Pseudonereis masalacensis, (GRUBE, 1878)

Bakken, Torkild, 2007, Revision of Pseudonereis (Polychaeta, Nereididae), Zoological Journal of the Linnean Society 150 (1), pp. 145-176 : 159-171

PSEUDONEREIS MASALACENSIS ( GRUBE, 1878)

Remarks: Hutchings & Glasby (1985) examined the type material and found it to be in too poor condition to redescribe. They suggested this taxon should be regarded as indeterminable until material could be obtained from the type locality, especially due to lack of complete chaetae. They commented that no enlargement of dorsal notopodial lobes was observed in the holotype of P. masalacensis , suggesting that this species does not belong to Pseudonereis .

Type locality is Masolac , Philippines .

PSEUDONEREIS MULTISETOSA HARTMANN- SCHRÖDER, 1992

( FIG. 8 )

Pseudonereis multisetosa Hartmann-Schröder, 1992: 64–65 , figs 42–52.

Material examined: Holotype, Rangiroa , French Polynesia, sample 9, 11.ix.1982, ZMH P-20706 (1). Size range of material examined 67 chaetigers, 10 mm long, 0.7 mm wide.

Description: Frontal antennae present, 1 pair, cirriform. Prostomium with entire anterior margin, longer than wide. Eyes present, 2 pairs. One apodous anterior segment, greater than length of chaetiger 1. Tentacular cirri with distinct cirrophores, longest tentacular cirri extend back to chaetiger 5. Jaws with dentate cutting edge, translucent yellow-brown, 11–12 teeth. Maxillary ring of pharynx with paragnaths, arranged in discrete areas, Areas II–IV arranged in regular comb-like rows. Area I = 2 conical paragnaths; Area II = 17–18 conical paragnaths in 3 rows; Area III = 20 conical paragnaths in 2 rows; Area IV = 29–33 including conical paragnaths in 3 rows and a patch of cones towards jaws. Oral ring paragnaths present, Area V and VI present as distinct groups. Area V = 0; VI = 4 conical paragnaths arranged in transverse lines, bar-shaped paragnaths absent; VII–VIII = 8 conical paragnaths in a single row, present only as a ventral band.

Notopodium with dorsal notopodial ligule rounded, shorter than ventral notopodial ligule in anterior 10– 12 chaetigers ( Fig. 8A ), as long as posteriorly, not markedly elongate on posterior chaetigers, markedly broader on last about 10 posterior chaetigers ( Fig. 8D ). Prechaetal notopodial lobe absent, acicular process absent. Dorsal cirrus simple, lacking basal cirrophore but basis somewhat inflated anteriorly ( Fig. 8A ), 3–4 times length of ventral notopodial ligule at chaetigers 10–20 ( Fig. 8B ), basally attached to dorsal notopodial ligule on anterior and mid-body chaetigers, subterminally attached on posterior chaetigers ( Fig. 8D ).

Neuropodial inferior lobe prominent in anterior chaetigers, a small superior lobe present throughout ( Fig. 8C ). Neuropodial postchaetal lobe present throughout, a low rounded lobe, projecting level with end of acicular ligule. Ventral neuropodial ligule of anterior chaetigers present, short and rounded, similar in length to or slightly shorter than acicular neuropodial ligule, on posterior chaetigers up to half length of acicular neuropodial ligule.

Notochaetae: homogomph spinigers present, in midbody chaetigers (about 20–40) with short very pointed blades and very broad proximally ( Fig. 8H ); notopodial homogomph falcigers present first present in chaetiger 40–50, notopodial homogomph falcigers serrated; notopodial homogomph spinigers replaced by homogomph falcigers in posterior chaetigers. Neurochaetae, dorsal fascicle: homogomph spinigers and heterogomph falcigers present on anterior ( Fig. 8F ) and posterior chaetigers, blades serrated, homogomph falcigers ( Fig. 8E ) on anterior chaetigers present to about chaetiger 15, on posterior chaetigers absent. Neurochaetae, ventral fascicle: heterogomph spinigers present, heterogomph falcigers with short blades present throughout ( Fig. 8G ).

Pygidium ventrally incised, anal cirri cirriform reaching back 6 chaetigers.

Remarks: This taxon is known only from the holotype, which is in good condition, but may be a juvenile. Paragnath numbers presented here are different from those in the original description, as the pharynx had not been dissected.

