Revision of Chloeia Savigny in Lamarck, 1818 (Annelida, Amphinomidae) Author Salazar-Vallejo, Sergio I. text Zootaxa 2023 2023-02-07 5238 1 1 134 http://dx.doi.org/10.11646/zootaxa.5238.1.1 journal article 53418 10.11646/zootaxa.5238.1.1 751096f2-4b5b-43c3-9748-4d07afe044c3 1175-5326 7621793 768E9932-2D18-4115-8359-3FF800328BCD Chloeia mezianei sp. n. urn:lsid:zoobank.org:act: 2A871517-83EF-4CB6-90A4-734E671D5357 Figs 3 , 39 , 40 Chloeia flava : Grube 1877: 509 ( Cape Vert only); Malaquin 1894: 415 ( non (Pallas, 1766)). Chloeia modesta : Fauvel 1913: 32 ( non Ehlers, 1887 ). Chloeia viridis: Monro 1930: 30 ; Fauvel & Rullier 1957: 68–71 , Fig. 2 ; Fauvel & Rullier 1959a: 507 ; Fauvel & Rullier 1959b: 155 ; Guy 1964: 178 , Figs 5 , 7 ; Rullier 1964: 144–145 ; Amoureux 1973a: 51 ; Hartmann-Schr̂der 1974: 96 ( non Schmarda, 1861 ). Type material . Gulf of Guinea . Holotype ( MNHN IA-TYPE 2052 ), RV Gérard Tréca , Sta. 56, Loos Archipelago , between Tamara (= Fotoba) Island (09°29´08″ N, 13°49´24″ W) and Coral Island ( 09°26´15.00″ N , 13°49´09.12″ W ), 12 m , 16 Mar. 1953 , J. Forest , coll. FIGURE 39 . Chloeia mezianei sp. n . , holotype (MNHN IA-TYPE 2052). A. Anterior region, dorsal view. B. Anterior end, dorsal view. C. Chaetigers 10-14, dorsal view. D. Chaetiger 3, notochaetal tips. E. Same, neurochaetae. F. Chaetiger 15, harpoon notochaetae. G. Same, neurochaetal tips. H. Posterior region, dorsal view. Scale bars: A, 2.4 mm; B, 0.6 mm; C, 0.8 mm; D, E, 30 μm; F, 130 μm; G, 50 μm; H, 1.4 mm. Additional material . Islamic Republic of Mauritania . Five specimens ( MNHN A54.4 ), juveniles, Mission Gruvel , 16 km SW off Nouakchott , 20 m , M. Gruvel , coll. (data used for variation) . Senegal . Four specimens ( MNHN A54.3 ), Mission Gruvel , Port , Dakar , May 1904 (specimens stiff, markedly bent ventrally; most notochaetae broken; middorsal spots visible, lateral ones barely visible in a few anterior chaetigers; bipinnate branchiae from chaetiger 4; body 26–40 mm long, 7–11 mm wide, 29–32 chaetigers). One specimen ( MNHN A414.1 ), no depth or date data, M. Parfait , coll. (bent ventrally; middorsal band visible; branchial stems pale, branches brownish; caruncle brownish, median ridge blackish; body 27 mm long, 8 mm wide, 30 chaetigers). One specimen ( ZMH V647 ), Gorée Island , Dakar , no depth or date data, H. Hupfer , coll. (brownish, bent ventrally, without posterior end; middorsal band and lateral bands visible throughout body; dorsal cirri blackish; branchial stems dark, branches pale; chaetae dirty yellowish; body 15.5 mm long, 5 mm wide, 21 chaetigers) . Gambia . One specimen ( NHMD 915070 ), RV Atlantide , Sta. 161 (no coordinates), off Bathurst , 18 m , 24 Apr. 1946 , J.B. Kirkegaard , coll. (colorless; body 9 mm long, 3.5 mm wide, 23 chaetigers). Cape Vert . One specimen ( MNHN 367.14 ), São Tiago Island , off Praia, RV Calypso, Sta. 14 ( 14°53´45″ N , 23°30´24″ W ), 25–30 m , 16 Nov. 1959 (anterior fragment; midventral band and lateral extensions visible, lateral bands faded off; dorsal cirri dark purple; branchiae with stems centrally pale, branches purplish; anterior eyes 2–3× larger than posterior ones; fragment 15 mm long, 6 mm wide, 16 chaetigers). Three specimens ( MNHN 367.70 ), São Tomé , RV Calypso, Sta. 70, Western coast, diving, 3–10 m , 16 Jun. 1956 (bent ventrally; middorsal bands visible along body, lateral bands visible along anterior body third; dorsal cirri dark purple; branchiae purplish; body 6–12 mm long, 2–3 mm wide, 16–21 chaetigers). One specimen ( MNHN 367.75 ), Boa Viata Island , RV Calypso, Sta. 75 ( 16°04´20″ N , 22°58´10″ W ), 45 m , 25 Nov. 1959 (middorsal band barely visible; lateral bands faded off; dorsal cirri dark purple; branchiae colorless; anterior eyes 3 larger than posterior ones; right parapodium of chaetiger 10 previously removed; body 11 mm long, 3.5 mm wide, 23 chaetigers). Three specimens ( NHMD 915066 ), RV Atlantide , Sta. unnumbered (no coordinates), Bay of Praia , 22 m , 13 Dec. 1945 , J.B. Kirkegaard , coll. (colorless; one bent ventrally; body 4–11 mm long, 1.5–3.5 mm wide, 14–20 chaetigers) . Ghana . One specimen ( BMNH 