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Dipolydora spinosa sp. nov. http://zoobank.org:act: CE6B878F-E52F-4B92-866F-456FF0FEA34A (Figs 4−10) Type material. South-China Sea, Vietnam, Gulf of Thailand, Phu Quoc Is., 0.5 m, in shell of saddle oyster Placuna ephippium (Philipsson), 3 Feb 1986, MIMB 42712 (paratype). Côn Dao Is., 5 m, in shell of the oyster Lopha cristagalli (Linnaeus), 19 Feb 1986, MIMB 42713 (paratype). Nha Trang Bay: Dung Is., 10 m, in shells of barnacle, Chama sp. and Millepora dichotoma Forsskål, 2 Jun 2006, MIMB 42714 (8 paratypes); Mun Is., 10 m, in hump coral Porites lutea Milne Edwards & Haime, 7 Jun 2006, 42715 (paratype); Dung Is., 10 m, in empty shell of the oyster Lopha cristagalli, 10 Jun 2006, 42716 (paratype); Dung Is., 10 m, in hump coral Porites lutea, 20 Jun 2006, 42717 (18 paratypes); Mun Is., 10 m, in hump coral Porites lutea, 30 Jun 2006, 42718 (21 paratypes); Tre Is., 5 m, in shell of saddle oyster Placuna ephippium, 26 Apr 2021, 42719 (2 paratypes); Tre Is., 3 m, in dead coral encrusted by coralline alga, 30 Apr 2021, 42720 (2 paratypes), 42721 (holotype). Complete data on material examined is provided in Supplementary Table S2. Adult morphology. Adults up to 35 mm long, 1 mm wide for 140 chaetigers. Holotype originally 25 mm long, 0.9 mm wide for 107 chaetigers; anterior and posterior parts fixed in formalin; 35 middle chaetigers fixed in ethanol for molecular analysis. Body light tan in life. Juveniles to about 70-chaetiger stage with remains of larval pigmentation on 15−22 anterior chaetigers, comprising paired dorsal and dorso-lateral melanophores, median dorsal vesiculate melanophores from chaetiger 3, lateral melanophores from chaetiger 2, and small paired ventral melanophores from chaetiger 3; black pigment between eyes, and one pair of melanophores in lateral lips of peristomium (Figs 4A, B, 10D). Prostomium anteriorly bifurcated (Figs 4A, 5A, B), posteriorly extending to middle of chaetiger 5 as a low caruncle, shorter in small individuals (Fig. 6A) (over 1/3 of chaetiger 5 in holotype). Low occipital antenna present on caruncle at level of anterior part of chaetiger 1 (Fig. 7A). One to two small black eyes present in small juveniles; large individuals without eyes. Palps as long as 15−20 chaetigers, with frontal longitudinal groove lined with short cilia, latero-frontal motile compound cilia, short compound non-motile cilia arising directly from palp surface and sparsely scattered on lateral and abfrontal palp surfaces. Chaetiger 1 with capillary chaetae and postchaetal lamellae in both rami; lamellae same as on chaetiger 2; notochaetae as long as but fewer than those on chaetiger 2; neurochaetae shorter than those on chaetiger 2. Juveniles to 30−50-chaetiger stage with only capillary chaetae in posterior notopodia; larger individuals usually with three, occasionally two or four awl-like spines in addition to 1−5 notopodial capillaries in a tuft from chaetigers 28−52 (Figs 6B, 7 E−H, 8K, L). Spines almost straight, with distal end slightly protruding out of body wall, smooth surface and basal end blunt (Fig. 8K). Chaetiger 5 twice as large as chaetigers 4 and 6, on each side with up to six geniculate dorsal superior capillaries (Fig. 8F), six falcate spines alternating with bilimbate-tipped companion chaetae (Fig. 8 A−D) and arranged in oblique slightly curved row, and six ventral capillaries (Fig. 8E). Dorsal superior and ventral capillaries winged, shorter and fewer than those chaetae on chaetigers 4 and 6. Falcate spines with spoon-like hollow on subdistal concave side and narrow transverse shelf on subdistal convex side (Fig. 8B, D); transverse shelf apparently worn and therefore not seen on spines of some individuals. Hooded hooks in neuropodia from chaetiger 7, up to six in a series, accompanied by 1−2 inferior winged capillaries (Fig. 8G) until chaetigers 10−12, and alternating with 1−2 hair-like alimbate capillaries (Fig. 8I) in 3−10 posterior neuropodia. Hooks bidentate, with slightly curved shaft without constriction (Fig. 8H, J). Branchiae usually from chaetiger 8 (on chaetigers 8−88 in holotype), occasionally from chaetiger 9 (3 of 26 examined paratypes), full-sized from chaetigers 10−11, absent from posterior 1/3−1/4 part of body (Fig. 6B), free from notopodial postchaetal lamellae, flattened, with surfaces oriented parallel to body axis. Nototrochs in juveniles and females from chaetigers 9−13, in males from chaetiger 1, all composed by single rows of ciliated cells. Intersegmental transverse ciliation present on anterior chaetigers in males as single rows of ciliated cells along anterior edge of chaetigers, from chaetiger 2 to chaetigers 10−12. Pygidium fleshy, cup-shaped with dorsal gap to incision, white due to numerous fusiform glandular cells (Figs 7 E−G, 9A). Glandular pouches in neuropodia from chaetiger 6, large from chaetiger 7 to chaetigers 10−12 and then gradually diminishing in size. Digestive tract with gizzard-like structure beginning from chaetigers 13−29 (from chaetiger 21 in holotype) and extending through 1−2 chaetigers (Figs 5C, 6A). Gizzard composed of anterior transparent muscular part and posterior white, apparently secretory, part. Some grain-like structures present in wall of muscular part of some individuals (Fig. 5D; these may represent fragmented “teeth” of the gizzard as discussed in Blake 1971, Fig. 14g therein). Rectum white in 5−15 posterior most chaetigers. Nephridia from chaetiger 7, opening to exterior via two nephridiopores situated on lateral sides in anterior sterile chaetigers and on dorsal side in fertile chaetigers (Fig. 10E). MG staining. Paired blotches (gatherings of strongly stained, probably glandular, cells) appeared on the dorsal side from chaetigers 12–17 onwards (Fig. 10A, B, E), except 10−30 posterior most chaetigers. Narrow transverse bands were weakly stained on the ventral side from chaetiger 7 to chaetigers 10–12 (Fig. 10C). Prostomium and the peristomium did not absorb the dye, but the foregut was evenly colored (Fig. 10C, D). Habitat. Dipolydora spinosa sp. nov. is an opportunistic borer, making U-shaped burrows in shells of the saddle oyster Placuna ephippium, cockscomb oyster Lopha cristagalli (Linnaeus), jewel box clam Chama sp., barnacles, in the hump coral Porites lutea Milne Edwards & Haime, and the net fire coral Millepora dichotoma Forsskål. Up to three worms were found per one square cm of coral surface and hundreds of worms were occasionally present in corals of about 20 cm in diameter. Reproduction. Dipolydora spinosa sp. nov. is dioecious. Paired gonads are attached to segmental blood vessels from chaetigers 23−42 to chaetigers 50−82 (n=15) (Fig. 9A). Smallest mature individuals were about 17 mm long for 70 chaetigers. Holotype is a female with oocytes in chaetigers 28−82. Oogenesis is mainly intraovarian. Developed oocytes accumulate in the coelomic cavity prior to spawning. The coelomic oocytes were subspherical to slightly oval, 100−115 µm in diameter. In males, spermatogonia proliferate in testes; spermatogenesis occurs in the coelomic cavity. Spermatids are joined in octads (Fig. 8M). Spermatozoa are introsperm about 1 µm in diameter, with a pointed acrosome 1.5 µm long, nucleus 7.5 µm, middlepiece 6 µm, head+middlepiece 15 µm long, and flagellum 50 µm long. A small colony of about 20 individuals in the hump coral Porites sp. collected from Nha Trang Bay in June 2006 contained only males. One of them had seven chaetigers regenerating anteriorly, middle stock chaetigers, and a few chaetigers regenerating posteriorly; regenerating chaetigers were slightly smaller and more transparent than fully developed stock chaetigers. This colony likely appears to have developed due to asexual reproduction by architomy from a single settled larva of male gender. It seems likely that after settlement, individuals first undergo a series of architomic divisions and only after that mature and start reproducing sexually. Such colony formation has been found in other polydorin species that exhibit architomy (Radashevsky, pers. obs.) and will be the subject of future studies. Larval development. Females lay eggs into transparent capsules which join to each other in a string and each attach by two short stalks to the inner wall of the burrow (Fig. 9C). Up to 60 eggs were deposited in one capsule and up to 55 capsules with about 3000 eggs were produced by one female per brood; small females had smaller broods. The laid eggs were 105−120 µm in diameter, with smooth envelope less than 1 µm thick. Females brooding larvae in capsules had the next generation of vitellogenic oocytes developing in the ovaries. Most of the eggs in the broods developed into larvae, but about 10% of eggs were abortive (it is not known whether these result from sperm limitation or represent nurse egg development; see Smith & Gibson 1999). The egg capsules with trochophores and abortive eggs were found in Nha Trang Bay in June 2006 (Fig. 9C, D). The trochophores had a few very small trochoblasts bearing short cilia of the prototroch, and one pair of rounded ventral ciliary patches. They had no eyes and moved inside capsules due to active beating of the ciliary patches. The larvae had a small mouth and a short foregut lined with numerous fine cilia (Fig. 9D). The posterior end of the foregut was joined to a spherical mass of large yolky endodermal macromeres occupying most of the inner space of larvae. Lumen of midgut, posterior gut, and anus were not yet developed. The duration of larval development inside the capsules remains unknown, but compared to other polydorins producing similar broods (see Blake & Arnofsky 1999), D. spinosa sp. nov. larvae probably hatch after growing 3−5 chaetigers. Remarks. By having an occipital antenna on the prostomium, D. spinosa sp. nov. is similar to D. tentaculata (Blake & Kudenov, 1978) from New South Wales, Australia, and D. anatentaculata Delgado-Blas, 2008 from the Gulf of Mexico, Florida. Dipolydora spinosa sp. nov. and D. tentaculata have caruncles of similar maximal length, extending to the middle of chaetiger 5 (for D. anatentaculata, Delgado-Blas 2008: 8, fig. 2A described the “caruncle extending posteriorly to segment 4” but illustrated it as reaching chaetiger 5). Dipolydora spinosa sp. nov. has a cup-shaped pygidium with only dorsal incision (which is comparatively rare among Dipolydora spp. that mainly have lobate pygidia with three or four lobes) while the pygidial shapes are unknown for D. tentaculata and D. anatentaculata. Dipolydora spinosa sp. nov. and D. anatentaculata are shell-borers while D. tentaculata lives in tubes in sediment. Dipolydora spinosa sp. nov. differs from these two species in the structure of the falcate spines of chaetiger 5. In D. tentaculata and D. anatentaculata, the spines are simple, without accessory structures, whereas in D. spinosa sp. nov. the spines have a spoon-like hollow on the concave side and a narrow transverse shelf on the convex side. Blake & Kudenov (1978: 250) noted that in D. tentaculata the notochaetae of the posterior chaetigers “including long slender capillaries and a stouter spine.” On the illustration (Blake & Kudenov 1978: fig. 39c), however, the chaeta resembles thick anterior-row capillary notochaeta rather than a genuine spine (see Discussion below). Adults of D. spinosa sp. nov. have 2−4 (usually three) awl-like spines in addition to 1−5 capillaries in a tuft in the posterior notopodia; the spines are almost straight, with basal end blunt. Etymology. The species name, feminine for Latin spinosus (spiny, thorny), refers to the awl-like spines in the posterior notopodia of adults. Distribution. South-China Sea, Vietnam (see Table S2, Fig. 1A, B)., Published as part of Radashevsky, Vasily I., 2022, Three new species of shell-boring Dipolydora (Annelida: Spionidae: Polydorini) from the South China Sea and the Gulf of Thailand, Vietnam, with comments on the modified spines in posterior notopodia and sperm morphology in polydorins, pp. 101-119 in Zootaxa 5162 (2) on pages 106-111, DOI: 10.11646/zootaxa.5162.2.1, http://zenodo.org/record/6809879, {"references":["Blake, J. A. (1971) Revision of the genus Polydora from the east coast of North America (Polychaeta: Spionidae). Smithsonian Contributions to Zoology, 75, 1 - 32.","Smith, H. L. & Gibson, G. D. (1999) Nurse egg origin in the polychaete Boccardia proboscidea (Spionidae). Invertebrate Reproduction and Development, 35 (3), 177 - 185. https: // doi. org / 10.1080 / 07924259.1999.9652384","Blake, J. A. & Arnofsky, P. L. (1999) Reproduction and larval development of the spioniform Polychaeta with application to systematics and phylogeny. Hydrobiologia, 402, 57 - 106. https: // doi. org / 10.1023 / A: 1003784324125","Blake, J. A. & Kudenov, J. D. (1978) The Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera. Memoirs of the National Museum of Victoria, 39, 171 - 280.","Delgado-Blas, V. H. (2008) Polydora and related genera (Polychaeta: Spionidae) from the Grand Caribbean region. Journal of Natural History, London, 42 (1 - 2), 1 - 19. https: // doi. org / 10.1080 / 00222930701831240"]}

