Piromis capulata ( Moore, 1909 ) Moore, 1909

Salazar-Vallejo, Sergio I., 2011, Revision of Piromis Kinberg, 1867 and Pycnoderma Grube, 1877 (Polychaeta: Flabelligeridae), Zootaxa 2819, pp. 1-50 : 14-15

Piromis capulata ( Moore, 1909) n. comb.

Figure 4

Trophonia capulata Moore, 1909a:284 –286, Pl. 9, Figs. 60–61; Stasek, 1966:12; Loi, 1980:138.

Pherusa capulata: Hartman, 1969:295 –296, Figs. 1–2 ; Fauchald, 1972:414; Blake, 2000:1314, Fig. 1.5. Piromis roberti: Spies, 1975:188 , 191, 192, Pl. 3, Fig. 5 (non Hartman, 1951).

Piromis capulata: Spies, 1975:188 , 191, 192, Pl. 4, Figs. 12–13 , Pl. 5, Figs. 15–16 ; Pl. 7, Figs. 23, 25, 26 (informal n. comb.).

Type material. Eastern Pacific Ocean. Holotype (CAS-19721) collected in San Diego Bay, San Diego, California, intertidal, Dec. 1902, E.C. Starks, coll.

Additional material. Eastern Pacific Ocean. Two anterior fragments (LACM-AHF-2510), R/V Velero III, Sta. 1030 (27°39ʹ0 5ʺ N, 114°54ʹ47ʺ W to 27°39ʹ12ʺ N, 114°54ʹ12ʺ W), off Turtle Bay, Baja California, México, 26–31 fathoms, gray sand, mud, 18 Jan. 1940 (dorsal tubercles in chaetigers 2–6). One specimen (ECOSUR-7/ 1987), complete, Punta Las Rosas, Todos Santos Bay, Ensenada, Baja California, in eelgrass ( Phyllospadix scouleri Hooker ) root masses, low intertidal, 30 Jul. 1987, SISV, coll. (anterior end exposed, pale). One specimen (ECO- SUR-12/1981), complete, Bahia San Quintín, Baja California, 4 Dec. 1981., L.E. Calderon, coll. Five specimens (ECOSUR-15/1985), Laguna Ojo de Liebre, Baja California Sur, in Catarina scallops ( Argopecten circularis (Sowerby)) bottoms, 15 Apr. 1985, E. Baqueiro, coll. Five specimens ( USNM [unnumbered]), dried out, Catalina Harbor & Monterey (sic), 26-32-45C, 1874, H.H. Dall, coll. Fifteen anterior fragments ( USNM 40484), off Balboa, San Diego, California, Nov. 1932, G.E. McGinitie, coll. (paired dorsal lobes in chaetigers 2–8; size-dependent).

Description. Holotype (CAS-19721) complete, some parapodia previously removed, anterior end ventrally dissected; body cylindrical, long, tapering posteriorly ( Fig. 4 A); tunic papillated, with small sediment particles adhered over most of the body, especially dorsally, posterior region without sediment. Long capitate papillae arranged in longitudinal rows, two dorsally, four ventrally, over the first 5–6 chaetigers, forming low, eroded dorsal tubercles in the holotype ( Fig. 4 B, C); better preserved in other specimens ( Fig. 4 D), from chaetiger 2. Holotype 111 mm long, 4.5 mm wide, cephalic cage 9 mm long, 134 chaetigers.

Anterior end not exposed, hardly seen through the dissection cut; features based on a non-type specimen (ECOSUR). Prostomium dark, elevated; eyes large, black. Caruncle well developed, separating branchial filaments into two lateral groups ( Fig. 4 E); median keel pale, lateral ridges pale, with two black longitudinal bands. Palps large, corrugated; palp bases darker than surrounding areas. Dorsal and lateral lips thin, slightly darker than surrounding areas; ventral lip reduced. Branchiae cirriform, arising on a tongue-like protuberance, separated in two lateral groups, each group with filaments arranged in transversal rows, each row with 6–8 filaments, with about 50 filaments per group. Longest branchial filaments about as half as long as palps. Nephridial lobes not seen.

Cephalic cage chaetae less than 1/20 body length, or twice as long as body width. Chaetigers 1–3 involved in the cephalic cage; chaetae arranged in short rows, dorsolateral in chaetiger 1, lateral in chaetigers 2–3. Chaetiger 1 with 10 notochaetae and 8 neurochaetae, chaetigers 2–3 with 8 notochaetae and 6 neurochaetae per bundle. Anterior dorsal margin of chaetiger 1 with a median trifid lobe projected anteriorly (damaged). Anterior chaetigers with long papillae forming anteriorly directed, elongate conical lobes. Chaetigers 1–3 of similar length. Chaetal transition from cephalic cage to segmental chaetae abrupt; bidentate compound neurohooks from chaetiger 4, shorter from chaetiger 6. Gonopodial lobes present in chaetigers 5–6, larger in the former ( Fig. 4 F).

