taxonID	type	description	language	source
F72C87EBFF8BFFB7057EFBA988823705.taxon	description	4. Polydora carinhosa sp. nov.	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF88FFA2057EFE998EED3635.taxon	description	Polydora ecuadoriana Blake (1983, pp. 257 – 258, Fig. 26).	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF88FFA2057EFE998EED3635.taxon	materials_examined	Material ECUADOR, Manabí, Bahia de Santa Elena, 200 ms offshore, 1 / 2 mile NE of Punta Ballenita, ANTON BRUUN R / V, cruise 16, st. SE- 6669, 02 ° 12.33´S, 80 ° 52.17´W, 8 – 10 m, 8 May 1966, coll. S. A. Earle, USNM 59920 (20 paratypes); st. SE- 6670, 02 ° 11.47´S, 80 ° 56.52´W, 8 – 9 m, 8 May 1966, USNM 60549 (2 paratypes).	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF88FFA2057EFE998EED3635.taxon	description	MEXICO, Caribbean Sea: Veracruz, Tamiahua Lagoon, st. E- 6, 25 ° 35´N, 97 ° 35´W, 1 m, from oyster Crassostrea sp., 10 Jan 1993, coll. V. H. Delgado-Blas, ECOSUR SPIO- 75 (7); Quintana Roo, st. CP 32 - 4 41 - 3, ECOSUR SPIO- 76 (1). BRAZIL, Espírito Santo: Espírito Santo Bay: Ilha do Frade, 20 ° 18´S, 40 ° 17´W, intertidal, from shells of the oyster C. rhizophorae, coll. R. C. Nalesso and V. I. Radashevsky, 29 May 2003, MZSP 167 (9), 16 Jan 2004, MZSP 171 (24), 25 Feb 2004, MZSP 315 (6), MZSP 313 (23); Maria Ortiz, 20 ° 15´S, 40 ° 18´W, intertidal, 20 Feb 2004, coll. K. M. Machado, MZSP 168 (16). São Paulo: São Sebastião, Praia das Cigarras, 23 ° 43.7´S, 43 ° 24´W, intertidal, from shells of various gastropods inhabited by hermit crabs, 24 Mar 2004, coll. V. I. Radashevsky (10 +, notes on live material, not fixed). São Sebastião, Praia do Saco Grande, 23 ° 49.7´S, 45 ° 25.5´W, 1 – 5 m, from shells of gastropods Stramonita haemastoma (Linnaeus, 1767), Strombus pugilis (Linnaeus, 1758), and Tegula viridula (Gmelin, 1791) inhabited by hermit crabs Paguristes tortugae Schmitt, 1933 and Pagurus brevidactylus (Stimpson, 1859), 30 Jun 2004, coll. V. I. Radashevsky, (10 +, notes on live material, not fixed). São Sebastião Island, Praia do Curral, 23 ° 51.5´S, 45 ° 25.9´W, 3 – 5 m, from shells of the gastropod S. haemastoma inhabited by hermit crabs, 13 May 2004, coll. V. I. Radashevsky, (10 +, notes on live material, not fixed). Paraná: Paranaguá Bay: Praia Grande of Ilha do Mel, 25 ° 32.8´S, 48 ° 17.6´W, intertidal, from shell of the barnacle Megabalanus sp., 13 Aug 1998, coll. V. I. Radashevsky and P. C. Lana, USNM 1022187 (9). Pontal do Sul, mouth of Perequê tidal creek, 25 ° 33.8´S, 48 ° 21.4´W, intertidal, from shells of live gastropod S. haemastoma and oyster C. rhizophorae, 22 Aug 1998, coll. V. I. Radashevsky, USNM 1022185 (32), 1022186 (1). 25 ° 28´S, 48 ° 27´W, 2 m, from shells of the cultured oyster C. rhizophorae, 13 Sep 2001, coll. V. I. Radashevsky, MZSP 166 (1). Brasilia village of Ilha do Mel, 25 ° 33´S, 48 ° 19´W, 0.5 m, from shells of gastropods Pugilina morio (Linnaeus, 1758) and S. haemastoma inhabited by the hermit crab Clibanarius vittatus (Bosc, 1802), 25 Aug 2001, coll. V. I. Radashevsky, MZSP 172 (5). Guaratuba Bay, 25 ° 53´S, 48 ° 34´W, intertidal, from shells of the oysters C. brasiliena, C. gigas, and C. rhizophorae, 3 Jun 2002, coll. V. I. Radashevsky, MZSP 169 (22). Santa Catarina, Florianópolis: North Bight of Ilha de Ratones Grande, 27 ° 28´S, 48 ° 33´W, 1 m, from shell of the cultured oyster C. gigas, coll. Y. M. B. Neptune, 24 Sep 1998, SMF 13967 (3); Praia da Ponta de Sambaqui, 27 ° 28.5´S, 48 ° 33.7´W, 1 m, from shell of the cultured oyster C. gigas, coll. Y. M. B. Neptune, 25 Apr 2003, SMF 14010 (100 +), 15 Aug 2003, MZSP 170 (55), SMF 13930 (45).	