On the proposed conservation of Polydora websteri Hartmann in Loosanoff & Engle, 1943 (Annelida, Polychaeta) by a ruling that it is not to be treated as a replacement name for P. caeca Webster, 1879, and designation of a lectotype for P. websteri

Volume: 57Start Page: 110End Page: 11

Polydora websteri Hartman in Loosanoff & Engle, 1943 (Annelida, Polychaeta): proposed conservation of the specific name by a ruling that it is not to be treated as a replacement for P. caeca Webster, 1879, and designation of a lectotype for P. websteri

Volume: 55Start Page: 212End Page: 21

Molecular analysis of six Rhynchospio Hartman, 1936 species (Annelida: Spionidae) with comments on the evolution of brooding within the group

Radashevsky, Vasily I., Pankova, Victoria V., Nuzhdin, Sergey V. (2016): Molecular analysis of six Rhynchospio Hartman, 1936 species (Annelida: Spionidae) with comments on the evolution of brooding within the group. Zootaxa 4127 (3): 579-590, DOI: 10.11646/zootaxa.4127.3.1

Pseudopolydora (Annelida: Spionidae) from the Arabian Gulf, Kuwait

Seven species of Pseudopolydora are described and illustrated from the Arabian Gulf, Kuwait: P. achaeta Radashevsky & Hsieh, 2000, P. antennata (Claparède, 1868), P. arabica Radashevsky & Al-Kandari, 2020, P. auha sp. nov., P. kuwaiti sp. nov., P. melanopalpa sp. nov., and P. multispinosa sp. nov. The morphology of the developed planktonic larvae is described for P. antennata and P. kuwaiti sp. nov. Adults of all species live in tubes in soft sediments, while adults of P. kuwaiti sp. nov. also bore in shells of gastropods and dead corals encrusted by coralline algae. Pseudopolydora antennata and P. arabica form dense settlements up to 50 000 individuals per 1 m2, while other species are comparatively rare. The phylogenetic relationships between the examined species and other Pseudopolydora (18 species in total) were assessed in an analysis of sequence data of four gene fragments: mitochondrial 16S rDNA, nuclear 18S rDNA and 28S rDNA, and Histone 3 (2473 bp in total). Three species complexes are distinguished involving P. antennata, P. diopatra Hsieh, 1992 and P. paucibranchiata (Okuda, 1937). Sets of adult morphological features shared by species of each complex and an identification key to Pseudopolydora species from the Arabian Gulf are provided

Pseudopolydora melanopalpa Radashevsky & Malyar & Pankova 2021, sp. nov.