Pseudonereis multisetosa differs from all other Pseudonereis taxa by the presence of a homogomph falciger in the neuropodial dorsal fascicle, which is rare feature in Nereidinae , but occurs in Leonnates and resembles those described for L. indicus ( Qui & Qian, 2000) . The short, very pointed spiniger with broad lower part observed in mid-body chaetigers (∼20–40) is a different kind of spiniger ( Fig. 8H ) only illustrated before in Leonnates jousseaumei ( Gravier, 1899) , which was in turn synonymized with L. indicus ( Qui & Qian, 2000) . This chaeta is here called a spiniger due to its thin whip-like tip. Hartmann-Schröder (1992) described it as having both spiniger and falciger characteristics.

Hartmann-Schröder (1992) stated that the presence of paragnaths in regular comb-shaped rows (termed pectinate in the original description) on the maxillary ring and the conical paragnaths on the oral ring justified the placement of this taxon in Pseudonereis . Another character pointing to a placement in Pseudonereis is the expansion in breadth of the dorsal notopodial ligule in posterior chaetigers. There are indications of this expansion in the type specimen as well, but it is only slightly expanded in breadth and only in the last about ten chaetigers.

As there are several distinctive morphological features pointing in different directions, further analyses including this taxon should be undertaken when more material becomes available. The form of the dorsal parapodial ligules in posterior chaetigers should in that case be redescribed as well as the types and distribution of chaetae.

Distribution: Rangiroa, Tuamoto Islands, French Polynesia (type locality).

PSEUDONEREIS NOODTI (HARTMANN- SCHRÖDER, 1962)

( FIG. 9 )

Neanthes noodti Hartmann-Schröder, 1962a: 129–130 , pl. 11, figs 65, 66, pl. 12, fig. 68, pl. 20, fig. 67. – 1962b: 395–398, figs 7–10.

Material examined: Lectotype, Chimbote , Peru 25.iv.1956, coll. Noodt, ZMH P-24716 (1) ; Paralectotype ZMH P-14380 (1). Size range: lectotype 13 chaetiger anterior fragment +38 chaetigers posterior fragment; paralectotype 31 chaetigers anterior fragment +37 chaetigers posterior fragment. Both specimens 1.5 mm wide .

Description: Frontal antennae present, 1 pair, cirriform; palpostyles conical. Prostomium with entire anterior margin, longer than wide. Eyes present, 2 pairs. One apodous anterior segment, greater than length of chaetiger 1. Tentacular cirri with distinct cirrophores, longest tentacular cirri extend back to chaetiger 2–3. Jaws with dentate cutting edge, brown/ black with 6 teeth. Maxillary ring of pharynx with paragnaths, arranged in discrete areas, Areas II–IV arranged in regular comb-like rows. Area I = 1 conical paragnath; Area II = 25–27 p-bar paragnaths in 3 diagonal rows; Area III = 64 p-bar and conical paragnaths in 4 rows; Area IV = 70–75 including p-bar paragnaths in four comb-shaped rows, additional cones and p-bars towards the jaws. Area V and VI present as distinct groups. Area V = 1, a large cone; Area VI = 1, a large triangular shield-shaped bar present, cones absent; Area VII–VIII = 16–17, two alternating rows of similar sized paragnaths, cones and p-bars interspersed.

Notopodium with dorsal notopodial ligule short and rounded as long as ventral notopodial ligule ( Fig. 9A, B ), markedly elongate and markedly broader on posterior chaetigers ( Fig. 9C, D ), merging to dorsal cirrus over chaetigers 15–20. Prechaetal notopodial lobe absent, acicular process absent. Dorsal cirrus simple, lacking basal cirrophore, 2–3 times length of ventral notopodial ligule at chaetigers 10–20 ( Fig. 9B ), basally attached in anterior chaetigers, terminally attached to dorsal notopodial ligule on posterior chaetigers commencing from about chaetiger 25. Ventral notopodial ligule short and rounded, similar throughout.

Neuropodial inferior lobe prominent in anterior chaetigers, a small superior lobe gives the acicular ligule a bilobed appearance ( Fig. 9B–D ). Neuropodial postchaetal lobe present throughout as a low rounded flattened lobe, not projecting beyond end of acicular ligule, level with or lower than the acicular ligule. Ventral neuropodial ligule of anterior chaetigers present, short rounded, similar in length to acicular neuropodial ligule; on posterior chaetigers present up to half length of acicular neuropodial ligule. Ventral cirri almost as long as acicular ligule throughout.

