Laubieriopsis hartmanae ( Levenstein, 1970 )

Salazar-Vallejo, Sergio I., Zhadan, Anna E. & Rizzo, Alexandra E., 2019, Revision of Fauveliopsidae Hartman, 1971 (Annelida, Sedentaria), Zootaxa 4637 (1), pp. 1-67 : 51-56

Laubieriopsis hartmanae ( Levenstein, 1970) reinstated

Figures 31–34

Fauveliopsis hartmani Levenstein 1970: 229–234 , Figs. 2 –3 .

Fauveliopsis hartmanae: Levenstein 1972: 173 , Table 2, 175 Textfig. 1 (n. comb.).

Fauveliopsis glabra: Fauchald 1972b: 219–220 (partim, non Hartman in Hartman & Barnard, 1960).

Fauveliopsis brevis: Levenstein 1975: 133 (non Hartman, 1965).

Laubieriopsis hartmanae: Petersen 2000: 504 , Fig. 2 L–M (n. comb.).

Laubieriopsis brevis japonica Imajima, 2009: 125–127 , Fig. 4 (n. syn.).

TL: Not defined. NTL: Neotype locality: abyssal plain adjacent to the Kurile-Kamtchatka Trench.

D: Kurile-Kamchatka trench, Japan trench, to Western Mexico and Peru, in Atakamsky trench, 384–6835 m.

Neotype material. Northern Pacific , abyssal plain adjacent to the Kurile-Kamtchatka Trench. Neotype ( ZMMU WS11255 ), preserved in EtOH96, R/ V Sonne, Cruise 223 (KuramBio1), Sta. 11-9 (40°12.49’ – 40°12.37’ N, 148°05.40’ – 148°05.43’ E), 5263–5362 m, mud, epibenthic sledge, 29 Aug. 2012, I. Alalykina (det.) . Paraneotypes. Northern Pacific , abyssal plain adjacent to the Kurile Kamtchatka Trench, preserved in EtOH96, R/ V Sonne, Cruise 223 (KuramBio1), I. Alalykina (det.). 11 paraneotypes ( ZMMU WS11237 ), Sta. 11-9 (40°12.49– 40°12.37’ N, 148°05.40’ – 148°05.43’ E), 5263–5362 m, mud, epibenthic sledge, 29 Aug. 2012 . 16 paraneotypes ( ZMMU WS11240 ), Sta. 4-3 (46°58.34’– 46°58.46’ N, 154° 33.03’– 154° 33.39’ E), 5681-5780 m, mud, epibenthic sledge, 6 Aug. 2012

