Pista nonatoi, Nogueira, João Miguel De Matos, Harris, Leslie, Hutchings, Pat & Fukuda, Marcelo Veronesi, 2011

Nogueira, João Miguel De Matos, Harris, Leslie, Hutchings, Pat & Fukuda, Marcelo Veronesi, 2011, Four terebellines (Polychaeta, Terebellidae) with problematic taxonomic histories, Zootaxa 2995, pp. 1-26 : 6-16

Pista nonatoi spec. nov.

Figures 3–6

Pista corrientis . Blankensteyn 1988: 51 –54, Fig. 10 ; Nogueira 2000: p. 180–183, Figs. 42–43; Alves 2008: 70 –74, Figs. 21B, 22–23.

? Pista corrientis . Hartman 1966: 97 –99, Figs. 4–9 .

Material examined. Brazil, State of Rio de Janeiro, offshore: 1 spec., coll. 2 Mar 1998, 21°51'S 40°07'W, 110 m; 23 specs, coll. 14 Feb 1998, 24°02'S 43°30'W, 147 m. Brazil, State of São Paulo, offshore: 1 spec., coll. 12 Jan 1998, 24°46'S 45°11'W, 99 m; 5 specs, coll. 16 Dec 1997, 25°11'S 47°08'W, 157 m. – Ubatuba, offshore: 8 specs, coll. 17 Mar 2001, 23°25'S 44°46'W, 35 m; 1 spec., coll. 10 Jun 2001, 23°25'S 44°46'W, 35 m; 1 spec., coll. 21 Mar 2002, 23°31'S 45°05'W, 8 m; 1 spec., coll. 23 Mar 2002, 23°32'S 45°05'W, 15.5 m; 1 spec., coll. 23 Mar 2002, 23°33'S 45°04'W, 23.4 m; 1 spec., coll. 20 May 2002, 23°48'S 45°10'W, 30.1 m. Ubatuba, Praia de Picinguaba, 23°22'S 44°50'W, in algae: 6 specs, coll. 8 Jun 2001; 1 spec., coll. 8 Oct 2001; 2 specs, coll. 18 Oct 2001. – Caraguatatuba, offshore: 1 spec., coll. 22 Apr 2001, 23°42'S 45°11'W, 25 m; 1 spec., coll. 27 Nov 2002, 23°43'S 45°18'W, 11 m; 1 spec., coll. 15 Feb 2001, 23°53'S 45°30'W, 25 m. Caraguatatuba, Praia de Martim de Sá, 23°37'S 45°22'W, in algae: 1 spec., coll. 16 Mar 2001. – São Sebastião, offshore: 2 specs, coll. 13 Feb 2001, 23°42'S 45°11'W, 25.2 m; 1 spec., coll. 27 Jun 2001, 23°46'S 45°10'W, 39.3 m. São Sebastião, Praia de São Francisco, 23°44'S 45°24'W, rocky shore, intertidal: 1 spec., coll. 24 Jul 2005. São Sebastião, Praia da Baleia, 23°46'S 45°39'W, rocky shore, intertidal: 1 spec., coll. 23 Jul 2005; 1 spec., coll. 17 Oct 2005. Guarujá, Ilha das Palmas, 24°00'S 46 ° 19'W, rocky shore, intertidal: 1 spec., coll. 6 Mar 2004; 1 spec., coll. 5 Oct 2005. Brazil, State of Paraná, Paranaguá Bay, 25 ° 33'S 48°25'W: 2 specs (MCEM-BPO 282), coll. 27 Feb 1986, Paranaguá Bay. Brazilian southeastern/southern continental shelf: 2 specs (MCEM-BPO 271), coll. 26 Mar 1984, 24°16'S 46°01'02''W, 45 m, sand; 1 spec. (MCEM-BPO 274), coll. 22 Mar 1984, 25°04'S 46°26'00''W, 65 m, muddy-sand; 1 spec. (MCEM- BPO 278), coll. 13 Nov 1985, 25°04'S 46°26'00''W, 65 m, muddy-sand; 1 spec. (MCEM-BPO 277), coll. 4 Nov 1985, 25°55'S 47°52'03''W, 49 m, muddy-sand; 1 spec. (AM W34750), coll. 3 Nov 1985, 26°22'S 48°19'08''W, 49 m, muddy-sand; 2 specs (MCEM-BPO 276), coll. 3 Nov 1985, 26°22'S 48°19'08''W, 49 m, muddy-sand; 2 specs (MCEM-BPO 273), coll. 17 Mar 1984, 26°22'S 48 ° 19'08''W, 49 m, muddy-sand.

