Chloeia incerta de Quatrefages, 1866

Salazar-Vallejo, Sergio I., 2023, Revision of Chloeia Savigny in Lamarck, 1818 (Annelida, Amphinomidae), Zootaxa 5238 (1), pp. 1-134 : 70-74

Chloeia incerta de Quatrefages, 1866 reinstated, restricted

Figs 31 , 32

Chloeia incerta de Quatrefages, 1866: 388 ; Hartman 1959: 131; Solís-Weiss et al. 2004: S2; Salazar-Vallejo et al. 2014: 11 (list).

Chloeia parva Baird, 1868: 233–234 , Pl. 4, Fig. 8a, b ; Horst 1886: 167–168; Horst 1912: 19–20, Pl. 7, Fig. 4 , Pl. 8, Figs 1–3 ; Ehlers 1920: 15; Fauvel 1932: 56; Monro 1934: 355; Fauvel 1935: 293; Fauvel 1939: 273; Fauvel 1953: 96, Fig. 46f ; Gallardo 1968: 56, Pl. 10, Figs 8–12 ; Barroso & Paiva 2011: 422, Tab. 1; Wang et al. 2019: 5–8, Figs 3–5 , Tables 1, 2.

Chloeia merguinensis Beddard, 1889: 258–261 , Pl. 21, Figs 2 , 8 , 9 (Mergui Archipelago, Burma)

Type material. Indonesia, Celebes Sea, Sulawesi. Lectotype ( MNHN IA-TYPE 249 ), Leclancher, coll. (no further data), herein designated.

Additional material. Indonesia. One specimen ( SMF 1985 ), Aru & Kei Islands, Sungi Kololobo , 5 May 1908, H. Merton, coll. (bent ventrally; middorsal spots barely visible, better defined along posterior chaetigers; chaetae white; body 28 mm long, 7 mm wide, 30 chaetigers) . India. One specimen ( BMNH 1925.1.28.142 ), Madras, T.Y. Chen, coll., no further data (dorsal spots T-shaped; lateral bands thinner from the posterior segmental margin to anterior notopodial surfaces, wider along notopodia; most notochaetae of anterior region lost; body whitish, slightly bent laterally, 70 mm long, 15 mm wide, 37 chaetigers). One specimen ( MNHN 399.1 ), from Indian Museuum collection, Vizagaputaun , May 1926, Rao & Varugio, coll. (dorsal spots barely pigmented; median antenna purplish, without tip; body straight, 28 mm long, 7 mm wide, 29 chaetigers) . Singapore. One specimen ( ZMH V652 ), no depth or date data, G. Ŵlber, coll. (partially dried out; middorsal spots triangular, wider anteriorly; lateral bands paler, continued along anterior notopodial surfaces; dorsal cirri barely purple; branchiae colorless; venter cream; body sinuous, 60 mm long, 15 mm wide, 35 chaetigers) . China, Amoy . One specimen ( BMNH 1933.3.2.7 ), no further data (dorsal spots T-shaped; lateral bands thinner from the posterior segmental margin to anterior notopodial surfaces, wider along notopodia; most notochaetae of anterior region lost; body brownish, slightly bent laterally, 76 mm long, 16 mm wide, 38 chaetigers) . Hong Kong. One specimen ( ECOSUR), Tolo Harbor, site 3-9, trawling, 17 Aug. 2018 (straight, body ends bent ventrally, slightly dehydrated; body grayish, most notochaetae broken, neurochaetae pale; dorsal pigmentation as T-shaped bars, longitudinal one wider anteriorly; oblique divergent bands running along anterior parapodial surfaces; dorsal cirri blackish; caruncle with median ridge blackish; branchiae grayish with blackish stems; 88 mm long, 19 mm wide, 40 chaetigers). Two specimens ( ECOSUR), Tolo Harbor , June 2020 (no further data) (fixed in ethanol; smaller with body pale, bent ventrally; larger with body yellowish, straight; chaetae whitish; dorsal pigmentation as T-shaped bars, longitudinal one wider anteriorly; oblique divergent bands running along anterior parapodial surfaces; dorsal cirri blackish; caruncle with median ridge blackish; smaller with branchiae colorless; yellowish with blackish stems in larger one; 14–25 mm long, 4.5–7.0 mm wide, 28–29 chaetigers) . Papua New Guinea. One specimen ( UF 3998 ), Madang Province, Rempi Area, N. Tadwai Islet (-4.98, 145.79; 04°58´48.00″ S, 145°47´24.00″ E), 10–15 m, 20 Nov. 2012, J. F. Barazer, coll. (dorsal spots fading, lateral bands indistinct; some notochaetae with reddish bases; posterior dorsal cirri and branchiae reddish, other appendages pale; body bent laterally, 41 mm long, 5.5 mm wide, 32 chaetigers) .