Notochaetae: homogomph spinigers. Neurochaetae, dorsal fascicle: homogomph spinigers present ( Fig. 9H ), heterogomph falcigers on anterior ( Fig. 9F ) and posterior chaetigers ( Fig. 9G ) present, blades serrated. Neurochaetae, ventral fascicle: heterogomph spinigers absent, heterogomph falcigers with short blades in anterior and posterior chaetigers present.

Anal cirri present.

Remarks: The type material consists of two anterior and two posterior ends, which due to the size of the specimens is possible to match; the smaller specimen has a complete posterior end. It was only possible to count paragnaths on the oral ring in the larger specimen. The larger specimen (anterior fragment and adjoining posterior fragment) is designated lectotype, and the smaller specimen as paralectotype.

Pseudonereis noodti is most similar to P. gallapagensis . They can be separated by P. noodti having a higher number of paragnaths in Area II and in Area IV (38–57 in P. gallapagensis vs. 70–75 in P. noodti ). Also in P. noodti the dorsal cirrus becomes terminally attached about chaetiger 25 as opposed to about chaetiger 15 in P. gallapagensis . Heterogomph spinigers in neuropodial ventral fascicle were not observed in the two specimens examined in P. noodti , but they are present in P. gallapagensis . Absence of heterogomph spinigers in the neuropodial ventral fascicle was also noted in the original description ( Hartmann-Schröder, 1962a).

The striking similarity of P. noodti to P. gallapagensis was not recognized by Hartmann- Schröder (1962a,b) who examined several specimens and gave a detailed description including of epitokes and reproductive modes ( Hartmann-Schröder, 1962b). The additional specimens included in the original treatment of this species ( Hartmann-Schröder, 1962a,b) were unavailable for study.

Distribution: South American Pacific coast from Chimbote Peru (type locality) in the north and south along the Chilean coast to Puerto Montt (Hartmann- Schröder, 1962b). Intertidally and in rockpools.

PSEUDONEREIS PALPATA ( TREADWELL, 1923)

( FIG. 10 )

Nereis (Neanthes) palpata Treadwell, 1923: 1239− 1243 , figs 6–15.

Material examined: Paratype, Ilha dos Alcatrazes , Sao Paulo, Brazil, USNM 19031 (1). Size range of material examined 67 chaetigers (incomplete), 75 mm long, 4 mm wide .

Description: Frontal antennae present, 1 pair, cirriform, as long as palps. Prostomium with entire anterior margin. Eyes present, 2 pairs. One apodous anterior segment, greater than length of chaetiger 1. Tentacular cirri with distinct cirrophores, longest tentacular cirri extend back to chaetiger 4. Jaws with dentate cutting edge, brown to black, plate-like finely serrated at proximal end, no distinct teeth. Maxillary ring of pharynx with paragnaths, arranged in discrete areas, Areas II–IV arranged in regular comb-like rows. Area I = 2, conical paragnaths; Area II = 39–40 in transverse rows; Area III = 109 in four transverse rows; Area IV = 108–120 in four diagonal lines, additional towards the jaws. Area V and VI present as distinct groups. Area V = 1 conical; VI = 1 conical and 1 bar present (one bar on left and right side, one cone on right side); VII–VIII = 19 paragnaths.

Notopodium with dorsal notopodial ligule rounded as long as ventral ligule ( Fig. 10A ), markedly elongate and broader on posterior chaetigers ( Fig. 10B ). Prechaetal notopodial lobe absent; acicular process absent. Dorsal cirrus basally attached to dorsal notopodial ligule on anterior and mid-body chaetigers, terminally attached on posterior chaetigers, 3–4 times as long as ventral notopodial ligule at chaetiger 10–20. Ventral notopodial ligule rounded.

Neuropodial inferior lobe rounded distinct throughout; a small superior lobe present throughout; neuropodial postchaetal lobe absent. Ventral neuropodial ligule up to half as long as acicular neuropodial ligule throughout. Ventral cirrus as long as acicular ligule in anterior chaetigers, becoming shorter, 0.5 times as long in posterior chaetigers.

Notochaetae: homogomph spinigers. Neurochaetae, dorsal fascicle: homogomph spinigers and heterogomph falcigers ( Fig. 10C ) present, blades serrated, blades having teeth only slightly longer proximally than distally. Neurochaetae, ventral fascicle: heterogomph spinigers present, with blades finely serrated proximally, homogomph spinigers absent; heterogomph falcigers with short blades present.

Remarks: The description presented here is based on a single specimen in poor quality. A more detailed redescription of this species based on a quantity of material from Brazil is being prepared and will be presented elsewhere. In the data matrix ( Table 3) the neuropodial postchaetal lobe is scored as absent, but due to the poor condition of the single specimen examined, this character should be verified from additional material.