Type series. Information on each specimen of the type series is provided in Table 1. Slides: Holotype ( MZUSP 00950): notochaetae from segments 5 and 17; neurochaetae from segments 8, 10, 11, 21, and 63.

Comparative material examined. Pista alonsae Santos, Nogueira, Fukuda & Christoffersen, 2010 . Holotype ( MZUSP 01036), paratypes 1 (CIPY-POLY-1396), 2 (CIPY-POLY-1397), 3 (CIPY-POLY-1398), and 4 ( LACM- AHF POLY 2263) coll. 9 Nov 2007, Brazil, Rio Grande do Norte, Macau, 05°04’S 36°27’W; slides: holotype: notochaetae from segments 8 and 17, neurochaetae from segments 5, 10, 11, 13, and region with neuropodia only; paratype 2: neurochaetae from segments 5, 10, 11, and 20. Paratype 5 (LACM-AHF POLY 2258) coll. 10 Nov 2007, Brazil, Rio Grande do Norte, Macau, 05°04’S 36°28’W. Paratypes 6 ( MZUSP 01037) and 7 (CIPY-POLY- 1399) coll. 9 Nov 2007, Brazil, Rio Grande do Norte, Macau, 05°04'S 36°27'W. Paratypes 8 ( MZUSP 01038) and 9 ( ZMUC Pol 2109) coll. 9 Nov 2007, Brazil, Rio Grande do Norte, Macau, 05°04’S 36°26’W.

Pista cetrata ( Ehlers, 1887, as Terebella cetrata ). Holotype ( MCZ 834), coll. USA, Florida, Key West, 2–4 m, Blake Expedition, by A. Agassiz, 1877–1878; incomplete spec., in relatively good state of preservation. Slides: notochaetae from segment 18; neurochaetae from segments 6, 13, 23, 61.

Pista corrientis McIntosh, 1885 . Holotype (BMNH 1885.12.1.348): coll. South Atlantic, Argentina, off the mouth of Rio de la Plata (37o17’S 53o52’W), in sea bottom with green sand, ~ 1100 m (600 fm), Challenger Expedition, 14 Feb 1876; complete spec., in poor state of preservation but most of important characters still visible; all notochaetae broken. Slides: neurochaetae from segments 9, 19, 23 and 83.

Pista cristata ( Müller, 1776) . Non-type material. ZMUC Pol 707: coll. North Atlantic Ocean, Denmark, off Laesø, W edge of Kummelbankens, 39–43 m, stones and mud, R/V Ophelia dredge 72, by C. Nielsen, 7 Jul 1960; 2 specs in relatively good state of preservation, all posteriorly incomplete; slides: notochaetae from segments 6 and 19; neurochaetae from segments 5–11, 14, and anterior segment of region with neuropodia only. ZMUC Pol 2054: coll. North Atlantic Ocean, Denmark, Frederikshavn, soft bottom, by H. Ditlevsen, 15 Jul 1927; 3 specs in relatively good state of preservation, all posteriorly incomplete and partially inside the tubes. ZMUC Pol 2055: coll. North Atlantic Ocean, Denmark, Laesø Rende, 30 m, Biological field course, 29 Jul 1947; 3 specs in poor state of preservation, 2 of them inside the tubes. ZMUC Pol 2056: coll. North Atlantic Ocean, Denmark, Frederikshavn, Biological field course, 1 Aug 1947; 6 specs in relatively good state of preservation, 4 of them partially inside the tubes.

