Pherusa papillata ( Johnson, 1901 )

Salazar-Vallejo, Sergio I., 2014, Revision of Pherusa Oken, 1807 (Polychaeta: Flabelligeridae), Zootaxa 3886 (1), pp. 1-61 : 33-36

Pherusa papillata ( Johnson, 1901)

Figure 12

Trophonia papillata Johnson, 1901:416 , Pl. 12, Figs 111, 123.— Moore 1908:356, 1909a:284.

Stylarioides papillata .— Moore 1923:221–222 (partim).— Hartman 1938:14.

Pherusa papillata .— Hartman 1961:120–121, Pl. 24, Figs 1–7 (syn.), 1969:303–304, Figs 1–6 .— Blake 2000:19–20, Fig. 1.8 .

Type material. Northeastern Pacific Ocean. Two syntypes ( MCZ 1900 ), Port Orchard, Puget Sound , Washington, 2 Jul. 1898, A. Robertson, coll.

Additional material. Northestern Pacific Ocean. Twenty specimens ( USNM 43821 ), 16 complete, San Juan Archipelago, Washington (no further data), M.H. Pettibone, coll. (complete ones 25–61 mm long, 2.5–6.0 mm wide, cephalic cage 5–8 mm long, 70–89 chaetigers; first neurohooks from chaetiger 4 (invariable); gonopodial lobes in chaetigers 5–6; 6–7 notochaetae per bundle, 5 in the smallest one; 10–14 transverse series of dorsal papillae (mean 12) per segment). One specimen ( USNM 1260294 ), SW off mouth of Columbia River (46°08.6' N, 124°13.9' W), Oregon, anchor dredge, 92 m, 21 May 1964, M.S. Alton, coll. (anterior end almost completely everted) .

Description. Largest syntype (MCZ 1900) cylindrical, pale, posteriorly tapered ( Fig. 12A ); 74 long, 4 mm wide, cephalic cage 7 mm long, 87 chaetigers. Body papillae with very little sediment, filiform, globose due to adherent sediment, completely covering the segments ( Fig. 12B, C ), about 7 rows per segment on chaetigers 2–3, then up to 12 rows per segment (up to 8 on anterior fragment).

Cephalic hood barely exposed, margin papillated. Prostomium low cone; eyes barely visible ( Fig. 12C ); caruncle reduced. Palps as long as branchiae; palp keels low. Lateral lips well developed; ventral and dorsal lips reduced.

Branchiae cirriform, distorted (observed in non-type specimen), arranged in two rows ( Fig. 12D ), all of the same length and width, resembling palps. Nephridial lobes not seen.

Cephalic cage chaetae as long as 1/12 body length, almost twice as long as body width. Chaetigers 1–3 forming cephalic cage; chaetae arranged in short series. Chaetigers 1–2 with longest chaetae, those in chaetiger 3 about twice as long as following ones, but about half as long as those in previous chaetigers. First notopodia with 12 chaetae, first neuropodia with 10 chaetae, following chaetigers with fewer chaetae.

Anterior dorsal margin of first chaetiger smooth (anterior fragment), or with a short lobe (complete specimen). Chaetigers 1–3 of similar length. Chaetal transition from cephalic cage to body chaetae abrupt; anchylosed, falcate blunt neurohooks start from chaetiger 4. Gonopodial lobes reduced to flat rounded areas, visible on chaetigers 5 and 6 (only in complete syntype).

Parapodia slightly developed; chaetae emerge from low transverse lobes. Parapodia lateral; medial neuropodia ventrolateral. Notopodia and neuropodia without long papillae.

Medial notochaetae arranged in transverse series; all multiarticulated capillaries, articles short basally, medium-sized medially, long distally, 6–8 per fascicle ( Fig. 12F , insets), as long as 1/5 body width. Neurochaetae multiarticulated capillaries in chaetigers 1–3; anchylosed, slightly falcate, reddish-brown neurohooks from chaetiger 4, arranged in transverse series ( Fig. 12E, F inset), generally 4 per fascicle, reduced to 3 in prepygidial chaetigers.

Posterior end tapered into a cone; pygidium with anus dorsoterminal, without anal cirri.

Remarks. Pherusa papillata ( Johnson, 1901) is a member of the group of species without sediment cover on their body papillae. This and the fact that their neurohooks start on chaetiger 4, means that P. papillata resembles P. hobsonae , P. moorei , P. mikacae , and P. rullieri n. sp. However, as both species have neurohooks falcate, and 10–20 transverse series of papillae per segment, P. papillata is most similar to P. mikacae ; these two species differ in the integument pigmentation, relative shape and number of transverse rows of papillae, and number of neurohooks in medial chaetigers. In P. papillata the integument is pale, dorsal papillae are filiform, globose, arranged in 10–12 transverse series per segment, and there are 3–4 neurohooks per bundle, whereas in P. mikacae integument is brownish, dorsal papillae are depressed and arranged in more series (18–20) per segment, and there are more (5) neurohooks per bundle.

Distribution. Northeastern Pacific Ocean, off British Columbia to California, in subtidal shallow depths (up to 92 m). The record by Monro (1933:1058) is questionable.

