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Nereis edentata Hadiyanto 2023, n. sp.
Nereis edentata n. sp. Fig. 9A–I Material examined. Holotype. Western Australia: Port Denison, 29°16′33.59″S 114°54′45.68″E, 29 December 2020, 1 specimen (WAM V11651). Paratypes: Kalbarri, 27°43′17.09″S 114°09′11.39″E, 15 November 2020, 2 specimens (WAM V11652). Horrock, 28°23′23.32″S 114°25′48.98″E, 18 November 2020, 2 specimens (WAM V11653). Glenfield Beach, 28°41′01.32″S 114°36′21.65″E, 17 November 2020, 2 specimens (WAM V11654). Dynamite Bay, Green Head, 30°04′15.62″S 114°57′44.93″E, 28 December 2020, 2 specimens (WAM V11655). Yanchep Beach, 31°33′02.36″S 115°37′24.86″E, 14 January 2021, 3 specimens (WAM V11656). Beaumaris Beach, 31°44′23.57″S 115°43′17.82″E, 18 January 2021, 1 specimen (WAM V11657). Cape Peron, 32°15′59.14″S 115°41′06.74″E, 12 January 2021, 1 specimen (WAM V11658). Hunters Beach, Green Head, 30° 4′27.54″S 114°58′11.46″E, 29 July 2016, 1 specimen (WAM V11659). Description. Holotype a complete specimen with 54 chaetigers, body 19.1 mm long and 1.3 mm wide; cream yellow in alcohol. Prostomium wider than long. Eyes black, two pairs, equal sizes, in trapezoidal arrangement. Palps one pair, palpophores globose, palpostyles conical. Antennae one pair, extending to level of palps. Tentacular cirri four pairs with basal articulation, longest one extending to chaetiger 5. Pharyngeal jaws translucent reddish black, curved at tips, with nine teeth on each jaw. Paragnaths reddish black, conical, present on maxillary and oral rings, arranged as follows: Area I= small, 1; Area II= small, 7 on left and 8 on right, in a triangle in each side; Area III= small, 2 in one straight transverse row; Area IV= small, 10 on left and 11 on right, in a curved cluster in each side; Area V= 0; Area VI= small, 6 on left in 2 rows and 5 on right in a circle; Areas VII–VIII= 12 in one row (8 large paragnaths widely spaced and 4 small paragnaths in ventromedial row) (Fig. 9A–B). Apodous segment slightly longer than first chaetiger. First two chaetigers uniramous. Notopodia present with digitiform dorsal and ventral ligules (Fig. 9C–E), basal dorsal ligules enlarged in middle (Fig. 9D) and posterior chaetigers (Fig. 9E). Dorsal cirri cirriform, attached on middle of dorsal parapodia, twice longer than dorsal ligules, similar length throughout chaetigers. Neuropodia with digitiform ventral ligules, acicular ligules, and triangular postchaetal lobes extending below acicular ligule tips (Fig. 9C–E). Ventral cirri cirriform, attached basally on ventral parapodia, slightly shorter than ventral ligules in all chaetigers. Notochaetae present with homogomph spinigers in anterior chaetigers, homogomph falcigers (Fig. 9F) from chaetiger 20, two falcigers in each notopodium. Notopodial falcigerous blades short, smooth, slightly curved at tips. Neurochaetae present with heterogomph falcigers (Fig. 9G) and homogomph spinigers (Fig. 9H) in dorsal fascicles, heterogomph spinigers (Fig. 9I) and falcigers in ventral fascicles. Neuropodial falcigerous blades medium size, with fine serrations. All spinigerous blades long, with fine serrations. Acicula black. Pygidium with anus on dorsal side, anal cirri cirriform, as long as last four chaetigers. Variations in paratypes. Complete specimens with 39–69 chaetigers, body 6.2–37.3 mm long and 0.4–1.5 mm wide; cream yellow in alcohol. Incomplete specimens with 29–44 chaetigers, posterior end missing, remaining body 5.7–32.9 mm long and 0.7–1.5 mm wide; cream yellow in alcohol. Longest tentacular cirri extend to chaetiger 2–5. Paragnaths small but larger in Areas VII–VIII, arranged as follows: Area I= 0–1, Area II= 3–8 in a triangle, Area III= 0–5 in one straight transverse row, Area IV= 5–12 in a curved cluster, Area V= 0, Area VI= 2–7 in an oval, Areas VII–VIII= 6–10 in one row, including 1–3 smaller paragnaths in ventromedial row in some specimens. Notopodial homogomph falcigers present from chaetiger 18– 24, two falcigers (occasionally 3) in each notopodium or 1–2 falcigers in small specimens (<15 mm long). Anal cirri as long as last three