2013.100 ), postlarva, Tema Bay , without date or depth, W. People , coll. (colorless; branchiae from chaetiger 4, pinnate, non-bipinnate; mouth in chaetiger 1–2; notochaetae furcates, major tines 3× longer than minor ones; neurochaetae furcates, major tines 3–4× longer than minor ones, and aciculars; body fusiform, 3 mm long, 1.4 mm wide, 10 chaetigers). Eight specimens ( BMNH 2013.300 – 307 ), juveniles, Tema Bay , without date or depth, W. Pople , coll. (middorsal purple band visible in all specimens, lateral bands faded off; dorsal cirri dark purple; branchiae purplish; median antenna ½–4/5 as long as caruncle; anterior eyes 2× larger than posterior ones; bipinnate branchia from chaetiger 4; body 11–19 mm long, 2.5–5.0 mm wide, 20–25 chaetigers). Three specimens ( NHMD 915069 ), RV Atlantide , Sta. 75 ( 04°43´N , 01°41´W ), 46 m , 23 Jan. 1946 , J.B. Kirkegaard , coll. (colorless, bent ventrally; largest specimen with pharynx barely exposed and middorsal depression coincident with position of middorsal band; body 10–19 mm long, 4–5 mm wide, 25–28 chaetigers) . Guinea Bissau . Two specimens ( NHMD 915067 ), RV Atlantide , Sta. 44 ( 10°22´N , 16°22´W ), 49–55 m , 17 Dic. 1945 , J.B. Kirkegaard , coll. (almost colorless, bent ventrally; middorsal band visible along a few chaetigers in largest specimen; both with very long chaetae; anterior eyes slightly larger than posterior ones; body 8–12 mm long, 3–4 mm wide, 19–24 chaetigers). One specimen ( NHMD 915068 ), RV Atlantide , Sta. 44 ( 10°22´N , 16°22´W ), 49–55 m , 17 Dec. 1945 , J.B. Kirkegaard , coll. (bent ventrally, colorless, with a middorsal depression coincident with middorsal band position; body 18 mm long, 6 mm wide, 26 chaetigers) . Ivory Coast . One specimen ( SMF 14586 ), 1958–1961, A. Guy , coll. (soft, including chaetae; middorsal band barely visible along a few median chaetigers; body 43 mm long, 9 mm wide, 32 chaetigers). One specimen ( SMF 14559 ), Abidjan , Guy Station 80 (no coordinates), sand, 35 m , 1958–1961, A. Guy , coll. (body pinkish, soft, including chaetae; some anterior segments with a diffuse middorsal spot, resembling an inverted T, with all bars wide; chaetae of chaetigers 3 and 15 removed for observation; chaetae mostly acicular, some furcates with tiny spurs; harpoon-chaetae without denticles, likely dissolved; body soft, 40 mm long, 12 mm wide, 31 chaetigers) . Equatorial Guinea . One specimen ( BMNH 1930.10.8.413 ), off Pyramid Rock , Annobon , RSS Discovery , Sta. 283 (no coordinates), 18–30 m , 14 Ago. 1927 (data used for variation). One specimen ( MNHN 367.78 ), São Tomé & Principe Island , RV Calypso , Sta. 78 ( 00°25´15″ N , 06°43´05″ E ), 30 m , 21 Jun. 1956 (partial histolysis; middorsal band barely visible along a few anterior chaetigers; dorsal cirri dark purple; branchiae purplish; body 21 mm long, 4 mm wide, 23 chaetigers). Four specimens ( MNHN 367.94 ), RV Calypso , Sta. 94 ( 01°38´25″ N , 07°22´05″ E ), 31 m , 27 Jun. 1956 (three with middorsal band visible, other bands faded off; dorsal cirri dark purple; branchiae purplish; anterior eyes 2× larger than posterior ones; body 12–23 mm long, 3.5–5.0 mm wide, 20–28 chaetigers) . Democratic Republic of Congo . One specimen ( NHMD 915071 ), RV Galathea , Sta. 88 ( 06°24´S , 12°01´E ), 75 m , 8 Dec. 1950 , J.B. Kirkegaard , coll. (bent ventrally, colorless; body 11 mm long, 3 mm wide, 24 chaetigers). Four specimens ( MMSUCO Amp 5), Bay of Pointe-Noire , night lamp catch, 27 May 1964 , A. Stauch , coll. (data used for variation) . Angola . Five specimens ( NHMD 915072 ), RV Galathea Expedition , Sta. 102, Loanda , Harbor , 8–10 m , 15 Dec. 1946 , J.B. Kirkegaard , coll. (colorless; median antennae 4/5 as long as caruncle; 2–3× longer than median antennae; median antenna 2× longer than palps; branchiae from chaetiger 4; 21–50 mm long, 8–12 mm wide, 29–32 chaetigers). Two specimens ( NHMD 915073 ), RV Galathea Expedition , Sta. 121 ( 12°20´S , 18°40´E ), 27 m , 20 Dec. 1950 , J.B. Kirkegaard , coll. (middorsal band reddish to pale brown, visible along anterior third of body; dorsal cirri purple; body bent ventrally, 19–23 mm long, 5–6 mm wide, 