Pista palmata ( Verrill, 1873, as Scionopsis palmata ). Non-type material. USNM 090608: coll. North Atlantic Ocean, Gulf of Mexico, USA, Texas, off Port Isabel (26o10’N 97o08’W), 15 m, BLM, Sta IV-4, STOCS, winter 1977; incomplete spec., in relatively poor state of preservation. USNM 090611: coll. North Atlantic Ocean, Gulf of Mexico, USA, Texas, off Port Isabel (26o10’00”N 97o08’00”W), 15 m, BLM, Sta S-53, IXTOC, Dec 1980; fragment in very poor state of preservation. USNM 090610: coll. North Atlantic Ocean, Gulf of Mexico, USA, Texas, off Port Isabel (26o10’N 97o08’W), 15 m, BLM, Sta IV-4, ves: STOCS, fall 1983; 2 specs, one complete and tiny, other incomplete, both in very poor state of preservation. Slides: spec. USNM 0 90610 (incomplete spec.): notochaetae from segments 6 and 18; neurochaetae from segments 5, 11, 27.

Pista sombreriana McIntosh, 1885 . Holotype (BMNH 1885. 12.1.344): coll. North Atlantic Ocean, Caribbean, off Sombrero and St. Thomas, Challenger Expedition; incomplete spec. in very poor state of preservation, in three pieces, anterior piece further cut in two halves along longitudinal midline. Slides: notochaetae from segment 14; neurochaetae from segments 6, posterior segment of region with biramous parapodia and anterior segment of region with neuropodia only (the state of preservation of the specimen precludes counting of segments).