Dipolydora echinata sp. nov. http://zoobank.org:act: E9F81CFA-BD86-433A-A1AF-7BC235544D5D (Figs 2 & 3) Type material. South China Sea, Vietnam, Nha Trang Bay: Dung Is., 10 m, in shell of jewel box clam Chama sp., 19 May 2006, MIMB 42700 (paratype); Tam Is., 10 m, in fire coral Millepora platyphylla Hemprich & Ehrenberg, 22 May 2006, MIMB 42701 (2 paratypes); Dung Is., 10 m, in shell of saddle oyster Placuna ephippium (Philipsson), 10 Jun 2006, MIMB 42702 (holotype). Complete data on material examined is provided in Supplementary Table S1. Adult morphology. Adults up to 20 mm long, 0.4 mm wide for 116 chaetigers; holotype 15 mm long, 0.3 mm wide for 113 chaetigers. Body light tan in life; distinct pigmentation absent. Prostomium bifurcated anteriorly, with shallow transverse fissure at level of chaetiger 1 separating narrow posterior caruncle extending to end of chaetiger 3 (Fig. 2A); caruncle shorter in small individuals. Anterior part of caruncle, just behind fissure, inflated, but occipital antenna absent. Eyes absent. Palps as long as 15−20 chaetigers, with frontal longitudinal groove lined with short cilia, latero-frontal motile compound cilia, and short compound non-motile cilia arising directly from palp surface and scattered on lateral and abfrontal palp surfaces. Chaetiger 1 with short capillaries and small postchaetal lamellae in both rami. Posterior notopodia (from chaetiger 36 in holotype) with up to six awl-like spines in addition to 1−5 slender capillaries in a tuft (Figs 2 M−O, 3A, B). In live individuals, spines entirely embedded into body; in fixed specimens, distal tips often slightly protruding out of notopodia. Distal ends of spines pointed and slightly curved. Partially developed spines with basal end truncate (Fig. 2M); fully developed spines double-edged, with basal and distal ends pointed (Fig. 2N). Chaetiger 5 larger than chaetigers 4 and 6, on each side with up to four geniculate dorsal superior capillaries (Fig. 2H), six heavy falcate spines alternating with bilimbate-tipped companion chaetae (Fig. 2 B−F) and arranged in an oblique slightly curved row, and seven ventral capillaries (Fig. 2G). Dorsal superior and ventral capillaries winged, shorter than those chaetae on chaetigers 4 and 6. Falcate spines with thin transverse subdistal collar on convex and lateral sides; lateral parts of collar usually prominent, resembling small lateral teeth or spurs (Fig. 2C, E, F). Hooded hooks in neuropodia from chaetiger 7, up to seven in a series, accompanied by 1−2 inferior winged capillaries (Fig. 2I) until chaetigers 8−10, and alternating with 1−2 hair-like alimbate capillaries (Fig. 2K) in 10−20 posterior neuropodia. Hooks bidentate, with slightly curved shaft without constriction; upper tooth well developed, at about right angle to main fang in anterior neuropodia (Fig. 2J), greatly reduced and closely applied to main fang in posterior neuropodia (Fig. 2L). Branchiae from chaetiger 7 to chaetiger 50 in 113-chaetiger holotype, to chaetiger 37 in a 85-chaetiger individual, and to chaetiger 42 in a 116-chaetiger individual; full-sized from chaetigers 11−14, free from notopodial postchaetal lamellae, flattened, with surfaces oriented parallel to body axis. Of two paratypes (MIMB 42701), one individual with branchiae from chaetiger 9, and another with branchiae from chaetiger 10. In these worms, first ten anterior chaetigers slightly smaller than succeeding chaetigers, suggesting recent regeneration likely after asexual reproduction by architomy or after a partial predation event. Branchiae from chaetiger 9 or 10 also in some other examined but not fixed individuals. Nototrochs from chaetigers 9−13, composed of single rows of ciliated cells. Intersegmental ciliation absent. Pygidium cup-shaped, with dorsal gap to incision, white due to numerous fusiform glandular cells (Fig. 3A, C). Glandular pouches in neuropodia from chaetiger 7, full-sized from chaetigers 7−8 and diminishing in size after chaetigers 10−15. Digestive tract with gizzard-like structure beginning from chaetigers 16−21 and extending through 1−2 chaetigers. Gizzard composed of anterior transparent muscular part and posterior white, apparently secretory, part. Rectum white in 3−5 posterior most chaetigers. Nephridia from chaetiger 7, opening to exterior via two nephridiopores situated on lateral sides in anterior sterile chaetigers and on dorsal side in fertile chaetigers. Habitat. Dipolydora echinata sp. nov. is an opportunistic borer, making U-shaped burrows in shells of the saddle oyster Placuna ephippium (Philipsson), razor clam Pinna cf. bicolor Gmelin, jewel box clam Chama sp., vase snail Vasum turbinellus (Linnaeus), spindle snail Polygona infundibulum (Gmelin) occupied by hermit crab, in fire coral Millepora platyphylla Hemprich & Ehrenberg, and under crust of coralline algae on shells of the penguin’s wing oyster Pteria penguin (Röding). Reproduction. Two mature individuals (holotype and a not fixed specimen) were males with sperm from chaetiger 38 through most of the body. Spermatogonia proliferated in testes; spermatogenesis occurred in the coelomic cavity. Spermatids were joined in tetrads (Fig. 2P). Spermatozoa were introsperm with elongated straight head about 1 µm in diameter, head+middlepiece 12 µm long, acrosome 3 µm, nucleus 5 µm, middlepiece 4 µm, and flagellum 38 µm long. Egg morphology and larval development of the species remain unknown. Remarks. Dipolydora echinata sp. nov. co-occurs in shells with D. spinosa sp. nov. (see below). Both species have bifurcated prostomia, awl-like spines in the posterior notopodia, and cup-shaped pygidia with only dorsal incision. They differ in that D. echinata sp. nov. has no occipital antenna, caruncle extending to the end of chaetiger 3, branchiae beginning on chaetiger 7, awl-like spines in the posterior notopodia double-edged, up to six in a series, and spermatids joined in tetrads, while D. spinosa sp. nov. has an occipital antenna, caruncle extending to the middle of chaetiger 5, branchiae beginning on chaetigers 8−9, awl-like spines in the posterior notopodia with basal part blunt, up to three in a series, and spermatids joined in octads. Moreover, D. spinosa sp. nov. has chaetiger 5 falcate spines with spoon-like hollow on subdistal concave side and a thin transverse shelf on convex side, while spines of D. echinata sp. nov. have no spoon-like hollow on concave side and have a wide collar on convex side with lateral parts prominent, resembling small teeth or spurs. By having a cup-shaped pygidium with only dorsal incision, D. echinata sp. nov. is similar to D. protuberata (Blake & Kudenov, 1978) from Victoria, Australia, and D. pilikia (Ward, 1981) from Hawaii. The species differ in that D. echinata sp. nov. has caruncle extending to the end of chaetiger 3, branchiae beginning on chaetiger 7 and lacking from the posterior half of the body, and chaetiger 5 falcate spines with transverse subdistal collar with lateral parts prominent, resembling small teeth or spurs, while D. protuberata and D. pilikia have caruncles extending beyond chaetiger 3, branchiae beginning on chaetigers 8−9 and continuing beyond the middle of the body, and chaetiger 5 falcate spines with a subterminal protuberance. Blake & Kudenov (1978: 250) and Ward (1981: 724) noted that D. protuberata and D. pilikia have “short acicular spines” in addition to capillaries in the posterior notopodia. These spines, however, were not illustrated for D. pilikia. Ward (1987: 354) also noted about D. pilikia: “This very common bioeroder in coral rock … has been collected from coral rock, coral sediment blocks, and sediment in Kaneohe Bay, from sand and rubble on the reef flat at Fort Kamehameha, and from sediment in Pearl Harbor, Oahu.” This description may be interpreted as some worms were living in tubes in soft sediments while others bored into corals. Dipolydora pilikia has never been redescribed and not reported again from Hawaii. Dipolydora protuberata was found only in tubes in soft sediments while D. echinata sp. nov. was found only in burrows in mollusc shells. Etymology. The species name, feminine for Latin echinatus (echinate, prickly), refers to the awl-like spines in the posterior notopodia of adults. Distribution. South-China Sea, Vietnam (see Table S1, Fig. 1A, B)., Published as part of Radashevsky, Vasily I., 2022, Three new species of shell-boring Dipolydora (Annelida: Spionidae: Polydorini) from the South China Sea and the Gulf of Thailand, Vietnam, with comments on the modified spines in posterior notopodia and sperm morphology in polydorins, pp. 101-119 in Zootaxa 5162 (2) on pages 103-105, DOI: 10.11646/zootaxa.5162.2.1, http://zenodo.org/record/6809879, {"references":["Blake, J. A. & Kudenov, J. D. (1978) The Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera. Memoirs of the National Museum of Victoria, 39, 171 - 280.","Ward, L. A. (1981) Spionidae (Polychaeta: Annelida) from Hawaii, with descriptions of five new species. Proceedings of the Biological Society of Washington, 94 (3), 713 - 730.","Ward, L. A. (1987) Family Spionidae. In: Devaney, D. M. & Eldredge, L. G. (Eds.), Bailey-Brock, J. H., Phylum Annelida. Bishop Museum Press, Honolulu, pp. 340 - 369."]}

Dipolydora vietnamita sp. nov. http://zoobank.org:act: EB418B1C-0919-4E41-AAF0-624C42743768 (Figs 11−14) Type material. South-China Sea, Vietnam, Gulf of Thailand, An Thoi Archipelago, Thom Is., 5 m, in shell of the gastropod Vasum turbinellus (Linnaeus) occupied by hermit crab, 4 Feb 1986, MIMB 42722 (paratype). Nha Trang Bay: Tam Is., 10 m, in dead coral encrusted by coralline alga, 22 May 2006, MIMB 42723 (4 paratypes); Dung Is., 10 m, in shell of Chama sp., 2 Jun 2006, MIMB 42724 (26 paratypes); Tre Is., eastern side, 15 m, in dead coral, 14 Jun 2006, MIMB 42725 (40 paratypes), 42726 (holotype); Nha Trang, pier fouling, 3 m, in coral on pillar, 11 Jul 2006, MIMB 42727 (25 paratypes); under crust of coralline alga on shell of Arca avellana Lamarck bought in Nha Trang market, 30 Apr 2021, MIMB 42728 (5 paratypes); in oyster shell bought in Nha Trang market, 30 Apr 2021, MIMB 42729 (paratype). Complete data on material examined is provided in Supplementary Table S3. Adult morphology. Adults up to 21 mm long, 0.55 mm wide for 125 chaetigers. Pigmentation absent on body; some individuals with fine black pigment diffused on sides of palp grooves. Prostomium anteriorly bifurcated, posteriorly extending to end of chaetiger 4 as a low caruncle, shorter in small individuals (Figs 11A, 12A). Eyes and occipital antenna absent. Palps as long as 15−20 chaetigers, with frontal longitudinal groove lined with fine cilia, latero-frontal motile compound cilia on sides of groove, short compound non-motile cilia arising directly from palp surface and scattered on lateral and abfrontal palp surfaces. Chaetiger 1 with short capillaries and small postchaetal lamellae in both rami. Posterior notopodia with only capillary chaetae. Chaetiger 5 larger than chaetigers 4 and 6, with up to five dorsal superior capillaries (Fig. 13F), five falcate spines alternating with bilimbate-tipped companion chaetae (Fig. 13 A−D) and arranged in oblique slightly curved row, and six ventral capillaries (Fig. 13E). Dorsal superior and ventral capillaries winged, shorter and fewer than those chaetae on chaetigers 4 and 6. Falcate spines with spoon-like hollow on subdistal concave side (Fig. 13B, D). Hooded hooks in neuropodia from chaetiger 7, up to five in a series, accompanied by 1−2 winged inferior capillaries until chaetigers 9−10 (Fig. 13G), and alternating with 1−2 hair-like alimbate capillaries in 5−10 posterior neuropodia (Fig. 13I). Hooks bidentate, with slightly curved shaft without constriction. Branchiae usually from chaetiger 9, occasionally from chaetigers 8 or 10, full-sized from chaetigers 12−14, very short after middle of body and absent from posterior 1/3−1/8 part of body (Fig. 12B), free from notopodial postchaetal lamellae, flattened, with surfaces oriented parallel to body axis. Nototrochs from chaetigers 6−7 in small individuals with up to about 40 chaetigers, from chaetigers 10−16 in larger worms, composed of single rows of ciliated cells in both sexes. Intersegmental ciliation absent. Pygidium usually thin, flaring, disc-like to cup-shaped, up to two times wider than posterior most chaetigers, trilobed, with one ventral lobe and two smaller dorsal lobes, white due to numerous fusiform glandular cells. Lateral clefts shallow and indistinct in some individuals, thus pygidium resembling a cup or a disc with only dorsal incision (Fig. 11D, E). Glandular pouches in neuropodia from chaetiger 6, full-sized from chaetigers 7−8, diminishing in size after chaetigers 10−13. Digestive tract with gizzard-like structure beginning from chaetigers 9−18 (Fig. 12A) and extending through 1−2 chaetigers. Gizzard composed of anterior transparent muscular part and posterior white, apparently secretory, part. Rectum white in 3−5 posterior most chaetigers. Nephridia from chaetiger 7, opening to exterior via two nephridiopores situated on lateral sides in anterior sterile chaetigers and on dorsal side in fertile chaetigers. MG staining. Weak staining appeared on the lateral and ventral sides of the peristomium; the foregut was evenly colored (Fig. 11F, G). Paired blotches (gatherings of strongly stained, probably glandular, cells) appeared on the dorsal side from chaetigers 19–22 onwards (Fig. 11C, F), except 10−20 posterior most chaetigers. Narrow transverse bands were weakly stained on the ventral side of chaetigers 6 and 11; wider bands of stronger staining were present on chaetigers 8–10 (Fig. 11B). In most specimens, peristomium and ventral side of chaetigers 6 and 11 were not stained, whereas in some specimens the dye was not absorbed on the ventral side of anterior chaetigers. In 20 of 21 stained specimens, the prostomium and about 20 anterior chaetigers did not absorb the dye (Fig. 11F), whereas in one specimen intense staining appeared on the dorsal side of prostomium and eight anterior chaetigers (Fig. 11A). Glandular pouches in neuropodia absorbed the dye and were well seen in lateral view from chaetigers 8−11 to chaetigers 25−30 (Fig. 11G). Habitat. Dipolydora vietnamita sp. nov. is an opportunistic borer, making burrows in corals, coralline algae, shells of the jewel box clam Chama sp., hazelnut ark Arca avellana Lamarck, and vase snail Vasum turbinellus (Linnaeus) occupied by hermit crabs. This species is common in the coastal waters of Vietnam, co-occurring with other shell-boring polydorins in the same shells and corals. Sometimes the worms occur in dense aggregations up to ten individuals per square centimeter, forming colonies of hundreds of worms in one shell and thousands of worms in a coral. Two worms were found boring in a shell of the date mussel Lithophaga sp. which, in turn, perforated the shell of the hazelnut ark Arca avellana (Fig. 14 A−C). Reproduction. Dipolydora vietnamita sp. nov. is dioecious. Paired gonads are attached to segmental blood vessels from chaetigers 21−24 through 2/3−3/4 of the body. Spermatogonia proliferate in testes; spermatogenesis occurs in the coelomic cavity. Spermatids are joined in tetrads (Fig. 13K). Spermatozoa are introsperm about 1 µm in diameter, with a pointed acrosome 3 µm long, nucleus 5 µm, middlepiece 4 µm, head+middlepiece 12 µm long, and flagellum 54 µm long. Egg morphology and larval development of the species remain unknown. Asexual reproduction. Many fragments of worms regenerating anterior and posterior chaetigers were found in shells from Nha Trang Bay in June 2006. Examination of those fragments has allowed for determining the course of the architomic asexual reproduction of D. vietnamita sp. nov. In the beginning, individuals divide into two or three stock fragments, each of which subsequently regenerates lost parts. The smallest stock fragment had six middle chaetigers. The anterior regeneration starts with a bud that grows and differentiates into head and body anlage. Palps begin to develop immediately after head differentiation. The body anlage elongates gradually and becomes segmented, probably simultaneously, into eight segments; no more segments are added anteriorly. Chaetae, postchaetal lamellae and other external segment attributes, as well as mouth and foregut, develop only when the segmentation is completed. Eyes never develop in the regenerated head. The posterior regeneration also starts with a bud that gives rise to the pygidium and the pre-pygidial growth zone. New posterior stolon segments develop successively following the formation of the growth zone. Similar records of architomy have been found in other spionids such as Pygospio elegans Claparède, 1863 (Rasmussen 1953) and Dipolydora armata (Radashevsky & Nogueira 2003). Remarks. Dipolydora vietnamita sp. nov. is similar to D. socialis (Schmarda, 1861) from Chile, and D. protuberata from New South Wales, Australia, by having subdistal protuberance on the falcate spines of chaetiger 5. However, the two latter species are tube-dwellers in soft sediments while D. vietnamita sp. nov. is a shell- and coral-borer. Dipolydora vietnamita sp. nov. and D. socialis have trilobed pygidia but differ in the arrangement of branchiae and composition of the gizzard-like structures. Dipolydora socialis has branchiae usually beginning from chaetiger 8 and gizzard with four chitinous plates attached to the inner wall of muscular part, while D. vietnamita sp. nov. has branchiae usually beginning from chaetiger 9 and gizzard without chitinous plates. Dipolydora vietnamita sp. nov. differs from D. protuberata by having trilobed pygidium instead of disc-like pygidium with a dorsal notch. Etymology. The species name refers to the type locality of the species in Vietnam. Distribution. South-China Sea, Vietnam (see Table S3, Fig. 1A, B)., Published as part of Radashevsky, Vasily I., 2022, Three new species of shell-boring Dipolydora (Annelida: Spionidae: Polydorini) from the South China Sea and the Gulf of Thailand, Vietnam, with comments on the modified spines in posterior notopodia and sperm morphology in polydorins, pp. 101-119 in Zootaxa 5162 (2) on pages 112-115, DOI: 10.11646/zootaxa.5162.2.1, http://zenodo.org/record/6809879, {"references":["Claparede, E. (1863) Beobachtungen uber Anatomie und Entwicklungsgeschichte wirbelloser Thiere an der Kuste von Normandie Angestellt. Wilhelm Engelmann, Leipzig, vii + 120 pp.","Rasmussen, E. (1953) Asexual reproduction in Pygospio elegans Claparede (Polychaeta sedentaria). Nature, 171, 1161.","Radashevsky, V. I. & Nogueira, J. M. M. (2003) Life history, morphology and distribution of Dipolydora armata (Polychaeta: Spionidae). Journal of the Marine Biological Association of the United Kingdom, 83 (2), 375 - 384. https: // doi. org / 10.1017 / S 0025315403007227 h","Schmarda, L. K. (1861) Neue wirbellose Thiere beobachtet und gesammelt auf einer Reise um die Erde 1853 bis 1857. Erster Band. Turbellarien, Rotatorien und Anneliden. Zweite Halfte. Verlag von Wilhelm Engelmann, Leipzig, 164 pp."]}

In this study we report a new record of a cryptogenic polychaete from southern Africa. The species was found inhabiting sand tubes in intertidal sand flats in the Knysna Estuary on the southern coast of South Africa. Morphological comparisons using light and scanning electron microscopy showed extensive taxonomic similarities with Dipolydora socialis described from other localities and from museum vouchers. In addition, 18S rRNA and COI barcodes were generated for the species. Genetic analysis of the assembled polydorid dataset corroborated the morphological data in delineating the species as a taxonomic unit with >99% genetic similarity to available sequences of D. socialis in the GenBank database. Dipolydora socialis has been reported as having a widespread distribution, and since it can reside within tubes associated with fouling communities or as a shell borer, several vectors may have been responsible for its global spread and introduction to southern Africa. Finally, considering the many cryptic complexes that are currently being uncovered within polychaetes, including spionids, future taxonomic studies should incorporate additional genetic data from other regions of the world to determine whether D. socialis may also be part of a larger species complex.