This taxon is unique among Pseudonereis in possessing both bar-shaped and conical paragnaths in Area VI. The type or shape in bar-shaped paragnaths presented as new characters here was not recorded from the paratype and will have to be verified from additional material.

Distribution: South America, Atlantic coast, type locality: Estado de S. Paulo, Brazil.

PSEUDONEREIS PSEUDONOODTI ( FAUCHALD, 1977)

( FIGS 11 , 12 )

Neanthes pseudonoodti Fauchald, 1977: 27–29 , fig. 7.

Material examined: Holotype, Paitilla Beach , Panama, Pacific, intertidal, USNM 53090 ; Paratypes, Paitilla Beach , Panama, Pacific, intertidal, Tetraclita zone USNM 53091 (4). Additional material: Paitilla Beach, Panama, Pacific, coll. A.A. Reimer, 20.i.1971 USNM 065983 (1) ; Paitilla Beach, Panama, Pacific, intertidal, coll. A.A. Reimer, 20.i.1971 USNM 065984 (5) . Size range of material examined 29–74 chaetigers (only one specimen complete), 4–18 mm long, 0.5–1 mm wide.

Description: Frontal antennae present, 1 pair, cirriform; palpostyles conical. Prostomium with entire anterior margin, wide as long. Eyes present, 2 pairs. One apodous anterior segment, greater than length of chaetiger 1. Tentacular cirri with distinct cirrophores, longest tentacular cirri extend back to chaetiger 3–4. Jaws with dentate cutting edge, translucent brown, with 8 teeth. Maxillary ring of pharynx with paragnaths, arranged in discrete areas, Areas II–IV arranged in regular comb-like rows. Area I = 4 conical paragnaths, one in front of a transverse row of 3; Area II = 15–27 p-bar paragnaths in 4 rows in a triangular patch; Area III = 28–40 p-bar and conical paragnaths in 4 rows; Area IV = 20–61 p-bar and conical paragnaths in 4–5 rows, additional cones and p-bar paragnaths towards the jaws absent. Oral ring paragnaths present, Area V and VI present as distinct groups. Area V = 3 conical paragnaths present arranged in a triangular pattern ( Fig. 11A ); Area VI = 1 shieldshaped bar with pointed tip present, cones a bsent; Area VII–VIII = 69–100, in 2–3 rows with similar sized conical paragnaths, with larger p-bars interspersed ( Fig. 11B ).

Notopodium with dorsal notopodial ligule digitiform rounded in anterior chaetigers ( Fig. 12A, B ), markedly elongate on posterior chaetigers ( Fig. 12D, E ), markedly broader on posterior chaetigers ( Fig. 12E ), expansion commencing at about chaetiger 30–35. Prechaetal notopodial lobe absent, acicular process absent. Dorsal cirrus simple, lacking basal cirrophore, 2–3 times length of ventral notopodial ligule at chaetiger 10–20 ( Fig. 12B ), basally to middorsally attached to dorsal notopodial ligule on anterior chaetigers ( Fig. 12B ), terminally attached to dorsal notopodial ligule on posterior chaetigers, from about chaetiger 40 ( Fig. 12D, E ).

Neuropodial inferior lobe prominent in anteriormost chaetigers only; superior lobe absent. Neuropodial postchaetal lobe present throughout, a low rounded lobe projecting level with or lower than end of acicular ligule, most prominent in posterior chaetigers ( Fig. 12D, E ). Ventral neuropodial ligule of anterior chaetigers up to half length of acicular neuropodial ligule, on posterior chaetigers reduced ( Fig. 12E ).

Notochaetae: homogomph spinigers. Neurochaetae, dorsal fascicle: homogomph spinigers present; heterogomph falcigers on anterior ( Fig. 12F ), and posterior chaetigers present, blades serrated. Neurochaetae, ventral fascicle: heterogomph spinigers present commencing in chaetiger about 5–10, heterogomph falcigers with short blades present throughout ( Fig. 12G ).

Remarks: The vial with the paratypes ( USNM 53091) with nine specimens included five specimens of Nereis riisei . The description is based on the remaining four specimens.

The neuropodial postchaetal lobe was difficult to detect in the smaller specimens examined (<1 mm body width). Fauchald (1977) stated in the original description that there was a single large paragnath in Area V and two in each of Area VI. This is reinterpreted here to be three widely spread cones in Area V in a triangular pattern, and a single in Area VI ( Fig. 12A ). The paragnaths in Area VI are high, flat and pointed triangular bars (shield-shaped) as seen in most other Pseudonereis species.