Pseudopolydora melanopalpa sp. nov. urn:lsid:zoobank.org:act: DA98F3F8-54B1-4CCC-BC82-16C3ABF0DC34 Figs 1D, 17–19 Diagnosis Prostomium anteriorly incised, with two short rounded lobes. Caruncle to end of chaetiger 5. Fine black pigment scattered on distal part of palps. Etymology The species name refers to fine black pigment scattered on the distal part of palps in adults. Material examined Holotype KUWAIT • 1 spec.; Arabian Gulf, Sulaibikhat Bay, desalinization station outfall area; 29.36157° N, 47.81915° E; 6 Feb. 2019; Vladimir A. Grintsov leg.; low intertidal; mud; MIMB 40908. Paratypes KUWAIT • 5 specs; same collection data as for holotype; MIMB 40909. Description All types: anterior fragments of females in good condition with palps; one 25-chaetiger fragment with a few small newly regenerated posterior achaetous segments with a small bilobed pygidium. Holotype: 39-chaetiger anterior fragment about 10 mm long, broken in the middle (Fig. 17A). Specimens up to 10 mm long, 1 mm wide with 45 chaetigers; when complete, worms apparently up to 15 mm long with at least 60 chaetigers. No pigmentation on body; fine black pigment scattered on distal part of palps (Figs 17A, C, E, 18C). Prostomium anteriorly incised, with two short rounded lobes (Figs 17C–E, 18A–B), posteriorly extending to end of chaetiger 5 (end of chaetiger 4 in holotype) as a low caruncle. Peristomium with two lateral lips as thick folds in front of large ventral lip (Fig. 17A). Short occipital antenna present on caruncle between palps (Fig. 18C). Two pairs of black eyes arranged trapezoidally, comprising one pair of median eyes, and one pair of slightly larger lateral eyes situated anteriorly and set wider apart. Palps as long as 10–15 chaetigers, with frontal longitudinal groove lined with fine cilia, and short compound non-motile cilia arising directly from palp surface sparsely arranged in line on sides of groove and sparsely scattered on lateral and abfrontal palp surfaces. Chaetiger 1 with short capillaries in neuropodia, small notopodial lamellae and well-developed neuropodial postchaetal lamellae; notochaetae absent. Chaetiger 2 notochaetae all slender capillaries with narrow limbation. Anterior-row notopodial capillaries on chaetiger 3 with slightly enlarged wing; capillaries on chaetiger 4 with wing slightly larger than on chaetigers 3. Anterior-row capillaries on chaetigers 3, 4, 6 and 7 arranged in J-shaped series; posterior-row capillaries on these chaetigers arranged in vertical series (Figs 17A–B, 18C). Posterior notopodia with a few long alimbate capillary chaetae. Chaetiger 5 same in size as chaetigers 4 or 6, with dorsal superior capillaries, two kinds of notopodial spines arranged in a double U-shaped row, and ventral capillaries; notopodial postchaetal lamellae absent; neuropodial lamellae same as on chaetigers 4 and 6 (Figs 17A–B, 18C). Dorsal superior capillaries shorter and fewer than those capillaries on chaetigers 4 and 6. Ventral capillaries same in size, number and arrangement (in three groups) as those on chaetigers 4 and 6. Upper posterior part of double U-shaped row of spines slightly lower than upper anterior part. Newly developed spines in posterior upper part of U-shaped row slightly larger than older spines in anterior upper part of row. Outer (anterior-row) notopodial spines up to 12 in a series, with distal part of stem enlarged, with concavity on top and large triangular tooth on its side directed upwards and facing towards the inside of the U-shaped row of spines; fine bristles arising from concavity forming long