Aulacomya atra populations of the San Jose gulf, Northern Patagonia, Southwestern Atlantic Ocean, are infested by two polydorin species, Polydora rickettsi and Dipolydora cf. giardi. The infestation by these boring polychaetes causes the formation of pearls which is evidenced by the presence of capsules containing polydorin tissue debris and the elemental composition of organic material inside the pearls. Moreover, a positive relationship between the abundance of perforations of polydorin polychaetes and abundance of pearls was found by applying generalized lineal model analysis. These results constitute the first evidence of pearls formation due to infestation by polychaete. Fil: Diez, María Emilia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Centro Nacional Patagónico. Instituto de Biología de Organismos Marinos; Argentina Fil: Vázquez, Nuria Natalia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Centro Nacional Patagónico. Instituto de Biología de Organismos Marinos; Argentina Fil: Cremonte, Florencia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Centro Nacional Patagónico. Instituto de Biología de Organismos Marinos; Argentina

Genus Dipolydora Verrill, 1881 Dipolydora armata (Langerhans, 1880) DISTRIBUTION: NA. LITERATURE RECORDS: Fauvel 1938; Katzmann 1972; Lardicci 1989; Požar-Domac 1994; Casellato & Stefanon 2008; Castelli et al. 2008. OTHER REPORTED NAMES: Polydora armata Langerhans, 1880. Dipolydora coeca (��rsted, 1843) DISTRIBUTION: NA, CA. LITERATURE RECORDS: Cantone & Di Pietro 2002; Aleffi et al. 2003; Solis-Weiss et al. 2004; Castelli et al. 2008. OTHER REPORTED NAMES: Polydora coeca ��rsted, 1843 Dipolydora flava (Clapar��de, 1870) DISTRIBUTION: NA, CA, SA. LITERATURE RECORDS: Zavodnik et al. 1985; Aleffi et al. 1988; Lardicci 1989; Crema et al. 1991; Požar-Domac 1994; Crema et al. 2000, 2001; Fraschetti et al. 2002; Aleffi et al. 2003; Solis-Weiss et al. 2004, 2007; Simonini et al. 2007; Castelli et al. 2008. NEW RECORDS: BM 28, BM 52, BM 53, BM 54, BM 65, BM 66, BM 77, BM 78, BM 91, BM 92, BM 96, BM 98, BM 99, BM 101. OTHER REPORTED NAMES: Polydora flava Clapar��de, 1870. �� Dipolydora giardi (Mesnil, 1893) DISTRIBUTION: NA. NEW RECORDS: BM 91. REMARKS: This is the first record from the Adriatic Sea. DEPOSITED MATERIAL: BM 91, 0 8.07. 2010., 1 spec. (PMR- 17635). Dipolydora quadrilobata (Jacobi, 1883) DISTRIBUTION: NA. LITERATURE RECORDS: Fauvel 1934; Vatova 1935; Katzmann 1972; Marcuzzi 1972; Lardicci 1989; Požar-Domac 1994; Castelli et al. 2008., Published as part of Mikac, Barbara, 2015, A sea of worms: polychaete checklist of the Adriatic Sea, pp. 1-172 in Zootaxa 3943 (1) on pages 117-118, DOI: 10.11646/zootaxa.3943.1.1, http://zenodo.org/record/244663, {"references":["Fauvel, P. (1938) Annelida Polychaeta della Laguna di Venezia. Bollettino. R. Comitato Talassografico Italiano, 246, 1 - 27.","Katzmann, W. (1972) Die Polychaeten Rovinjs (Istrien / Jugoslawien). Zoologischer Anzeiger, 188, 116 - 144.","Lardicci, C. (1989) Censimento dei Policheti dei mari Italiani: Spionidae Grube, 1850. Atti della Societa Toscana di Scienze Naturali di Pisa, Memorie, Serie B, 96, 121 - 152.","Pozar-Domac, A. (1994) Index of the Adriatic Sea Polychaetes (Annelida, Polychaeta). Natura Croatica, 3, (Supplement 1), 1 - 23.","Casellato, S. & Stefanon, A. (2008) Coralligenous habitat in the northern Adriatic Sea: an overview. Marine Ecology, 29, 321 - 341. http: // dx. doi. org / 10.1111 / j. 1439 - 0485.2008.00236. x","Castelli, A., Bianchi, C. N., Cantone, G., Cinar, M. E., Gambi, M. C., Giangrande, A., Iraci Sareri, D., Lanera, P., Licciano, M., Musco, L. & Sanfilippo, R. (2008) Annelida Polychaeta. In: Relini, G. (Ed), Checklist della flora e della fauna dei mari italiani (Parte I). Biologia Marina Mediterranea, 15 (Supplement 1), pp. 327 - 377.","Cantone, G. & Di Pietro, N. (2002) Policheti bentonici della Fossa di Pomo (Medio Adriatico). Biologia Marina Mediterranea, 9 (1), 494 - 500.","Aleffi, F., Bettoso, N. & Solis - Weiss, V. (2003) Spatial distribution of soft - bottom polychaetes along western coast of the northern Adriatic Sea (Italy). Anali za istarske in mediteranske studije Series Historia Naturalis, 13 (2), 211 - 222.","Zavodnik, D., Vidakovic, J. & Amoureux, L. (1985) Contribution to sediment macrofauna in the area of Rovinj (North Adriatic Sea). Cahiers de Biologie Marine, 26, 431 - 444.","Aleffi, F., Orel, G., Vio, E. & Del Piero, D. (1988) Popolamenti bentonici e fenomeni di anossia nel Golfo di Trieste (Alto Adriatico): Dati. Nova Thalassia, 9, 165 - 231.","Crema, R., Castelli, A. & Prevedelli, D. (1991) Long Term Eutrophication Effects on Macrofaunal Communities in Northern Adriatic Sea. Marine Pollution Bulletin, 22 (10), 503 - 508.","Crema, R., Prevedelli, D., Valentini, A. & Castelli, A. (2000) Recovery of the macrozoobenthic community of the Comacchio lagoon system (Northern Adriatic Sea). Ophelia, 52 (2), 143 - 152. http: // dx. doi. org / 10.1080 / 00785236.1999.10409423","Crema, R., Prevedelli, D. & Castelli, A. (2001) Effect of submerged structures on the diversity of macrozoobenthos in the Northern Adriatic Sea. In: Faranda, F. M., Guglielmo, L. & Spezie, G. (Eds), Mediterranean Ecosystems: Structures and Processes. Springer - Verlag Italia, pp. 367 - 374. http: // dx. doi. org / 10.1007 / 978 - 88 - 470 - 2105 - 1 _ 48","Simonini, R., Ansaloni, I., Bonini, P., Grandi, V., Graziosi, F., Iotti, M., Massamba - N'Siala, G., Mauri, M., Montanari, G., Preti, M., De Nigris, N. & Prevedelli, D. (2007) Recolonization and recovery dynamics of the macrozoobenthos after sand extraction in relict sand bottoms of the Northern Adriatic Sea. Marine Environmental Research, 64, 574 - 589. http: // dx. doi. org / 10.1016 / j. marenvres. 2007.06.002","Fauvel, P. (1934) Annelides Polychetes de Rovigno d'Istria. Thalassia, 1 (7), 1 - 78.","Vatova, A. (1935) Ricerche preliminari sulle biocenosi del Golfo di Rovigno. Thalassia, 2 (2), 1 - 30.","Marcuzzi, G. (1972) Le collezioni dell'ex Istituto di Biologia marina di Rovigno conservate presso la stazione Idrobiologica di Chioggia. Atti e Memorie dell'Accademia Patavina di Scienze Lettere ed Arti, 84 (2), 169 - 219."]}

Dipolydora Verrill, 1881 Dipolydora Verrill, 1881: 320; Blake 1996: 181, resurrected and redefined. Type-species. Polydora concharum Verrill, 1879. Designated by Verrill (1881), by monotypy. Remarks. Dipolydora Verrill, 1881 currently comprises about forty species of polydorin spionids (tribe Polydorini Benham, 1896 sensu Radashevsky, 2012) that occupy diverse habitats from the intertidal to deep water. The name Dipolydora was not in use after its designation by Verrill (1881) until Blake (1996) resurrected it and assigned to it a series of Polydora Bosc, 1802 species that, in contrast to species of Polydora, had notochaetae on chaetiger 1 and lacked a constriction or manubrium on the shaft of the hooded hooks. Nine Dipolydora species were reported from Australian waters by Blake & Kudenov (1978) (as Polydora species), McDiarmid et al. (2004) and Sato-Okoshi et al. (2008)., Published as part of Radashevsky, Vasily I., 2015, Spionidae (Annelida) from Lizard Island, Great Barrier Reef, Australia: the genera Aonides, Dipolydora, Polydorella, Prionospio, Pseudopolydora, Rhynchospio, and Tripolydora, pp. 635-694 in Zootaxa 4019 (1) on page 643, DOI: 10.11646/zootaxa.4019.1.22, http://zenodo.org/record/235906, {"references":["Verrill, A. E. (1881) New England Annelida. Part I. Historical sketch, with annotated lists of the species hitherto recorded. Transactions of the Connecticut Academy of Arts and Sciences, 4, 285 - 324.","Blake, J. A. (1996) Family Spionidae Grube, 1850. Including a review of the genera and species from California and a revision of the genus Polydora Bosc, 1802. In: Blake, J. A., Hilbig, B. & Scott, P. H. (Eds.), Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Vol. 6. The Annelida Part 3 - Polychaeta: Orbiniidae to Cossuridae. Santa Barbara Museum of Natural History, Santa Barbara, California, pp. 81 - 223.","Radashevsky, V. I. (2012) Spionidae (Annelida) from shallow waters around the British Islands: an identification guide for the NMBAQC Scheme with an overview of spionid morphology and biology. Zootaxa, 3152, 1 - 35.","Blake, J. A. & Kudenov, J. D. (1978) The Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera. Memoirs of the National Museum of Victoria, 39, 171 - 280.","McDiarmid, H., Day, R. & Wilson, R. (2004) The ecology of polychaetes that infest abalone shells in Victoria, Australia. Journal of Shellfish Research, 23, 1179 - 1188.","Sato-Okoshi, W., Okoshi, K. & Shaw, J. (2008) Polydorid species (Polychaeta: Spionidae) in south-western Australian waters with special reference to Polydora uncinata and Boccardia knoxi. Journal of the Marine Biological Association of the United Kingdom, 88, 491 - 501. http: // dx. doi. org / 10.1017 / S 0025315408000842"]}

Dipolydora armata (Langerhans, 1880) (Fig. 5) Polydora armata Langerhans, 1880: 93 ���94, pl. 4, fig. 5 a���c; Blake & Kudenov 1978: 255 ���258, fig. 43 a���e. Dipolydora armata.��� Blake 1996 (Part.): 196���198, fig. 4.36; Bick 2001: 178 ���186, figs. 1���7; Williams 2001: 438 ���442, figs 5, 6; Radashevsky & Nogueira 2003: 377���381, figs 1���7; Sato-Okoshi et al. 2008: 495���496, fig. 4. Dipolydora cf. armata.��� Radashevsky & Fauchald 2000: fig. 6 F. Material examined. Queensland: AM W. 45506, MI QLD 2447 (20); MIMB 28114, MI QLD 2447 (7). Adult morphology. Up to 2.5 mm long, 0.3 mm wide for 30 chaetigers. Body and palp pigmentation absent. Prostomium incised anteriorly, extending posteriorly to end of chaetiger 2 as a low caruncle. Occipital antenna and eyes absent. Chaetiger 1 with capillaries and small postchaetal lamellae in both rami. Winged notopodial capillaries gradually becoming longer, thinner and less numerous along midbody chaetigers. Notopodia from chaetigers 18��� 24 onwards with up to three thin alimbate capillaries and 15 awl-like spines in a bundle (Fig. 5 E, F). Chaetiger 5 with up to three dorsal superior winged capillaries, two heavy falcate spines, and four ventral winged capillaries (Fig. 5 A, B). Falcate spines with central main fang, large lateral tooth and apical structure appearing as wide cowling or third tooth on convex side of main fang; numerous fine bristles present on basis of teeth (Fig. 5 B). Hooks in neuropodia from chaetiger 7, up to three in a series, accompanied by 1���2 inferior capillaries throughout body and 1���3 alternating capillaries in 5���10 posterior chaetigers (Fig. 5 C, D). Hooks bidentate; shaft slightly curved, without constriction (Fig. 5 C, D). Branchiae from chaetiger 7 to chaetiger 11, fewer in small individuals, flattened, with surfaces oriented parallel to body axis, basally fused to notopodial postchaetal lamellae. Pygidium small, cup-shaped, with only narrow dorsal gap or bilobed, divided by dorsal gap and ventral incision into two rounded lateral lobes. Remarks. Dipolydora armata was originally described from Madeira Island by Langerhans (1880) as a borer of calcareous algae and has since been reported world-wide. In Australia, D. armata was reported from Western Australia (Augener 1914; Hartmann-Schr��der 1979; Sato-Okoshi et al. 2008), Victoria and South Australia (Blake & Kudenov 1978; Hartmann-Schr��der 1987; McDiarmid et al. 2004). Worms from Lizard Island appear identical to D. armata described from California (Hartman 1941; Blake 1996), the Marshall Islands (Woodwick 1964), Mediterranean (Bick 2001), Philippines (Williams 2001), Brazil and other localities (Radashevsky & Nogueira 2003, type material re-examined). Dipolydora armata has been considered cosmopolitan, widespread in tropical and subtropical waters, and rare in boreal and temperate waters. Such a wide distribution of the same species seems, however, suspicious taking into account small size of adults (usually 3���4 mm long for 30���35 chaetigers), low fecundity (up to 100 eggs per brood), adelphophagy (larvae feeding on the nurse eggs inside egg capsule), low larval production, and very short time that larvae swim in the water (see Lewis 1998 for details of larval development in the species). Molecular investigation is certainly needed to confirm the conspecificity of the remote populations. Habitat. In this study, adult D. armata were found boring into the shell of a gastropod Turbo sp. at about 14 m depth. Distribution. Widespread in tropical and subtropical waters. Intertidal to 100 m. The species is herein reported for the first time from Queensland and the Great Barrier Reef., Published as part of Radashevsky, Vasily I., 2015, Spionidae (Annelida) from Lizard Island, Great Barrier Reef, Australia: the genera Aonides, Dipolydora, Polydorella, Prionospio, Pseudopolydora, Rhynchospio, and Tripolydora, pp. 635-694 in Zootaxa 4019 (1) on pages 643-645, DOI: 10.11646/zootaxa.4019.1.22, http://zenodo.org/record/235906, {"references":["Langerhans, P. (1880) Die Wurmfauna von Madeira. III. Zeitschrift fur wissenschaftliche Zoologie, 34, 87 - 143.","Blake, J. A. & Kudenov, J. D. (1978) The Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera. Memoirs of the National Museum of Victoria, 39, 171 - 280.","Blake, J. A. (1996) Family Spionidae Grube, 1850. Including a review of the genera and species from California and a revision of the genus Polydora Bosc, 1802. In: Blake, J. A., Hilbig, B. & Scott, P. H. (Eds.), Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Vol. 6. The Annelida Part 3 - Polychaeta: Orbiniidae to Cossuridae. Santa Barbara Museum of Natural History, Santa Barbara, California, pp. 81 - 223.","Bick, A. (2001) The morphology and ecology of Dipolydora armata (Polychaeta, Spionidae) from the western Mediterranean Sea. Acta Zoologica, 82, 177 - 187. [Stockholm] http: // dx. doi. org / 10.1046 / j. 1463 - 6395.2001.00078. x","Williams, J. D. (2001) Polydora and related genera associated with hermit crabs from the Indo-West Pacific (Polydora: Spionidae), with descriptions of two new species and a second polydorid egg predator of hermit crabs. Pacific Science, 55, 429 - 565. http: // dx. doi. org / 10.1353 / psc. 2001.0037","Radashevsky, V. I. & Nogueira, J. M. M. (2003) Life history, morphology and distribution of Dipolydora armata (Polychaeta: Spionidae). Journal of the Marine Biological Association of the United Kingdom, 83, 375 - 384. http: // dx. doi. org / 10.1017 / S 0025315403007227 h","Sato-Okoshi, W., Okoshi, K. & Shaw, J. (2008) Polydorid species (Polychaeta: Spionidae) in south-western Australian waters with special reference to Polydora uncinata and Boccardia knoxi. Journal of the Marine Biological Association of the United Kingdom, 88, 491 - 501. http: // dx. doi. org / 10.1017 / S 0025315408000842","Radashevsky, V. I. & Fauchald, K. (2000) Chaetal arrangement and homology in spionids (Polychaeta: Spionidae). Bulletin of Marine Science, 67, 13 - 23.","Augener, H. (1914) Polychaeta II, Sedentaria. Die Fauna Sudwest-Australiens. Ergebnisse der Hamburger sudwestaustralischen Forschungsreise 1905 herausgegeben von Prof. Dr. W. Michaelsen und Dr. R. Hartmeyer, 5, 1 - 172.","Hartmann-Schroder, G. (1979) Teil 2. Die Polychaeten der tropischen Nordwestkuste Australiens (zwischen Derby im Norden und Port Hedland im Suden). Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 76, 77 - 218.","Hartmann-Schroder, G. (1987) Teil 13. Die Polychaeten der antiborealen Kuste von Victoria (Australien) (zwischen Warrnambool im Western und Port Welshpool im Osten). Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut, 84, 27 - 66.","McDiarmid, H., Day, R. & Wilson, R. (2004) The ecology of polychaetes that infest abalone shells in Victoria, Australia. Journal of Shellfish Research, 23, 1179 - 1188.","Hartman, O. (1941) Some contributions to the biology and life history of Spionidae from California. With keys to species and genera and descriptions of two new forms. Allan Hancock Pacific Expeditions, 7, 289 - 323.","Woodwick, K. H. (1964) Polydora and related genera (Annelida, Polychaeta) from Eniwetok, Majuro, and Bikini Atolls, Marshall Islands. Pacific Science, 18, 146 - 159.","Lewis, J. B. (1998) Reproduction, larval development and functional relationships of the burrowing, spionid polychaete Dipolydora armata with the calcareous hydrozoan Millepora complanata. Marine Biology, 130, 651 - 662. [Berlin] http: // dx. doi. org / 10.1007 / s 002270050287"]}