25–27 chaetigers). One specimen ( NHMD 915074 ), RV Galathea Expedition , Sta. 122 (no coordinates), NW off Lobito Lighthouse, 20 m , 20 Dec. 1950 , J.B. Kirkegaard , coll. (juvenile, bent ventrally; right parapodium of chaetiger 14 previously removed; pharynx fully exposed; middorsal band visible along most chaetigers; anterior eyes 2 larger than posterior ones; branchiae from chaetiger 4; body 12 mm long, 2 mm wide, 23 chaetigers). Four specimens ( ZMH P13828 ), Lobito , sand, 0.5 m , 1 Jul. 1967 , G. Hartmann-Schr ̂der, coll. (colorless; most notochaetae broken, accumulated along a few anterior chaetigers; body 23–42 mm long, 6–7 mm wide, 27–30 chaetigers) . Diagnosis . Chloeia with bipinnate branchiae from chaetiger 4, progressively smaller posteriorly; median antenna as long as caruncle; anterior eyes 2–3× larger than posterior ones; middorsal band T-shaped, slightly tapered in median segments, lateral bands parallel; harpoon notochaetae without spurs; neurochaetae spurred and acicular. Description . Holotype ( MNHN IA-TYPE 2052), with body fusiform, bent ventrally, 33 mm long, 9 mm wide, 31 chaetigers. Holotype cream; anterior prostomial margin pale. Middorsal band visible along most segments, slightly wider anteriorly, tapered posteriorly on each segment; lateral marks paler, fused along posterior segmental margin, divergent anteriorly, extended laterally along notopodial anterior surface ( Fig. 39A ); another lateral band, more diffuse, connecting branchial and dorsal cirri bases. Dorsal cirri dark purple. Branchial stems purple along external surface slightly extended along branches bases ( Fig. 39C ), branches pale. Venter cream, midventral band paler, better defined along posterior body half. Prostomium anteriorly entire. Eyes blackish ( Fig. 39B ), anterior eyes 2× larger than posterior ones. Median antenna inserted at anterior caruncular margin, 2/3 as long as caruncle, 2× longer than lateral antennae. Lateral antennae bases close to each other, 2× longer than palps. Mouth ventral on chaetiger 2. Pharynx not exposed. Caruncle pale, sigmoid, trilobed, tapered, reaching chaetiger 5. Median ridge plicate, purple, with about 28 vertical folds, partially concealing lateral lobes. Lateral lobes narrow, with about 26 vertical folds. Bipinnate branchiae from chaetiger 4, parallel throughout body, progressively larger to chaetiger 13–14, smaller posteriorly; in median segments branchiae with 6–7 lateral branches. Parapodia biramous, notopodia with cirriform branchiae along chaetigers 1–3, progressively shorter than dorsal cirri. Dorsal cirri 2× longer than bipinnate branchiae along median chaetigers, 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 complete, hoods variably eroded. Notochaetae in anterior chaetigers spurred ( Fig. 39D ), major tines 32—38× longer than minor ones, or aciculars (without spurs). Median chaetigers with two types of notochaetae: aciculars and harpoon-chaetae without spurs or smooth tines ( Fig. 39F ). Neurochaetae spurred or aciculars, anterior chaetigers with major tines 9–18× longer than minor ones ( Fig. 39E ); in median chaetigers minor tines reduced, major tines 15—20× longer than minor ones ( Fig. 39G ) or acicular (without spurs). Posterior end tapered ( Fig. 39H ); pygidium with anus terminal; anal cirri whitish, subcylindrical, blunt, 3–4× longer than wide. Live pigmentation . Unknown; likely more intense than holotype pattern (previous records have confused a deep red, deeper water species, and this banded, shallow water one). Etymology . The species name is derived after Dr. Tarik Meziane, curator of Annelida in the Muséum National d’Histoire Naturelle, Paris, in recognition of his long-term support of my research activities, and of my former and current students. The specific epithet is a noun in the genitive case ( ICZN 1999 , Art. 31.1.2). Variation . Juveniles (MNHN A54.4) complete, all with longitudinal dark purple bands, one with lateral bands, paler, along a few anterior chaetigers ( Fig. 40A ); anterior eyes 2× larger than posterior ones ( Fig. 40B ); most with pharynx partially exposed; bipinnate branchiae from chaetiger 