Pseudonereis pseudonoodti is very similar to P. cortezi , but can be distinguished by absence of the neuropodial superior lobe, which is present in P. cortezi , and the number of paragnaths in Area V. In P. pseudonoodti the paragnaths in Area III are more widely spaced ( Fig. 11B ) than in other species, although still in tight regular rows, the notopodial dorsal ligule is greatly expanded in length but not so much in breadth, and it is almost impossible to see clearly where the dorsal cirrus starts on the expansion from the notopodial ligule.

Distribution: Type locality Paitilla Beach (Pacific), Panama, intertidal rocky substrates ( Fauchald, 1977).

PSEUDONEREIS TRIMACULATA HORST, 1924

( FIG. 13 )

Nereis (Pseudonereis) trimaculata Horst, 1924: 187– 188 , pl. XXXVI, figs 8, 9.

Pseudonereis rottnestiana forma costatodentata Hartmann-Schröder, 1979: 119, figs 220–222.

Material examined: Syntype, between Gisser and Ceram, Indonesia, Siboga Expedition, stn 172, reef exploration, ZMA Vpol-954 (1); Broome, WA, probe 6, 9/91975, HZM P-16567 (7); south-west tip of West Lewis Island, Western Australia, Australia, 20°36.38′S 116°36.06′E, WA624, intertidal, coll. P.Hutchings & L. Avery 27.vii.2000, AM W29016 (1); North-west of West Lewis Island , Western Australia, Australia, 20°33.52′S 116°38.21′E, WA621, under boulders and in crevices, depth 0 m, coll. P.Hutchings & L. Avery 26.vii.2000, AM W29017 (2). Size range of material examined 60–94 chaetigers for 17–41 mm long, 1–2 mm wide .

Description: Frontal antennae present, 1 pair, cirriform. Prostomium with entire anterior margin. Eyes present, 2 pairs. One apodous anterior segment, greater than length of chaetiger 1. Tentacular cirri with distinct cirrophores, longest tentacular cirri extend back to chaetiger 5–7. Jaws with dentate cutting edge, brown, with 5–6 teeth. Maxillary ring of pharynx with paragnaths, arranged in discrete areas, Areas II–IV arranged in regular comb-like rows. Area I = 1–2 conical paragnaths, in a longitudinal row when 2; Area II = 23–38 p-bar paragnaths in 4 rows; Area III = 51–69 p-bar paragnaths in 4 rows; Area IV = 50– 78 including p-bar paragnaths in 4 rows additional cones and 2–4 p-bars towards jaws. Oral ring paragnaths present, Area V and VI present as distinct groups. Area V = 1–3 large conical paragnaths present, arranged in a triangular pattern when 3; Area VI = 1 shield-shaped bar present, cones absent; Area VII–VIII = 20–24 large conical paragnaths of similar size and interspersed p-bars present in two alternating rows.

Notopodium with dorsal notopodial ligule digitiform rounded as long as ventral neuropodial ligule in anterior chaetigers ( Fig. 13A, B ), markedly elongate on posterior chaetigers commencing about chaetiger 55–60, markedly broader on posterior chaetigers commencing about chaetiger 50–55 ( Fig. 13D, E ). Prechaetal notopodial lobe absent, acicular process absent. Dorsal cirrus simple, lacking basal cirrophore, 2 times length of ventral notopodial ligule at chaetiger 10–20 ( Fig. 13B ), basally attached in anterior chaetigers, subterminally attached in mid-body chaetigers ( Fig. 13D ), and terminally attached to dorsal notopodial ligule on posterior chaetigers in the last quarter of the body ( Fig. 13E ).

Neuropodial inferior lobe prominent in anterior chaetigers, a small superior lobe present ( Fig. 13A ), less developed from mid-body chaetigers, visible throughout. Neuropodial postchaetal lobe present throughout, a low rounded lobe with a small pointed tip, projecting level with the end of acicular ligule ( Fig. 13D ). Ventral neuropodial ligule rounded, well developed throughout, up to half length of acicular neuropodial ligule on anterior chaetigers, on posterior chaetigers similar to length of acicular neuropodial ligule. Ventral cirri 0.5 times as long as neuropodial acicular ligule.

Notochaetae: homogomph spinigers present ( Fig. 13I ). Neurochaetae, dorsal fascicle: homogomph spinigers present ( Fig. 13H ), heterogomph falcigers on anterior and posterior ( Fig. 13G ) chaetigers present, blades serrated. Neurochaetae, ventral fascicle: heterogomph spinigers present from about chaetiger 30 ( Fig. 13F ), heterogomph falcigers with short blades present throughout.