to four chaetigers. Remarks. Nereis edentata n. sp. most closely resembles Nereis species having one row of paragnaths on Areas VII–VIII and smooth notopodial homogomph falcigers including: Nereis coutieri Gravier, 1899, Nereis eugeniae (Kinberg, 1865) and Nereis posidoniae Hutchings & Rainer, 1979. Nereis edentata n. sp. and N. coutieri have enlarged dorsal notopodial ligules in posterior chaetigers, but they differ in the ornamentation of the dorsal chaetiger 2 and the arrangement of paragnaths on Areas III and VII–VIII. Nereis coutieri has a brown band dorsally on chaetiger 2 (Gravier 1899a; Gravier 1902; Fauvel 1911; Yousefi et al. 2011), while Nereis edentata n. sp. lacks a band. Nereis coutieri has more paragnaths on Area III (5–7) (Gravier 1899a; Gravier 1902; Yousefi et al. 2011) and those paragnaths are arranged in a curved row with a deep concavity ahead in specimens from the Red Sea (type locality) (Gravier 1899a; Gravier 1902) or an irregular group in specimens from South Africa and Mozambique (Day 1967), instead of in a straight transverse row as for Nereis edentata n. sp. Paragnaths on Areas VII–VIII are present in a similar size for N. coutieri (Gravier 1899a; Gravier 1902; Day 1967; Bonyadi-Naeini et al. 2017) but in two sizes for some specimens of Nereis edentata n. sp. In addition, N. coutieri has only one uniramous chaetiger in the first parapodium (Fauvel 1911) although this is not mentioned in the original description (Gravier 1899a) and previous studies (Gravier 1902; Day 1967; Yousefi et al. 2011), while Nereis edentata n. sp. has two uniramous chaetigers in first two parapodia. Nereis edentata n. sp. also differs from materials from Lizard Island, Queensland that were identified by Glasby (2015) as Nereis cf. coutieri. Dorsal notopodial ligules of the latter species are shorter than ventral notopodial ligules (Glasby 2015). Hartmann-Schröder (1982, 1983) found N. coutieri from south-west Australia (Yallingup and Margaret River). These specimens need to be assessed further as there was no description of the ornamentation of the dorsal chaetiger 2 and the arrangement of paragnaths on Area III. Nereis edentata n. sp., N. eugeniae, and N. posidoniae do not have a pigmented band on the dorsal chaetiger 2, but they differ in the size of the eyes, the length of the tentacular cirri, the presence of paragnaths on Area VI, and the enlargement of dorsal notopodial ligules in posterior chaetigers. The eyes of N. eugeniae are small and occasionally absent (Hartman 1964; Day 1967; Darbyshire 2014). The tentacular cirri of N. eugeniae and N. posidoniae are shorter than those of Nereis edentata n. sp., with the longest one extending to only the apodous segment (Kinberg 1865; Hartman 1964; Day 1967; Hutchings & Rainer 1979; Darbyshire 2014). Paragnaths on Area VI are absent for N. posidoniae (Hutchings & Rainer 1979) but present for Nereis edentata n. sp. and N. eugeniae (3–6) (Hartman 1964; Day 1967; Darbyshire 2014). Dorsal notopodial ligules in posterior chaetigers are not expanded in N. eugeniae (Hartman 1964; Day 1967; Darbyshire 2014) and not described in N. posidoniae (Hutchings & Rainer 1979) but enlarged in Nereis edentata n. sp. Etymology. The new species is named after the Latin adjective, edentates, meaning toothless, referring to the smooth-bladed homogomph falcigers. Distribution. South-west Australia. Distribution records indicate that the new species appears to be transitional between the tropical and temperate Australia realms (Table 2). Habitat. Intertidal, rocky shores (Table 2).Published as part of Hadiyanto, Hadiyanto, 2023, Nereididae (Annelida: Phyllodocida) from intertidal macroalgae in Western Australia, pp. 151-203 in Zootaxa 5239 (2) on pages 168-170, DOI: 10.11646/zootaxa.5239.2.1, http://zenodo.org/record/762412
Nereididae (Annelida: Phyllodocida) from intertidal macroalgae in Western Australia
Hadiyanto, Hadiyanto (2023): Nereididae (Annelida: Phyllodocida) from intertidal macroalgae in Western Australia. Zootaxa 5239 (2): 151-203, DOI: 10.11646/zootaxa.5239.2.1, URL: http://dx.doi.org/10.11646/zootaxa.5239.2.