4; body 6.5–11.0 mm long, 2.8–3.5 mm wide, 18–22 chaetigers. Specimens caught with light trap (MMSUCO Amp 5) were complete, three bent ventrally, largest one almost without notochaetae; middorsal band brownish tapering posteriorly along each segment; anterior eyes 2–3× larger than posterior ones ( Fig. 40C ), lateral bands barely visible along a few anterior segments; bipinnate branchiae from chaetiger 4; body 20–50 mm long, 5.5–10.5 mm wide, 29–34 chaetigers. A larger specimen (BMNH 1930.10.8.413) was complete, straight; anal cirri lost; middorsal band purple brownish, better defined along anterior body third, lateral bands visible only in a few anterior chaetigers ( Fig. 40D ); anterior eyes 2× larger than posterior ones ( Fig. 40E ); bipinnate branchiae from chaetiger 4; body 32 mm long, 7 mm wide, 29 chaetigers; anal cirri lost ( Fig. 40F ). Remarks . Chloeia meziane sp. n. is described with specimens collected from Mauritania to Angola ; it has a complex pigmentation pattern, and bippinate branchiae from chaetiger 4, progressively smaller posteriorly, and because of these features it belongs in the group viridis. Because its middorsal bands are not homogeneous, and there are additional parallel bands, C. meziane resembles C. pseudeuglochis Augener, 1922 described from the Eastern Pacific and redescribed recently elsewhere (Yáñez-Rivera & Salazar-Vallejo 2o22). These two species differ in several features such as the relative size of eyes, the median antenna to caruncle, and the type of neurochaetae. In C. mezianei the anterior eyes are 2–3× larger than posterior ones, the median antenna is as long as caruncle, or slightly shorter than it, and the neurochaetae are acicular or spurred, whereas C. pseudeuglochis has anterior eyes slightly larger than posterior ones, median antenna half as long as caruncle, and its neurochaetae are acicular or furcates. FIGURE 40 . Chloeia mezianei sp. n . , non-type specimens. A. Juvenile (MNHN 54.4), dorsal view. B. Same, anterior end, oblique right lateral view. C. Another specimen (MMSUCO 5), anterior end, dorsal view. D. Another specimen (BMNH 1930.10.8.413), anterior region, dorsal view. E. Same, anterior end, oblique left lateral view. F. Same, posterior region, dorsal view. Scale bars: A, C, 1.5 mm; B, 0.5 mm; D, 1.7 mm; E, 0.8 mm; F, 1 mm. Some previous records from the Gulf of Guinea given as C. viridis or C. venusta might correspond with this species. Those records indicating a reddish pigmentation have been referred to above as C. gilleti sp. n. , whereas those indicating three blackish longitudinal bands are herein regarded as belonging to C. mezianei . Further, because the lateral bands fade off soon, and in some specimens all bands are faded out, identifications must be confirmed by chaetal features. As indicated above, C. mezianei and C. gilleti sp. n. described above from Western Africa, differ in chaetal features. Thus, C. mezianei has spurred notochaetae in anterior chaetigers, and acicular or spurred neurochaetae in median chaetigers, whereas in C. gilleti the anterior notochaetae are furcates, and neurochaetae of median segments are furcates with small minor tines. Malaquin (1894) reported specimens collected in 1890 by Chevreux in Dakar (Sta. 84, 88, 108). Monro (1930: 30) followed Augener (1925) about the synonymy of Western Atlantic species ( C. viridis Schmarda, 1861 and C. euglochis Ehlers, 1887 ). He even included C. parva Baird in the synonymy. Monro’s specimen from the Gulf of Guinea has dorsum with T-shaped markings with slender faded lines; 28 chaetigers and caruncle reaching chaetiger 6. Tebble (1955: 82) noted branchiae start in chaetiger 5, which differs from the Western Atlantic species. Kirkegaard (1983: 202) indicated that Guy (1964: 178) “separates the two species because of some differences in the colour and small differences in the bifurcate setae.” Amoureux (1973: 51) indicated he did not find furcate notochaetae. Distribution . Cape Vert, Mauritania , Senegal , Ghana , Guinea Bissau , Ivory Coast , Equatorial Guinea , Democratic Republic of Congo to Angola , in sediments from the intertidal to 75 m water depth.