Description. Thick mucous tube, with embedded sand, coarse gravel and fragments of shells ( Fig. 3 A). Complete specimens up to 61 mm long, 3 mm wide, up to 92 segments. Preserved body beige to light brown ( Fig. 3 A– K), live specimens frequently with darker pigmentation on lobes and around neuropodia. Body not dorsally inflated anteriorly ( Figs. 3 A–F; 4B–D, H–I); anterior segments compact, about same length, segments progressively longer on segments 8–20, elongation more evident from segment 11 ( Figs. 3 A–F; 4A–D, H–K). Ventral shields on segments 2–20, smooth, rectangular, progressively wider until segment 4, then progressively narrower until segment 8, about same width on segments 9–16, then progressively narrower again until segment 20; shields progressively longer on segments 6–16, about same length on segments 17–19, last shield distinctly shorter ( Figs. 3 A–B, I–J; 4A, J–K); after segment 20 shields replaced by mid-ventral groove extending posteriorly. Prostomium at base of upper lip; distal part forming shelf-like process from which buccal tentacles originate ( Figs. 3 D, F, J–K; 4D–I); no eyespots on basal part of prostomium. Peristomium restricted to lips; upper lip short, distinctly wider than high; lower lip short, cushion-like, partially covered by lobes on segment 1 ( Figs. 3 B, D, F, I–K; 4A, E–G, J–K). Segment 1 narrow, with one pair of large, distally rounded lobes originating dorso-laterally, extending anteriorly and ventrally, reaching beyond level of upper lip, connected to each other by lower mid-ventral membrane partially exposing lower lip ( Figs. 3 A–F, I–K; 4A–K). Segment 2 dorsally and laterally short, longer ventrally, with one pair of short and rounded ventro-lateral lobes, connected to each other by low membrane across ventrum ( Figs. 3 B–C, E, I–K; 4A–K). Segment 3 with one pair of large, distally rounded lobes originating dorsally to level of notopodia on segment 4, near dorsal mid-line, and terminating at level of lateral margin of ventral shields ( Figs. 3 A–F, I–K; 4A–K); anterior margin of segment 3 forming mid-dorsal crest ( Fig. 4 C, I). Segment 4 with short lateral lobes, as thin flaps, connected to each other by dorsal crest on anterior margin of segment ( Figs. 3 A–F, I–K; 4A–E, G–K); segments 5– 6 with short ventro-lateral lobes as low flaps between ventral shields and neuropodia ( Figs. 3 E, I–J; 4A–B, D–E, H, J–K). Two pairs of branchiae on segments 2–3, with long, irregularly annulated basal stem and secondary branches originating all at same level and about same length ( Figs. 3 B–F, I, K; 4B–I, K). Seventeen pairs of notopodia, starting from segment 4 and extending until segment 20, all about same size; notopodia of segments 4–7 inserted progressively more laterally, then vertically aligned ( Figs. 3 A–F, K; 4B–E, H–I, K). Broadly winged notochaetae on both tiers, wing distinctly broader on one margin, those on posterior tier with limbation starting from mid-length of chaetae ( Fig. 5 A, D); anterior notopodia with notochaetae of posterior tier about twice as long as those on anterior tier ( Fig. 5 A, D). Neuropodia starting from segment 5, as low rectangular ridges slightly raised from surface of body until segment on which notopodia terminate ( Figs. 3 A–F, I–K; 4B, D, H, K), as short pinnules on anterior part of region with neuropodia only ( Figs. 3 G; 4L), longer on posterior body segments ( Fig. 3 H); neuropodial pinnules situated laterally on body, well separated from mid-ventral groove, internal neuropodial shafts present ( Figs. 3 G– H; 5C, F). Long-handled uncini on segments 5–10, with thin handle originating from heel ( Fig. 5 E), short-handled thereafter ( Fig. 5 B–C, F–G); uncini with distally rounded prow, dorsal button situated at mid-length between base of main fang and tip of prow, and crest with ~4 rows of secondary teeth ( Figs. 5 E; 6A–B), on anterior neuropodia, 3–4 rows on segments 11–20, only visible under SEM ( Figs. 5 B; 6C); uncini arranged in completely intercalated double rows from segment 11 until segment on which notopodia terminate ( Figs. 5 B; 6C); from segment 21, uncini similar to those from anterior neuropodia, but short-handled and with inconspicuous dorsal button ( Figs. 5 C, F–G; 6D). One pair of large nephridial papillae on segment 3, inserted dorsally to bases of lobes, genital papillae situated posteriorly and dorsally to notopodia on segments 6–7 ( Figs. 3 C–F; 4C, H–I, K). Pygidium smooth.

Variation. Branchiae are easily lost and regenerated in this species. Frequently at least one of the branchiae is missing and they often differ in size within the same pair ( Fig. 3 B–F, K). Most specimens have the first pair of branchiae longer than those of the second pair, but some specimens have the second pair of branchiae longer, possibly due to regeneration of the first pair. Most specimens have branchiae as described above, with secondary branches originating together and about the same length, but paratype 2 has arborescent branchiae, with secondary branches originating at different points along the main stem and unequal in length ( Table 1).

Remarks. Santos et al. (2010) made an extensive comparison between Pista alonsae Santos, Nogueira, Fukuda & Christoffersen, 2010 and all other species of Pista previously reported for Brazilian and Caribbean waters. Therefore, to avoid repetitions, in this paper we include a comparative table for all those species of Pista discussed by Santos et al. (2010) ( Table 2 ), and only provide further discussion on the similarities and differences between P. nonatoi spec. nov., P. alonsae and P. corrientis .