Six native bivalves of commercial interest, Aequipecten tehuelchus, Mytilus sp., Aulacomya atra, Ostrea puelchana, Protothaca antiqua and Pododesmus rudis, were examined on account of shell-boring spionid polychaetes in northern Patagonia, Argentina. Adults of Polydora rickettsi were found boring into shells of all molluscs but Mytilus sp., whereas Dipolydora cf. giardi was found only in shells of A. atra. The intensity of infestation of molluscs was low, and the prevalence of infestation by P. rickettsi varied from 20 to 54%, depending on the host species. Both boring spionids are reported for the first time from the Patagonian coast, and also as shell-borers of the examined molluscs.

The polychaete Dipolydora commensalis is an obligate symbiont of hermit crabs and produces a burrow along the columella of the gastropod shells they inhabit. Adults of D. commensalis have short palps that they use to feed on particles dropped or brought in by the respiratory currents of hosts. To determine whether hermit crabs influence palp length, specimens of D. commensalis were isolated in glass capillary tubes and the growth of palps was measured over a 3-week period. Palp length was also measured in worms isolated in gastropod shells with or without hermit crabs for 2 weeks. In addition, to determine whether adults of D. commensalis have regeneration capabilities like those of free-living relatives, worms were cut at the fifth or 15th setiger and then monitored for 35 d. Worms extracted from shells and placed into capillary tubes had initial palp lengths of 1.0±0.4 mm (n=17); after isolation, palps were 40% longer (1.4±0.4 mm, n=17). Worms in gastropod shells with hermit crabs had an average palp length of 0.9±0.4 mm (n=31), whereas worms in shells without hermit crabs had palps that were 33% longer (1.2±0.5 mm, n=40). Adults of D. commensalis are capable of regeneration; 35 d after ablation at setigers 5 or 15, the average number of anterior setigers regenerated was 5 (n=15) and 9±1.3 (n=13), respectively. The average number of posterior setigers regenerated from the 15 setiger anterior fragments was 11±6 (n=10). The findings suggest that the palps (and sometimes anterior ends) of the worms are exposed during feeding and are cut during movement of the hermit crab. In the laboratory worms can live for >4 years, considerably longer than the functional life span of most gastropod shells inhabited by hermit crabs.

Wild molluscs from five sites and farmed oysters and abalone from one site each along the south coast of South Africa were examined for shell-boring Polydora and Dipolydora. One new species each of Dipolydora and Polydora are described. D. keulderae sp. n. belongs to the D. barbilla / D. bidentata group and was present at all the sites sampled. P. dinthwanyana sp. n. belongs to the P. ciliata / P. websteri group and was recorded only at Haga Haga, the most easterly site sampled. In total, five Dipolydora species (D. cf. armata, D. capensis, D. cf. giardi, D. keulderae and D. normalis) and three Polydora species (P. cf. ciliata, P. dinthwanyana and P. cf. hoplura) were recorded. All but D. normalis bored into mollusc shells and while the rest of the worms showed no host preference, the larger, subtidal Haliotis spp. and Turbo sarmaticus usually hosted the largest number of species at each site. D. normalis and P. dinthwanyana were recorded only at Haga Haga, P. cf. ciliata only at Port Elizabeth and the rest at all the sites. This study extends the known distributions of D. normalis, D. cf. armata and P. cf. ciliata in South Africa.

The isozyme patterns of glucose-6-phosphate isomerase (GPI) have been analyzed in ten species of polychaetes of the genera Polydora and Dipolydora (Polychaeta: Spionidae). The GPI patterns of these species have been found to have some specific characteristics that cannot be explained in terms of the generally accepted views on the nature of isozymes. The patterns are represented by two hybridizing isozymes with different expression specificities that exhibit coordinated allozymic variation in most individuals of each species studied. Involvement of alternative splicing in the expression of the GPI gene is considered to be the most probable mechanism of the formation of the unusual GPI isozyme patterns in polydorids.

Identification of tracemakers is of primary importance for evaluating the biotic interactions inferred from bore holes in fossil shell assemblages. Domicile bore holes in the subapical whorls of gastropods produced by spionid polychaeteDipolydorasp., supposed to be commensal with hermit crabs, are common in dead gastropod assemblages from deepwater habitats in the Philippines. These holes exhibit unique features and support a new criterion for the interpretation of nonpredatory borings in fossil gastropods. Diagnostic of these bore holes are: small circular to elliptical outer opening, the presence of weak dissolution of the columella beneath the bore hole, and the presence of a hollowed tube composed of detritus held together with mucus within some gastropod whorls anterior to the hole. The two selection factors of subapical whorls and elongate shells are supplementary criteria for recognition of these holes. Bore holes are recognized here in a deepwater gastropod assemblage from the upper Pliocene Shinzato Formation of Okinawa, Japan, and namedPolydorichnus subapicalisn. igen. and isp. These holes are identical to modern examples exhibiting similar site and species selectivity.P. subapicalishas its oldest fossil record in the upper Miocene of the Philippines, was common in offshore assemblages from the Miocene onward, and is a good indicator of occupation by a hermit crab and for commensalism between polychaetes and hermit crabs.

A study was conducted to identify and quantify the spionid polychaetes that infest cultured abalone Haliotis midae at aquaculture facilities on the west (Farm A), south (Farms B and C) and east (Farm D) coasts of South Africa. The relationship between total intensity of infestation by the polydorids and the condition of their hosts at Farms A, B and D were also measured. The abalone were infested by Dipolydora capensis, Polydora hoplura and a Boccardia sp. However, the intensity of infestation differed among sites, suggesting that both geographical and farm-specific conditions may influence species composition. Total intensity of infestation has a negative impact on the condition of abalone at the three farms tested, but at Farms A and B, this effect was reduced in larger abalone.

Dipolydora paracaulleryi sp. nov. (Figures 7–10) Polydora (Polydora) caullery.— Hartmann-Schröder, 1979: 88. Holotype: NE Atlantic Ocean, Expedition DIVA 3, cruise METEOR ME 79 - 1: Great Meteor Seamount, 29 ° 44.13 ’N 028° 24.91 ’W, station 656, epibenthic sledge, depth 283.3 m, 18 -Aug-2009, 96% ethanol, 1 af (ZSRO- P 2305). Paratypes: NE Atlantic Ocean, Expedition DIVA 3, cruise METEOR ME 79 - 1: Great Meteor Seamount: 29 ° 44.13 ’N 028° 24.91 ’W, station 656, epibenthic sledge, depth 283.3 m, 18 -Aug-2009, 96% ethanol, 1 af SEM (ZSRO-P 2304); 29 ° 41.84 ’N 028° 22.55 ’W, station 645, grab, depth 295 m, 18 -Aug-2009, 96% ethanol, 1 af (ZSRO-P 2306). Little Meteor Seamount: 29 ° 36.29 ' N 028° 59.68 ' W, station 624, grab, depth 274.2 m, 17 -Aug- 2009, 96% ethanol, 1 af (ZSRO-P 2303). Atlantic Seamount cruise, METEOR cruise 9 c, Great Meteor Seamount: 29 ° 57.7 ’N 028° 35.1 ’W, station 151 a, Agassiz trawl, depth 305–316 m, 16 -Jul- 1967, formalin, 1 af (ZMH-P 17290). Additional material examined. NW Atlantic Ocean: Great Bank of Newfoundland, depth 121–148 m, 12 - Sep- 1993 – 15 -Sep-1993, 2 af (SMF 5133); North Sea: German Bight, Borkum Riff, depth 27 m, 23 -Apr-1968, 3 af (SMF 6186), depth 26 m, 22 -Apr-1968, 2 af (SMF 6187); German Bight, Langeoog, 1967, 1 af (SMF 6189); Mediterranean Sea: Gulf of Marseille, 5 -Nov-1982, 2 af (SMF 16028). Diagnosis. Anterior margin of prostomium truncated, not or only slightly projecting beyond the peristomium. Caruncle large and triangular, encompassing bases of first notopodia, extending posteriorly to chaetiger 2 (Figures 7 A, 8 A–B). Occipital antenna absent. Eyes absent in adults. Nuchal organ as ciliary bands on either side of caruncle (Figure 8 A). First notopodia with few capillary chaetae (Figures 7 A, 8 A–B). Chaetiger 5 with dorsal fascicle of fringed geniculate chaetae, bushy-topped modified spines with accessory lateral flange, and ventral fascicle of fringed capillaries (Figures 7 C, F, 8 C, 9). Dorsal branchiae from chaetiger 7 to maximally chaetiger 12. Bidentate hooded hooks from chaetiger 7, accompanied by one capillary in inferiormost position (Figure 7 C, E). Posterior notopodia with bundles of thick acicular spines (Figure 7 D). Pygidium with 4 lobes, ventral lobes larger, more expanded, touching each other, dorsal lobes smaller, distinctly separated (Figure 7 D). Description. Small species; holotype anterior fragment with 14 chaetigers, about 2 mm in length and 0.35 mm wide. Other examined anterior fragments with 12–14 chaetigers, 1.1–1.2 mm in length and 0.1–0.3 mm wide. One complete specimen 1.2 mm long and 0.2 mm wide for 29 chaetigers. Prostomium with truncated, indistinctly incised anterior margin, not or only slightly projecting beyond the peristomium anteriorly, laterally separated from peristomium by a narrow furrow (Figure 7 A–B, 8 A–B); posterior part of prostomium with slight elevation, extending posteriorly to chaetiger 2 as large and triangular caruncle; caruncle encompassing bases of first notopodia (Figures 7 A, 8 A–B). Anterior fragments without eyes; one complete specimen with three black eyes, anterior eyes paired, posterior eye in the middle of prostomium directly behind the anterior pair. Occipital antenna absent. Palps arising at the end of the furrow between prostomium and peristomium, extending posteriorly for about 10 to 15 chaetigers (Figure 10 A). Nuchal organ as ciliary bands laterally to caruncle until the second chaetiger (Figures 7 A, 8 A–B). First parapodia with 2–4 notopodial capillary chaetae, neuropodial capillaries more numerous, parapodia 2 to 4 and 6 with fringed capillaries; notopodial capillaries of superior longitudinal row or fascicle distinctly longer than those of transverse row (Figures 7 A, 8 A); medium and posterior notopodia with acicular spines, on medium chaetigers only one or few spines additionally to capillaries, spines replacing capillaries on posterior notopodia completely and forming a curved row (Figure 7 D). Bidentate hooded hooks from chaetiger 7, numbering up to 5 per fascicle, hooded hooks accompanied throughout by one thin capillary in inferiormost position, hooks without constriction on shaft (Figure 7 E). Both notopodial and neuropodial lamellae relatively small, on anterior chaetigers notopodial lamellae triangular and neuropodial lamellae oval (Figure 8 B); in posterior parapodia both neuropodial and notopodial lamellae narrow; praechaetal lamellae not developed. Dorsal, flattened branchiae from chaetiger 7 (Figure 8 D), reaching full size by chaetiger 8, continuing to chaetigers 11 or 12, reaching midline dorsally and touching from chaetiger 8 to 11 or 12; basally fused to notopodial postchaetal lamellae. Chaetiger 5 modified, but not larger than chaetigers 4 and 6 (Figure 7 A–C), lacking postchaetal lobes (Figure 8 C); superior dorsal fascicle of two broad, fringed, geniculate chaetae, two major spines, and ventral fascicle of 4 to 6 fringed capillaries; major spines with large, beak-like curved end bearing crest of bristles and lateral accessory flange, companion capillaries absent (Figures 8 C, 9). Glandular pouches present on chaetigers 6 and 7, ventrolaterally; most striking after methyl green staining (Figure 10 B–C). Gizzard-like structure absent in anterior part of digestive tract. Pygidium with 4 lobes, ventral lobes larger, more expanded, touching each other, dorsal lobes smaller, distinctly separated (Figure 7 D). Pigmentation. Body generally opaque white in ethanol. Complete specimen with dorsolateral and lateral intersegmental brown pigment on anterior chaetigers up to segment 10. Additionally, neuropodia of chaetiger 6 brown-rimmed. Methyl green staining pattern. Anterior rim of the prostomium, peristomium and anal cirri most intensively stained; bands of scattered dots both on borders of chaetiger 6 and 7 ventrally (Figure 10 A–C), expanded in the regions of the glandular pouches on chaetiger 6 and 7 (Figures 7 B, 10 B); in addition, border from between chaetiger 9 / 10 to about 12 / 13 deep blue ventrally, some deep blue spots scattered on posterior part of prostomium and dorsum anteriorly (Figure 10 A). Ecology. Dipolydora paracaulleryi sp. nov. was found on the Great and Little Meteor Seamounts in coral and crushed shell gravel in about 300 m depth. Geographical distribution. NE Atlantic: Great Meteor Seamount (type locality) and Little Meteor Seamount. Etymology. The species name is derived from D. caulleryi, the closest relative to D. paracaulleryi sp. nov. Remarks. Dipolydora paracaulleryi sp. nov. is a member of the D. armata / D. caulleryi group owing to the presence of distinct modifications of the major spines of the fifth chaetiger and the heavy, protruding spines in posterior notopodia (see Blake 1996). Seven species belong to this group. But Dipolydora paracaulleryi sp. nov. closely resembles only D. caulleryi (Mesnil, 1897). The two species possess unidentate, falcate, bristle-topped spines on chaetiger 5, and branchiae and hooded hooks start on chaetiger 7. Another very similar species is D. blakei (Maciolek, 1984) but this species possesses 3–4 falcate spines on chaetiger 5 with a large lateral tooth, rather than only 2 spines with a lateral flange, and a pygidium with 2 lateral lobes and a cup-shaped pygidium respectively, rather than a pygidium with one pair of dorsal and one pair of ventral lobes (Maciolek 1984, Radashevsky & Simboura 2013). The number of superior geniculate chaetae (4–6) on chaetiger 5 of D. blakei is greater compared with D. paracaulleryi sp. nov. (only 2). Dipolydora paracaulleryi sp. nov. differs from D. caulleryi in having 2 unidentate, falcate, bristle-topped spines rather than 3 or 4, and is distinctly smaller (8 mm long for D. caulleryi opposed to about 2 mm for D. paracaulleryi sp. nov.). Several important diagnostic characters such as the number of chaetigers, the length of the caruncle, the shape of postchaetal lobes, and number and shape of pygidial lobes have not been described in the original description of D. caulleryi. It is unknown whether type material still exists, and if so, where it has been deposited. For this reason, further potential distinctions between D. caulleryi and D. paracaulleryi sp. nov. cannot be indicated. Moreover, descriptions of specimens assumed belonging to D. caulleryi vary conspicuously. Pettibone (1954) described D. caulleryi specimens of 50 mm length and 2.2 mm width, with a caruncle extending to chaetiger 4 to 6, and 0 to 6 eyes, branchiae to first appear on chaetiger 7 or 8, exhibiting 3–12 major spines on chaetiger 5 without companion capillaries, and 2 to 3 neuropodial hooded hooks. Specimens described by Blake (1971) are of comparable size, possess a caruncle extending to chaetiger 3 or 4, possess about 13 neuropodial hooded hooks, and major spines of chaetiger 5 are arranged in an elevated semicircle with capillary companion chaetae. The investigated specimens from the North Sea and NW Atlantic Ocean correspond with Blake’s (1971) description, though companion chaetae on chaetiger 5 are absent. As the descriptions by the two authors differ significantly both from each other and from the original description it is assumed here that descriptions by Mesnil (1897), Pettibone (1954) and Blake (1971) refer to different species. Dipolydora caulleryi specimens from the Mediterranean Sea, also investigated during this study, resemble D. paracaulleryi sp. nov. in having 2–3 major spines with lateral flange; length and shape of the caruncle are also similar. But these slightly larger specimens possess 3 geniculate notochaetae and 8–10 ventral capillaries on chaetiger 5 (opposed to 2 and 4–6 chaetae respectively in D. paracaulleryi sp. nov.) and only 3–4 neuropodial hooded hooks (4–5 in D. paracaulleryi sp. nov.). The identity of the Mediterranean specimens cannot be clarified based on current knowledge. A comparative study of specimens of D. caullery and morphologically similar species from different localities is required to learn about their infraspecific variability and to identify reliable distinguishing characters for species of this species complex.