flag-like pointed transparent tip which usually broken in worn old spines in anterior upper part of row (Fig. 19A–B, D). Inner (posterior-row) notopodial spines up to nine in a series, falcate, with short rounded distal part geniculate, with subdistal bulbous swelling bearing very short fine bristles and facing towards the inside of the U-shaped row of spines (Fig. 19A, C–D). Hooks in neuropodia from chaetiger 8, up to 30 in a series, not accompanied by capillaries. Hooks bidentate, with upper tooth closely applied to main fang; upper part of shaft with constriction; lower part of shaft bent at right angle (Fig. 19E–F). Branchiae from chaetiger 7 to chaetiger 42, up to 36 pairs, on chaetiger 7 1.5–2 times as short as those on chaetiger 8, full-sized from chaetigers 10–11, fewer in small individuals, free from notopodial postchaetal lamellae, flattened, with surfaces oriented perpendicular to body axis, with longitudinal ciliation (extension of nototroch) on inner edge. Pygidium bilobed, with two semi-oval lateral lobes, white due to great number of spindle-shaped glandular cells with striated content. Subspherical to irregular shape glandular cells with striated content present on dorsal side of chaetigers. Cells few on anterior and posterior chaetigers, forming distinct paired gatherings from chaetiger 12 to chaetigers 30–35 (Figs 17D, 18A). Glandular pouches in neuropodia from chaetiger 1, largest and paired in each neuropodium in chaetigers 6 and 7, single in other neuropodia (Fig. 19E). Digestive tract without ventral buccal bulb and gizzard-like structure, without pigmentation. Nephridia from chaetiger 4 onwards. Distal parts of nephridia on female fertile chaetigers enlarged, containing glandular cells which absorbing MG and therefore well seen in fixed specimens from chaetiger 13 (Fig. 18A, E–F). Nephridia of chaetiger 13 used to release gametes from chaetiger 12. In female fertile chaetigers, paired nephridia on each chaetiger opening to exterior via common middorsal nephridiopore anteriorly to nototroch (Fig. 18F). MG staining Intensely stained anterior part of prostomium in front of eyes, dorso-lateral sides of peristomium, scattered glandular cells on dorsal side of chaetigers 7–11, and paired gatherings of glandular cells on dorsal side from chaetiger 12 to chaetigers 30–35. Weakly stained lateral sides of chaetigers; scattered glandular cells on ventral side of chaetigers (Figs 17–18). Habitat Adult P. melanopalpa sp. nov. inhabit silty tubes in the muddy intertidal. Reproduction All six type specimens of P. melanopalpa sp. nov. are females with oocytes from chaetiger 12 to chaetigers 32–40. Paired ovaries are attached to the segmental blood vessels in fertile chaetigers (Fig. 19G). The developed coelomic oocytes are about 120 µm in diameter, with smooth thin envelope less than 1 µm thick. Remarks Adult P. melanopalpa sp. nov. share a series of characters with members of the P. antennata -complex (see below in the Discussion). They differ, however, from other species of this complex by having black pigment on the distal part of palps, and the prostomium with two rounded lobed instead of pointed fronto-lateral extensions (see Simon et al. 2019: table 1). Distribution Arabian Gulf: Kuwait (Fig. 1D).Published as part of Radashevsky, Vasily I., Al-Kandari, Manal, Malyar, Vasily V. & Pankova, Victoria V., 2021, Pseudopolydora (Annelida: Spionidae) from the Arabian Gulf, Kuwait, pp. 120-168 in European Journal of Taxonomy 773 (1) on pages 152-156, DOI: 10.5852/ejt.2021.773.1519, http://zenodo.org/record/554446