Holotype Paratype 1 Paratype 2 Paratype 3 Paratype 4 Paratype 5 Paratype 6 Paratype 7 Paratype Paratype 9 Paratype MZUSP ZUEC- MZUSP USNM ZUEC- LACM- MZUSP ZUEC- 8 LACM- 10 USNM 0 0 950 POL 9606 0 0 951 1155161 POL 9607 AHF Poly 0 0 952 POL 9608 MZUSP AHF Poly 1155160

2695 0 0 953 2696

Collection Praia de Praia de Ilha das Pal- Offshore Uba- Offshore Offshore Offshore Praia de Praia de Ilha das Praia de data Martim de Picingu- mas tuba Ubatuba Ubatuba Ubatuba São Fran- Picingu- Palmas Picinguaba Sá (23°37'S aba (24°00'S (23º32'539''S (23º25'785'' (23º25' 785'' (23º48' 240' cisco aba (24°00'S (23°22' S 45°22'W), (23°22'S 46 ° 19'W), 45º05'409''W), S S 'S (23°44'S (23°22'S 46 ° 19'W), 44°50'W), Caraguata- 44°50'W), Santos, state state of São 44º46' 738'' 44º46' 738'' 45º10'124'' 45°24'W), 44°50'W), Guarujá, Ubatuba, tuba, state of Ubatuba, of São Paulo, fine W), state of W), state of W), state São Ubatuba, state of São state of São São Paulo, state of Paulo, inter- sand, 15.4 m, São Paulo, São Paulo, of São Sebastião, state of Paulo, Paulo, shalshallow sub- São Paulo, tidal, on 23 Mar 2002 medium medium Paulo, fine state of São São Paulo, intertidal, low subtidal, in shallow rocky shore, sand, 35 m, sand, 35 m, sand, 30 m, Paulo, shallow on rocky tidal, in algae, 16 subtidal, 0 5 Oct 2005 17 Mar 2001 17 Mar 20 May intertidal, subtidal, shore, 0 6 algae, 18 Mar 2001 in algae,0 8 2001 2002 on rocky in algae, Mar 2004 Oct 2001

Jun 2001 shore, 24 0 8 Jun

Jul 2005 2001

Size (mm) 38.5; 3/ 53 24.5; 2/ 41 11.2; 1/ 60 19.2; 2/ 40 13; 1.7/ 32 17.5; 1.8/ 11; 1/ 71 14.5; 1.5/ 14; 2/ ~38 25; 2.2 /46 11.5; 2.1/ (length; (incomp.) (incomp.) (incomp.) ~50 ~50 (incomp.) (incomp.) ~25 width/ (incomp.) (incomp.) (incomp.) number of

segments)

Branchiae

(left/right)

1st pair Missing; Missing; Very short; Missing; large Missing; Missing; Missing; Large; Large; Large; Missing; short missing short large large missing short large large large

Missing; miss-

2nd pair Missing; Short; large ing Large; short Large; Short; Large; Large; Short; large Large; Large; short short missing large large large short Additional Incomplete Incom- Complete Incomplete Incomplete Incomplete Complete Incomplete Incom- Incom- Incominforma- spec., in plete spec., spec. in spec., in good spec., in spec., in spec., in spec., in plete spec., plete spec., plete spec., tion good state of in good good state state of preser- good state of good state good state good state in good in good in good preservation state of of preserva- vation preservation of preserva- of preser- of preser- state of state of state of