Bick. A. 2001. The morphology and ecology of Dipolydora armata (Polychaeta, Spionidae) from the western Mediterranean Sea. — Acta Zoologica (Stockholm) 82 : 177‐187 The cosmopolitan spionid Dipolydora armata is a dominant species of epifauna on Thais haemastoma shells in the western Mediterranean Sea. Some characters, particularly number, thickness and arrangement of notopodial spines, were extremely variable among specimens. Three morphological forms were distinguished based on the general appearance of the bundles of spines; one form possesses at least one bundle of stout spines, whereas bundles of medium spines or thinner acicular spines characterise the other forms. Differences in prostomial shape, as well as in the extension and form of the caruncle, were artefacts (preservation or angle of observation). The present study supports D. rogeri as a junior synonym of D. armata. D. armata was regularly found on shells inhabited by hermit crabs ( Calcinus tubularis , Clibanarius erythropus ) and on damaged, empty shells. Abundance, dominance, size frequency, occurrence of larvae and occurrence of regenerated specimens varied, possibly as a result of fouling strategy and recurrent settlements.

The palp morphology of Dipolydora coeca, D. quadrilobata, Polydora ciliata, P. cornuta and Pseudopolydora pulchra were studied with light and scanning electron microscope. Two new palp characters were found on the two species of Dipolydora : randomly scattered motile cirri and glandular holes. The palp morphology of the investigated species is compared with earlier reports on other members of the Polydora complex. Congeneric palp similarities are found in Polydora and Dipolydora. A single exception is the aberrant Dipolydora commensalis , which has a distinctive palp morphology not found in any other spionid. The palp morphology of the two closely related genera Dipolydora and Polydora is significantly different, which supports the validity of the recently resurrected genus Dipolydora . Comparative studies indicate that the palp morphology of Pseudopolydora pulchra closely resembles that of Dipolydora . The recently studied palp morphology of Boccardia polybranchia has many similarities with that of Polydora.

Dipolydora quadrilobata (Jacobi, 1883) Figure 4 Polydora quadrilobata Jacobi, 1883: 3, 2 pls; Mesnil 1897: 87 ���88, pl. 3, figs. 9���11; Fauvel 1927: 54, fig. 18 l���r; Uschakov 1955: 272, fig. 94 A���D; Blake 1969 a: 37 ���51, figs. 27���37 (larvae); 1971: 13���15, fig. 9 (synonymy); Rasmussen 1973: 111 ���112; Radashevsky 1993: 18���21, fig. 9 (synonymy); Sato-Okoshi 2000: 445. Polydora (Polydora) quadrilobata. Hartmann-Schr��der 1996: 319 ���320, fig. 144 a���f. Dipolydora quadrilobata. Blake 1996: 198, fig. 4.32 I���N. Material examined. Bulgaria: Sta. 190 ��� 4 /VA 2, Varna (43 ��08.17��N, 28 �� 14.41 ��E), 33.7 m, muddy seabed, 23 Mar 2008, coll. A. Teac��, MNINGA PLY044 (46); Romania: Sta. 89, Eforie Nord, 10 m, medium-grained sand, 20 Aug 2003, coll. V. Abaza (1); Sta. 91, Tuzla, 10 m, sand mixed with shell debris, 20 Aug 2003, coll. V. Abaza (6); Sta. BS07/MA07 (43 �� 46.47 ��N, 28 �� 31.05 ��E), 33.5 m, silty mud, 20 Apr 2007, coll. A. Teac��, MNINGA PLY045 (2); Sta. BS05/ 66 (44 �� 13.37 ��N, 29 �� 38.03 ��E), 60.3 m, mud with Modiolula phaseolina (Philippi, 1844), 10 Aug 2007, coll. A. Teac��, MNINGA PLY046 (2); Sta. BS05/01 (44 �� 19.29 ��N, 29 �� 30.27 ��E), 56 m, greenish-grey mud with Mytilus galloprovincialis and M. phaseolina shells, 11 Aug 2007, coll. A. Teac��, MNINGA PLY047 (99); Sta. BS05/03 (44 �� 19 �� 27 �� N, 29 �� 33 �� 35 �� E), 54.5 m, mud with M. phaseolina, 11 Aug 2007, coll. A. Teac��, MNINGA PLY048 (357); Sta. BS05/ 28 (44 �� 11.29 ��N, 29 �� 40.57 ��E), 60.6 m, mud with M. phaseolina, 11 Aug 2007, coll. A. Teac��, MNINGA PLY049 (3); Sta. S���RO 1 ���02 Portiţa (44 �� 38.482 ��N, 29 �� 39.845 ��E), 50.1 m, black muds with Melinna palmata and Mya arenaria, 7 Apr 2008, coll. A. Teac��, MNINGA PLY050 (114); Sta. BS08/ 91 ��� 1 (43 �� 50 �� 29.43 �� N, 29 �� 15 �� 27.68 �� E), 62.5 m, mud with M. phaseolina, 21 Jun 2008, coll. A. Teac��, MNINGA PLY051 (2); Sta. BS08/05 (43 �� 59 ��31.00�� N, 29 �� 21 �� 31.18 �� E), 56 m, greenish-grey mud with Mytilus and Modiolula shells, 22 Jun 2008, coll. A. Teac��, MNINGA PLY052 (92); Sta. BS08/ 45 ��� 1 (43 �� 55 �� 29.71 �� N, 29 �� 21 �� 24.84 �� E), 59.8 m, greenish-grey mud with Mytilus and Modiolula shells, 22 Jun 2008, coll. A. Teac��, MNINGA PLY053 (9); Sta. BS08/ 22 (43 �� 57 �� 31 �� N, 29 �� 17 �� 81 �� E), 55.3 m, greenish-grey mud with Mytilus and Modiolula shells, 23 Jun 2008, coll. A. Teac��, MNINGA PLY054 (5); Ukraine: Sta. 140 ��� 3 /Phy 4 a (45 �� 39.11 ��N, 31 �� 20.78 ��E), 46.1 m, pelitic sand with shell rubble and Phyllophora, 15 Mar 2008, MNINGA PLY055 (3); Sta. 156 ��� 2 /DN 10 (45 �� 26.95 ��N, 30 �� 59.28 ��E), 45.5 m, pelitic sand with shell rubble and Phyllophora, 17 Mar 2008, MNINGA PLY056 (287); Sta. 159 ��� 4 /Phy 7 (45 �� 34.13 ��N, 30 �� 17.38 ��E), 23.4 m, silty mud with Mytilus shells, 18 Mar 2008, MNINGA PLY057 (2). Description. Up to 15 mm long, 0.75 mm wide at chaetiger 5, with up to 90 chaetigers. Colour of preserved worms pale yellowish. Body without dorsal pigmentation. Prostomium more or less distinctly incised anteriorly, forming two rounded lobes; caruncle indistinct, extending to posterior end of chaetiger 3 (Fig. 4 A). Occipital antenna absent. Eyes usually absent in adults; juveniles with four eyes arranged in nearly straight transverse row with the outer pair larger and irregular in shape. Nuchal organs as long, narrow ciliary bands on lateral sides of caruncle. Palps long, flexible, with prominent ciliated groove, extending posteriorly to chaetiger 15. Chaetiger 1 biramous, with well-developed podial lobes, carrying both short notopodial capillaries and more numerous neuropodial capillaries (Fig. 4 A). Chaetigers 2���4 and 6 with noto- and neuropodial fascicles of winged capillary chaetae (Fig. 4 A���B). Notochaetae arranged in two vertical rows of shorter and thicker capillaries plus a dorsal superior tuft of longer and thinner capillaries. Neurochaetae also as two vertical rows of thick capillaries and a ventral inferior tuft of thinner and shorter capillaries. Number of capillary notochaetae diminishing toward posterior end. In far posterior notopodia capillaries completely replaced by stout, sharply pointed spines arranged in semicircular row (Fig. 4 E���F); number of spines increasing from 1���3 in a tuft up to 16, providing a spinous appearance to posterior end. Chaetiger 5 modified, larger than adjacent segments, without postchaetal lamellae and carrying a horizontal curved row of 4���6 large spines (Fig. 4 A���B). Modified spines of chaetiger 5 falcate, distally curved, with two nearly equal teeth and a tuft of fine bristles in concavity between teeth. Companion chaetae absent (see Remarks). Dorsal superior fascicle of chaetiger 5 with up to seven geniculate limbate chaetae arranged in a vertical or oblique row (Fig. 4 C���D); ventral inferior fascicle as compact tuft of up to 9 winged capillaries. Neuropodial bidentate hooded hooks, up to six in vertical row from chaetiger 7. Main fang of hook at obtuse angle to shaft and at reduced angle to distal smaller tooth. Shafts of hooks strongly curved, without constriction and manubrium. In anterior neuropodia up to four inferior capillaries present ventral to hooks. In posterior neuropodia inferior capillaries diminishing in number, becoming narrow and non-winged. Branchiae from chaetiger 7, short at first, reaching full size by chaetiger 10���12, absent from posterior third of body, not connected to notopodial lobes. Pygidium with four subequal lobes (Fig. 4 F). Habitat and ecology. On the Romanian coast D. quadrilobata is found in soft sediments (fine sand, muddy sand, sandy mud, silty mud, and mud) at 10��� 100 m. However, the species is most common and abundant between 40���60 m in mud between beds of Modiolula phaseolina, with densities up to 2600 ind. m��� 2. The worm constructs permanent tubes that stand erect in the substrate, extending 15���20 mm. Tubes straight, up to 40 mm long and 1.5 mm wide, built of fine sand grains bound by silt and detrital particles. The upper part of tube is smooth and greyish-brown in colour, while the lower part, buried in sediment, is coarser and rusty-coloured. Reproduction and development. Not known to date in the Black Sea. Distribution. Arctic-boreal species, reported from the Pacific (Blake 1996) and Atlantic coasts of North America (Blake 1971), Sea of Japan and Sea of Okhotsk (Sato-Okoshi 2000), Bering Sea (Uschakov 1955; Radashevsky 1993), White Sea (Khaitov et al. 1999); North Sea (Rasmussen 1973; Hartmann-Schr��der 1996), British Channel, Kiel Bay, Baltic Sea, Atlantic coast of Europe (Fauvel 1927), Adriatic Sea (Castelli et al. 1995; Požar-Domac 1978) and Black Sea (Todorova & Panayotova 2006). Remarks. There is little confusion regarding the taxonomic position of Dipolydora quadrilobata. This species is closely related to D. caulleryi in having heavy, protruding, awl-shaped notopodial spines in far posterior segments and a pygidium divided into four nearly equal lobes, both falling into the �� armata/ caulleryi �� subgroup (sensu Blake 1996). S��derstr��m (1920) even considered D. quadrilobata to be a juvenile of D. caulleryi. Nevertheless, Dipolydora quadrilobata is unique among species of the Polydora complex by having modified spines of chaetiger 5 expanded distally into two nearly equal teeth with a bushy tuft of fine bristles between them, and by four eyes arranged in a nearly straight transverse row (Blake 1996). The Black Sea specimens match the morphological descriptions of specimens from northern Europe (Hartmann-Schr��der 1996) or from northern Pacific (Radashevsky 1993). However, brown reticulated pigmentation on anterior end of body reported by many authors (Fauvel 1927; Blake 1971, 1996; Radashevsky 1993; Hartmann-Schr��der 1996) was absent in the Black Sea specimens. Sato-Okoshi (2000), however, indicates that in Japan this pigmentation was present only in some specimens. Also, in one individual were observed two limbate capillaries near the bases of the major spines of chaetiger 5, like normal companion chaetae (Fig. 4 D)., Published as part of Surugiu, Victor, 2012, Systematics and ecology of species of the Polydora - complex (Polychaeta: Spionidae) of the Black Sea, pp. 45-65 in Zootaxa 3518 on pages 53-56, DOI: 10.5281/zenodo.208946, {"references":["Jacobi, R. (1883) Anatomisch-histogische Untersuchung der Polydoren der Keiler Bucht. Inaugural Dissertation, Keil: 1 - 35.","Mesnil, F. (1897) Etudes de morphologie externe chez les Annelides. II. Remarques complementaires sur les Spionidiens, la famille nouvelle de disomidiens. La place des Aonides (sensu Tauber, Levinsen). Bulletin Scientifique de la France et de la Belgique, 30, 83 - 100, pl. 1.","Fauvel, P. (1927) Polychetes sedentaires. Addenda aux Errantes, Archiannelides, Myzostomaires. Faune de France 16, Paul Lechevalier, Paris, 494 pp.","Uschakov, P. V. (1955) Polychaeta of the Far Eastern Seas of the U. S. S. R. Akademia Nauk SSSR, Moscow-Leningrad, 445 pp.","Blake, J. A. (1969 a) Reproduction and larval development of Polydora from northern New England (Polychaeta: Spionidae). Ophelia, 7, 1 - 63.","Rasmussen, E. (1973) Systematics and ecology of the Isefjord marine fauna (Denmark). Ophelia, 11, 1 - 507.","Sato-Okoshi, W. (2000) Polydorid species (Polychaeta: Spionidae) in Japan, with descriptions of morphology, ecology and burrow structure. 2. Non-boring species. Journal of the Marine Biological Association of the United Kingdom, 80, 443 - 456.","Hartmann-Schroder, G. (1996) Annelida, Borstenwurmer, Polychaeta. Die Tierwelt Deutschlands. Vol. 2. Neubearbeitete Auflage, Gustav Fischer Verlag, Jena, 58, 648 pp.","Blake, J. A. (1996) Family Spionidae Grube, 1850. Including a review of the genera and species from California and a revision of the genus Polydora Bosc, 1802. In: Blake, J. A., Hilbig, B. & Scott, P. H. (Eds.), Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Vol. 6. The Annelida Part 3. Polychaeta: Orbiniidae to Cossuridae. Santa Barbara Museum of Natural History, Santa Barbara, California, pp. 81 - 223.","Blake, J. A. (1971) Revision of the Genus Polydora from the East Coast of North America (Polychaeta: Spionidae). Smithsonian Contributions to Zoology, 75, 1 - 32.","Khaitov, V. M., Fokin, M. V. & Nicolaeva, A. M. (1999) Structure of communities associated with dense assemblages of the tube-dwelling polychaete Polydora quadrilobata Jacobi (Spionidae) in the White Sea. Hydrobiologia, 393, 221 - 226.","Castelli, A., Abbiati, M., Badalamenti, F., Bianchi, C. N., Cantone, G., Gambi, M. C., Giangrande, A., Gravina, M. F., Lanera, P., Lardicci, C., Somaschini, A. & Sordino, P. (1995) Annelida Polychaeta, Pogonophora, Echiura, Sipuncula. In: Minelli, A., Ruffo, S. & La Posta, S. (Eds.), Checklist delle specie della fauna italiana. Vol. 19. Annelida: Polychaeta, Pogonophora, Echiura, Sipuncula. Edizioni Calderini, Bologna, pp. 1 - 45.","Pozar-Domac, A. (1978) Catalogue of the polychaetous annelids of the Adriatic Sea. 1. Northern and central Adriatic. Acta Adriatica, 19, 2 - 59.","Todorova, V. R. & Panayotova, M. D. (2006) Alien aquatic fauna from the Bulgarian Black Sea and coastal lakes - an inventory of introductions and impacts. NEOBIOTA. From Ecology to Conservation. 4 th European Conference on Biological Invasions, Vienna (Austria), 27 - 29 September 2006. Book of Abstracts, pp. 251 - 252.","Soderstrom, A. (1920) Studien uber die Polychaetenfamilie Spionidae. Inaugural-Dissertation. Almquist & Wicksells, Uppsala, 286 pp. [not seen]"]}