Nereis yuedensis Hadiyanto 2023, n. sp.
Nereis yuedensis n. sp. Fig. 11A–J Nereis cf. jacksoni Hartmann-Schröder, 1982: 78. Not Kinberg, 1865. Material examined. Holotype. Western Australia: Dynamite Bay, Green Head, 30°04′15.62″S 114°57′44.93″E, 28 December 2020, 1specimen (WAM V11661). Paratypes: Dynamite Bay, Green Head, 30°04′15.62″S 114°57′44.93″E, 28 December 2020, 9 specimens (WAM V11662). Dynamite Bay, Green Head, 30°04′15.62″S 114°57′44.93″E, 28 December 2020, male epitoke, 1 specimen (WAM V11663). Description. A holotype specimen incomplete, posterior end missing, with 45 chaetigers, remaining body 9.9 mm long and 0.6 mm wide; cream yellow in alcohol. Prostomium as long as wide. Eyes black, two pairs, equal sizes, in rectangular arrangement. Palps one pair, palpophores globose, palpostyles conical. Antennae one pair, shorter than palps. Tentacular cirri four pairs with basal articulation, longest one extending to chaetiger 3. Pharyngeal jaws reddish brown, curved at tips, with seven teeth on each jaw. Paragnaths small, translucent to reddish, conical, present on maxillary and oral rings, arranged as follows: Area I= 0, Area II= 1 on left and 2 on right, Area III= 0, Area IV= 3 on left and 6 on right, in one curved row each side, Area V= 0, Area VI= 0, Areas VII–VIII= 10 in one row (Fig. 11A–B). Apodous segment as long as first chaetiger. First two chaetigers uniramous. Notopodia present with strongly reduced dorsal ligules from first chaetiger and conical ventral ligules. Dorsal cirri digitiform, attached on middle of dorsal parapodia, about twice longer than ventral ligules, similar length throughout chaetigers. Neuropodia with conical ventral ligules, acicular ligules, and triangular postchaetal lobes extending to same level of acicular ligule tips. Ventral cirri cirriform, attached basally on ventral parapodia, as long as ventral ligules, similar length throughout chaetigers (Fig. 11C–E). Notochaetae present with homogomph falcigers from chaetiger 3, one for each notopodia.Notopodial falcigerous blades short, with terminal and three large lateral teeth in anterior chaetigers (Fig. 11F), two large lateral teeth and two smaller teeth basally in posterior chaetigers (Fig. 11G). Neurochaetae present with heterogomph falcigers and homogomph spinigers (Fig. 11H) in dorsal fascicles, heterogomph spinigers (Fig. 11I) and falcigers (Fig. 11J) in ventral fascicles. Neuropodial falcigerous and spinigerous blades short, with fine serrations. Acicula reddish brown to black. Variations in paratypes. Complete specimens present with 32–44 chaetigers, body 3.9–6.8 mm long and 0.3–0.4 mm wide; cream yellow in alcohol. Incomplete specimens present with 35–38 chaetigers, posterior end missing, remaining body 4.7–9.3 mm long and 0.4–0.5 mm wide; cream yellow in alcohol. An incomplete, male epitokous specimen present with 51 chaetigers, anal cirri missing, body 10.2 mm long and 0.6 mm wide; cream yellow in alcohol. Longest tentacular cirri extending to chaetiger 1–3. Area VI present with 2–3 cones, Areas VII–VIII present with 8–10 cones in one row, other Areas difficult to see. Notopodial falcigerous blades short, with terminal and two large lateral teeth and one smaller tooth basally. Pygidium present with anus on dorsal side, anal cirri cirriform, as long as last three chaetigers. A male epitokous specimen present with black, enlarged, overlapping paired eyes. Body divided into 11 prenatatory chaetigers, 23 natatory chaetigers, and 17 post-natatory chaetigers. Parapodia elongated, neuropodial postchaetal lobes modified into large, flat lamella in natatory chaetigers. Dorsal cirri swollen basally with elongated lateral papillae in first seven pre-natatory chaetigers, slender and longer with shorter and more papillae in natatory chaetigers, unmodified in post-natatory chaetigers. Ventral cirri swollen basally with lateral papillae in