preserva- tion, but tion vation, vation, pos- preserva- preserva- preserva-

tion smaller than possibly a sibly a tion tion tion

other specs. juvenile juvenile

Arborescent

branchiae,

with second-

ary branches

originating

at different

levels and of

different

lengths

Specimens of Pista nonatoi spec. nov., have previously been assigned to P. corrientis by several Brazilian researchers. That species differs from P. nonatoi spec. nov., in having short lobes on segment 1 that do not reach the level of the tip of upper lip; distinctly smaller lobes on segment 3 that originate at the level of the notopodia on segment 4 and terminate at the level of the ventral edge of the neuropodia, well separated from ventral shields; uncini throughout with 3 rows of secondary teeth; long-handled uncini on segments 5–20; and nephridial papillae on segment 3 inserted dorsal to the bases of the branchial stems. Pista nonatoi sp. nov., by contrast, has lobes on segment 1 that extend beyond the level of the upper lip and cover the anterior end; lobes on segment 3 that originate dorsal to the level of the notopodia on segment 4, near the dorsal mid-line, and terminate at the level of the lateral margin of the ventral shields (compare Fig. 1 A–C, F–L with Figs. 3 A–F, I–K; 4A–K); uncini with 3–4 rows of secondary teeth on segments 11–20 and ~4 rows on segments 5–10 and from 21 onwards; long-handled uncini present only on segments 5–10; and nephridial papillae on segment 3 inserted dorsal to the bases of the lobes on segment 3 and ventral to the bases of branchial stems.

Pista alonsae is the species most similar morphologically to Pista nonatoi spec. nov., differing from the latter species by its dorsolaterally indented lobes on segment 3; ventral shields present on segments 2–16; uncini throughout with 3–4 rows of secondary teeth; and long-handled uncini on segments 5–20. In contrast, P. nonatoi spec. nov., has ventral shields on segments 2–20, uncini on segments 11–20 with fewer rows of secondary teeth than on the remaining parts of the body, and long-handled uncini present only on segments 5–10.

P. cretacea coded after Fauvel 1927; P. fasciata , P. quadrilobata , Pista sp. A and B coded after Kritzler 1984; P. herpini after Fauvel

1928).

..... continued on the next page ..... continued on the next page It is uncertain if Hartman’s description of P. corrientis for specimens from Antarctica corresponds to P. nonatoi spec. nov. The author did not mention several characters currently considered important for the taxonomy of the group. For example, Hartman (1966) did not mention the presence of lobes on segments 2 and 4–6, nor were they illustrated in the figure of the anterior end in ventral view ( Hartman 1966: 98, Pl. XXXIII, fig. 4). In addition, she did not provide any information on the morphology of the lobes on segment 3, nor mention the number of secondary rows of teeth on the uncinial crest on each part of the body. Also, according to the figure cited above, the lobes on segment 1 do not reach beyond the level of the upper lip.

Etymology. This species is dedicated to Professor Edmundo Ferraz Nonato, from Instituto Oceanográfico, Universidade de São Paulo, who first identified this taxon in Brazilian waters, and in honor of his role as the foremost Brazilian polychaete researcher. Professor Nonato is the academic father, grandfather, and even great-grandfather of every Brazilian polychaete researcher working today.

Figures

FIGURE 1. Pista corrientis, holotype (BMNH 1885.12.1.348). A: anterior end, ventral view; B: anterior end, right lateral view; C: anterior end, left lateral view; D: anterior parapodia of region with neuropodia only; E: posterior end, left lateral view; F: detail of anterior end, dorsal view; G: detail of anterior end, left lateral view; H: detail of anterior end, ventral view; I: detail of anterior end, right lateral view; J: close up view of anterior end, dorsal view; K: close up view of anterior end, left lateral view; L: close up view of anterior end, ventral view. Numbers refer to segments; unspecified arrows show nephridial or genital papillae; ll = lower lip; P = prostomium; po = pharyngeal organ; ul = upper lip. Scale bars: A = 0.8 mm; B – C = 0.6 mm; D = 0.3 mm; E = 0.2 mm; F – L = 0.5 mm.

FIGURE 3. Pista nonatoi sp. nov., holotype (MZUSP 00950). A: worm inside the tube, anterior end, right lateral view; B: anterior end, right ventrolateral view; C: anterior end, right lateral view; D: anterior end, right dorsolateral view; E: detail of the anterior end, right lateral view; F: detail of the anterior end, right dorsolateral view; arrows point to genital papillae; G: neuropodia from anterior region with neuropodia only; H: neuropodia from posterior region with neuropodia only; I – J: close up views of anterior end, ventral view; K: close up view of anterior end, dorsal view. Numbers refer to segments; ll = lower lip; P = prostomium; ul = upper lip. Scale bars: A = 2 mm; B = 1.5 mm; C – F, I = 1 mm; G = 0.4 mm; H, J = 0.5 mm; K = 0.7 mm.