Pherusa rullieri n. sp.

Figure 13

Stylarioides plumosa .— Rullier, 1965:47 (non Müller, 1776).

Type material. Western Africa, Gulf of Guinea. Holotype ( MNHN 1556 ), Benin , Sta. 1 (06º10' N, 02º10' E), 55 m, greyish muddy sand, 8 Oct. 1963, A. Crosnier, coll.

Description. Holotype (MNHN 1556) soft, apparently preserved in ethanol or already rotting when fixed with formalin ( Fig. 13A ); body cylindrical, tapered posteriorly, broken into two parts, some medial segments missing. Anterior fragment 23 mm long, 6 mm wide, cephalic cage 12.5 mm long, 22 chaetigers; posterior fragment 20 mm long, 3 mm wide, 25 chaetigers. Body papillae larger over few anterior chaetigers, smaller throughout the body; most papillae depressed, about 20–26 transverse rows per segment.

Cephalic hood papillose, barely exposed ( Fig. 13B, C ); not dissected to avoid further damage. Cephalic cage chaetae twice as long as body width. Chaetigers 1–2 forming cephalic cage but second chaetiger with chaetae less than half as long as those present in chaetiger 1. Chaetiger 3 with chaetae half as long as those present in chaetiger 2, but twice as long as those present in following chaetigers. Chaetiger 1 with 18 noto- and 10 neurochaetae, arranged in a sinusoid series; chaetiger 2 with 11–12 noto- and 14 neurochaetae, arranged in a curved series.

Anterior dorsal margin of first chaetiger without large papillae, crossed by first notochaetae. Anterior chaetigers (2–4) with larger papillae arranged in 2–3 irregular, transverse rows. Chaetigers 1–3 decreasing progressively in length. Chaetal transition from cephalic cage to body chaetae abrupt; long, slightly curved, aristate neurohooks start in chaetiger 4. Gonopodial lobes not seen.

Parapodia poorly-developed, slightly larger along anterior region, chaetae emerge fom low lobes. Parapodia lateral, medial neuropodia ventrolateral. Notopodia and neuropodia, with 3–4 digitate, large interramal papillae.

Medial notochaetae arranged in transverse or oblique series; all multiarticulated capillaries, articles short basally, long articles medially, slightly decreasing in size towards distal region ( Fig. 13D–F ); 10–12 per fascicle, as long as ¼–1/3 body width Neurochaetae long, multiarticulated capillaries in chaetigers 1–3; anchylosed, falcate, tapered, aristate, light-brown neurohooks from chaetiger 4, arranged in transverse series, mostly 8 per fascicle throughout the body, aristae often broken ( Fig. 13E ).

Posterior end unknown.

Remarks. Pherusa rullieri n. sp. resembles those other species with body papillae without sediment cover and with neurohooks from chaetiger 4: P. hobsonae , P. mikacae , P. moorei , and P. papillata ( Johnson, 1901) . However, P. rullieri is most similar to P. moorei and to P. sibogae n. comb. by having medial and posterior neurohooks slightly curved, aristate. However, P. moorei has a cephalic cage 4 x longer than body width, whereas in P. rullieri and P. sibogae it is only twice as long as body width. These two latter species differ because in P. rullieri body papillae are abundant, without sediment particles whereas in P. sibogae body papillae are scarce and there are sand particles over papillae and tunic, although they do not form tubercles.

Etymology. This species is named after the late Dr François Rullier in recognition of his many publications on polychaetes, and especially because he reported this specimen in one of his contributions. The epithet is a noun in the genitive case.

Type locality. Benin, SE off Cotonou , 55 m depth .

Distribution. Only known from the type locality in the Gulf of Guinea.

Figures

FIGURE 1. Pherusa plumosa (Müller, 1776). A. Neotype (USNM 23257), complete specimen, dorsal view. B. Anterior end, dorsal view. C. Another specimen (USNM 8952), head, frontal view (branchiae and palps removed; BS: branchial scar, LL: lateral lip, PS: palp scar, VL: ventral lip). D. Neotype, anterior end, right side, ventral view. E. Same, chaetiger 9, notochaetae. F. Same, neurochaetae. Scale bars: A: 7.3 mm, B: 1.2 mm, C: 1 mm, D: 0.67 mm, E: 0.33 mm, F: 0.3 mm.

FIGURE 2. Pherusa affinis (Leidy, 1855), paraneotypes (USNM 1251742). A. Anterior end, dorsal view. B. Same, ventral view. C. Non-type specimen (USNM 40214), anterior end, oblique dorsal view. D. Same, head, frontal view (branchiae and palps removed; BS: branchial scar, LL: lateral lip, PS: palp scar). E. Another paraneotype (USNM1251742), chaetiger 10, right notopodium (insets: basal and medial chaetal regions). F. Same, chaetiger 21, right neuropodium (insets: neurochaetal tips). Scale bars: A: 0.80 mm, B: 0.76 mm, C: 1.2 mm, D: 0.70 mm, E: 110 µm, F: 100 µm.