Anal cirri reaching back about 10 chaetigers.

Remarks: The original material ( ZMA V-pol 954) labelled as ‘type’ and given as syntype in the museum record ( ZMA) consisted of two specimens, not three as stated in the original description. Of the two specimens the smaller one is a Platynereis sp. and the other was used to prepare the original description. This specimen is designated as lectotype, and confirmed well with the Australian material examined and illustrated ( Fig. 13 ). The lectotype is a complete specimen and measures 41 mm in length for 94 chaetigers, 2 mm wide. Paragnaths were not possible to count except in Area I = 1; Area VI = 1/1; Area VII–VIII = 20.

Pseudonereis trimaculata is similar to P. gallapagensis , P. variegata , P. noodti and P. pseudonoodti ; it most closely resembles P. pseudonoodti ( Table 1). The other taxa in this group have a large single conical paragnath in Area V while P. pseudonoodti and P. trimaculata have three. Apart from this character there are differences in paragnath numbers, length of dorsal cirrus and development and length of the ventral neuropodial ligule separating these species ( Table 1).

Horst (1924) gave only a brief description but included illustrations of two parapodia, one anterior and the other which appears to be from a mid-body chaetiger, although the figure legend stated it was from a posterior parapodium. He drew attention to black spots on the parapodia that he used to distinguish his new taxon from other described Pseudonereis taxa. These spots are glands often observed in parapodia of nereidids, but now only vaguely visible in the lectotype.

Hartmann-Schröder (1979) described specimens representing this taxon as a variety of P. rottnestiana , using the term ‘form’ (‘ Pseudonereis rottnestiana forma costatodentata ’). The label with these specimens ( HZM P-16567) did not have the name as such but a comment was added to distinguish the specimens as separate from P. rottnestiana (see P. anomala for further comments).

Distribution: Type locality Geser, Indonesia, known from the type locality and Western Australia. This is the first record of this taxon from Australia. All records are from intertidal and low littoral environments.

PSEUDONEREIS VARIEGATA ( GRUBE, 1857)

( FIGS 14 , 15 )

Nereilepas variegata Grube, 1857: 164–165 .

Nereis ferox Hansen, 1882: 14 , pl. IV, figs 34–39. – Augener 1934: 133.

Nereis variegata Ehlers, 1901: 112 –11118, pl. XIV, figs 1–21.

Pseudonereis variegata Hartman, 1949: 69–70 . – Hartmann-Schröder 1962b: 434–435. – Fauchald 1977: 33, fig. 9d–e.

Material examined: Syntypes Nereis ferox Hansen, 1882 , Rio de Janeiro, ZMUB 2130 (3); Rio de Janeiro, v. Beneden, ZMUC-POL-1804 (2). Size range of material examined 64–94 chaetigers (62 incomplete specimen), 26–46 mm long (35 incomplete specimen), 1.5–3 mm wide .

Description: Frontal antennae present, 1 pair, cirriform. Prostomium with entire anterior margin, longer than wide. Eyes present, 2 pairs ( Fig. 14A ). One apodous anterior segment, greater than length of chaetiger 1. Tentacular cirri with distinct cirrophores, longest tentacular cirri extend back to chaetiger 3–5. Jaws with dentate cutting edge, black/brown platelike serrated with 4–5 teeth. Maxillary ring of pharynx with paragnaths, arranged in discrete areas. Areas II– IV arranged in regular comb-like rows. Area I = 1–2 conical paragnaths; Area II = 13–34 p-bar paragnaths in three rows; Area III = 59–76 p-bar paragnaths in four rows, anteriormost row half as long as the others; Area IV = 63–87 p-bar paragnaths in 4–5 rows, additional cones and 2–4 p-bars towards jaws. Area V and VI present as distinct groups. Area V = 1 conical paragnath. Area VI = 1 large shield-shaped bar present ( Fig. 14B, C ); Area VII–VIII = 18–21, conical paragnaths and large p-bars in two alternating rows, p-bars being more posteriorly placed ( Fig. 14B ).