Diagnosis. Chloeia with bipinnate branchiae from chaetiger 4, decreasing in size posteriorly; middorsal bands Y-shaped to triangular, lateral bands oblique; harpoon notochaetae without spurs; neurochaetae spurred and furcates.

Description. Lectotype (MNHN IA-TYPE 249) complete (Fug, 31A); body fusiform, 60 mm long, 19 mm wide, 36 chaetigers.

Lectotype brownish, chaetae yellowish; anterior prostomial margin brownish, lateral lips brownish. Dorsum brownish, from chaetiger 5 with a middorsal Y-shaped spot per segment ( Fig. 31B ); lateral bands visible from chaetiger 4, running along anterior notopodial surfaces, progressively paler posteriorly; dorsal cirri brownish; venter brownish.

Prostomium anteriorly entire. Eyes blackish, anterior eyes about 4× larger than posterior ones. Median antenna inserted at anterior caruncular margin, half as long as caruncle, slightly longer than lateral antennae. Lateral antennae bases separated from each other, bent laterally, slightly longer than palps. Mouth ventral on chaetiger 3. Pharynx not exposed.

Caruncle brownish, contracted, trilobed, tapered, reaching chaetiger 4 ( Fig. 31C ). Median ridge plicate, vertical folds not counted for avoiding further damage, concealing right lateral lobe. Lateral lobes narrow, with about 22 vertical folds.

Bipinnate branchiae from chaetiger 4, continued throughout body, parallel along most segments, divergent in a few posterior ones; largest by chaetiger 12, progressively smaller posteriorly. Median segments with 7–9 lateral branches per branchia.

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

Chaetae most complete. Complete chaetae with distal fragile hoods. Notochaetae in anterior chaetigers furcates and spurred ( Fig. 31D ), tines wide, major tines 4–12× longer than minor ones. Median chaetigers with one type of notochaetae: acicular harpoon-chaetae ( Fig. 31F ). Neurochaetae spurred, or furcates, major tines 3—4× longer than minor ones, blunt in anterior chaetigers ( Fig. 31E ), median chaetigers with sharper neurochaetae ( Fig. 31G ), major tines 4–6× longer than minor ones

Posterior end tapered ( Fig. 31H ); pygidium with anus terminal; anal cirri brownish, digitate, 4× longer than wide.

Variation. One-hundred years old specimens (BMNH 1925.1.28.142) with middorsal and oblique lateral bands continuous ( Fig. 32A ), with purplish dorsal cirri ( Fig. 32A, B ). Caruncle with a darker median ridge ( Fig. 32C ). Chaetae variably damaged, and hoods poorly defined in harpoon notochaetae ( Fig. 31D ), or neurochaetae ( Fig. 32E ). Other specimens (BMNH 1933.3.2.7) with continuous middorsal and oblique lateral bands ( Fig. 32F ), but bands thinner. Anal cirri digitate, whitish. A relaxed, ethanol preserved, 25 mm long specimen from Hong Kong (ECOSUR) shows middorsal spots are unconnected; the middorsal band is roughly triangular, progressively thinner along each segment, and the anterior transverse band is connected with the oblique, lateral bands running along anterior parapodial surfaces. Another larger specimen from the same locality (ECOSUR) shows middorsal, transverse and oblique lateral bands fused.

Live pigmentation (after Wang et al. 2019: 7). Dorsum with a middorsal band; each segment with band expanded anteriorly, decreasing posteriorly, resembling a solid Y, often with two lateral transverse bands continued through notopodial anterior surfaces, converging towards posterior segmental margin. Branchiae with pale stems, lateral branches pigmented. Dorsal cirri blackish. Chaetae whitish. There are some videos available in internet ( Mylife 2015, Krohn 2017, Healthy 2018a, 2018 b, Senja 2021, 1:10).