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF88FFA2057EFE998EED3635.taxon	materials_examined	Adult morphology (based on material from Brazil) Up to 40 mm long and 1.2 mm wide at chaetiger 7 for 125 chaetigers. Body pale or light tan in life. Remains of larval black pigment often present on dorsal side of anterior chaetigers and on pygidium in individuals with less than 30 chaetigers; bigger individuals without pigmentation or fine black pigment scattered on ventral side of posterior chaetigers. Pigmentation on palps greatly variable, including fine black lines along longitudinal ciliated groove (Fig. 2 A), up to 7 separate black bars (Fig. 6 A) or lacking. Presence of pigmentation and its pattern not strongly correlated with total number of chaetigers in an individual (Fig. 3 B). Bars as paired gatherings of pigment on either side of groove, not extending all along palp periphery. Number of bars on left and right palps often different in same individual. Light yellow pigment present along ciliated groove between black bars on palps in some live individuals. Black pigment usually retained but yellow pigment absent in formaldehyde-fixed specimens. Prostomium anteriorly incised and in large individuals curved downwards (Figs 4 A, 5 A), thus incision visible only in ventral view (Fig. 5 B). Caruncle extending to end of chaetiger 3, shorter in small individuals (Fig. 3 C). Occipital antenna absent. One to four black eyes present or eyes absent; their number not correlated with total number of chaetigers (Fig. 3 A). Palps as long as 10 – 20 chaetigers, with longitudinal groove lined with fine frontal cilia, latero-frontal motile compound cilia bordering groove, short lateral papillae with non-motile cirri arranged along either side of groove, and short compound non-motile cilia arising directly from palp surface and scattered on lateral and abfrontal palp surfaces. Chaetiger 1 with short capillaries in neuropodia and well developed cirriform postchaetal lamellae in both rami; notochaetae absent. Capillary chaetae in notopodia gradually becoming fewer, shorter and alimbate in posterior chaetigers. Chaetiger 5 greatly enlarged, overlapping chaetiger 6 dorsally, with 3 – 5 dorsal superior winged capillaries, up to 8 major modified spines alternating with bilimbatetipped companion chaetae and arranged in slightly curved, diagonal row, and 5 – 8 winged ventral capillaries; postchaetal lamellae absent (Fig. 5 C). Dorsal superior and ventral capillaries shorter and fewer than those on chaetigers 4 and 6. Major spines falcate, with accessory lateral tooth or flange and thin subdistal longitudinal flange located laterally on main fang above lateral tooth (Fig. 2 D); subdistal flange often worn and absent on spines in large individuals. Hooks in neuropodia from chaetiger 7, bidentate, with constriction on shaft, up to 9 in a series (Figs 2 B, C, 5 D). Branchiae from chaetiger 7, full-sized from chaetigers 9 – 10, gradually diminishing in size along posterior half of body, distributed to almost end of body or absent on posterior one fourth to one half of body. Pygidium large, scoop-shaped in juveniles and adults, same color as body, with wide dorsal gap, dorsal anus, postanal ciliary band and very few secretory cells (Figs 4 B – E, 5 E, F). Lateral ciliated organs as small pits between noto- and neuropodia on all chaetigers except 4 and 5 (Figs 4 A, 5 C). Glandular pouches from chaetiger 7, large in chaetigers 7 to 9 – 10, then gradually diminishing in size, single throughout. Large, flask-shaped secretory cells of pouches opening to exterior individually and appearing externally as small papillae below vertical rows of neurochaetae (Fig. 5 D). Digestive tract without gizzard-like structure. Metanephridial segmental organs from chaetiger 7, opening to exterior laterally on anterior, sterile segments and dorsally on gametogenic segments; paired nephridia opening separately on all segments. Distal part of segmental organs in gametogenic segments in females inflated, whitish in life, in formaldehyde-fixed specimens usually absorb methyl green stain intensely. Middle part of segmental organs in gametogenic segments in males formed by large urn-shaped cells and expanded. Habitat	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF88FFA2057EFE998EED3635.taxon	description	Polydora ecuadoriana is a borer in various calcareous substrata. In Ecuador it was found in shallow water boring into coralline algae and mollusk shells (Blake 1983). In Brazil it occurred intertidally and in shallow water boring into shells of live oysters Crassostrea brasiliana, C. gigas, C. rhizophorae, the barnacle Megabalanus sp. and empty shells of the