Chloeia pseudeuglochis Augener, 1922

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 : 524-526

Chloeia pseudeuglochis Augener, 1922

Figs 1C , 11 , 12

Chloeia pseudeuglochis Augener, 1922: 39 .

Chloeia flava: Treadwell 1923: 2 (non ( Pallas, 1766)).

Chloeia euglochis: Treadwell 1928: 450 ; Treadwell 1941: 18 (non Ehlers, 1887).

Chloeia viridis: Monro 1928: 79–80 ; Monro 1933: 9–11, Text fig. 4; Berkeley & Berkeley 1939: 322–323; Hartman 1940: 205; Berkeley & Berkeley 1958: 399; Reish 1968: 73; Fauchald & Reimer 1975: 82; Fauchald 1977b: 11; Engle & Richards 2001: 188–189, Fig. 2A (non Schmarda, 1861).

Chloeia entypa: Hartman 1939: 8 (juvenile; non Chamberlin, 1919).

Chloeia conspicua: Fauvel 1943: 7–8 (non Horst, 1910).

? Chloeia rosea: Fauvel 1943: 7 (question mark in the publication; non Potts, 1909).

Type Material. Eastern Pacific. [ ZMH V-7466] Costa Rica, Puerto Culebra , Panalu (Bahía Panamá?), 16 Jul. 1902, R. Paessler, coll.

Additional material. Eastern Pacific, México. Nayarit, Playa Carreyeros (24°45’ N, 105°30’ W), intertidal rocks, Mar. 2008, P. Salazar, coll. [ LACM 516-36 ] Gulf of California, E San Francisco Island , 216–270 m, 25 Feb. 1936. [ LACM 661-37 ), Gulf of California,Agua Verde Bay, anchorage, night sampling, attracted to electric night, 10 Mar. 1937 (5). [ LACM 662-37 ], Gulf of California, Agua Verde Bay, off San Marcial Reef, 11 m, dredged, 11 Mar. 1937. [ LACM 7118 ], Gulf of California, 13 km SSW off Puerto Peñasco, 22–25 m, 4 Dec. 1970, J.R. Hendrickson & F. Mendoza-López, coll. [ LACM 10.3.1971), Gulf of California, Puerto Peñasco, 54 m, 10 Mar. 1971, R. Brusca, coll. (13). [ LACM 5.13.2000], California, Santa Catalina Island, W side Empire Landing, 13-20 m, 13 May 2000, J. Engle, coll. (3). [ MNHN A54.12 ), Gulf of California, no depth data, 1898, L. Diget, coll. [ UMAR 224 ] Oaxaca, Puerto Angel, Playa Panteón, 21 Jan. 2008, T.F. Villalobos-Guerrero, coll. [ UMAR 225 ] Oaxaca, Playa Panteón, 21 Apr. 2007, O. Valencia Méndez, coll. (5); [ UMAR 1501 ] Oaxaca, Puerto Angel, playa Panteón, Apr. 2005, Coll. G.R.C. [ UMAR 1502 ] Oaxac, no further locality data, May 2006, A.H.O. coll. [ UMAR 1503 ] Oaxaca, Puerto Angel, Playa Panteón, Feb. 2008, T. F. Villalobos-Guerrero, coll. [ USNM 15815 ] Baja California, Puertecitos, R / V Albatross, Sta. 3035 (30°21’N, 114°25’15”W), 55 m, Mar. 1889. [ USNM 15816 ] Baja California, San Jose Island, R / V Albatross, Sta. 2999 (24°54’30” N, 110°39’ W), 71 m, Mar. 1889. [ USNM 15817 ] Baja California, Los Angeles Bay, Sep. 1876. [ USNM 35061 ] Baja California, Ventana Bay, Stranger M /S, 9–18 m, Apr. 1937, W. Williams, coll. (6). [ USNM 35063 ] Baja California, San Lucas Bay, M /S, Stranger, Apr. 1937, F. Lewis, coll.

Diagnosis. Chloeia with ventral cirri of similar size throughout body; bipinnate branchiae from chaetiger 4; dorsum with three dark longitudinal bands separated by two paler bands; chaetal bundles often with a wide reddish band.