Notopodium with dorsal notopodial ligule stout rounded as long as ventral notopodial ligule in anterior chaetigers ( Fig. 15A, B ), markedly elongate and markedly broader on posterior chaetigers, expanded in breadth from about chaetiger 30 ( Fig. 15C ), in length from about chaetiger 40 ( Fig. 15D, E ). Prechaetal notopodial lobe absent, acicular process absent. Dorsal cirrus simple, lacking basal cirrophore, 2 times length of ventral notopodial ligule at chaetiger 10–20, basally attached in anterior chaetigers ( Fig. 15B ), subterminally attached from about chaetiger 30 ( Fig. 15D, E ), terminally attached to dorsal notopodial ligule on posteriormost chaetigers ( Fig. 15F ). Ventral notopodial ligule stout rounded, in posterior chaetigers digitiform.

Neuropodial inferior lobe prominent in anterior chaetigers; small superior lobe present throughout ( Fig. 15E, F ). Neuropodial postchaetal lobe present throughout, low rounded not projecting beyond end of acicular ligule, in posterior chaetigers with a small triangular tip ( Fig. 15D, F ). Ventral neuropodial ligule of anterior chaetigers present, well developed, similar in length to acicular neuropodial ligule throughout. Ventral cirrus 0.5–1 times as long as neuropodial acicular ligule.

Notochaetae: homogomph spinigers. Neurochaetae, dorsal fascicle: homogomph spinigers ( Fig. 15G ) and heterogomph falcigers present throughout, blades serrated. Neurochaetae, ventral fascicle: heterogomph spinigers present from about chaetiger 40, with blades finely serrated; heterogomph falcigers with short blades ( Fig. 15H ) present throughout.

Pygidium with multiple ventral incisions, anal cirri reaching back five chaetigers.

Remarks: Description and illustrations presented here are based on the syntypes of Nereis ferox and variation given from additional material examined. Type material of Nereilepas variegata Grube was not available for study. Nereis ferox has been synonymized in the literature by Ehlers (1901) and Augener (1934). Ehlers apparently examined the types of N. ferox which he found to be identical to P. variegata . Type specimens of N. ferox and P. variegata should be compared but could not be located in this study.

Pseudonereis variegata resembles most closely P. gallapagensis ( Table 1) but is distinguished by the length of the ventral neuropodial ligule, which in P. variegata is of similar length to the acicular ligule but up to half as long in P. gallapagensis ; and by the number of conical paragnaths in Area IV (68–87 in P. variegata ; 38–57 in P. gallapagensis ). The length of the dorsal cirrus in anterior chaetigers is also different being longer in P. gallapagensis . Pseudonereis variegata has generally a higher number of paragnaths in all areas than P. gallapagensis , although these differences are small and may be insignificant when more material is examined. Similarities to P. atopodon are also noted ( Table 1; see this taxon for further comments).

Paragnaths in Area VII–VIII in several rows have been used as a diagnostic feature to distinguish P. variegata from P. gallapagensis (Hartmann- Schröder, 1962b; Fauchald, 1977). The material examined here indicates this character is not useful. The same sources also used the attachment of the dorsal cirrus on the dorsal notopodial ligule in posterior chaetigers as another distinctive feature, and have been given as distally attached in posterior chaetigers in P. gallapagensis and subdistally in P. variegata . In the data matrix the dorsal cirrus is coded as terminally attached in both taxa, but the distinction may be applied in identifications, especially when anterior fragments only are available or at least not complete specimens. In P. variegata the dorsal cirrus is terminally attached only in the posteriormost chaetigers, while in P. gallapagensis the dorsal cirrus is terminally attached from anterior chaetigers, from about chaetiger 15.

Other taxa mentioned in the literature in association with P. variegata are three taxa ( Nereis coerulea , Nereis obscura , and Phyllonereis benedeni ) Hansen (1882) described from Brazil and later examined by Ehlers (1901) and Augener (1934) who found them to be identical with P. variegata . They did not mention P. gallapagensis . The material collected by van Beneden in Rio de Janeiro, Brazil, and examined by Hansen is obviously split between different museums, and part of this material could not be located and examined.

Hartman (1949) placed Paranereis elegans Kinberg, 1865 in synonymy with P. variegata although she stated the single specimen was damaged. The same specimen has been examined ( SMNH Type-459) and the fragment left is found to be of no scientific value, and hence is not included in the material examined above. Extensive synonymies are listed in Ehlers (1901) (see also Augener, 1934); those included above represent only those specimens examined by me.

Distribution: Type locality of Nereis ferox is Rio de Janeiro, Brazil. South America, Atlantic coast. Habitat is inshore and shelf. According to Grube (1857) specimens he used to describe Nereilepas variegata were collected by Kröyer in Valparaiso ( Chile) and Callao (probably Peru).