Pseudopolydora achaeta Radashevsky & Hsieh 2000

Pseudopolydora achaeta Radashevsky & Hsieh, 2000 Fig. 3 Pseudopolydora achaeta Radashevsky & Hsieh, 2000: 223−226, figs 4–5, 11a. Pseudopolydora achaeta – Lana et al. 2006: 50. — Zvyagintsev et al. 2011: 53. — Abe et al. 2014: 3−5; 2016: 654−656, fig. 3; 2019: 6−11. — Bogantes et al. 2021: 581, fig. 2a–b. Pseudopolydora aff. achaeta – Abe & Sato-Okoshi 2021: 56−57, fig. 9a–b (larval morphology). Pseudopolydora sp. A – Radashevsky & Migotto 2006: fig. 1c. Description Two specimens were found in Sulaibikhat Bay, Kuwait, comprising 20-chaetiger anterior fragment of a small juvenile and a 70-chaetiger complete female about 16 mm long and 0.8 mm wide (MIMB 40934; Fig. 3A–B). Transverse bands of diffused black pigment present on dorsal side of up to 15 anterior chaetigers; small middorsal melanophores present from chaetigers 4−6 to chaetigers 10−12. Prostomium anteriorly weakly incised, notched or almost blunt, posteriorly extending to end of chaetiger 2 as a low caruncle. Occipital antenna present. Chaetiger 1 reduced, weakly separated from peristomium, with small notopodial and well developed neuropodial lamellae; notochaetae absent; neurochaetae comprising 1−5 very fine, hair-like capillaries. Chaetiger 5 same in size as chaetigers 4 or 6, with dorsal superior and ventral capillaries same in shape and number as those chaetae on chaetigers 4 or 6; two kinds of heavy spines arranged in a vertical slightly curved double row; noto- and neuropodial postchaetal lamellae present (Fig. 3 C−E). Anterior-row spines pennoned, with curved pointed tip, without subdistal constriction (Fig. 3F), up to 22 in a series; posterior-row spines simple falcate (Fig. 3G), up to 19 in a series. Bidentate hooded hooks in neuropodia from chaetiger 8, up to 15 in a series. Branchiae from chaetiger 7 to chaetiger 15. Pygidium flaring disc with wide dorsal gap and dorso-lateral processes (Fig. 3H–I). Glandular pouches in neuropodia from chaetiger 1, largest and paired in each neuropodium in chaetigers 6 and 7, single in other neuropodia. MG staining Intensely stained ventral and lateral sides and notopodial postchaetal lamellae of 15–16 anterior chaetigers, outer edges of branchiae (Fig. 3C); narrow transverse bands on dorsal side of branchiate chaetigers. Remarks Pseudopolydora achaeta was originally described from the South China Sea, Taiwan, as a common polychaete inhabiting tubes in soft sediments in brackish-water environments (Radashevsky & Hsieh 2000). Since then, the species was reported from Paraná and São Paulo (Brazil) (Lana et al. 2006; Radashevsky & Migotto 2006), Sea of Japan (Russia) (Zvyagintsev et al. 2011), Pacific side of Honshu Island (Japan) (Abe et al. 2014, 2016, 2019; Abe & Sato-Okoshi 2021), and from Florida (USA) (Bogantes et al. 2021). Here, for the first time, we report it for the Arabian Gulf (Kuwait). The only female had small oocytes up to 65 µm in diameter developing from chaetiger 15 onwards (Fig. 3J). The morphology of the specimens fits the diagnostic characters of P. achaeta. The same pattern of methylene green staining was observed in the type specimens of P. achaeta from Taiwan (MIMB 3401). The wide distribution of this species outside of its native area in the Northwest Pacific can be explained by unintentional human-mediated transportations of larvae with ballast water of ships, followed by successful invasions. Distribution South China Sea: Taiwan; Japan: Pacific side of Honshu Island; Brazil: Paraná, São Paulo; Atlantic USA: Florida; Arabian Gulf: Kuwait.Published as part of Radashevsky, Vasily I., Al-Kandari, Manal, Malyar, Vasily V. & Pankova, Victoria V., 2021, Pseudopolydora (Annelida: Spionidae) from the Arabian Gulf, Kuwait, pp. 120-168 in European Journal of Taxonomy 773 (1) on pages 128-130, DOI: 10.5852/ejt.2021.773.1519, http://zenodo.org/record/554446

