Chloeia viridis Schmarda, 1861

Yáñez-Rivera, Beatriz & Salazar-Vallejo, Sergio I., 2022, Revision of Chloeia Savigny in Lamarck, 1818 from tropical American seas (Annelida, Amphinomidae), Zootaxa 5128 (4), pp. 503-537 : 526-530

Chloeia viridis Schmarda, 1861 restricted

Figs 1D , 5A, B , 13 , 14

Chloeia viridis Schmarda, 1861: 144–146 , Pl. 35, Textfigs 295–305; Augener 1925: 20–21; Hartman 1949: 37–38; Nonato & Luna 1970: 65, Figs 1 , 2 ; Amaral & Nonato 1994: 372–374, Figs 15, 16; Barroso & Paiva 2011: 422, Tab. 1 ; Humann et al. 2013: 85 (alive specimens).

Chloeia euglochis: Treadwell 1902: 194 (pigm.); Treadwell 1939: 176–177, Fig. 10 (non Ehlers, 1887).

Type material. Jamaica. Two syntypes ( NHMW 442 ), without field data .

Additional material. Western Atlantic. Gulf of Mexico. México, Campeche Bank IMCA IV, Sta. 58 (18°49’ N, 92°16’ W), Oct. 1989, 16 m, 26° C, 34‰, 1.250 OM, mud. Florida. [ USNM 4532 ] Pensacola (30°25’ N, 87°13’ W), S. Stearns, coll. (7) . [ USNM 16490 ], Miami , R/ V Eolis, 110 m, 1912, J.B. Henderson, coll. Georgia. [ USNM 19643 ], Savannah , Tompkins, I. R., coll. [ USNM 23926 ], Cumberland Island, Camden County, no further data. [ USNM 32385 ], Sapelo Sound , 11 m, 31 Jan. 1962, M. Gray, coll. Bahamas. [ USNM 24534 ] Bimini Island , W. Andrew, coll. [ USNM 24535 ], Bimini Island , W. Andrew, coll. Puerto Rico [ USNM 15919 ], Playa de Ponce, R/ V Fish Hawk, Sta. 3170 (18°13’ N, 66°32’ W), 1898 (2). [ USNM 41565 ], Aguadilla, Crashboat Basin , 12 m, 24 Jul. 1965, J. Randall, coll. [ USNM 42757 ] Puerto Rico, 23 m, 1963, N.C. Hulings & D.E. Feray, coll. (7) . [ USNM 52575 ] Puerto Rico, Magueyez Island, Boat Pier , 29 Aug. 1974, R.J. Larson, coll. Colombia [ UMML 22.913 ], R/ V Pillsbury, Cruise 6607, Sta. 365 (09°31’ N, 76°16’ W), 57 m, Jul. 1966, (3). [ USNM 98522 ], Cartagena, 25 m, J.J. Laverde-Castillo, coll.

Diagnosis. Chloeia with ventral cirri of similar size throughout body; bipinnate branchiae from chaetiger 4; dorsum with a dark longitudinal band, with an anterior expansion (T-shaped) in each segment; chaetal bundles homogeneously pale.

Description. Syntypes (NMW 442) complete, 39–43 mm long, 10–12 mm wide; pigmentation pattern visible, including purple dorsal cirri, and dark spots along anterior parapodial surfaces. Largest syntype ( Fig. 13A ) with all parapodia; middorsal longitudinal band better defined along anterior and posterior chaetigers; anterior end slightly bent ventrally, caruncle slightly twisted to the right ( Fig. 13B ). Smallest syntype without a right parapodium of an anterior chaetiger ( Fig. 13C ); middorsal longitudinal band visible throughout body; anterior end with caruncle slightly twisted to the left; prostomium visible with large, blackish eyes ( Fig. 13D ).