Figures

Figure 8. Pseudonereis multisetosa Hartmann-Schröder, 1992 holotype ZMH P-20706. A, parapodium 5th chaetiger anterior view. B, parapodium 12th chaetiger anterior view. C, parapodium 20th chaetiger posterior view. D, parapodium 56th chaetiger posterior view. E, homogomph falciger neuropodial dorsal fascicle 12th chaetiger. F. heterogomph falciger dorsal fascicle 12th chaetiger. G, heterogomph falciger ventral fascicle 56th chaetiger. H, notopodial homogomph spiniger 42nd chaetiger. Scale bar in A–D 0.1 mm, E–G 0.01 mm. H is taken from Hartmann-Schröder (1992).

Figure 9. Pseudonereis noodti (Hartmann-Schröder, 1962) holotype HZM P-14380. A, parapodium 3rd chaetiger anterior view. B, parapodium 10th chaetiger anterior view. C, parapodium 20th chaetiger anterior view. D, parapodium 58th chaetiger anterior view. E, parapodium 42nd chaetiger posterior view. F, heterogomph falciger neuropodial dorsal fascicle 10th chaetiger. G, heterogomph falciger neuropodial dorsal fascicle 58th chaetiger. H, homogomph spiniger neuropodial dorsal fascicle 58th chaetiger. End-view drawings of parapodia in A–C are not to scale. Scale bar in A–E 0.1 mm, F–H 0.01 mm.

Figure 10. Pseudonereis palpata (Treadwell, 1923) paratype USNM 19031. A, parapodium 10th chaetiger anterior view. B, parapodium 65th chaetiger anterior view. C. Heterogomph falciger neuropodial dorsal fascicle 41st chaetiger. Scale bar in A and B 0.5 mm, C 0.01 mm. End-view drawings of parapodia in A and B are not to scale.

Figure 11. Pseudonereis pseudonoodti (Fauchald, 1977) paratype USNM 53091. A, anterior end dorsal view. B, anterior end ventral view. A and B are of the same specimen measuring 0.8 mm body width at chaetiger 10 excluding parapodia.

Figure 12. Pseudonereis pseudonoodti (Fauchald, 1977) paratype USNM 53091. A, parapodium 3rd chaetiger anterior view. B, parapodium 10th chaetiger anterior view. C, parapodium 30th chaetiger anterior view. D, parapodium 40th chaetiger posterior chaetiger. E, parapodium 50th chaetiger anterior view. F, heterogomph falciger neuropodial dorsal fascicle 50th chaetiger. G, heterogomph falciger neuropodial ventral fascicle 30th chaetiger. End-view drawings of parapodia in A– E are not to scale. Scale bar in A–E 0.1 mm, F and G 0.01 mm.

Figure 13. Pseudonereis trimaculata Horst, 1924 ZMH P-16567. A, parapodium 3rd chaetiger anterior view. B, parapodium 11th chaetiger anterior view. C, parapodium 30th chaetiger anterior view. D, parapodium 50th chaetiger posterior view. E, parapodium 65th chaetiger anterior view. F, heterogomph spiniger neuropodial ventral fascicle 30th chaetiger. G, heterogomph falciger neuropodial dorsal fascicle 30th chaetiger. H, homogomph spiniger neuropodial dorsal fascicle 30th chaetiger. I, homogomph spiniger notopodium 3rd chaetiger. End-view drawings of parapodia in A–D are not to scale. Scale bar in A–E 0.1 mm, F–I 0.01 mm.

Figure 14. Pseudonereis variegata (Grube, 1857) syntypes of Nereis ferox Hansen, 1882 ZMUB 2130. A, anterior end dorsal view. B, anterior end ventral view. C, anterior end lateral view. The laterally flattened shield-shaped paragnaths protruding from the tissue are demonstrated in C. A–C are of the same specimen measuring 3 mm body width at chaetiger 10 excluding parapodia.

Figure 15. Pseudonereis variegata (Grube, 1857) syntypes of Nereis ferox Hansen, 1882 ZMUB 2130. A, parapodium 3rd chaetiger anterior view. B, parapodium 11th chaetiger anterior view. C, parapodium 30th chaetiger anterior view. D, parapodium 40th chaetiger posterior view. E, parapodium 56th chaetiger anterior view. F, parapodium 78th chaetiger anterior view. G, homogomph spiniger neuropodial dorsal fascicle 11th chaetiger. H, heterogomph falciger neuropodial ventral fascicle 11th chaetiger. End-view drawings of parapodia in A–F are not to scale. Scale bar in A 0.01 mm, B–F 0.5 mm, G and H 0.01 mm.