Pseudopolydora Czerniavsky 1881

Genus Pseudopolydora Czerniavsky, 1881 Pseudopolydora Czerniavsky, 1881: 362. Carazzia Mesnil, 1896: 227. Polydora (Carazzia) – Fauvel 1927: 48; 1953: 316. Polydora (Pseudopolydora) – Hartmann-Schröder 1971: 317; 1996: 322. Pseudopolydora – Fauchald 1977: 25. — Blake & Kudenov 1978: 267. — Blake 1996: 202. — Blake et al. 2020: 79−80. List of species of Pseudopolydora from the Arabian Gulf and their type localities Pseudopolydora achaeta Radashevsky & Hsieh, 2000, South China Sea, Taiwan. Pseudopolydora antennata (Claparède, 1868). Tyrrhenian Sea, Gulf of Naples, Italy. Pseudopolydora arabica Radashevsky & Al-Kandari, 2020. Arabian Gulf, Al-Salmiya, Kuwait. Pseudopolydora auha sp. nov. Arabian Gulf, Auha Is., Kuwait. Pseudopolydora kuwaiti sp. nov. Arabian Gulf, Al-Judailiat, Kuwait. Pseudopolydora melanopalpa sp. nov. Arabian Gulf, Sulaibikhat Bay, Kuwait. Pseudopolydora multispinosa sp. nov. Arabian Gulf, Al-Salmiya, Kuwait. Identification key to Pseudopolydora from the Arabian Gulf 1. Chaetiger 5 distinctly larger than chaetiger 4. Chaetiger 5 spines arranged in a curved diagonal double row. Occipital antenna absent. Yellow bands regularly arranged on palps (invisible after fixation). Caruncle to end of chaetiger 2. Tube-dwelling or shell-boring............ P. kuwaiti sp. nov. – Chaetiger 5 same in size as chaetiger 4. Chaetiger 5 spines arranged in a slightly curved, J- or U-shaped vertical double row. Occipital antenna present. Yellowish chromatophores present or absent on palps in live. Caruncle to end of chaetiger 2 or longer. Tube-dwelling............................ 2 2. Chaetiger 5 spines arranged in a straight or slightly curved vertical double row. Caruncle to end of chaetiger 2. Black pigment present on dorsal side of 10−15 anterior chaetigers............................................................................................................................ P. achaeta Radashevsky & Hsieh, 2000 – Chaetiger 5 spines arranged in a J- or U-shaped vertical double row. Caruncle extending beyond chaetiger 2. Black pigment absent on dorsal side of anterior chaetigers.......................................... 3 3. Prostomium anteriorly entire, rounded. Chaetiger 5 spines arranged in a J-shaped double row. Yellowish chromatophores regularly arranged on palps (invisible after fixation). Caruncle to end of chaetiger 4. Pygidium disc-like to cup-shaped with only dorsal gap....................................................................................................................................... P. arabica Radashevsky & Al-Kandari, 2020 – Prostomium anteriorly incised to bifurcated. Chaetiger 5 spines arranged in a U-shaped double row. Yellowish chromatophores absent on palps. Caruncle to end of chaetiger 4 or longer. Pygidium bilobed with dorsal and ventral clefts............................................................................................... 4 4. Fine black pigment scattered on distal end of palps. Prostomium anteriorly incised, with two rounded lobes............................................................................................................. P. melanopalpa sp. nov. – No pigmentation on palps. Prostomium with two rounded lobes or long pointed antero-lateral processes........................................................................................................................................... 5 5. Chaetiger 5 with up to 32 spines in the outer (anterior) row of notochaetae, and 27 spines in the inner (posterior) row of notochaetae; notopodial postchaetal lamellae same as on chaetiger 4 or 6. MG staining of formalin-fixed specimens drastically intense on ventral side from chaetiger 8 onwards. Notopodial anterior-row spines of chaetiger 5 with distal part of stem curved, gradually narrowing, with wide, distally pointed limbation.......................................................... P. multispinosa sp. nov. – Chaetiger 5 with fewer spines in notopodia; notopodial postchaetal lamellae reduced. MG staining on ventral side of formalin-fixed specimens comparatively weak, gradually changing along body. Notopodial anterior-row spines of chaetiger 5 with distal part of stem enlarged, with concavity on top and large triangular tooth on its side directed upwards; fine bristles arising from concavity forming long flag-like pointed geniculate transparent tip................................................................ 6 6. Caruncle to end of chaetiger 4......................................................... P. antennata (Claparède, 1868) – Caruncle to end of chaetiger 6.................................................................................. P. auha sp. nov.Published as part of Radashevsky, Vasily I., Al-Kandari, Manal, Malyar, Vasily V. & Pankova, Victoria V., 2021, Pseudopolydora (Annelida: Spionidae) from the Arabian Gulf, Kuwait, pp. 120-168 in European Journal of Taxonomy 773 (1) on pages 127-128, DOI: 10.5852/ejt.2021.773.1519, http://zenodo.org/record/554446