Remarks. Solís-Weiss et al. (2004: S2) gave the China Sea as the type locality for C. incerta de Quatrefages, 1866 , with the syntypes MNHN 246–252, but the type locality must be the set of localities for all the included syntypes ( ICZN 1999, Arts 73.2.3, 76.1). The syntype series includes one without locality (MNHN 246), three from the Bay of Bengal (MNHN 247, 248, 252), one from Sulawesi (MNHN 249), and another one from the “Mers de la Chine ” (MNHN 250). Further, the syntypes belong to four different species: C. flava (MNHN 247, 252), C. parva Baird, 1868 (MNHN 249), C. pulchella Baird, 1868 (MNHN 248 (beheaded), 250), and a very damaged specimen resembling C. tumida Baird, 1868 (MNHN 246).

As herein restricted, C. incerta de Quatrefages, 1866 matches C. parva Baird, 1868 . Because there is no type locality for C. parva , the type specimen is lost (E. Sherlock 2021 in litt.), and the species is apparently widespread, the syntype MNHN 249 is herein designated as lectotype of C. incerta . Further, C. incerta is a senior synonym of C. parva , both species defined by a particular pigmentation pattern along their dorsum, and the lectotype is designated for defining the status of the species name group ( ICZN 1999 Arts 71.3, 74.7.3, Recomm. 73C,1–6), and for providing a type locality for a rather well-known, widespread species, such that any future studies might clarify if more than one species is involved, and the type locality is designated after the lectotype. The remaining specimens are not regarded as paralectotypes because they are not conspecific, and consequently cannot be regarded as paralectotypes ( ICZN 1999 Art. 74.1.3). This designation includes a description and illustrations ( ICZN 1999 Art. 74.4)

de Quatrefages (1866: 389) was unable to differentiate between the syntype series and the other species, and hence the specific epithet. He noted, however, his new species was very similar to C. flava (Pallas, 1766) , but differed by the shape of the bifurcate neurochaetae by having tines of similar length. Because different species were involved, this difference must be reassessed by a comparison of some specimens of different localities previously identified as C. flava (see above).

Baird (1868: 230–231) concluded that C. incerta might be available for those specimens from the China Sea, and he proposed C. quatrefagessi for the same specimens. A careful study about type and topotype specimens will help solve this confusion and to clarify if more than one species has circular middorsal spots (see above). Chloeia parva Baird, 1868 was described in the same publication. The holotype was about 25 mm long, narrow and tapered in both ends, and the most remarkable feature was the presence of a series of dorsal T- or Y-shaped dots and lateral marks running “across each of the segments, and another encircling the ventral setiferous tubercle or foot. The branchiae are small, simply branched, and are of a dark colour.” Regretfully, the specimen did not have a locality indicated for it. Fauvel (1953) included it for the IndoPacific (incl. C. merguinensis Beddard, 1888 from the Mergui Archipelago).

Horst (1886: 167) noted for C. parva the median antennae were “not longer than once and a half the length fo the pair in front of it”. He also confirmed both the narrow body shape and the pigmentation pattern after he identified a 90 mm long specimen being 12 mm wide medially, with up to 36 segments; he also noted dorsal cirri were “blackish and extends beyond the bristle-tuft.” For the chaetae, Horst noted that notochaetae had “a horny-yellow shaft and a transparent, vitreous tip; there is a boundary visible between these both divisions, as if the tip was joined on the extremity of the shaft.”

Beddard (1888) described C. merguiensis with hesitation because it was very similar to C. parva including body shape, pigmentation pattern, and size of dorsal cirri. Beddard also indicated that branchial color resembled the dorsal body wall and “the main stem is slightly pigmented.” And regarding the size of branchiae, Beddard noted the first branchiae were rudimentary in comparison to subsequent ones. The only difference Beddard found was about the notochaetal pigmentation because his specimen did not show a difference between the shaft and the tip. Horst (1912) added that chaetae “have a yellow-red hue, especially those of the neuropodium.” The other illustrations available by Gallardo (1968) were of chaetae only without referring body shape, pigmentation pattern, size of dorsal cirri or chaetal pigmentation.

Wang et al. (2019) made a complete description of Hong Kong specimens of C. parva . Their specimens were 38–97 mm long, 8–20 mm wide, with 33–39 chaetigers, the pigmentation pattern matches the original description, but the body shape was indicated as fusiform, which might be size-dependent, dorsal cirri are not longer than notochaetae, and chaetae were shown as golden, not reddish. Further, branchiae are very short anteriorly, becoming larger posteriorly with a few long, tapered filaments.