FIGURE 4. Pista nonatoi sp. nov., (spec. AM W 34750). A: anterior end, ventral view; B: anterior end, right dorsolateral view; C: anterior end, dorsal view; D: anterior end, right lateral view; E: anterior end, oral area and left lateral, superior view; F: oral area, superior view; G: close up view of oral area, ventral view; H: close up view of anterior end, left lateral view; I: close up view of anterior end, dorsal view; J: close up view of anterior end, ventral view; K: close up view of anterior end, left ventrolateral view; L: posterior segments, left lateral view. Numbers refer to segments; unspecified arrows point to genital papillae; ll = lower lip; P = prostomium; ul = upper lip; arrows point to nephridial and genital papillae. Scale bars: A – B, D, G – J = 0.4 mm; C, E – F, K, L = 0.3 mm.

FIGURE 5. Pista nonatoi sp. nov., holotype (MZUSP 00950). A: notochaetae, segment 5; B: uncini, segment 11; C: uncini, segment 21; D: close up view of some notochaetae from segment 5; E: uncini, segment 8; F: uncini, segment 63; G: uncinus, segment 21. Scale bars: A = 80 µm; B = 30 µm; C, E – F = 20 µm; D = 40 µm; G = 10 µm.

FIGURE 6. Pista nonatoi sp. nov., (spec. AM W 34750). A: uncini, segment 5; B: uncini, segment 6; C: uncini, segment 13; D: uncini from region with neuropodia only. Scale bars: A – B, D = 6 µm; C = 10 µm.

FIGURE 7. Pista sombreriana, holotype (BMNH 1885.12.1.344). A: entire worm, dorsal view; B; entire worm, ventral view; C: notochaetae from anterior tier, segment 14; D: uncini, segment 6; E: uncini from posterior region with biramous parapodia; F: uncini from anterior region with neuropodia only. Scale bars: A – B = 1.5 mm; C = 50 µm; D – E = 20 µm; F = 15 µm.

FIGURE 8. Pista sombreriana, holotype (BMNH 1885.12.1.344). A: anterior end, left lateral view; B: anterior end, right lateral view; C: anterior end, dorsal view; D: anterior end, ventral view; E: close up view of anterior end, left lateral view; arrows point to genital papillae; F: close up view of anterior end, right lateral view. Numbers refer to segments; ll = lower lip; P = prostomium; ul = upper lip. Scale bars: A – B = 0.8 mm; C – D = 1 mm; E – F = 0.4 mm.

FIGURE 9. Eupolymnia turgidula, holotype (MCZ 846). A: Ehlers’ drawing of the holotype, entire worm, ventral view (Ehlers 1887: Taf. 52, Fig. 1); B: entire worm, dorsal view; C: entire worm, left ventrolateral view; D: anterior end, ventral view; E – F: close up views of anterior end, left ventrolateral view; G: anterior end, left lateral view; H: anterior end, dorsal view; I: transition between regions with biramous parapodia and neuropodia only, right lateral view; J: close up view of anterior segments of region with neuropodia only, right lateral view; K: piece of tube. Numbers refer to segments, uspecified arrows point to nephridial papillae; * = lower lip; ** = prostomium, basal part; P = prostomium; P (dp) = prostomium, distal part; ul = upper lip. Scale bars: B – C = 1.5 mm; D – I = 1 mm; J = 0.6 mm; K = 2 mm.

FIGURE 10. Eupolymnia turgidula, holotype (MCZ 846). A: notochaetae, segment 18; B: tips of some notochaetae from posterior tier, segment 18; C: uncini, segment 7; D – E: uncini, segment 12; F: uncini, segment 23. Scale bars: A, E = 80 µm; B = 40 µm; C = 15 µm; D = 10 µm; F = 30 µm.