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Additional material. Eastern Pacific. Western Mexico. Nine specimens ( LACM 7493 ), off Natividad Island, R / V Velero IV, Sta. 7228 (27°37’17” N, 115°49’16” W to 27°36’35” N, 115°48’00” W), 4395– 3726 m, red and green clay, Menzies small dredge, 30 Dec. 1960 (6.7–11.8 mm long, 0.4–0.7 mm wide, 16 chaetigers; GP on posterior margin of chaetiger 6 or just before chaetae of chaetiger 7; anal papillae visible in some specimens, two dorsal, one ventral; oocytes 100 µm in chaetigers 4–8 in a macerated female) . 14 specimens ( LACM 7495 ), off Natividad Island , R / V Velero IV, Sta. 7229 (27°54’25” N, 115°40’00” W), 1751 m, green mud, piston corer, 31 Dec. 1960 (most broken; body 4.8–8.0 mm long, 0.3–0.4 mm wide, 16 chaetigers; GP on posterior margin of chaetiger 6; oocytes not seen) . Seven specimens ( LACM 4378 ), off Natividad Island , R / V Velero IV, Sta. 7231 (27°24’00” N, 115°12’15” W), 551 m (in label; 2480– 2401 m in Fauchald 1972b), green mud, piston corer, 1 Jan. 1961 (3 for SEM; body 7.5–9.8 mm long, 0.5–0.7 mm wide, 16 chaetigers; GP on posterior margin of chaetiger 6, just before chaetae of chaetiger 7; oocytes 100 µm, in chaetigers 5–7; portion of a fibrous tube) . Seven specimens ( LACM 7494 ), off Natividad Island , R / V Velero IV, Sta. 7249 (27°36’25” N, 115°56’25” W), 3752– 3709 m, red clay and rock, Menzies small dredge, 4 Jan. 1961 (3 for SEM; body 4.7–11.6 mm long, 0.3–0.6 mm wide, 16 chaetigers; GP paired, on posterior margin of chaetiger 6, just before chaetae of chaetiger 7; oocytes 100 µm, in chaetigers 5–7; pygidium fully everted in some specimens, with three anal papillae, two dorsal and one ventral) . One specimen ( LACM 7496 ), off Punta San Telmo, R / V Velero IV, Sta. 13744 (18°12.0’ N, 104°00.0’ W), 2297 m, mud, Campbell grab, 16 Jan. 1970 (6.5 mm long, 0.5 mm wide, 16 chaetigers; GP on posterior margin of chaetiger 6, just before chaetae of chaetiger 7; oocytes not seen, body wall opaque) . Colombia. One specimen ( LACM 7487 ), juvenile (as revealed by longer chaetae), without posterior region, R/V Vema, Sta. 15-38 (05°00’ N, 79°04’ W), 3021–3266 m, small bottom trawl, green to black mud, 15 Nov. 1958 . Seven specimens ( LACM 7486 ), four complete, R/V Vema, Sta. 15-65 (07°35’ S, 81°24’ W), 5817–5834 m, small bottom trawl, green to black mud, 6 Dec. 1958 (6.3–11.1 mm long, 0.4–0.9 mm wide, 16 chaetigers; GP halfway sometimes between chaetal lobes of chaetigers 6 and 7, other times closer to chaetal lobe of chaetiger 7; oocytes not seen) . Peru. Two specimens ( LACM 7490 ), partly driedout, W off Pacasmayo, R/V Vema, Sta. 15-75 (07°33.5’ S, 81°22.0’ W), 5894 m, small bottom trawl, green to black mud, 20 Dec. 1958 (7.7–9.0 mm long, 0.4 mm wide, 16 chaetigers; GP or oocytes not seen). Six specimens ( LACM 7489 ), two complete, WSW off Ciudad Edén, R/V Vema, Sta. 15-76 (07°31’ S, 81°27’ W), 5821– 5817 m, small bot- tom trawl, green to black mud, 20 Dec. 1958 (complete specimens 6.0– 9.1 mm long, 0.5 mm wide, 16 chaetigers; GP or oocytes not seen). Chile. Eight specimens ( LACM 7483 ), six complete, all with integument detached, corrugated, SW off Valparaíso, R/ V Vema, Sta. 17-2 (34°29’ S, 74°21’ W), 4030–4036 m, standard bottom trawl, 14 Mar. 1961 (chaetiger number difficult to assess because integument was detached; one has 16 chaetigers). One specimen ( LACM 7484 ), SSW off Concepción, R / V Vema, Sta. 17-7 (40°32’ S, 75°08’ W), 3272– 3081 m, standard bottom trawl, gray-brown clay, 22 Mar. 1961 (6 mm long, 0.8 mm wide, 16 chaetigers; GP paired, posterior margin of chaetiger 6, before chaetal lobes of chaetiger 7; oocytes in chaetigers 5–8, about 100 µm). Antarctic. Two specimens ( USNM 46829 ), U.S.C.G.C. Glacier, cruise 2, Sta. 22 (73°20.4’ S, 30°26.9’ W), 3111 m, 13 Mar. 1969, J.S. Rankin, coll. (8.3–10 mm long, 0.7–0.8 mm wide; GP transverse, fusiform, before chaetiger 7). 16 specimens ( USNM 46830 ), U.S.C.G.C. Glacier, Cruise 2, Weddell Sea, Sta. 23 (72°49.6’ S, 30°29.7’ W), 3697 m, 14 Mar. 1969, J.S. Rankin, coll. (4.5–13.5 mm long, 0.3–1.0 mm wide, 16 chaetigers; anterior end exposed in some specimens; prostomium without appendages; pharynx tubular, thick. Six specimens ( USNM 46831 ), U.S.C.G.C. Glacier, Cruise 2, Weddell Sea, Sta. 27 (64°46.2’ S, 41°30.1’ W), 4575 m, 19 Mar. 1969, J.S. Rankin, coll. (one strange specimen with caespitose or tufted aciculars; small specimens, 3.3–6.4 mm long, 0.3–0.5 mm wide, GP not seen).

Diagnosis. Laubieriopsis with 16 chaetigers ( Figs 31A , 32B , 34A ). First 4 chaetigers with two aciculars and two capillaries per bundle ( Figs 33A , 34B, C ). Median and posterior chaetigers with one acicular and one capillary per ramus ( Figs 33A , 34F, G ). GP on both sides of posterior margin of chaetiger 6, short, rounded, barely projected ( Figs 31 B-D, 32A-C, 33F, 34D, E). Pygidium with two middorsal small papillae ( Figs 33H , 34F ). Oocytes ( Figs 31A, D , 34F ) about 100 µm in diameter.

Description. Neotype (ZMMU WS11255) complete. Body subcylindrical, blunt anteriorly, truncated posteriorly, pale throughout ( Figs 31A , 34A ); 8.6 mm long, 0.55 mm wide, 16 chaetigers. Integument iridescent, smooth, transparent, inner organs visible ( Fig. 31D ).