Best preserved specimen (USNM 42757) 9 mm long, 4 mm wide, 21 chaetigers,. Body fusiform with one longitudinal, middorsal violet band surrounded by a yellowish area, and violet dorsal cirri ( Fig. 14C, D ). Each longitudinal violet band expanded anteriorly in each segment, T-shaped; anterior part of each band sometimes hidden after body contraction.

Live dorsal pigmentation pattern on each segment includes two brownish or dark orange bands directed laterally, two brownish diverging bands, enlarged posteriorly, and a middorsal blackish band, Y- or T-shaped; branchiae pale, antennae, palps and dorsal cirri violet or blackish ( Fig. 14A ). Other specimens with a similar pattern but with middorsal band bluish, lateral bands transverse, olive green, and lateral bands dark purple or blackish; branchiae greenish, or paler along anterior branchial surface and darker, brownish to blackish along posterior branchial surface ( Fig. 1D ).

Prostomium semicircular with four black eyes, anterior ones 2× larger than posterior ones. Median antenna arising from anterior margin of caruncle (1 mm long), longer than lateral antennae (0.6 mm) and palps (0.4mm). Mouth ventral on chaetiger 3.

Caruncle tapered into a narrow posterior margin (1 mm long, 0.5 mm wide). Median lobe with about 11 folds. Roundish projections between each pair of folds along longitudinal mid-line ( Fig. 14B ), mid-line brown in live specimens. Lateral basal lobes narrow. Branchiae from chaetiger 4, continuing throughout body.

Parapodia biramous, notopodium with cirriform branchiae along chaetigers 1–3, as long as cirrophores; in chaetiger 4 dorsal cirri reduced, cirriform branchiae violet papilla continued to chaetigers 7 or 8. Second ventral cirri (1.1 mm long) about 2× longer than adjacent ones (0.4 to 0.5 mm).

Noto- and neurochaetae furcates in first chaetigers; proportion between tines 1:2–1:3 ( Fig. 14E ). Neurochaetae in median and posterior chaetigers include long and short furcates; shorter chaetae with a small, spur-like tine, proportion between tines 1:7 ( Fig. 14F ); longer chaetae with size proportion of tines about 1:3 ( Fig. 14G ). Notochaeta of posterior chaetigers include furcates and harpoon chaetae ( Fig. 14H–I ).

Juveniles. Specimens 10 mm long already have a middorsal band running throughout body ( Fig. 5A ), and the dorsal pattern starts along a few anterior or distal segments, and median segments have two posteriorly convergent bands ( Fig. 5B ); antennae and palps dark purple, and anterior surfaces of parapodia have a blackish hue; chaetae are homogeneously pale.

Remarks. Hartman (1938) overlooked Schmarda’s (1861) original description and illustrations ( Fig. 14C ), and based her identifications of C. viridis after Monro’s (1933) characterization. As indicated above, Monro followed Augener (1925) conclusion. Hartman, consequently, described the pigmentation pattern as consisting of three longitudinal dorsal bands (this pattern corresponds to C. pseudeuglochis ; see above). In addition, she emphasized the presence of violet cirri, which is shared by several species in the genus ( Table 1 ). As indicated above, C. viridis differs from C. euglochis even in small specimens (see figure 5C–F). In the original description, Schmarda (1861) emphasized that the greenish pigmentation in live specimens turned violet when preserved in alcohol. In fact, the white greenish pigmentation together with the orange-brown coloration are lost in preserved specimens, but the main violet middorsal band remains. The other reason for regarding C. viridis a juvenile was because the type was 29 mm long and would presumably turn into the colorful adult C. euglochis was rejected after finding 10 mm long specimens of C. euglochis having red-banded notochaetae and complex dorsal pigmentation. Additional evidence was provided by Nonato & Luna (1970) by describing larger specimens. They noted their material, being 28–52 mm long, had (translated) a middorsal wide (violet) band, expanded in the anterior margin of each segment, and the rest of the back was uniformly white-yellowish. Schmarda (1861) described and illustrated notochaeta from median chaetigers as serrated, while in anterior chaetigers the notochaetae and neurochaetae are furcate.