Dipolydora commensalis (Andrews, 1891) Polydora commensalis Andrews, 1891: 291 ���292, plate 14, fig. 27.��� Foster, 1971 a: 20 ���22, figs 1���12. Dipolydora commensalis: Blake, 1996: 199, fig. 4.31 A���D. Type locality. Beaufort, North Carolina, USA. Type material. Holotype (USNM 4909), paratypes (USNM 4905, 4906, 4907). Records. Cura��ao (Foster 1971 a)., Published as part of Delgado-Blas, Victor Hugo & Salazar-Silva, Patricia, 2011, Taxonomic catalogue of the Spionidae (Annelida: Polychaeta) of the Grand Caribbean, pp. 39-66 in Zootaxa 2782 on page 44, DOI: 10.5281/zenodo.206833, {"references":["Andrews, E. A. (1891) Report upon the Annelida Polychaeta of Beaufort, North Carolina. Proceeding of the United States National Museum, 14, 277 - 302.","Foster, N. M. (1971 a) Spionidae (Polychaeta) of the Gulf of Mexico and the Caribbean Sea. Studies on the Fauna of Curacao and other Caribbean Islands, 129, 1 - 183.","Blake, J. A. (1996) Family Spionidae Grube, 1850, including a review of the genera and species from California and a revision of the genus Polydora Bosc, 1802 In: Blake JA, B Hilbig & PH Scott (eds.), The Annelida, Part 3. Polychaeta: Orbiniidae to Cossuridae, Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and the Western Santa Barbara Channel. Vol. 6. Santa Barbara Museum Natural History, Santa Barbara, pp. 81 - 223."]}

Dipolydora flava (Clapar��de, 1870) Type locality: Gulf of Naples, Italy; associated with Hyalinoecia rigida. Type material: Unknown. None deposited? Location and author of most recently described material: Channel to north of Quarantine Island, Otago, New Zealand (Rainer 1973), as Polydora dorsomaculata n.sp., synonymized in Blake and Kudenov (1978); no Australian material examined. Recorded distribution: Australia: Lower east coast, Bass Strait. New Zealand; Indian Ocean; Europe, Ceylon, Sumatra, Japan., Published as part of Walker, Lexie M, 2011, A review of the current status of the Polydora - complex (Polychaeta: Spionidae) in Australia and a checklist of recorded species, pp. 40-62 in Zootaxa 2751 on page 51, DOI: 10.5281/zenodo.203565, {"references":["Claparede, E. (1870) Les Annelides Chetopodes du Golfe de Naples. Supplement. Memoires de la Societe de Physique et d'Histoire Naturelle de Geneve, 20, 365 - 542, pls 361 - 316.","Rainer, S. (1973) Polydora and related genera (Polychaeta: Spionidae) from Otago waters. Journal the Royal Society of New Zealand, 3, 545 - 564.","Blake, J. A. & Kudenov, J. D. (1978) The Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera. Memoirs of the National Museum of Victoria, 39, 171 - 280."]}

Dipolydora notialis (Blake and Kudenov, 1978) Type locality: Tipara, South Australia from Haliotis roei shell. Type material: Holotype: NMV G 2877. Location and author of most recently described material: Blake and Kudenov (1978). Recorded distribution: Known only from type locality. Not reported since original description., Published as part of Walker, Lexie M, 2011, A review of the current status of the Polydora - complex (Polychaeta: Spionidae) in Australia and a checklist of recorded species, pp. 40-62 in Zootaxa 2751 on page 51, DOI: 10.5281/zenodo.203565, {"references":["Blake, J. A. & Kudenov, J. D. (1978) The Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera. Memoirs of the National Museum of Victoria, 39, 171 - 280."]}

Dipolydora contoyensis Delgado-Blas, 2008 Dipolydora contoyensis Delgado-Blas, 2008: 9 ���11, fig. 3. Type locality. Caribbean Sea, Contoy Island, Quintana Roo, Mexico (21 �� 29��N, 86 �� 48��W), burrowing in rock fragments. Type material. Holotype (LACM-AHF POLY 2192), paratypes (ECOSUR���0083, LACM-AHF POLY 2193). Records. Only known from type locality., Published as part of Delgado-Blas, Victor Hugo & Salazar-Silva, Patricia, 2011, Taxonomic catalogue of the Spionidae (Annelida: Polychaeta) of the Grand Caribbean, pp. 39-66 in Zootaxa 2782 on page 45, DOI: 10.5281/zenodo.206833, {"references":["Delgado-Blas, V. H. (2008) Polydora and related genera (Polychaeta: Spionidae) from the Grand Caribbean region. Journal of Natural History, 42, 1 - 19."]}

Genus Dipolydora Verrill, 1881 Dipolydora Verrill, 1881: 320.��� Blake, 1996: 181. Type species: Polydora concharum Verrill, 1881, by original designation., Published as part of Delgado-Blas, Victor Hugo & Salazar-Silva, Patricia, 2011, Taxonomic catalogue of the Spionidae (Annelida: Polychaeta) of the Grand Caribbean, pp. 39-66 in Zootaxa 2782 on page 43, DOI: 10.5281/zenodo.206833, {"references":["Verrill, A. E. (1881) New England Annelida, 1. Historical sketch, with annotated list of the species hitherto recorded. Connecticut Academy of Arts and Sciences, 4, 285 - 324.","Blake, J. A. (1996) Family Spionidae Grube, 1850, including a review of the genera and species from California and a revision of the genus Polydora Bosc, 1802 In: Blake JA, B Hilbig & PH Scott (eds.), The Annelida, Part 3. Polychaeta: Orbiniidae to Cossuridae, Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and the Western Santa Barbara Channel. Vol. 6. Santa Barbara Museum Natural History, Santa Barbara, pp. 81 - 223."]}

Dipolydora socialis (Schmarda, 1861) Leucodore socialis Schmarda, 1861: 64, fig. a���c, plate 26, fig. 209. Polydora socialis.��� Hartman, 1951: 83. Johnson, 1984: 6.28���6.30, fig. 6.19���6.20.��� Gobin, 1990: 43. Dipolydora socialis. ��� Blake, 1996: 189 ���192, fig. 4.34.��� Bone & Vi��itez, 2002: 70 ���71. Type locality. Vi��a del mar, Chile. Type material. Not found Records. USA: Alligator Harbor, Franklin Co., Florida, 1.8 m, silt-covered tube and mud flats (Hartman 1951); off Florida, 10���106 m, coarse to very fine sand, silty fine sand, clayey sand, sandy clayey silt (Johnson 1984). Trinidad and Tobago: Oropuche (Gobin 1990). Venezuela: Morrocoy (Bone & Vi��itez 2002). Remarks. Description of the specimens from Gulf of Mexico given by Hartman (1951) is brief and without illustrations, the identity of the records needs to be confirmed against the type material. Among specimens recorded in the Grand Caribbean by Johnson (1984), Gobin (1990) and Bone & Vi��tez (2002), their identity is based on published descriptions and their identity should be confirmed against the type material., Published as part of Delgado-Blas, Victor Hugo & Salazar-Silva, Patricia, 2011, Taxonomic catalogue of the Spionidae (Annelida: Polychaeta) of the Grand Caribbean, pp. 39-66 in Zootaxa 2782 on page 45, DOI: 10.5281/zenodo.206833, {"references":["Schmarda, L. K. (1861) Neue wirbellose Thiere beobachtet und gesammelt auf einer Reise um die Erde 1853 bis 1857. Leipzig, Vol. 1. Turbellarien, Rotatorien und Anneliden, 165 pp.","Hartman, O. (1951) The littoral marine annelids of the Gulf of Mexico. Publications of the Institute of Marine Science, Texas University, 2, 7 - 124.","Johnson, P. G. (1984) Family Spionidae Grube, (1850) pp. 6.1 - 6.69 In: Uebelacker, J. M. & P. G. Johnson (eds.), Taxonomic Guide to the Polychaetes of the Northern Gulf of Mexico. Barry A. Vittor & Associates, Inc. Mobile, AL. pp. 6 - 1 to 6 - 69.","Gobin, J. (1990) A checklist of marine polychaetous annelids (Polychaeta) of the Gulf of Paria, Trinidad, West Indies. Caribbean Marine Studies, 1, 37 - 47.","Blake, J. A. (1996) Family Spionidae Grube, 1850, including a review of the genera and species from California and a revision of the genus Polydora Bosc, 1802 In: Blake JA, B Hilbig & PH Scott (eds.), The Annelida, Part 3. Polychaeta: Orbiniidae to Cossuridae, Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and the Western Santa Barbara Channel. Vol. 6. Santa Barbara Museum Natural History, Santa Barbara, pp. 81 - 223.","Bone, D. & Vieitez, J. M. (2002) Spionidae (Annelida: Polychaeta) from the Parque Nacional Morrocoy, Falcon, Venezuela. Revista de Biologia Tropical, 50, 69 - 75."]}

Dipolydora quadrilobata (Jacobi, 1883) Polydora quadrilobata Jacobi, 1883: 3, plate 1, figs 1 ���9, 13���15, 17, 20 b.��� Taylor, 1973. Dipolydora quadrilobata: Blake, 1996: 198, fig. 4.32 I���N. Type locality. Kiel Canal, Germany. Type material. (ZMH V��� 1546). Records. USA: Tampa Bay, Florida (Taylor 1973). Remarks. The identity of the records from the Grand Caribbean need to be confirmed against the type material., Published as part of Delgado-Blas, Victor Hugo & Salazar-Silva, Patricia, 2011, Taxonomic catalogue of the Spionidae (Annelida: Polychaeta) of the Grand Caribbean, pp. 39-66 in Zootaxa 2782 on page 45, DOI: 10.5281/zenodo.206833, {"references":["Jacobi, R. (1883) Anatomisch-histologische Untersuchung der Polydoren der Kieler Bucht. Weissenfels, 1 - 37.","Taylor, J. L. (1973) Biological studies and inventory for the Tampa Harbor, Florida Project. A report submitted to the U. S. Army Corps of Engineers Jackson-ville District, Florida, 101 pp.","Blake, J. A. (1996) Family Spionidae Grube, 1850, including a review of the genera and species from California and a revision of the genus Polydora Bosc, 1802 In: Blake JA, B Hilbig & PH Scott (eds.), The Annelida, Part 3. Polychaeta: Orbiniidae to Cossuridae, Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and the Western Santa Barbara Channel. Vol. 6. Santa Barbara Museum Natural History, Santa Barbara, pp. 81 - 223."]}

Dipolydora tentaculata (Blake and Kudenov, 1978) Type locality: Botany Bay, Towra Beach, New South Wales, Australia. Type material: Holotype: NMV G 2885; Paratypes: NMV G 2886; QM G 11597. Location and author of most recently described material: Botany Bay, Towra Beach, New South Wales; Moreton Bay, Middle Banks, QLD, Australia (Blake & Kudenov 1978). Recorded distribution: Australia: Queensland (Moreton Bay), New South Wales (Botany Bay, Hawkesbury River)., Published as part of Walker, Lexie M, 2011, A review of the current status of the Polydora - complex (Polychaeta: Spionidae) in Australia and a checklist of recorded species, pp. 40-62 in Zootaxa 2751 on page 52, DOI: 10.5281/zenodo.203565, {"references":["Blake, J. A. & Kudenov, J. D. (1978) The Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera. Memoirs of the National Museum of Victoria, 39, 171 - 280."]}