first four pre-natatory chaetigers, slender and longer in natatory chaetigers, unmodified in post-natatory chaetigers. Paddleshaped notochaetae present in natatory chaetigers. Remarks. The number and arrangement of paragnaths were evaluated based on the holotype and two paratype specimens (including an epitokous specimen). Other paratype specimens had retracted pharynx and were too small to dissect ventrally. Nevertheless, important taxonomic characters, including parapodia and chaetae, were consistent with those of the holotype and paratype specimens with everted pharynx. Nereis yuedensis n. sp. resembles those Nereis species having one row of paragnaths on Areas VII–VIII and notopodial homogomph falcigers with at least one large lateral tooth, including N. bifida Hutchings & Turvey, 1982; N. denhamensis Augener, 1913; N. falcaria (Willey, 1905); N. heirissonensis Augener, 1913; N. panamensis Fauchald, 1977; N. spinigera Hutchings & Turvey, 1982; N. taitungensis Hsueh, 2020; N. taiwanensis Hsueh, 2020; and N. usticensis Cantone, Catalono & Badalamenti, 2003. However, notopodial homogomph falcigers of the latter species often begin after chaetiger 10 or from median chaetigers. Nereis yuedensis n. sp. also differs in the distinct arrangement of lateral teeth of the notopodial falcigers, the absence of notopodial dorsal ligules, and the arrangement of paragnaths (Table 3). Nereis yuedensis n. sp. differs from N. cockburnensis. The first notopodial homogomph falcigers of both species appear at chaetiger 3, but those of the latter species are present with serrated lateral teeth and the remaining falcigers are present with a long terminal tooth and much smaller teeth basally. In addition, N. cockburnensis has more paragnaths on both maxillary and oral rings and well-developed dorsal notopodial ligules throughout chaetigers. Hartmann-Schröder (1982) assigned a nereidid specimen having notopodial homogomph falcigers at least from chaetiger 5 with 3–4 teeth from Jurien Bay (about 25 km from the type locality of N. yuedensis n. sp.) to Nereis cf. jacksoni, yet the arrangement of paragnaths of the specimen was not described. The characteristics of notopodial homogomph falcigers of that specimen is similar to those of N. yuedensis n. sp, and thus they are probably same species. Etymology. The species is named in honour of the Aboriginal people of the Dynamite Bay area, the Yued (Yued is also the name of the region and dialect group), who are part of the broader group of Noongar Aboriginal Australians who have lived in the southwest corner of Western Australia for approximately 40,000 years. The Yued name for the bay is Cuencandjaa, meaning ‘mouthful of sand’. Distribution. Dynamite Bay, Green Head, Western Australia. Although records are limited, at this stage the new species is possibly endemic to temperate Australia realm (Table 2). Habitat. Rocky shores, intertidal (Table 2).Published as part of Hadiyanto, Hadiyanto, 2023, Nereididae (Annelida: Phyllodocida) from intertidal macroalgae in Western Australia, pp. 151-203 in Zootaxa 5239 (2) on pages 172-173, DOI: 10.11646/zootaxa.5239.2.1, http://zenodo.org/record/762412
FIGURE 1 in Nereididae (Annelida: Phyllodocida) from intertidal macroalgae in Western Australia
FIGURE 1. Sampling sites of Nereididae on rocky intertidal shores of Western AustraliaPublished as part of Hadiyanto, Hadiyanto, 2023, Nereididae (Annelida: Phyllodocida) from intertidal macroalgae in Western Australia, pp. 151-203 in Zootaxa 5239 (2) on page 153, DOI: 10.11646/zootaxa.5239.2.1, http://zenodo.org/record/762412
FIGURE 17. Pseudonereis anomala Gravier, 1899 in Nereididae (Annelida: Phyllodocida) from intertidal macroalgae in Western Australia