Description. Holotype (ZMH V-7466) complete; body elongate; dorsal pigmentation pattern visible but yellow banding in dorsum and along anterior branchial surfaces faded out; chaetal lobes with dark brown spot along anterior surfaces. Chaetae with subtle remains of transverse banding ( Fig. 11A, B ). Prostomium with eyes blackish, anterior eyes 2× larger than posterior ones. Caruncle twisted, crest with brownish longitudinal band, lateral lobes pale ( Fig. 11C ).

Best preserved non-type specimen (UMAR 224) complete, 95 mm long, 32 mm wide, 34 chaetigers. Live dorsal pigmentation pattern consists of three longitudinal broad orange-red bands (sometimes the middorsal one darker), and two narrow white yellowish bands around the median wider band; at the posterior margins of each segment these bands diminish, pigmentation interrupted, continued in the anterior margin of following segment ( Figs 1C ; 12A ). Venter salmon pink with one pair of oval spots per segment, progressively smaller posteriorly, not visible along last three segments ( Fig. 12B ). Antennae and dorsal cirri deep violet, caruncle orange-red, anterior part darker, branchiae red and chaetae with an orange band medially ( Fig. 12C ). After preservation, pigmentation less intense, yellowish bands included being pigments disappear, and contrasting pale bands appeared ( Fig. 12E ).

Prostomium oval with two pairs of eyes, anterior eyes slightly larger, darker. Median antenna inserted before caruncle, centrally on prostomium (2.7 mm long), longer than lateral antennae (1.5 mm). Palps located laterally on buccal lips (1.3 mm), resembling antennae, with anterior brown-violet pigmentation ( Fig. 12C, D ). Mouth ventral on chaetiger 3.

Caruncle trilobed, oval (3 mm long, 1.5 mm wide). Median lobe plicate with about 25 vertical folds. Lateral basal lobes narrow, each with similar number of folds ( Fig. 12C, D ). First appearance of bipinnate branchiae on chaetiger 4, present throughout body, last three smaller.

Parapodia biramous, in the first three chaetigers notopodia with cirriform branchiae.All first ventral cirri similar in size (2.5–3 mm). Chaetal lobes with a dark reddish spot along their anterior surfaces.

Pointed noto- and neurochaetae similar along anterior, median and posterior chaetigers ( Fig. 12F, H ). Furcate neurochaetae scarce; proportion between tines 1:3–1:4 ( Fig. 12G ). Notochaetae with tips serrated, resembling harpoon chaetae with few to several distal denticles (4–28, Fig. 12I ), abundant after a few anterior chaetigers.

Anus dorsal in last two chaetigers; anal cirri digitate.

Variation. The five specimens attracted to lights ( LACM 661-37 ) were 41–43 mm long, 10–13 mm wide, 31–33 chaetigers, showed pigmentation variably faded including chaetal banding. Only two specimens retain the main longitudinal reddish bands dorsally. Median antenna ⅔–½ as long as caruncle. Eyes black, separate; anterior eyes slightly larger than posteriors. Chaetae include furcates in both parapodial rami, each with small shortest tines, resembling a spur; chaetae do not include long capillaries, or longer chaetae than other specimens as could be expected for reproductive specimens. A larger specimen ( LACM 7118 ) 63 mm long, 14 mm wide, 34 chaetigers, has longitudinal reddish bands; chaetae include notochaetae with tiny spurs, while neurochaetal spurs, or shortest tines, are smaller in anterior chaetigers (1:4–1:6), than in medians (1:5–1:6). A larger specimen ( LACM 5.13.2000), 75 mm long, 16 mm wide, 32 chaetigers, has acicular notochaetae and neurochaetae, no trace of spurs. Chaetael variation in three specimens ( LACM 10.3.1971), 53–82 mm long, 11–12 mm wide, 32–34 chaetigers, shows that notochaetal furcates transform into aciculars by reduction of the smallest tine, becoming a spur, and then becoming not visible, that harpoon chaetae appear in chaetiger 4 in specimens 63 mm long (11 mm wide, 33 chaetigers), whereas neurochaetal furcates show a less marked reduction in the smallest tine. In conclusion, furcates become aciculars in largest specimens, but no modifications in eye size or in their relative proximity was found in specimens attracted to lights ( LACM 661-37 ), in comparison with other specimens from the same locality and collection date ( LACM 662-37 ). Hartman (1939: 8) recorded a small juvenile from Cocos Island, having only 12 chaetigers, and she noted the longitudinal bands, but no other details were provided .