Chloeia incerta de Quatrefages, 1866 has a complex pigmentation pattern, and bippinate branchiae from chaetiger 4, becoming progressively smaller towards its posterior region; these features indicate it belongs in the group viridis. Further, by having additional dorsal oblique lateral bands, it resembles C. viridis Schmarda, 1861 originally described from the Caribbean Sea, because both species have lateral expansions from their middorsal spots along anterior segment margins. The main differences rely on the development of the middorsal spots; thus, in C. incerta they are Y-shaped to subtriangular, progressively thinner along each segment, whereas in C. viridis , they are Tshaped with the middorsal band of similar width along each segment.

Distribution. India, Indonesia, Singapore, Papua New Guinea to China, in shallow water (10–15 m) sediments.

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 2. Chloeia viridis Schmarda, 1861, non-type specimens (BMNH 1931.6.22.16-18). A. Black form, anterior region, dorsal view. B. Same, chaetigers 10-12, dorsal view (branchiae removed from chaetiger 11). C. Same, posterior region, dorsal view. D. Red form, anterior region, dorsal view. E. Same, chaetigers 12-14, dorsal view (branchiae removed from chaetiger 13). F. Same, posterior region, dorsal view. Scale bars: A, F, 0.9 mm; B, E, 0.7 mm; C, 1 mm; D, 1.2 mm.

FIGURE 3. Chloeia mezianei sp. n., non-type specimen (NHMD 915074). A. Left lateral view. B. Anterior end and pharynx, seen from above. C. Same, right lateral view. Scale bars: A, 1 mm; B, 0.6 mm, C, 0.5 mm.

FIGURE 4. Chloeia fusca M’Intosh, 1885, non-type specimen (UF 4368). A. Ventral view. B. Ventral view after body wall removed. C. Same, after Methyl Green staining. D. Same, lower half of digestive tube removed. (AI, anterior intestine; MB, midventral band (nerve chord); MR, muscular ridge; Oe, oesophagus; PI, posterior intestine; R, rectum; St, stomach). All scale bars: 1 mm.

FIGURE 5. Chloeia bemisae sp. n., paratype, anterior fragment (UF 4388b). A) Chaetiger 3, right parapodium, posterior view, cirriform branchia cut (insets matched: 1, branchial cut tip; 2, branchial basal region; 3, dorsal cirrophore; 4, dorsal cirrostyle tip). B) Chaetiger 5, right parapodium, posterior view (insets matched: 1 and 2, branchial filament tip; 3, dorsal cirrostyle tip) (Br: branchia, DCs: dorsal cirrostyle). Scale bars: A, 0.20 mm; B: 0.38 mm.

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 9. Chloeia amphora Horst, 1910, lectotype (ZMA V.Pol 149.4). A. Oblique dorsal view. B. Anterior region, dorsal view. C. Chaetigers 10-12, dorsal view. D. Chaetiger 3, notochaetae. E. Same, neurochaetal tips. F. Chaetiger 12, notochaetal tips (inset: spur). G. Same, neurochaetal tips. Scale bars: A, 4.2 mm; B, 2.4 mm; C, 0.6 mm; D, 130 μm; E, G, 30 μm; F, 90 μm; H, 1.7 mm.

FIGURE 31. Chloeia incerta de Quatrefages, 1866, reinstated, lectotype (MNHN IA-TYPE 249). A. Dorsal view, depressed with a glass slide. B. Anterior region, dorsal view. C. Anterior end, dorsal view. D. Chaetiger 3, notochaetal tips. E. Same, neurochaetal tips. F. Chaetiger 16, notochaetal tips. G. Same, neurochaetal tips. H. Posterior region, dorsal view. Scale bars: A, 6.5 mm; B, 2.5 mm; C, 0.8 mm; D, G, 60 μm; E, 45 μm; F, 190 μm; H, 2.4 mm.

FIGURE 32. Chloeia incerta de Quatrefages, 1866, reinstated, non-type specimens. A. BMNH 1925.1.28.142, dorsal view. B. Same, anterior region, dorsal view. C. Same, anterior end, dorsal view. D. Chaetiger 21, harpoon notochaetal tips. E. Same, neurochaetal tips (inset: neurochaetal tip). F. Another specimen (BMNH 1933.3.2.7), dorsal view. Scale bars: A, 7 mm; B, 2.1 mm; C, 1 mm; D, 200 μm; E, 150 μm; F, 0.8 mm.

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.