FIGURE 17. Pseudonereis anomala Gravier, 1899 (WAM V11690); A, proboscis, ventral dissection; B, parapodium, chaetiger 22, posterior view; C, notochaetae, homogomph spiniger, chaetiger 22; D, notochaetae, homogomph falciger, chaetiger 22; E, neurochaetae, dorsal fascicle, heterogomph falciger, chaetiger 22; F, neurochaetae, ventral fascicle, heterogomph spiniger, chaetiger 32. Scale: A, 0.5 mm; B, 0.1 mm; C–F, 0.01 mm.Published as part of Hadiyanto, Hadiyanto, 2023, Nereididae (Annelida: Phyllodocida) from intertidal macroalgae in Western Australia, pp. 151-203 in Zootaxa 5239 (2) on page 182, DOI: 10.11646/zootaxa.5239.2.1, http://zenodo.org/record/762412
FIGURE 7. Nereis cockburnensis Augener, 1913 in Nereididae (Annelida: Phyllodocida) from intertidal macroalgae in Western Australia
FIGURE 7. Nereis cockburnensis Augener, 1913 (WAM V11637); A, head, dorsal view; B, head, ventral view; C, parapodium, chaetiger 47, anterior view; D, notochaetae, homogomph spiniger, chaetiger 4; E, notochaetae, homogomph falciger, chaetiger 3; F, notochaetae, homogomph falciger, chaetiger 47; G, neurochaetae, ventral fascicle, heterogomph spiniger, chaetiger 47; H, neurochaetae, ventral fascicle, heterogomph falciger, chaetiger 47. Scale: A–B, 1 mm; C, 0.1 mm; D–H, 0.02 mm.Published as part of Hadiyanto, Hadiyanto, 2023, Nereididae (Annelida: Phyllodocida) from intertidal macroalgae in Western Australia, pp. 151-203 in Zootaxa 5239 (2) on page 165, DOI: 10.11646/zootaxa.5239.2.1, http://zenodo.org/record/762412
FIGURE 6 in Nereididae (Annelida: Phyllodocida) from intertidal macroalgae in Western Australia
FIGURE 6. Nereis bifida Hutchings & Turvey, 1982 (WAM V11631); A, head, dorsal view; B, head, ventral view; C, parapodium, chaetiger 18, anterior view; D, notochaetae, homogomph falciger, chaetiger 21; E, neurochaetae, dorsal fascicle, homogomph spiniger, chaetiger 18; F, neurochaetae, ventral fascicle, heterogomph spiniger, chaetiger 18; G, neurochaetae, ventral fascicle, heterogomph falciger, chaetiger 10. Scale: A–B, 1 mm; C, 0.05 mm; D–G, 0.02 mm.Published as part of Hadiyanto, Hadiyanto, 2023, Nereididae (Annelida: Phyllodocida) from intertidal macroalgae in Western Australia, pp. 151-203 in Zootaxa 5239 (2) on page 163, DOI: 10.11646/zootaxa.5239.2.1, http://zenodo.org/record/762412
FIGURE 8. Nereis denhamensis Augener, 1913 in Nereididae (Annelida: Phyllodocida) from intertidal macroalgae in Western Australia
FIGURE 8. Nereis denhamensis Augener, 1913 (WAM V11644); A, head, dorsal view; B, proboscis, ventral dissection; C, parapodium, chaetiger 41, posterior view; D, notochaetae, homogomph falciger, chaetiger 41; E, neurochaetae, dorsal fascicle, homogomph spiniger, chaetiger 41; F, neurochaetae, ventral fascicle, heterogomph spiniger, chaetiger 41; G, neurochaetae, ventral fascicle, heterogomph falciger, chaetiger 41. Scale: A–B, 1 mm; C, 0.1 mm; D–G, 0.02 mm.Published as part of Hadiyanto, Hadiyanto, 2023, Nereididae (Annelida: Phyllodocida) from intertidal macroalgae in Western Australia, pp. 151-203 in Zootaxa 5239 (2) on page 167, DOI: 10.11646/zootaxa.5239.2.1, http://zenodo.org/record/762412
FIGURE 10. Nereis heirissonensis Augener, 1913 in Nereididae (Annelida: Phyllodocida) from intertidal macroalgae in Western Australia
FIGURE 10. Nereis heirissonensis Augener, 1913 (WAM V11660); A, proboscis, ventral dissection; B, parapodium, chaetiger 34, posterior view; C, notochaetae, homogomph falciger, chaetiger 39; D, notochaetae, homogomph falciger, chaetiger 56; E, neurochaetae, dorsal fascicle, homogomph spiniger, chaetiger 38; F, neurochaetae, ventral fascicle, heterogomph spiniger, chaetiger 59; G, neurochaetae, ventral fascicle, heterogomph falciger, chaetiger 12. Scale: A, 0.25 mm; B, 0.1 mm; C–G, 0.02 mm.Published as part of Hadiyanto, Hadiyanto, 2023, Nereididae (Annelida: Phyllodocida) from intertidal macroalgae in Western Australia, pp. 151-203 in Zootaxa 5239 (2) on page 171, DOI: 10.11646/zootaxa.5239.2.1, http://zenodo.org/record/762412
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