Prostomium partly inverted, with round tip ( Figs 31B , 34B ).

Chaetigers 1–3 short, as long as wide, progressively longer; following chaetigers progressively longer than wide, posterior segments longer than wide. Chaetigers 1–3 with two slightly falcate aciculars and two slightly falcate capillaries per ramus ( Fig. 34C ). Interramal papillae globular, tiny, sessile.

Pygidium barely projected, with anus terminal, as a vertical slit separating it into two cushion-shaped lateral anal papillae ( Fig. 31E ).

Genital papillae paired, on both sides of posterior margin of chaetiger 6, each short rounded, surface granulose ( Figs 31C , 34D, E ). Oocytes 90–100 µm present in chaetigers 5 and 6 ( Figs 31D , 34H ).

Variation. There were 36 paraneotype specimens ( Figs 32 , 33 ). They were 6–13.5 mm long, 0.4–0.8 mm wide, all with 16 chaetigers ( Fig. 32A ). Genital papillae were paired, round to oval in shape, hardly visible in smaller specimens ( Figs 32B, C , 33F ). Interramal papillae more prominent in anterior segments ( Fig. 33D ), becomes smaller and round, ciliated in middle and posterior segments ( Fig. 33F, G ). Oocytes present in chaetigers 5 and 6; in males, testes present in chaetigers 5–8 ( Fig. 32A ). One specimen 12 mm long had a completely everted prostomium ( Fig. 32B, C ). Prostomium ovoid, fused with peristomium, with brown pigmentation on its whole dorsal surface, except an unpigmented distal spot; two darker spots were located on both sides of unpigmented area. Some specimens had 1–2 long pygidial papillae ( Fig. 33H ). Anterior aciculars are smooth, unidentate ( Fig. 33C ), in middle and posterior segments are also unidentate, but some have tips pilose or finely denticulate ( Fig. 33E ).

Remarks. Laubieriopsis hartmanae closely resembles L. brevis , as indicated in the key above. They differ, however, in subtle but significant features. In L. hartmanae , the genital papillae are globular, granulose and barely projected from body wall, and its ovaries extend along chaetigers 5 and 6, whereas in L. brevis genital papillae are oval, smooth, projected from body wall, and its ovaries extend along chaetigers 5–7 (or 8). These are small differences but the body features in the species provided with 16 chaetigers do not show marked variations, such that small differences must be used to separate similar species.

Laubieriopsis hartmanae was briefly described based upon 15 specimens, from four stations in Kurile-Kamchatka Trench and Japan Trench and one station in Peru-Chile (Atacama) Trench. These specimens were never deposited, and after being sent for evaluation, they were never returned such that they are lost.

Fauveliopsis hartmani Levenstein, 1970 , the original spelling, was modified to F. hartmanae ( Levenstein, 1970) . This species was later regarded as a junior synonym of L. brevis described from the Atlantic Ocean, first by Katzmann & Laubier (1974), despite the fact that one species was described from the Northern Pacific whereas the other was found in the Northern Atlantic. Further, the species was not recognized as distinct even by the original de- scriber ( Levenstein 1975:133), it was regarded as a junior synonym of L. brevis . Petersen (2000:514) made the most recent evaluation of the status of the species and had an ambiguous perspective. For example, she did not include it in her key to species ( Petersen 2000: 510), but in the same page indicated that she regarded it as a distinct species, and newly combined it into the current combination. However, she could not decide on the status of the Pacific species ( Petersen 2000:512), and later, as indicated above, recombined the name to Laubieriopsis hartmanae , probably on the basis of having found paired genital papillae and slightly bifid aciculars along anterior chaetigers, which she preferred to postpone until more specimens were available.

Levenstein reversed body ends in her description. Further, she thought there was a homonymy problem once Brada brevis Hartman, 1965 was regarded as belonging to Fauveliopsis by Hartman & Fauchald (1971:115), rendering Fauveliopsis brevis Hartman, 1967 a junior homonym. Hartman (1971:1422) renamed F. brevis Hartman, 1967 as F. brevipodus Hartman (1971) . Levenstein (1975:133) regarded her F. hartmanae as a junior synonym of F. brevis ( Hartman, 1965) , described from the Northwestern Atlantic. Petersen (2000:510) after the study of syntypes, concluded that L. hartmanae “differs from L. brevis in having paired GP.”