The type material is labeled as cotypes ( Fig. 12E ), and the term is equivalent to syntypes ( ICZN 1999, Art. 73.2.1). Further, the locality was indicated as Yamaika, but it was Jamaica, as indicated in the original description ( Schmarda 1861: 146). Some records do not have details about pigmentation, often because their specimens were small, and its specific affinities cannot be confirmed ( Rioja 1958; Day 1973; Gathof 1984).

On the other hand, a nomenclatural note is needed for clarifying the status of Chloeia candida Kinberg, 1857 , from the Virgin Islands, which could have priority over C. viridis , and of C. pallida Kinberg, 1867 , from northern Brazil.

Hartman (1949: 37) indicated that the type of C. candida is 13 mm long, with 22 chaetigers. The specimen is a juvenile which did not have any color at all, and it was described as having bipinnate branchiae from chaetiger 2, but Hartman noted them from chaetiger 4. In the original description ( Kinberg 1857: 11), no pigmentation was noted. The original description was repeated, and one plate was published later (Kinberg 1910: 33, Pl. 11, Fig. 2A ), and the whole specimen in dorsal view shows a hint of middorsal pigmentation, but because it was not indicated as a black longitudinal band, it might be the remains of the dorsal blood vessel. This feature, together with the fact that the chaetae do not show a reddish banding, has moved some authors to regard it as the same as C. viridis , including Hartman (1949: 38), and consequently C. candida would have priority over C. viridis . Regretfully, the smaller specimens Ehlers (1887) studied and informally named as C. modesta are of similar size as C. candida , being 12–16 mm long and with 22–25 chaetigers. These specimens also lack dorsal pigmentation pattern and their chaetae do not show the colorful bands typically seen in larger specimens. Consequently, on the basis of the information provided by the holotype, which is a juvenile, Chloeia candida Kinberg, 1857 must be regarded as indeterminable. A reversal of precedence, as indicated in the Code of Zoological Nomenclature ( ICZN 1999, Art. 23.9), cannot be applied because C. candida was recently used as a valid name ( Alós & Nüñez 2012: 53), and because they regarded it as the senior synomy of C. viridis and C. euglochis ( Parapar et al. 2012: 388) , they extended their distribution to the Eastern Pacific, Western and Eastern Atlantic ( Alós & Nüñez 2012: 53). We disagree with their conclusion and think that species have a more restricted distribution to the Western Atlantic. The Eastern Atlantic species belongs elsewhere.

On the other hand, Chloenea pallida Kinberg, 1867 is a nomen nudum because only the name was made available in the original publication ( ICZN 1999, Art. 12). Hartman (1949: 38) studied the holotype and noted it was another juvenile of a Chloeia species, having only 18 chaetigers. However, this does not make the name available.

Distribution. Caribbean region to northern Brazil, in shallow water.

Figures

FIGURE 1. Living specimens of Chloeia Savigny in Lamarck, 1818 from tropical American seas. A. Chloeia euglochis Ehlers, 1887, non-relaxed. B. Chloeia pinnata Moore, 1911, relaxed. C. Chloeia pseudeuglochis Augener, 1922, relaxed. D. Chloeia viridis Schmarda, 1861, non-relaxed (Photos: A, Paul Humann; B, Greg Rouse; C, Art Anker; D. Jean-Michel Sutour).

FIGURE 2. Chloeia pinnata Moore, 1911, non-type specimens (LACM 992-39), epitokes, anterior end after cropping chaetae of right parapodia of chaetigers 1–3, after Methyl Green staining for caruncle.A. Smallest specimen (11 mm long), dorsal view. B. Same, lateral view. C. Same, after Metyhl Green staining. D. Medium-sized specimen (18 mm long), dorsal view. E. Same, lateral view. F. Same, after Methyl Green staining. G. Largest specimen (23 mm long), dorsal view. H. Same, lateral view. I. Same, after Methyl green. Scale bars: A, 0.25 mm; B, 0.33 mm; C, 0.26 mm; D, G, 0.29; E, 0.44 mm; F, 0.35 mm.