Remarks. Chloeia pseudeuglochis Augener, 1922 was described from Pacific Costa Rica, but it was included in a paper dealing with Antillean polychaetes and was the only species included from the Pacific coast.. It seems that the title of the paper made some of our predecessors ignore its contents and this explains why the species name was not recorded again. The original description was very brief ( Augener 1922: 39) and only included a diagnosis (translated): “Pigmentation similar to (C.) euglochis , the chaetae with accordingly colored cross bandages. Differing from (C.) euglochis through the structure of the neurochaetae being tapered, or with one tip”. However, after its pigmentation pattern, some other similar species were recorded for the Eastern Pacific, and this explains why they are listed above.

The pigmentation was detailed by Monro (1933: 9–10) indicating that the progressively narrowing purplishbrown stripe, there were two longitudinal bands on each side, better defined along the posterior segmental area, and brown bands extended along parapodia. Monro (1933) characterized this species with Pacific Panamian material but identified them as C. viridis and regarded the Pacific specimen pattern as a color variant of the Atlantic C. viridis .

Treadwell (1941: 18) made another detailed characterization of the pigmentation pattern of C. pseudeuglochis , but he identified his specimens from Western Mexico to Panama as C. euglochis . Treadwell indicated his specimen 37,477 had, besides the pigmentation pattern noted by Monro, a series of slightly oblique yellow lines, which probably fade soon in ethanol. He also indicated chaetae were bright orange, and branchiae greenish-brown, with yellow stems.

Fauvel (1943: 8) recorded C. rosea Potts, 1909 after the pink pigmentation of one specimen.He noted notochaetae were very large and abundant, almost covering the back, but did not observe them. Chaetal features were noted by Berkeley & Berkeley (1939: 322–323; 1958: 399), indicating most chaetae were simple, not bifurcate, which rather matches C. pseudeuglochis .

Cortés (2012: 20) incorrectly indicated that the type of C. pseudeuglochis might be deposited in Copenhagen, and that it might be the same as C. pinnata . The type is in Hamburg, as shown by the label ( Fig. 11D ), and it was not collected by a Danish scientist but by a German Navy officer. The locality was indicated as Panalu but it probably referred to Bahía Panamá, in the Bahía (or Golfo de) Culebra, Costa Rica, as it was known by the end of the 1800’s ( Fradín 1892: 99), which belongs to the larger Golfo de Papagayo. It can be added that Augener (1912: 163) studied other specimens collected in Corinto, Nicaragua, by the same German officer, and named one species after him.

Chloeia pseudeuglochis differs from other congeneric species because of its pigmentation pattern. Further, C. pseudeuglochis shares the absence of furcate notochaetae with C. euglochis , but these two species differ. In C. pseudeuglochis there are no dorsal cross pigmentation patterns per segment, as in C. euglochis ; C. pseudeuglochis has only dorsal cirrophores violet, not the whole cirri as in C. euglochis , and it is found on the Atlantic side. Further, C. pseudoeuglochis has three red stripes, its dorsal cirri are completely violet and is a Pacific coast species. The first appearance of branchiae in chaetiger 4 is shared by several Chloeia species ( Table 1 ).

In the additional material [USNM 15817], J.E. Benedict added a note, in the early 1890s, indicating that this specimen corresponds to a new species; nevertheless, no publication followed.

Distribution. Gulf of California to Cocos and the Galapagos Islands, and Pacific Colombia; in sediments or mixed bottoms, common in shallow water, to 270 m depth. Occasionally recorded in Catalina Island, California, in 12-20 m water depth ( Engle & Richards 2001).

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 11. Chloeia pseudeuglochis Augener, 1922, holotype (ZMH V-7466). A. Dorsal view. B. Oblique, dorsal view. C. Anterior end, dorsal view. D. Labels. Scale bars: A, 0.5 mm; B, 0.4 mm; C, 0.7 mm (Photos: Alexandra Kerbl).

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).