Consequently, it is clear that there is a need to clarify the taxonomic status of the Pacific species, and a neotype is herein proposed to solve this situation ( ICZN 1999, Art. 75.3.1). A redescription and illustrations which will be ‘sufficient to ensure recognition of the specimen designated have been included to emphasize its body features and its differences with its more similar species have been indicated above ( ICZN 1999, Arts 75.3.2, 75.3.3). We have asked our colleagues in the Shirshov Institute of Oceanology, Moscow, about the original specimens and they have indicated they are lost ( ICZN 1999, Art. 75.3.4). We are certain that the specimen herein designated as neotype matches, as far as possible, the original description, and because no other species has been found in the region, we are sure they are conspecific ( ICZN 1999, Art. 75.3.5). The neotype was found from a similar depth and environmental conditions of the original specimens, being collected in the same locality (Kurile-Kamchatka Trench) and similar depth: 5263–5362 m, in original description vs. 4090-6860 m) ( ICZN 1999, Art. 75.3.6). As indicated above, the neotype and all paraneotypes have been deposited in the scientific collections of Zoological Museum of Moscow State University, White Sea branch and Zoological Institute, Russian Academy of Sciences, Saint-Petersburg, which are recognized institutions with proper facilities for preserving type material and for making them available for future studies ( ICZN 1999, Art. 75.3.7).

Laubieriopsis brevis japonica is herein regarded as conspecific with L. hartmanae , because both were described from the same region in the Pacific with specimens collected from similar depths.

Figures

FIGURE 2. Fauveliopsis armata Fauchald & Hancock, 1981, paratypes (LACM 1161). A. Methyl green stained, dorsal view, stereoscope. B. Same, dorsal view, compound microscope. C. Same, right side of anterior end. D. Same, posterior end, dorsal view. Scale bars: A, B: 250 µm, C: 60 µm, D: 80 µm.

FIGURE 3. Fauveliopsis brattegardi Fauchald, 1972a, holotype (ZMUB 52205). A. Dorsal view. B. Oblique, dorsal view. C. Anterior end, oblique dorsal view. D. Posterior end, dorsal view. Scale bars: A, B: 160 µm, C: 45 µm, D: 65 µm.

FIGURE 4. Fauveliopsis brevipodus Hartman, 1971, n. comb., holotype (USNM 55546), A. Dorsal view. B. Anterior region, dorsal view. C. Posterior region, dorsal view. Paratypes (USNM 55547). D. Oblique dorsal view. E. Dorsal view. Scale bars: A: 260 µm, B: 100 µm, C: 90 µm, D, E: 0.4 mm.

FIGURE 31. Laubieriopsis hartmanae (Levenstein, 1970), neotype (WS11255). A. Complete, oblique right lateral view. B. Anterior region, dorsal view, Methylene blue staining (arrowheads indicate genital papillae). C. Chaetiger 7, genital papillae and notochaeta. D. Anterior region, dorsal view; oocytes seen through body wall (arrowheads indicate genital papillae). E. Posterior end, dorsal view. Scale bars: A: 0.6 mm, B: 200 μm, C: 25 μm, D: 150 μm, E 100 μm.

FIGURE 32. Laubieriopsis hartmanae (Levenstein, 1970), arrowheads indicate genital papillae. A. Complete paraneotype (WS11237), male, dorsal view (t: testis in chaetigers 5-8). B. Another paraneotype (WS11240), anterior region, dorsal view, prostomium and peristomium fully everted (p: prostomium, pe: peristomium). C. Same, ventral view, after Methylene blue staining. Scale bars: A: 0.5 mm, B: 0.3 mm, C: 0.2 mm.

FIGURE 33. Laubieriopsis hartmanae (Levenstein, 1970), paraneotypes (WS11237).A, B.Anterior ends. C. Anterior neuropodia (a: aciculars, c: capillaries). D. Interramal papilla of anterior chaetiger. E.Acicular of chaetiger 6. F. Genital papilla and parapodia of chaetiger 7 (ip: interramal papilla). G. Parapodia of posterior region (ip: interramal papilla). H. Posteriormost chaetiger and pygidium with papillae. Scale bars: A: 130 μm, B: 110 μm, C: 20 μm, D: 15 μm, E, F: 30 μm, G: 40 μm, H: 100 μm.

FIGURE 34. Laubieriopsis hartmanae (Levenstein, 1970), non-type specimens (LACM 7331-61). A. Complete, dorsal view, asterisk indicates enlarged area (inset: chaetiger 6 showing paired genital papillae). B. Another specimen, anterior end, right lateral view. C. Chaetiger 3, neurochaetae, posterior view. D. Left genital papilla. E. Right genital papilla. F. Posterior region. G. Same, neurochaetae. H. Another specimen, longitudinally sliced showing oocytes (80–100 µm). Scale bars: A: 1 mm (inset: 250 µm.), B: 110 µm, C: 13 µm, D: 70 µm, E 50 µm, F: 360 µm, G: 35 µm, H: 220 µm.