Pseudopolydora arabica Radashevsky & Al-Kandari 2020

Pseudopolydora arabica Radashevsky & Al-Kandari, 2020 Figs 1B, 5D, 10 Pseudopolydora arabica Radashevsky & Al-Kandari, 2020: 2−10, figs 2−7. Pseudopolydora sp. B – Al-Kandari et al. 2019: 9. — Radashevsky et al. 2020: table 1, fig. 1. Pseudopolydora paucibranchiata – Swaleh & Mustaquim 1993: 204, fig. 2. — Not Okuda 1937. Adult diagnostic features Adults up to 20 mm long, 1 mm wide with 80 chaetigers (Fig. 10A). Live individuals with yellowishwhite pigment on lateral sides of prostomium and dorsal side of 3−5 anterior chaetigers (most intense on chaetigers 1−2), and up to 50 yellowish-white ramified chromatophores on each palp in (Fig. 10A– B); chromatophores not visible after fixation. Prostomium anteriorly narrow and rounded (Fig. 10B). Caruncle extending to end of chaetiger 4. Occipital antenna present. Chaetiger 1 with short capillaries in neuropodia; notochaetae absent. Anterior-row capillaries in notopodia from chaetiger 7 to chaetigers 10−20 with wide subtriangular, pennoned limbation. Chaetiger 5 almost as same in size as chaetigers 4 and 6 (Fig. 10B); dorsal superior capillaries slightly shorter and fewer than those capillaries on chaetigers 4 and 6; ventral capillaries as same as those on chaetiger 4; two kinds of notopodial spines arranged in a double J-shaped row; outer (anterior-row) notopodial spines with geniculate distal tip with wide limbation; inner (posterior-row) notopodial spines simple falcate (Fig. 10C). Bidentate hooded hooks in neuropodia from chaetiger 8. Branchiae from chaetiger 7 to chaetiger 30, up to 24 pairs. Pygidium small, cup-shaped, with dorsal gap. Glandular pouches in neuropodia from chaetiger 1, largest in chaetigers 6 and 7, single pouch in each neuropodium. MG staining Intensely stained dorsal side of the prostomium in front of eyes, dorsal side of the peristomium, and ventral side of chaetigers: complete transverse paired bands moderate intense on chaetigers 1–4, very intense complete transverse paired bands on chaetigers 5–7, and paired transverse bands split into three parts from chaetiger 8 onwards (Fig. 10D–G). Remarkably, the three-parts banding was associated with the start of the hooded hooks in neuropodia and was invariable in all examined specimens. Remarks Pseudopolydora arabica Radashevsky & Al-Kandari, 2020 was originally described from the Arabian Gulf as one of the common polychaetes inhabiting soft sediments. The population density of the species in some places reaches 50 000 individuals per 1 m 2. Distribution Arabian Gulf: Kuwait (Fig. 1B).Published as part of Radashevsky, Vasily I., Al-Kandari, Manal, Malyar, Vasily V. & Pankova, Victoria V., 2021, Pseudopolydora (Annelida: Spionidae) from the Arabian Gulf, Kuwait, pp. 120-168 in European Journal of Taxonomy 773 (1) on pages 138-140, DOI: 10.5852/ejt.2021.773.1519, http://zenodo.org/record/554446

Searching for a Home Port in a Polyvectic World: Molecular Analysis and Global Biogeography of the Marine Worm <i>Polydora hoplura</i> (Annelida: Spionidae)

The spionid polychaete Polydora hoplura Claparède, 1868 is a shell borer widely occurring across the world and considered introduced in many areas. It was originally described in the Gulf of Naples, Italy. Adult diagnostic features are the palps with black bands, prostomium weakly incised anteriorly, caruncle extending to the end of chaetiger 3, short occipital antenna, and heavy sickle-shaped spines in the posterior notopodia. The Bayesian inference analysis of sequence data of four gene fragments (2369 bp in total) of the mitochondrial 16S rDNA, nuclear 18S, 28S rDNA and Histone 3 has shown that worms with these morphological features from the Mediterranean, northern Europe, Brazil, South Africa, Australia, Republic of Korea, Japan and California are genetically identical, form a well-supported clade, and can be considered conspecific. The genetic analysis of a 16S dataset detected 15 haplotypes of this species, 10 of which occur only in South Africa. Despite the high genetic diversity of P. hoplura in South Africa, we tentatively propose the Northwest Pacific, or at the most the Indo–West Pacific, as its home region, not the Atlantic Ocean or the Eastern Pacific Ocean. The history of the discovery of P. hoplura around the world appears to be intimately linked to global shipping commencing in the mid-19th century, followed by the advent of the global movement of commercial shellfish (especially the Pacific oyster Magallana gigas) in the 20th century, interlaced with continued, complex dispersal by vessels and aquaculture. Given that P. hoplura has been detected in only a few of the 17 countries where Pacific oysters have been established, we predict that it may already be present in many more regions. As global connectivity through world trade continues to increase, it is likely that novel populations of P. hoplura will continue to emerge