Dipolydora armata (Langerhans, 1880) Polydora armata Langerhans, 1880: 93 ���94, fig. 4 D��� 5 A.��� Blake & Kudenov, 1978: 247.��� Hartman, 1951: 83.��� Lewis, 1998: 652. Dipolydora armata: Radashevsky & Nogueira, 2003: 377 ���380. Type locality. Madeira Type material. Syntypes (ZMH V��� 1547), paratypes (NHMW 7336). Records. USA: off southwestern Florida, shell of Carrier-Shell, Tugurium (Trochotugurium) longleyi (Bartsch, 1931), 214 m (Hartman 1951). Mexico: Xcalac, Quintana Roo; Yucatan (18 �� 16��N, 87 �� 49.6��W), coral reef, 15 m, from shell of deltoid rocksnail, Thais detoidea (Lamarck, 1822), occupied by hermit crab, Clibanarius cf. cubensis (de Saussure, 1858) (Radashevsky & Nogueira 2003). Belize: Carrie Bow Cay (16 �� 48��N, 88 ��05��W), 0.4���1.2 m, coral rubble, Dictyota, Thalassia in part live Porites, sand, reef crest rubble, encrusting red algae of Lithothamnion type, 2 m, from coralline algae encrusting gastropod shell occupied by hermit crab (Radashevsky & Nogueira 2003). Remarks. Dipolydora armata was described by Langerhans (1880) from Madeira, his original description and figures show bidentate spines on segment 5, without bristles, six pairs of branchiae, and the posterior 5���6 segments with spines. However, the description of the arrangement of posterior spines is vague. Subsequent reports of D. armata by a number of the other workers (e.g. Woodwick 1964, Blake & Kudenov 1978, Blake 1983, Martin 1996, Bick 2001, Radashevsky & Nogueira 2003) have extended its distribution and variously adjusted its diagnosis. Radashevsky & Nogueira (2003), re-described D. armata with specimens from Sao Paulo, Brazil although they examined material from many localities, ranging from Portugal to Belize and from Thailand to New Zealand. However, their re-description differs from the original by Langerhans. We do not agree with the diagnosis and the wide distribution of D. armata as presented by Radashevsky & Nogueira (2003). We think it is necessary that specimens from other localities are actually compared to specimens from the type locality. In fact one possible species, Polydora rogeri Martin, 1996, was synonymised with D. armata by Radashevsky & Nogueira (2003)., Published as part of Delgado-Blas, Victor Hugo & Salazar-Silva, Patricia, 2011, Taxonomic catalogue of the Spionidae (Annelida: Polychaeta) of the Grand Caribbean, pp. 39-66 in Zootaxa 2782 on page 44, DOI: 10.5281/zenodo.206833, {"references":["Langerhans, P. (1880) Die Wurmfauna von Madeira, 3. Zeitschrift fur wissenschaftliche Zoologie. Leipzig, 34, 87 - 143.","Blake, J. A & Kudenov, J. D. (1978) Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera. Memoirs of the National Museum of Victoria, 39, 171 - 280.","Hartman, O. (1951) The littoral marine annelids of the Gulf of Mexico. Publications of the Institute of Marine Science, Texas University, 2, 7 - 124.","Lewis, J. B. (1998) Reproduction, larval development and functional relationships of the burrowing, spionid polychaete Dipolydora armata with the calcareous hydrozoan Millepora complanata. Marine Biology, 130, 651 - 662.","Radashevsky, V. I. & Nogueira, J. M. M. (2003) Life history, morphology and distribution of Dipolydora armata (Polychaeta: Spionidae). Journal of Marine Biological Association United Kingdom, 83, 375 - 384.","Bartsch, P. (1931) Descriptions of new marine mollusks from Panama, with a figure of the genotype of Engina. Proceedings of the United States National Museum, 79 (15), 1 - 10.","Lamarck, J. B. (1822) Histoire Naturelle des Animaux sans Vertebres, 6 (2), 232 pp. Paris.","Woodwick, K. H. (1964) Polydora and related genera (Annelida, Polychaeta) from Eniwetok, Majuro, and Bikini Atolls, Marshall Islands. Pacific Science, 18, 146 - 159.","Blake, J. A. (1983) Polychaetes of the family Spionidae from South America, Antarctica, and adjacent seas and islands. Biology of Antartctic Seas XIV. Antarctic Research Series, 39, 205 - 288.","Martin, D. (1996) A new species of Polydora (Polychaeta, Spionidae) associated with the excavating sponge Cliona viridis (Porifera, Hadromerida) in the Northwestern Mediterranean Sea. Ophelia, 45 (3), 159 - 174.","Bick, A. (2001) The morphology and ecology of Dipolydora armata (Polychaeta, Spionidae) from the western Mediterranean Sea. Acta Zoologica (Stockholm) 82, 177 - 187."]}

Dipolydora giardi (Mesnil, 1896) Type locality: St Martin, La Manche, Northern France. Type material: No types designated, no depository indicated Radashevsky and Petersen (2005). Location and author of most recently described material: New material: Gulf of Naples, Italy; Recently discovered old material: ZMUC POL- 653 Mer de la Manche, France (Radashevsky & Petersen 2005); comments included on basis of description in Blake and Kudenov (1978). Recorded distribution: Australia: New South Wales, Victoria, south-western Western Australia. La Manche, France; Gulf of Naples, Italy; Cap de Creus, Mediterranean Spain; Lundy Island, Great Britain; California;; Chile; Ecuador; New Zealand (Otago and Wellington)., Published as part of Walker, Lexie M, 2011, A review of the current status of the Polydora - complex (Polychaeta: Spionidae) in Australia and a checklist of recorded species, pp. 40-62 in Zootaxa 2751 on page 51, DOI: 10.5281/zenodo.203565, {"references":["Radashevsky, V. I. & Petersen, M. E. (2005) On the morphology and distribution of Dipolydora giardi and status of D. trilobata (Polychaeta: Spionidae). Zootaxa, 25 - 36.","Blake, J. A. & Kudenov, J. D. (1978) The Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera. Memoirs of the National Museum of Victoria, 39, 171 - 280."]}

Dipolydora socialis (Schmarda, 1861) Type locality: Vi��a del Mar, Chile. Type material: Lost? Location and author of most recently described material: 1) Of type: Mesnil (1896); 2) California, USA (Blake 1996); no Australian material examined 3) Japan (Sato-Okoshi 2000). Recorded distribution: Australia: New South Wales, Victoria. Sea of Japan; western Pacific; Japan; east and west coasts North America; Gulf of Mexico; Chile; Faulkland Islands., Published as part of Walker, Lexie M, 2011, A review of the current status of the Polydora - complex (Polychaeta: Spionidae) in Australia and a checklist of recorded species, pp. 40-62 in Zootaxa 2751 on page 52, DOI: 10.5281/zenodo.203565, {"references":["Schmarda, L. K. (1861) Neue wirbellose thiere beobachtet und gesammelt auf einer reise un die erdr 1853 bis 1857. Vol. I. Turbellarian, Rotatorien und Anneliden. Leipzig: Wilhelm Engelmann.","Blake, J. A. (1996) Family Spionidae Grube, 1850 In: J. A. Blake, B. Hilbig & P. H. Scott (Eds), The Annelida: Part 3 Polychaeta: Orbiniidae to Cossuridae. Santa Barbara Museum of Natural History Santa Barbara, California, pp. 81 - 224."]}

Dipolydora protuberata (Blake and Kudenov, 1978) Type locality: Port Philip Bay, Victoria, Australia. Type material: Holotype: NMV G 2870; Paratypes: NMV G 2871. Location and author of most recently described material: Blake and Kudenov (1978). Recorded distribution: Known only from type locality. Not reported since original description., Published as part of Walker, Lexie M, 2011, A review of the current status of the Polydora - complex (Polychaeta: Spionidae) in Australia and a checklist of recorded species, pp. 40-62 in Zootaxa 2751 on page 52, DOI: 10.5281/zenodo.203565, {"references":["Blake, J. A. & Kudenov, J. D. (1978) The Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera. Memoirs of the National Museum of Victoria, 39, 171 - 280."]}

Dipolydora anatentaculata Delgado-Blas, 2008 Dipolydora anatentaculata Delgado-Blas, 2008: 7 ���8, fig. 2. Type locality. Rookery Bay, Florida, USA (26 �� 20��N, 81 �� 78 ���W), in gastropod shells, Cerithium eburneum. Type material. Holotype (USNM 1073344). Records. Only known from the type locality., Published as part of Delgado-Blas, Victor Hugo & Salazar-Silva, Patricia, 2011, Taxonomic catalogue of the Spionidae (Annelida: Polychaeta) of the Grand Caribbean, pp. 39-66 in Zootaxa 2782 on pages 43-44, DOI: 10.5281/zenodo.206833, {"references":["Delgado-Blas, V. H. (2008) Polydora and related genera (Polychaeta: Spionidae) from the Grand Caribbean region. Journal of Natural History, 42, 1 - 19."]}

Dipolydora caulleryi (Mesnil, 1897) Polydora caulleryi Mesnil, 1897: 87 ���89, plate 3, figs 12���16.��� Fauvel, 1927: 54 ���55, fig. 19.��� Hartman, 1951: 83 ���84.��� Johnson, 1984: 6.25, 6.28, figs 6.17���6.18. Type locality. France Type material. Not found Records. USA: Off South-western Florida, from mud and silt tubes in crevices of Carrier-Shell, Tugurium (Trochotugurium) longleyi, 214 m (Hartman 1951); off Texas and off Florida, 10���189 m, coarse to very fine sand, silty very fine sand, clayey sandy silt, clayey sand (Johnson 1984). Remarks. The specimens recorded in the Grand Caribbean as Dipolydora caulleryi are from different depths, their identity is based in published descriptions and should be confirmed against the type material., Published as part of Delgado-Blas, Victor Hugo & Salazar-Silva, Patricia, 2011, Taxonomic catalogue of the Spionidae (Annelida: Polychaeta) of the Grand Caribbean, pp. 39-66 in Zootaxa 2782 on page 44, DOI: 10.5281/zenodo.206833, {"references":["Mesnil, F. (1897) Etudes de morphologie externe chez les Annelides. II. Remarques complementaires sur les Spionidiens. La famille nouvelle des Disomidiens. La place des Aonides (sensu Tauber, Levinsen). Bulletin scientifique de la France et de la Belgique, 30, 83 - 100, plate 3.","Fauvel, P. (1927) Polychetes Sedentaires & Addenda aux Polychetes Errantes. Faune de France, 16, 1 - 494.","Hartman, O. (1951) The littoral marine annelids of the Gulf of Mexico. Publications of the Institute of Marine Science, Texas University, 2, 7 - 124.","Johnson, P. G. (1984) Family Spionidae Grube, (1850) pp. 6.1 - 6.69 In: Uebelacker, J. M. & P. G. Johnson (eds.), Taxonomic Guide to the Polychaetes of the Northern Gulf of Mexico. Barry A. Vittor & Associates, Inc. Mobile, AL. pp. 6 - 1 to 6 - 69."]}

Dipolydora aciculata (Blake and Kudenov, 1978) Type locality: Bourne Creek, Kilcunda, Victoria, Australia; in Haliotis shell. Type material: Holotype: NMV G 2872, missing prostomium and peristomium; Paratypes NMV 2873 Good. Location and author of most recently described material: Western Australia (Ocean Reef & Windy Harbour), in Haliotis roei, H. conicopora, Turbo torquatus, Patelloida alticostata (Sato-Okoshi et al. 2008). Recorded distribution: Australia: Kilcunda, Victoria; south-western Western Australia., Published as part of Walker, Lexie M, 2011, A review of the current status of the Polydora - complex (Polychaeta: Spionidae) in Australia and a checklist of recorded species, pp. 40-62 in Zootaxa 2751 on pages 50-51, DOI: 10.5281/zenodo.203565, {"references":["Blake, J. A. & Kudenov, J. D. (1978) The Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera. Memoirs of the National Museum of Victoria, 39, 171 - 280.","Sato-Okoshi, W., Okoshi, K. & Shaw, J. (2008) Polydorid species (Polychaeta: Spionidae) in south-western Australian waters with special reference to Polydora uncinata and Boccardia knoxi. Journal of the Marine Biological Association of the United Kingdom, 88, 491 - 501."]}

Dipolydora keulderae sp. n. Figs 3, 4 Etymology: This species is named for Felicia Keulder who helped with the sampling. Description: Small species, body equal width along length, narrows for last five chaetigers. Holotype 7.5 mm for 82 chaetigers and 0.4 mm wide at chaetiger 5 (Paratypes ranging from 4 mm for 45 chaetigers to 6 mm for 61 chaetigers). Prostomium notched, caruncle extends to posterior margin of chaetiger 3, but to end of chaetiger 2 in one specimen; no occipital tentacle; in holotype two pairs of eyes arranged in trapezoid, 0–2 pairs in others, posterior pair larger; no pigmentation (Fig. 3A). Palps extend back approximately to chaetiger 14. Chaetiger 1 with finger-like notopodial lobes that usually obscure single, fine notochaeta emerging anterior to lobe (Fig. 3A; lobes orientated posteriorly to show short notochaetae), neuropodial lobe small, inconspicuous, tuft of unilimbate neurochaetae; chaetigers 2–4, notochaetae arranged in three tiers, first row short, slightly bent and unilimbate, second row slightly longer than first row and spear-shaped, third row long and lanceolate; chaetigers 6–10 as on anterior chaetigers, but chaetae of first row more sharply bent; in subsequent chaetigers short chaetae fewer, with longer chaetae increasing in number; no modified posterior notochaetae. Neuropodial lobes of chaetigers 2–4, 6 small, neurochaetae unilimbate. Bidentate hooded hooks from chaetiger 7; 3, 5 and 1 per series on anterior, median and posterior chaetigers in holotype; 3–4, 6–7, 1 in paratypes; no constriction on shaft, in anterior hooks angle between main fang and shaft>90°, angle between teeth approximately 45°, accessory tooth decreasing in size until hooks in final eight (up to 11 in other specimens) chaetigers unidentate without hoods (Fig. 3F), number of chaetigers with unidentate hooks apparently not proportional to total number of chaetigers (Fig. 4); no companion chaetae with anterior hooded hooks in holotype, but present on one ramus each on chaetigers 7 and 8 in one specimen; last two chaetigers with fine capillary companion chaeta. Chaetiger 5 1.5× length of preceding and succeeding chaetigers; five modified spines alternating with bilimbate companion chaetae (Fig. 3B), modified spines falcate with collar on convex side of curved end of spine, in worn spines collar appears as a narrow ridge or cuff (Fig. 3C); dorsal fascicle of 3 or 4 geniculate bilimbate chaetae (Fig. 3D); fascicle of short unilimbate neurochaetae. Branchiae from chaetiger 7 (Fig. 3A), first two pairs very small, first obvious pair on chaetiger 9; finger-like, individual branchiae not meeting at dorsal midline; 15 branchiate chaetigers in holotype, 6–12 in paratypes, in specimens with 40 to 60 chaetigers, number of branchiate chaetigers generally proportional to total number of chaetigers (Fig. 4), with longest worm with 89 chaetigers not having more branchiate chaetigers. Pygidium reduced, with four lobes, dorsal pair smaller than ventral pair (Fig. 3E), in some specimens looks cuff-shaped. Comparison: D. keulderae sp. n. belongs to the D. bidentata / barbilla group (sensu Blake, 1996). The species resembles all other members of this group in having a notched or bilobed prostomium, modified spines with a collar on the convex side of the curved end of the spine, and hooded hooks from chaetiger 7. D. keulderae most closely resembles D. barbilla Blake, 1980 and D. pilocollaris Blake & Kudenov, 1978. Generally, the length of the caruncle is the same in all three species. Common to D. keulderae and D. barbilla are the maximum number of hooded hooks and the loss of the second tooth in the hooded hooks; however, D. barbilla has heavy posterior notopodial spines (Blake 1980) that are lacking in D. keulderae. D. keulderae and D. pilocollaris are similar in the start of the branchiae and the absence of posterior notopodial spines, but differ with respect to the maximum number of hooded hooks per fascicle and the presence of unidentate hooks posteriorly (Blake & Kudenov 1978). D. keulderae differs from both species with respect to the fine structure of the modified spines on chaetiger 5 and companion chaetae: in both D. barbilla and D. pilocollaris the falcate spines have bristled collars, while the latter also has hastate companion chaetae. D. keulderae also has companion chaetae accompanying the hooded hooks for only the last two chaetigers (the presence of a winged chaeta with only the hooded hooks on the right neuropodia on chaetigers 7 and 8 in one specimen suggests that this is an anomaly), while the other two species have companion chaetae with the anterior hooded hooks. D. pilocollaris also lacks notochaetae on chaetiger; in D. keulderae these notochaetae are present but very inconspicuous. Holotype: SOUTH AFRICA: Eastern Cape: SAMC A60060, Canon Rocks, Port Alfred, coralline algae, 28.i.2005, C.A. Simon. Paratypes: SOUTH AFRICA: Western Cape: SAMC A60061 (2), Struisbaai, host Turbo sarmaticus, 7.iii.2005, C.A. Simon; SAMC A60071 (6), Hermanus, host wild Haliotis midae, Anna Mouton (shells processed by Simone Neethling). Eastern Cape: SAMC A60062 (ethanol specimens), Port Elizabeth, host Crassostrea gigas, 21 x.2009, C.A. Simon. Distribution: Found at Haga Haga, Canon Rocks and Kowie Rocks (Port Alfred), Port Elizabeth, Grootbank (Tsitsikamma), Mossel Bay, Struisbaai and Hermanus. Habitat: Holotype found in coralline algae, but paratype found boring in the shells of Crassostrea gigas (Thunberg, 1793), Turbo sarmaticus L., 1758 and Haliotis midae L., 1758., Published as part of Simon, Carol A., 2011, Polydora and Dipolydora (Polychaeta: Spionidae) associated with molluscs on the south coast of South Africa, with descriptions of two new species, pp. 39 in African Invertebrates 52 (1) on pages 43-46, DOI: 10.5733/afin.052.0104, http://zenodo.org/record/7914609, {"references":["BLAKE, J. A. 1980. Polydora and Boccardia species (Polychaeta: Spionidae) from western Mexico, chiefly from calcareous habitats. Proceedings of the Biological Society of Washington 93: 947 - 962.","BLAKE, J. A. & KUDENOV, J. D. 1978. The Spionidae (Polychaeta) from south-eastern Australia and adjacent areas with a revision of the genera. Memoirs of the Natural Museum of Victoria 39: 171 - 280."]}