Tables

TABLE 2: Comparison between the species of Pista reported for the Atlantic coast of South America and the Caribbean (characters of

	Pista alonsae Santos, Nogueira, Fukuda & Christoffersen, 2010	Pista cetrata (Ehlers, 1887)	Pista corrientis McIntosh, 1885	Pista cretacea (Grube, 1860)
Type-locality	Macau, Rio Grande do Norte, northeastern Brazil, Atlantic Ocean	Southern Florida, Caribbean, Atlantic Ocean	Off mouth of La Plata River, Argentina, Atlantic Ocean	Mediterranean, Atlantic Ocean
Size (length x width [mm]; number of segments)	77 x 2.2; ~106 (incomplete)	25 x 2; 64 (incomplete)	27 x 1.5; 93	150–250 x 3–6; 120–180
Lobes on segment 1	Large, distally rounded, directed anteriorly and ventrally, connected to each other by membrane across ventrum partially exposing lower lip; distal tip reaching beyond level of tip of upper lip	Large, distally rounded, directed anteriorly and ventrally, connected to each other by membrane across ventrum partially exposing lower lip; distal tip reaching beyond level of tip of upper lip	Large, distally rounded, directed anteriorly and ventrally, connected to each other by membrane across ventrum partially exposing lower lip; distal tip below level of tip of upper lip	Short ventro-lateral lobes around the lower lip (considered as segment 2 by Fauvel)
Lobes on segment 2	Short, ventro-lateral, connected to each other by membrane across ventrum	Short, ventro-lateral, connected to each other by membrane across ventrum	Short, ventro-lateral, connected to each other by membrane across ventrum	Unknown, possibly absent
Lobes on segment 3	Large, distally rounded, originating dorsally to level of notopodia on segment 4, with deep dorso-lateral indentation, and terminating at level of lateral margin of ventral shields	Large, distally straight, originating dorsally to level of notopodia on segment 4 and terminating at level of lateral margin of ventral shields	Large, triangular, distally rounded, originating at level of notopodia on segment 4 and terminating at level of ventral edge of neuropodia	Relatively large, distally rounded, lateral lobes originating laterally to level of notopodia on segment 4 and terminating at level of lateral margin of ventral shields
Lobes on segment 4	Thin lateral flaps, connected to each other by crest across dorsum	Thin lateral flaps, terminating ventrally at level of ventral shields and dorsally by one pair of large triangular lobes covering base of lobes on segment 3, connected to each other by thick flap across dorsum, forming mid-dorsal pocket	Thin lateral flaps, connected to each other by crest across dorsum	Thin lateral flaps
Lobes on segments 5–6	Low flaps between ventral shields and neuropodia	Low flaps between ventral shields and neuropodia	Low flaps between ventral shields and neuropodia	Absent
Ventral shields on segments	2–16	2–13 (holotype poorly preserved, shields not visible after segment 13)	1–20	17 shields (possibly on segments 2–18)
Branchiae Pairs (number) Morphology	2 With long basal stems and all secondary branches originating together and about same length	2 With short basal stems and all secondary branches originating together and about same length (single branchia remaining in holotype)	2 Unknown, possibly with long basal stems and all secondary branches originating together and about same length	3 Arborescent, with short basal stems
Notopodia on segments	4–20	4–20	4–20	4–20
Uncini Long-handled on segments Number of rows of secondary teeth (anterior body; posterior region with biramous parapodia; anterior region with neuropodia only; posterior region with neuropodia only)	5–20 3–4 throughout	5–10 4; 3 onwards	5–20 3 throughout	5–7 5 throughout
Nephridial and genital papillae (segments)	3, 6–7	3, 6–8	3, 6–7	3–15
Additional information				Several characters, especially in regard to the morphology of the lobes, were re-interpreted from the drawing provided by Fauvel (1927; 267, Fig. 93h)