers). Gulf of Oman. One specimen ( BMNH 1936.2.8.20 ), John Murray Expedition , HEMS Mabahiss, Sta. 74 (25°17´00″ N, 56°45´00″ E), 155 m, 27 Nov. 1933 (whitish, pharynx partially exposed; some notopodia previously removed; dorsal cirri and branchial stems dark purple; branchiae from chaetiger 4; body bent ventrally, 10.5 mm long, 3 mm wide, 20 chaetigers) .

Diagnosis. Chloeia with bipinnate branchiae from chaetiger 4, progressively smaller posteriorly; middorsal spots as inverted T, posterior region with middorsal band wide, lateral branches short; notochaetae furcates and harpoon-chaetae with basal tines; neurochaetae furcates.

Description. Lectotype (ZMA V.Pol 161.3) complete ( Fig. 55A ), soft, left notopodia of chaetigers 4–8, 18 previously removed; most ventral cirri lost; body fusiform, 24 mm long, 5.5 mm wide, 26 chaetigers.

Lectotype pinkish, without dorsal pigmentation spots. Median antenna, dorsal cirri and branchial stems deep purple. Chaetae golden. Venter paler, midventral band barely distinct.

Prostomium blackish, with partial histolysis; anterior margin anteriorly entire, slightly eroded ( Fig. 55B ). Eyes blackish, anterior eyes well defined, posterior ones faded off. Median antenna inserted at anterior caruncular margin, as long as caruncle, 2× longer than lateral antennae. Lateral antennae bases separate from each other, 2× longer than palps. Mouth ventral on chaetiger 2. Pharynx not exposed.

Caruncle pale, trilobed, tapered, with partial histolysis, reaching chaetiger 4. Median ridge plicate, with about 16 vertical folds, almost completely concealing lateral lobes. Lateral lobes narrow, with about 12 vertical folds.

Bipinnate branchiae from chaetiger 4, continued throughout body, parallel in anterior and median region, divergent in posterior segments; becoming longer to chaetigers 9–10, smaller in posterior chaetigers, as long as or convergent; progressively smaller posteriorly, slightly longer than following chaetiger; median segments with 8–9 lateral branches per branchia.

Parapodia biramous, notopodia with cirriform branchiae along chaetigers 1–3, as long as dorsal cirri in chaetiger 1, progressively smaller in following chaetigers, dorsal cirri 2× longer than cirriform branchiae. Dorsal cirri as long as bipinnate branchiae along median chaetigers, 2–3× longer in posterior chaetigers. Second ventral cirri with cirrophores 2× longer and wider, and cirrostyle 2× longer than adjacent ones, directed dorsally. Other ventral cirri directed ventrolaterally, as long as one subsequent segment.

Chaetae most broken. Complete chaetae with distal fragile hoods, rarely eroded. Notochaetae in anterior chaetigers furcates ( Fig. 55C ), major tines 3–4× longer than minor ones. Median chaetigers with harpoon notochaetae with tiny denticles, looking smooth in low magnification ( Fig. 55E ), denticulatse tines 4–5× longer than smooth ones. Neurochaetae all furcates, major tines 4–6× longer than minor ones in anterior ( Fig. 55D ), and median chaetigers ( Fig. 55F ).

Posterior end tapered; pygidium with anus terminal ( Fig. 55G ); anal cirri pale, tapered, 3× longer than wide. Oocytes present in anterior chaetigers ( Fig. 55D ), each about 90 μm in diameter.

Live pigmentation (after Fig. 1E , and Poppe (2006) from the Philippines). Body pink; dorsum with inverted T middorsal brownish spots, transverse bands wider connecting branchial bases, longitudinal bars thinner, extended along all segment in chaetigers 4–7, following segments with anterior portion less defined. Median antenna, dorsal cirri and branchial stems deep purple to blackish. Chaetae golden to transparent.

Variation. Paralectotype (ZMA V.Pol 161.1) complete, better preserved; body fusiform, bent laterally, 10 mm long, 3.3 mm wide, 19 chaetigers. Dorsal pigmentation pattern faded off; dorsal cirri and branchial stems deep purple. Prostomium blackish. Eyes blackish, anterior eyes 3–4× larger than posterior ones. Median antenna almost as long as caruncle, almost 3× longer than lateral antennae. Lateral antennae 2× longer than palps. Mouth in chaetiger 2. Pharynx slightly exposed, basal incomplete rings, and lower pharynx smooth ring exposed. Bipinnate branchiae from chaetiger 4. Anal cirri pale, tapered, one broken, complete one 6× longer than wide.

A non-type specimen (ZMA V.Pol 2056) retains some pigmentation. Body fusiform, bent ventrally, 12 mm long, 2.8 mm wide, 19 chaetigers. Dorsum pinkish ( Fig. 56A ), venter cream. Dorsal spots better defined along posterior chaetigers, longitudinal bars wide, margins diffuse, better defined posteriorly, transverse bars almost reaching branchial bases, spots roughly resembling inverted T s along last two chaetigers ( Fig. 56C ). Purple bands along anterior notopodial surfaces better defined along last few chaetigers. Eyes blackish, anterior eyes 3–4× larger than posterior ones. Dorsal cirri and branchial stems deep purple. Ventral cirri of chaetiger 2 2–3× longer than adjacent ones ( Fig. 56B ). Chaetae transparent. Anal cirri whitish, tapered, 5× longer than wide.