Parapodia poorly developed, except for few anterior ones (1–9); chaetae emerging from body wall. Parapodia lateral, median neuropodia ventrolateral. Notopodia better developed in chaetigers 1–9, with long postchaetal papillae making elongate conical lobes, decreasing posteriorly. Median noto- and neuropodia with two short, capitate papillae and four postchaetal longer ones.

Median notochaetae arranged in short transverse row, 7–8 per bundle, 1/3 as long as body width; all notochaetae multiarticulated capillaries with articles short basally, becoming long medially, then slightly shorter distally ( Fig. 4 G). Neurochaetae multiarticulated capillaries in chaetigers 1–3; long bidentate compound neurohooks in chaetigers 4–5 becoming shorter in chaetiger 6, arranged in a J-shape with eight per neuropodium ( Fig. 4 H), neurochaetae becoming thinner, longer in posterior chaetigers.

Posterior region almost with few sediment particles; pygidium conical, anus dorsoterminal, without cirri.

Remarks. Piromis capulata ( Moore, 1909) n. comb. resembles P. robertsi ( Hartman, 1951) from the Gulf of Mexico, and P. fauchaldi n. sp. from the Gulf of California, since all have notopodial lobes. They differ because in P. capulata there are dorsal tubercles in addition to the notopodial lobes, and there are only fine sediment particles on the tunic while the other two species lack dorsal tubercles and their tunics include larger sediment grains.

Further, there might be some confused records for the Northeastern Pacific Ocean. The records of P. ro b e r t i or P. capulata by Spies (1975), and of P. eruca by Hobson & Banse (1981), are possibly conspecific and might belong to an undescribed species, which has large dorsal lappets over several anterior chaetigers. Further, the records by Hartman (1969) and Blake (2000) of Pherusa capulata suggest that their materials had four pairs of branchiae and bifid neurohooks, which would not fit the specific features of P. capulata . On the other hand, the informal new combination that was introduced by Spies (1975:189, ff) was adequate and it is herein confirmed.

Distribution. From San Diego, California to Ojo de Liebre (Scammon’s) lagoon in Baja California, in intertidal and shallow sandy bottoms. The records of Piromis eruca for the western border between the United States and Canada ( Hobson & Banse 1981:59, Fig. 11 j–l) differ by having series of two foliose dorsal lobes instead of only the bare tip of larger papillae. They do not belong to the Mediterranean species, and might be an undescribed species. The same might apply to the record by Blake (2000:13), since his specimen was collected in 91 m depth, and there is no indication of the posterior neurohooks, which separate Piromis from Trophoniella .

Figures

FIGURE 1. Piromis arenosus Kinberg, 1867. A. Syntypes in lateral view. B. Thicker syntype, anterior end, dorsal view. C. Same, posterior end, lateral view. D. Non-type specimen (LAMNH- 5210), anterior end dissected, slightly collapsed, palps and most branchiae removed (BS: branchial scars, Ca: caruncle, DL: dorsal lip, LL: lateral lip, Pr: prostomium, PS: palp scar). E. Thicker syntype, notochaetae portions, basal, medial and distal with an insert of the tip. F. Non-type specimen (LAMNH-AHF- 2506), neurochaetae from a median chaetiger with an insert of one tip. G. Same, neurochaetae from a posterior chaetiger with an insert of a tip. Scale bars: A: 6 mm, B: 1.5 mm, C – D: 1 mm, E: 25 µm, F: 90 µm, G: 40 µm.

FIGURE 2. Piromis amoureuxi n. sp. A. Holotype (MNHN- 900), twisted. B. Same, anterior end in dorsal view. C. Same, median segments in lateral view, showing sediment particles dorsally and laterally. D. Paratype (MNHN- 900 a), anterior end, palps and most branchiae detached (BD: branchial scars, LL: lateral lip, PS: palp scar). E. Holotype, anterior notopodium. F. Same, close-up of notochaetal regions, left, median and distal. G. Same, neurochaetae from a median chaetiger. H. Same, neurochaetae from a posterior chaetiger. Scale bars: A: 1 mm, B – C: 0.5 mm, D: 0.3 mm, E: 100 µm, F: 10 µm, G: 50 µm, H: 25 µm.

FIGURE 4. Piromis capulata (Moore, 1909) n. comb. A. Holotype (CAS- 19721), complete in lateral view. B. Same, anterior end, lateral view. C. Same, anterior end, dorsal view, notice the eroded areas. D. Non-type specimen (USNM- 40484), anterior end, dorsal view. E. Same, anterior end, palps and most branchiae detached (BS: branchial scars, LL: lateral lip). F. Same, right neuropodium showing the gonopodial lobe with an asterisk in the base. G. Holotype, notochaetal regions, basal, median, distal. H. Same, median neuropodia with an insert showing the tips. Scale bars: A: 5 mm, B: 2.3 mm, C: 1.5 mm, D – E: 2 mm, F: 325 µm, G: 100 µm, H: 50 µm.