FIGURE 5. Chloeia euglochis Ehlers, 1887, reinstated, and C. viridis Schmarda, 1861, restricted. A. Juvenile of C. viridis (UF 4308), dorsal view. B. Same, close-up of middorsal pigmentation pattern. C. Juvenile of C. euglochis (UF 4308a), dorsal view. D. Same, close-up of middorsal pigmentation pattern. E.Another juvenile of C. euglochis (UF 682), anterior region, dorsal view, most chaetae broken. F. Another juvenile of C. euglochis (UF 4350), greenish variant, anterior region, dorsal view. Scale bars: A, 1.3 mm; B, 0.5 mm, C, 1.4 mm; D, 0.7 mm; E, 1.5 mm; F, 1.2 mm.

FIGURE 10. Chloeia pinnata Moore, 1911, holotype (USNM 16842).A,Anteriorend, dorsal view. B. Segment 3. C.Prostomium and caruncle. D. Segments 3 and 4. E. Pigmentation pattern from median segments. F. Neurochaeta from first chaetiger. G. Long neurochaeta from chaetiger 10. H. Short neurochaeta from same chaetiger. I. Notochaeta from first chaetiger, insert shows the serrated inner border. J. Notochaeta from chaetiger 10. C. Paratype, all other holotype; B, C. White arrows show branchiae (Li: lips, LL: lateral lobes, MA: median antenna, ML: median lobe, Pa: palps, PA: paired antennae). Scale bars: A, C, D, 1 mm; B, E, 0.5 mm; F–H, 10 μm; I, 40 μm (inset 25 μm); J, 50 μm.

FIGURE 12. Chloeia pseudeuglochis Augener, 1922. A. Non-relaxed, living specimen from Oaxaca (UMAR 224), dorsal view. B. Same, ventral view. C. Anterior end, dorsal view. D. Prostomium and caruncle. E. Pigmentation pattern from median segments. F. Pointed neurochaeta from chaetiger 10. G. Furcate neurochaeta from same chaetiger. H. Pointed notochaeta from same chaetiger. I. Notochaeta from same chaetiger. A) Specimen from Gulf of California, B–H) holotype (Li: lips, LL: lateral lobes, MA: median antenna, ML: median lobe, Pa: palps, PA: paired antennae). Scale bars: A–C, 1 mm; D, 2 mm; E, F, 20 μm; G, H, 50 μm (Photos: A, B: Tulio Villalobos).

FIGURE 13. Chloeia viridis Schmarda, 1861, restricted, syntypes (NMW 442), most ethanol removed. A. Largest, complete syntype, dorsal view. B. Same, anterior region, dorsal view. C. Smallest complete syntype, dorsal view. D. Same, anterior region, dorsal view. E. Label. Scale bars: A, C, 5 mm; B, D, 1 mm (Photos: Pedro Frade).

FIGURE 14. Chloeia viridis Schmarda, 1861, restricted. A. Complete live specimen from Florida, dorsal view. B. Another specimen from Florida (USNM 4532), anterior end, dorsal view (LL: lateral (plicated basal) lobes, MA: median antenna, ML: median (plicate) lobe, PA: paired antennae). C. Illustration from Schmarda (1861). D. Specimen from Colombia (UMML 22.913), pigmentation pattern from median segments. E. Specimen from Puerto Rico (USNM 15919), neurochaeta from first chaetiger. F, G. Same, neurochaetae from median chaetigers. H, I. Same, notochaetae from median chaetigers. Scale bars: B, 1 mm; D, 2 mm; E, F, 20 μm; G, 50 μm; H, I, 10 μm (Photo A: Eric Vaandering).