Dipolydora socialis (Schmarda) Leucodore socialis Schmarda, 1861 Ecology: FT, CP Distribution: PA, NA, NT, AU Habitat: ES Comments: The widespread distribution of this species on both the east and west coasts of North America, South America, Japan and in Europe suggests that multiple species have been reported under this name and/or human-assisted introductions have occurred. The species is reported to contain both boring and non-boring populations (Blake 1971; Sato-Okoshi 2000). Additional references: D. Karlen, pers. comm. (Tampa Bay survey), Published as part of Glasby, Christopher J., Timm, Tarmo, Muir, Alexander I. & Gil, Jo��o, 2009, Catalogue of non-marine Polychaeta (Annelida) of the World, pp. 1-52 in Zootaxa 2070 on page 29, DOI: 10.5281/zenodo.187085, {"references":["Schmarda, L. K. (1861) Neue wirbellose Thiere beobachtet und gesammelt auf einer Reise um die Erde 1853 bis 1857. Vol. 1. Turbellarien, Rotatorien und Anneliden. Pt. 2. Verlag von Wilhelm Engelmann, Leipzig, 164 pp.","Blake, J. A. (1971) Revision of the genus Polydora from the east coast of North America (Polychaeta: Spionidae). Smithsonian Contributions to Zoology, 75, 1 - 32.","Sato-Okoshi, W. (2000) Polydorid species (Polychaeta: Spionidae) in Japan, with descriptions of morphology, ecology and burrow structure. 2. Non-boring species. Journal of the Marine Biological Association of the United Kingdom, 80, 443 - 456."]}

Dipolydora caulleryi (Mesnil) Polydora caulleryi Mesnil, 1897 Polydora brachycephala Hartman, 1936 b Ecology: FT Distribution: PA, NA Habitat: ES Comments: The widespread distribution (and habitat preferences) of this species on both the east and west coasts of North America, and in Europe, suggests that multiple species have been reported under this name and/or human-assisted introductions have occurred. Additional references: Hartman (1936 b), Wesenberg-Lund (1958), Blake (1971), Published as part of Glasby, Christopher J., Timm, Tarmo, Muir, Alexander I. & Gil, Jo��o, 2009, Catalogue of non-marine Polychaeta (Annelida) of the World, pp. 1-52 in Zootaxa 2070 on page 29, DOI: 10.5281/zenodo.187085, {"references":["Mesnil, F. (1897) Etudes de morphologie externe chez les Annelides. II. Remarques complementaires sur les Spionidiens. - La famille nouvelle des Disomidiens. - La place des Aonides (sensu Tauber, Levinsen). Bulletin Scientifique de la France et de la Belgique, 30 (1), 83 - 100.","Hartman, O. (1936 b) New species of Spionidae (Annelida Polychaeta) from the coast of California. University of California Publications in Zoology, 41 (6), 45 - 52.","Wesenberg-Lund, E. (1958) Lesser Antillean polychaetes, chiefly from brackish water, with a survey and a bibliography of fresh and brackish-water polychaetes. Studies on the Fauna of Curacao and other Caribbean Islands, 8, 1 - 41. [In Uitgaven \" Natuurwetenschappelijke Studiekring voor Suriname en de Nederlandse Antillen \", 17, 1 - 41]","Blake, J. A. (1971) Revision of the genus Polydora from the east coast of North America (Polychaeta: Spionidae). Smithsonian Contributions to Zoology, 75, 1 - 32."]}

A new genus and species of parasitic copepod (Clausiidae), Spionicola mystaceus, associated with the polychaete Dipolydora armata (Spionidae) is described and figured. The new copepod has an elongate body, 5-segmented antennule, 2-segmented rami on legs 1 and 2, 2 spines representing leg 3, no leg 4, leg 5 well developed and reduced armature elements on feeding limbs. The host is a mollusk-shell borer, collected off São Sebastião Island, State of São Paulo, Brazil.Novo gênero e nova espécie de Clausiidae (Crustacea, Copepoda) parasita de Dipolydora armata (Polychaeta, Spionidae) no Brasil: Descrição e ilustração de novo gênero e nova espécie de um copépodo parasita (Clausiidae), Spionicola mystaceus, associado ao poliqueto Dipolydora armata (Spionidae). O novo copépodo tem um corpo alongado, antenula penta articulada , ramos biarticulados nos pereiópodos um e dois, dois espinhos representando o pereiópodo 3, pereiópodo 4 ausente, pereiópodo 5 muito desenvolvido e estruturas dos apêndices bucais reduzidas. O hospedeiro é um perfurador de conchas de moluscos coletado ao largo da Ilha de São Sebastião, Estado de São Paulo, Brasil.

A study was conducted to identify and quantify the spionid polychaetes that infest cultured abalone Haliotis midae at aquaculture facilities on the west (Farm A), south (Farms B and C) and east (Farm D) coasts of South Africa. The relationship between total intensity of infestation by the polydorids and the condition of their hosts at Farms A, B and D were also measured. The abalone were infested by Dipolydora capensis, Polydora hoplura and a Boccardia sp. However, the intensity of infestation differed among sites, suggesting that both geographical and farm-specific conditions may influence species composition. Total intensity of infestation has a negative impact on the condition of abalone at the three farms tested, but at Farms A and B, this effect was reduced in larger abalone.Keywords: Boccardia, condition factor, Dipolydora, PolydoraAfrican Journal of Marine Science 2006, 28(1): 167–171

Dipolydora giardi (Mesnil, 1896) (Fig. 2) Polydora giardi Mesnil, 1896: 195 ���202, pl. 13, figs. 1���12. Fauvel, 1927: 50 ���52, fig. 17 h���m. Dipolydora giardi: Blake, 1996: 186. [New combination introduced. D. giardi record uncertain]. Material. FRANCE: La Manche, coll. & id. F. Mesnil, accessioned ZMUC 22 June 1896, ZMUC POL�� 653 (3 spec.). ITALY: Ischia Is., Gulf of Naples, 40 ��44.8' N, 13 ��56.7' E, 2 m, from shells of the whelks Stramonita haemastoma (L.) and Hexaplex trunculus (L.), coll. V. I. Radashevsky, 18 Jul 1994, BMNH 2002.620 ��� 629 (17 spec.). Comparative material. Dipolydora trilobata: RUSSIA: Sea of Japan, Vostok Bay of Peter the Great Bay, 42 ��53.6' N, 132 ��44.1' E, from shells of the oyster Crassostrea gigas (Thunberg), ZISP 1 / 46576 �� 3 / 46578 (holotype + 19 paratypes); IMBV 1 / 12171 (50 paratypes); USNM 183520 (30 + spec.). Description. Mesnil���s material consists of three complete individuals 3���4 mm long and 0.3���0.4 mm wide for c. 60, 70, and 75 chaetigers. One individual regenerating two anterior segments. Largest complete specimen from Italy c. 4 mm long and 0.3 mm wide for 60 chaetigers. Body pigmentation absent. Prostomium incised to bifid anteriorly. Caruncle extending posteriorly to end of chaetiger 3. Eyes and nuchal antenna absent (Fig. 2 A). Palps extending posteriorly for 10���15 chaetigers. Chaetiger 1 with postchaetal lamellae in both rami; both noto�� and neurochaetae short capillaries, fewer than in following chaetigers. Posterior notopodia with only capillaries, without spines. Chaetiger 5 greatly modified, larger than chaetigers 4 or 6, without postchaetal lamellae, with 2 or 3 dorsal superior geniculate capillaries (Fig. 2 H), 4 or 5 major spines alternating with bilimbate��tipped companion chaetae (Fig. 2 I), and a tuft of 4 or 5 ventral capillaries (Fig. 2 G). Major spines falcate, with lateral tooth on one side and fine spur or protuberance on the other side; spines arranged in a slightly curved diagonal row; lateral tooth occasionally broken in older spines positioned in anterior part of the row. Hooded hooks in neuropodia from chaetiger 7, up to 5 in a series, accompanied by single, winged inferior capillary chaetae in chaetigers 7���9 (Fig. 2 D) and by 1 or 2 hair��like capillaries situated between hooks in a few posterior chaetigers (Fig. 2 E). Hooks bidentate throughout, with angle between main fang and shaft wider in posterior chaetigers; hook shaft slightly curved, without constriction (Fig. 2 C, F). Branchiae from chaetiger 9���10, usually from 10 but one specimen with short branchial buds present on chaetiger 9 (Fig. 2 A); branchiae full��sized from chaetiger 11, absent from posterior half of body. Branchiae flattened, with surfaces oriented parallel to body axis, free from notopodial postchaetal lamellae. Pygidium small, cup��shaped, with one ventral lobe and two smaller dorsal lobes, appearing white because of large number of spindle��shaped mucous cells (Fig. 2 B). Glandular pouches from chaetiger 6, full��sized from chaetiger 7 or 8, diminishing in size after chaetiger 10 or 11. Gizzard��like structure present at about chaetiger 17. Hind gut white. Metanephridia opening middorsally via two separate nephridiopores on fertile segments apparent under methyl��green staining in a 40 ��segment posterior fragment from Italy which had oocytes up to 130 ��m in diameter in middle segments. No sperm were recognized in this fragment or in other specimens. Methyl��green staining patterns. Head and anterior chaetigers do not retain stain. Staining dorsally as paired spots, one on each half of chaetiger, usually from chaetiger 8 or 9, occasionally from chaetiger 7; size of spots and intensity of staining gradually increasing posteriorly. Remarks. No differences in size, morphology and life style (boring in calcareous substrata) could be detected between the Mesnil���s worms from northern France and those from the Mediterranean and thus we are confident they are D. giardi sensu stricto Mesnil as indicated. Two specimens from the Mesnil���s sample found in Copenhagen have an indistinct pygidium, but one of them clearly has a middorsal cleft and two dorso��lateral clefts that divide the cup into one ventral lobe and two smaller dorsal lobes. The same trilobed pygidium was also invariably present in all specimens collected by us from Italy. Dipolydora trilobata specimens examined for the comparison resemble D. giardi in that they also have an incised to bifid prostomium, caruncle extending to the end of chaetiger 3, pygidium with three small lobes, major falcate spines of chaetiger 5 with large lateral tooth on one side and a smaller accessory spur on the other side. Maximal size of worms and boring mode of life are also identical. Both species lack eyes, occipital antenna and modified spines in posterior notopodia. However, branchiae in D. giardi begin on chaetiger 10 (19 specimens in 20 examined), rarely on chaetiger 9 (1 specimen), whereas in D. trilobata they usually start on chaetiger 9 (46 specimens in 83 examined), occasionally on chaetiger 8 or 10 (23 and 13 specimens in 83 examined, respectively), and rarely on chaetiger 7 (1 specimen). Hooded hooks in D. giardi are distinctly bidentate in all neuropodia, whereas in D. trilobata they become unidentate in 2���5 posterior chaetigers (Radashevsky 1993: fig. 17 I). A more fundamental difference may be that Mesnil's D. giardi were reportedly simultaneous hermaphrodites (Mesnil 1896: p. 200) while D. trilobata is a gonochoristic species (Radashevsky unpublished). The description of gamete arrangement in D. giardi provided by Mesnil is, however, ambiguous and hermaphroditism in the species needs to be confirmed by further studies. Polydora anoculata Moore, 1907 from Massachusetts is also a similar species. Blake (1971) examined the holotype and emended the diagnosis. Subsequently Maciolek (1984) did not find differences between Moore���s and Mesnil���s species and placed P. anoculata into the synonymy of P. giardi. Blake (1971) and Maciolek (1984) reported that specimens from Massachusetts and North Carolina had the caruncle extending posteriorly to chaetiger 3���5, branchiae beginning from chaetiger 9���10, and the pygidium small and disclike, with only a dorsal notch. However, these characteristics indicate that Dipolydora anoculata is not only validly described, but also distinct from D. giardi and D. trilobata., Published as part of Radashevsky, Vasily I. & Petersen, Mary E., 2005, On the morphology and distribution of Dipolydora giardi and status of D. trilobata (Annelida: Spionidae), pp. 25-36 in Zootaxa 1086 on pages 28-31, DOI: 10.5281/zenodo.170484, {"references":["Mesnil, F. (1896) Etudes de morphologie externe chez les Annelides. I. Les Spionidiens des cotes de la Manche. Bulletin scientifique de la France et de la Belgique, 29, 110 - 287.","Fauvel, P. (1927) Polychetes sedentaires. Addenda aux Errantes, Archiannelides, Myzostomaires. Faune de France, 16, 1 - 494.","Blake, J. A. (1996) Family Spionidae Grube, 1850. Including a review of the genera and species from California and a revision of the genus Polydora Bosc, 1802. In: Blake, J. A., Hilbig, B. & Scott, P. H. (Eds.), Taxonomic Atlas of the Benthic Fauna of the Santa Maria Basin and Western Santa Barbara Channel. Volume 6. The Annelida Part 3 - Polychaeta: Orbiniidae to Cossuridae. Santa Barbara Museum of Natural History, Santa Barbara, California, pp. 81 - 223.","Radashevsky, V. I. (1993) Revision of the genus Polydora and related genera from the North West Pacific (Polychaeta: Spionidae). Publications of the Seto Marine Biological Laboratory, 36, 1 - 60.","Moore, J. P. (1907) Descriptions of new species of spioniform annelids. Proceedings of the Academy of Natural Sciences of Philadelphia. 59: 195 - 207.","Blake, J. A. (1971) Revision of the genus Polydora from the east coast of North America (Polychaeta: Spionidae). Smithsonian Contributions to Zoology, 75, 1 - 32.","Maciolek, N. J. (1984) A new species of Polydora (Polychaeta: Spionidae) from deep waters in the north-west Atlantic Ocean, and new records of other polydorid species. Sarsia, 69, 123 - 131."]}

The spionid polychaete Dipolydora giardi (Mesnil, 1896), originally described as a borer in coralline algae from northern France, was later reported world-wide boring into various shells, sponges and also inhabiting mud tubes on soft bottoms. The reported morphological variability and wide range of habitats suggested that more than one species might be involved. In the present study, D. giardi is redescribed based on some material from northern France sent by F. Mesnil in 1896 to the Zoological Museum, University of Copenhagen, and on new material collected in Italy. The species is characterized by its incised prostomium, caruncle to the end of chaetiger 3, branchiae on the anterior half of the body (usually beginning on chaetiger 10), pygidium with one ventral lobe and two smaller dorsal lobes, and heavy falcate spines of chaetiger 5 with a large lateral tooth on one side and a smaller accessory spur on the other side. Dipolydora trilobata (Radashevsky, 1993) is a closely similar species, but is maintained as distinct. Dipolydora anoculata (Moore, 1907) is probably a valid species and should be removed from the synonymy of D. giardi.

The functional morphology of the feeding palps and prostomium of the spionid polychaetes Streblospio benedicti and S. shrubsolii was studied. Three functional groups of cilia of the feeding palps were found on both species — frontal cilia, latero-frontal cirri and lateral cilia. Frontal cilia line the food groove and transport food particles to the pharynx, and have been reported for all spionid polychaetes except species of the genus Scolelepis. Laterofrontal cirri deflect particles onto the frontal surface and have been observed in several spionid genera including Paraprionospio, Streblospio, Polydora and Dipolydora. Lateral cilia beat in continuous metachronal waves creating lateral vortices that potentially entrain suspended particles, and are known in Paraprionospio and Streblospio. The two species of Streblospio did differ in the distribution of prostomial papillae. These papillae are eversible and thought to function in particle selection as particles on the pharynx come in contact with the papillae. Prostomial papillae were restricted to the peripheral surface of S. benedicti and were widely scattered on all surfaces of the prostomium of S. shrubsolii. A conical tentaculate structure occurs between the branchiae of the first setiger of S. benedicti, but only a low raised elevation is present on S. shrubsolii.