FIGURE 5. Piromis eruca (Claparède, 1868). A. Neotype (MNHN-A 507), complete in lateral view. B. Same, anterior end, lateral view. C. Same, anterior end, dorsal view. D. Paraneotype, head, palps and branchiae detached (BS: branchial scars, DL: dorsal lip, LL: lateral lip). E. Non-type specimen (MNHN-A 185), exposed anterior end, palps and branchiae detached (BS: branchial scars, Ca: caruncle, Pr: prostomium, PS: palp scar). F. Paraneotype, notochaetal regions, basal, median, distal and an enlargement of the tip. G. Same, neurochaetae from a median chaetiger, insert showing one tip. H. Same, neurochaetae from a posterior chaetiger, insert showing one tip. Scale bars: A: 3 mm, B: 1 mm, C: 1 mm, D – E: 0.5 mm, F: 50 µm, G: 200 µm, H: 25 µm.

FIGURE 11. Piromis websteri Day, 1973, st. n. A. Syntype (USNM 433), complete, lateral view. B – F: non-type specimen (USNM- 53883). B. Anterior end, dorsal view. C. Anterior end, ventral view, head exposed (Br: branchiae, CH: cephalic hood, DL: dorsal lip, LL: lateral lip, Pa: palp, PK: palp keel, VL: ventral lip). D. Notochaetae from chaetiger 20, showing basal, medial and distal regions. E. Neurochaetae from chaetiger 35, insert showing tips. G. Neurochaetae from chaetiger 125, insert showing tips. Scale bars: A: 2.5 mm, B – C: 0.5 mm, D: 70 µm, E: 150 µm, F: 50 µm.

FIGURE 12. Piromis wehei n. sp. A. Holotype (SMF- 15408) ,, complete, lateral view. B. Same, anterior end, dorsal view. C. Paratype, anterior end, ventral view. D. Same, head, frontal view; palps and branchiae removed (BS: branchial scars, LL: lateral lip, PS: palp scar). E. Holotype, notochaetae from chaetiger 19, basal, median and distal sections (insert: tip or notochaeta). F. Same, neurochaetae from chaetiger 10. G. Same, neurochaetae from chaetiger 71. Scale bars: A: 3 mm, B: 1 mm, C: 1.5 mm, D: 0.3 mm, E: 110 µm, F – G: 200 µm.

FIGURE 13. Pycnoderma congoense Grube, 1877. A. Syntype (ZMB 869), without posterior end, dorsal view; insert: anterior end, dorsal view. B. Syntype (ZMB 856), anterior end, dorsal view. C. Syntype (ZMB 4754), anterior end, lateral view. D. Nontype specimen (IRFA unnumb.), anterior end, dorsal view. E. Same, head exposed, slightly eroded, branchiae and palps removed (BS: branchial scars, Ca: caruncle, Pr: Prostomium, PS: palp scar). F. Syntype (ZMB 4754), parapodium from chaetiger 4. G. Same, neurochaetae from chaetiger 14. H. Syntype (ZMB 856), neurochaetae from chaetiger 37 (insert: chaetal tips). I. Syntype (ZMB 4754), neurochaetae from chaetiger 57. J. Non-type specimen (IRFA), neurochaetae from chaetiger 86 (insert: chaetal tips after pressing the cover slip). Scale bars: A: 5 mm, B – D: 1 mm, E: 0.5 mm, F: 300 µm, G: 225 µm, H: 90 µm, I: 180 µm, J: 75 µm.

FIGURE 15. Pycnoderma escobarae n. sp. Holotype (ECOSUR 0000). A. Lateral view. B. Same, anterior end, dorsolateral view. C. Same, terminal part, dorsal view. D. Parapodium from chaetiger 9, notochaetae detached. E. Parapodium of chaetiger 29. F. Same, notochaetae with long articles. G. Same, oligoarticulate neurohooks (insert: close-up of longest neurohook tip). Scale bars: A: 2 mm, B: 0.5 mm, C: 0.5 mm, D: 80 µm, E: 170 µm, F: 30 µm, G: 100 µm.

FIGURE 16. Pycnoderma ferruginea (Gallardo, 1968) n. comb. A. Holotype (LACM-AHF- 312), dorsal view. B. Same, anterior end, dorsal view. C. Same, close-up to show dorsally displaced chaetiger 1. D. Same, anterior end, lateral view. E. Nontype specimen (LACM-AHF- 2516), anterior end exposed, eroded, branchiae and palps lost (Ca: caruncle, BS: branchial scars, DL: dorsal lip, LL: lateral lip, PS: palp scar, VL: ventral lip). F. Holotype, parapodium from chaetiger 22 (insert: neurochaetal tips). Scale bars: A: 2.25 mm, B: 1.1 mm, C: 1 mm, D: 0.5 mm, E: 0.2 mm, F: 250 µm.