FIGURE 3. Pherusa andersonorum n. sp. A. Holotype (LACM 2602), oblique lateral view. B. Same, anterior end, ventral view showing ventral pharyngeal organ. C. Paratype (LACM 2603), anterior end, dorsal view, methyl green staining. D. Same, head, partly everted (branchiae and palps removed; BS: branchial scar, NL: nephridial lobe, PS: palp scar). E. Another paratype with one palp and all branchiae preserved (Pa: palp, VPO: ventral pharyngeal organ). F. Another paratype, cross section of a medial segment. G. Same, chaetiger 5, right neurochaetae. H. Same, chaetiger 20, right neurochaetae. Scale bars: A: 2.7 mm, B: 0.9 mm, C: 1.1 mm, D: 0.3 mm, E: 0.7 mm, F: 0.25 mm, G: 50 µm, H: 46 µm.

FIGURE 4. Pherusa aspera (Stimpson, 1854). A. Neotype (USNM 8889), anterior region, dorsal view. B. Same, ventral view. C. Another specimen (USNM 8957), head, frontal view (branchiae and palps removed; BS: branchial scar, LL: lateral lip, PS: palp scar). D. Neotype, chaetiger 10, right notopodium (inset: chaetal tips). E. Same, chaetiger 9, neurochaetae (two smaller neurochaetae not shown). Scale bars: A, C: 0.7 mm, B: 0.9 mm, D: 225 µm, E: 112 µm.

FIGURE 5. Pherusa hobsonae n. sp. A. Holotype (USNM 43803), lateral view. B. Anterior end, dorsal view. C. Same, right side, ventral view. D. Paratype (USNM 1131565), anterior end, dorsal view, tunic removed. E. Same, head, frontal view (branchiae and palps removed; BS: branchial scar, NL: nephridial lobe, PS: palp scar). F. Paratype, chaetiger 12, right parapodium, notochaetae (inset: basal and distal regions). G. Same, chaetiger 12, right parapodium, neurochaetae (one not shown). Scale bars: A: 3 mm, B: 2.2 mm, C: 1.7 mm, D: 1 mm, E: 0.5 mm, F: 0.25 mm, G: 45 µm.

FIGURE 6. Pherusa incrustata de Quatrefages, 1866, reinstated. A. Syntype (MNHN 382), dorsal view (inset: posterior region of dried syntype with large dorsal sediment particles). B. Anterior end, dorsal view (body wall damaged by removal of sediment particles). C. Anterior end, ventral view. D. Non-type specimen (MNHN 507), anterior end, dorsal view. E. Another non-type specimen (MNHN 183), head, frontal view (branchiae and palps removed; BS: branchial scar, LL: lateral lip, PS: palp scar; *: nephridial lobes base). F. Syntype (MNHN 381), chaetiger 10, basal, medial and distal notochaetal regions. G. Same, chaetiger 10, neurochaetae (2 broken). Scale bars: A: 3 mm, B: 2 mm, C: 1.9 mm, D: 1.4 mm, E: 0.8 mm, F: 0.3 mm, G: 0.4 mm.

FIGURE 7. Pherusa mikacae n. sp. A. Holotype (NHMR 17740), anterior end, dorsal view. B. Paratype (NHMR 17741), anterior end exposed, ventral view (palps corrugated, branchiae smooth). C. Paratype (ECOSUR), anterior end, dorsal view. D. Same, ventral view. E. Same, anterior region, ventral view. F. Parapodium 12, notochaetae (inset: basal, medial and distal regions). G. Same, parapodium 12, neurochaetae with eroded tips (inset: paratype NHMR 17742, parapodium 10, tip of neurohook). Scale bars: A: 0.9 mm, B: 0.8 mm, C, E: 0.7 mm, D: 1.3 mm, F: 160 µm, G: 80 µm.

FIGURE 12. Pherusa papillata (Johnson, 1901). A. Syntypes (MCZ 1900), dorsal view. B. Same, anterior fragment, anterior end, dorsal view. C. Same, anterior end, ventral view (branchiae and palps exposed). D. Non-type specimen (USNM 43821), head, frontal view (branchiae and palps removed; BS: branchial scar, NL: nephridial lobe, PS: palp scar). E. Same, chaetiger 9, right parapodium, neurochaetae. F. Same, chaetiger 25, right parapodium, posterior view (insets: basal and distal notochaetal regions, largest neurohook). Scale bars: A: 3.3 mm, B: 1.5 mm, C: 0.9 mm, D: 0.4 mm, E: 0.1 mm, F: 0.3 mm.

FIGURE 13. Pherusa rullieri n. sp., holotype (MNHN 1556). A. Anterior end, dorsal view. B. Anterior end, ventral view. C. Same, close up. D. Chaetiger 11, right parapodium, anterior view (insets: basal and medial notochaetal regions, aristate neurohooks). E. Chaetiger 25, right parapodium, anterior view. F. Chaetiger from posterior fragment, right parapodium, anterior view. Scale bars: A: 1.6 mm, B: 1.2 mm, C: 1.0 mm, D: 0.25 mm, E: 0.1 mm, F: 0.2 mm.