Remarks. Chloeia violacea Horst, 1910 was described from Indonesia; because it has a complex pigmentation pattern, and bipinnate branchiae from chaetiger 4, decreasing in size posteriorly, it belongs in the group viridis. Further, because its middorsal spots are as an inverted T per segment, it resembles C. gilleti sp. n. described above with Western Africa specimens. These two species differ especially after the width of the middorsal band, and in the type of harpoon notochaetae. Thus, C. violacea has narrow middorsal band, and its harpoon chaetae have smooth tines, whereas C. gilleti has wider middorsal band, especially along median and posterior regions, and its harpoon chaetae do not have spurs or tines.

The specific name was based on that “each segment has in the middle of its dorsum a violet spot, shaped as an inverted T, the horizontal limb of which just corresponds to the posterior border of the segment” ( Horst 1910: 174).

Monro (1937) recorded C. violacea for the Gulf of Oman from sediments at 155 m depth (3 specimens London), and he noted that fresh specimens had violet spots, becoming orange after fading. He also noted the median antennae is longer than the caruncle, and 3× longer than lateral antennae. Horst (1910: 174) did not indicate the size proportion between the median and the lateral antennae by stating that “the paired antennae too are rather long”. However, in the non-type specimen referred to above (ZMA V.Pol 2056), these size relationships match those observed by Monro.

Distribution. Indonesia to the Gulf of Oman, in sediments at 18–155 m water depth.

Figures

FIGURE 1. Living specimens of four Philippine species of Chloeia Savigny in Lamarck, 1818. A. C. amphora Horst, 1910, dorsal view. B. C. bemisae sp. n., pink variant, dorsal view. C. Same, reddish and brownish variants, dorsal view. D. C. flava (Pallas, 1766), oblique right dorsal view. E. C. violacea Horst, 1910, oblique left dorsal view (scales not available; photos: Philippe Poppe; © Philippe & Guido Poppe – www.poppe-images.com).

FIGURE 8. Chloeia amoureuxi sp. n., epitoke (MNHN 860.1). A. Anterior region, dorsal view. B. Anterior end, dorsal view. C. Anterior end, right lateral view. D. Posterior region. Scale bars: A, 1.8 mm; B, 0.5 mm; C, 0.6 mm; D, 1.5 mm.

FIGURE 13. Chloeia bemisae sp. n. A. Paratype (UF 4356), anterior region, dorsal view. B. Anterior end, dorsal view. C. Anterior end, right lateral view. D. Non-type specimen (NTM W17191), living, dorsal view. Scale bars: A, 1.8 mm; B, 0.8 mm; C, 0.7 mm; D, 2.4 mm (Photo D: Karen L. Gowlett-Holmes).

FIGURE 14. Chloeia bistriata Grube, 1868, neotype (SMF 19403). A. Dorsal view. B. Anterior end, dorsal view, after Methyl green staining. C. Same, posterior region, dorsal view. D. Paraneotype (SMF 30268), anterior end, right lateral view. E. Same, chaetiger 3, notochaetae. F. Same, neurochaetal tips. G. Chaetiger 12, notochaetae (inset: section of denticulate tine). H. Same, neurochaetae. Scale bars: A, 2.5 mm; B, 0.6 mm; C, 0.9 mm; D, 0.5 mm; E, 80 μm; F, 40 μm; G, 100 μm; H, 110 μm.

FIGURE 46. Chloeia pulchella Baird, 1868, reinstated.A. Holotype of C. p. pallida Baird, 1868 (BMNH 1846.10.14.60), anterior end, dorsal view, after Methyl green staining. B. Chaetigers 17-20, dorsal view. C. Posterior region, dorsal view. D. Syntype of C. incerta de Quatrefages, 1866 (MNMH IA-TYPE 248), beheaded, anterior end, dorsal view. E. Chaetigers 16-19, dorsal view. F. Posterior region, dorsal view. Scale bars: A, E, 0.9 mm; B, 1.1 mm; C, 1.5 mm; D, 1.4 mm; F, 1.9 mm.

FIGURE 55. Chloeia violacea Horst, 1910, lectotype (ZMA V.Pol 161.3). A. Dorsal view. B. Anterior end, dorsal view. C. Chaetiger 3, notochaetae. D. Same, neurochaetal tips and oocyte. E. Chaetiger 13, notochaetae (inset: denticulate section). F. Same, neurochaetal tips. G. Posterior region, dorsal view. Scale bars: A, 2.2 mm; B, 0.7 mm; C, 200 μm; D, 40 μm E, 170 μm; F, 50 μm; G, 1.1 mm.

FIGURE 56. Chloeia violacea Horst, 1910, non-type specimen (ZMA V.Pol 2056). A. Anterior region, dorsal view. B. Same, right lateral view, after Methyl Green staining. C. Posterior region, dorsal view. Scale bars: A, 0.5 mm; B, 0.3 mm; C, 0.8 mm.