gastropods Pugilina morio, Stramonita haemastoma, Strombus pugilis, and Tegula viridula inhabited by hermit crabs Clibanarius vittatus, Paguristes tortugae and Pagurus brevidactylus. The worms resided in U-shaped burrows (Fig. 6 C) within the shell. Each burrow opened to the exterior via two joined apertures forming a characteristic 8 - shaped opening (Fig. 6 B). Walls of the burrows were lined with fine silt and each aperture was extended by a smooth silty tube up to 10 mm high (Fig. 7 B). Paired palps of a worm can usually be seen protruding out of one of the tubes and collecting food particles suspended in the water and deposited on the surrounding surface (Fig. 6 A). The worms often caused the formation of dark blisters on the inner shell surface both in native and cultured oysters (Fig. 6 C). Severely infested mollusks had up to 50 % of the inner shell surface covered by blisters. Infestations were seasonal and their intensity varied substantially between locations. Up to 10 worms occurred per cm 2 of shell surface and more than one hundred worms were found in one oyster valve. Regeneration Worms regenerating lost anterior and posterior parts were occasionally found in shells. To investigate the regeneration process, various number of segments were cut off of live complete individuals and the fragments were maintained in dishes with filtered sea water in the laboratory at room temperature. In total, 45 worms were used in the experiment. The epithelium healed the same day as the worms were cut, and regenerating buds on both the anterior and the posterior fragments developed in 1 – 2 days. In 2 – 4 days, the anterior blastema differentiated into head and body anlage. Palps began to develop immediately on the head anlage. The body anlage elongated gradually and in 4 – 5 days after the cut became segmented, probably simultaneously, into a definite number of segments. Chaetae, postchaetal lamellae and other chaetiger attributes only developed once segmentation was completed. No more chaetigers were added anteriorly in further growth. The number of chaetigers regenerated anteriorly depended on the number of lost chaetigers. The two numbers were equal if eight or less chaetigers were cut off. For example, if the head and five anterior chaetigers were cut off in the experiment, the head and five chaetigers were differentiated from the anterior blastema. If, however, nine or more chaetigers were cut off (30 worms cut in the experiment), only eight chaetigers regenerated anteriorly. All the anterior regenerates developed nototrochs from chaetiger 3 onwards apart from chaetiger 5. The posterior blastema differentiated into pygidium and the prepygidial growth zone in 3 – 5 days after a cut. New posterior segments developed successively one by one following the formation of the growth zone. Reproduction Polydora ecuadoriana is gonochoristic. Of 46 mature individuals, 28 were females and 18 were males. Gametes developed along segmental blood vessels in middle segments, from chaetigers 21 – 47 to 35 – 101. The position of the first gametogenic chaetiger varied equally (from chaetiger 21 to 29) in small and large males (Fig. 3 D), whereas fertile chaetiger started more posteriorly (from chaetigers 25 – 47) in large females than in small ones (Fig. 3 E). In males, the testis contained only spermatogonia; individual primary spermatocytes, diads of secondary spermatocytes, tetrads of spermatids, and individual spermatozoa floated freely together in the coelomic cavity. The smallest male with spermatozoa in the coelomic cavity was about 7 mm long for 57 chaetigers. Spermatids were spherical, 3 µm in diameter. Spermatozoa were introsperm with an elongated straight head about 1 µm in diameter, head + middlepiece 12 µm long, acrosome 2 µm, nucleus 6 µm, middlepiece 4 µm, and flagellum 46 µm long. On one occasion in the laboratory, a male shed spermatophores through the nephridiopores on fertile segments. The spermatophores were fine, sinuous, 3 - 5 mm long and 8 µm in diameter (Fig. 7 A). They were composed of regularly packed spermatozoa enveloped by a fine matrix. In seawater, spermatophores broke apart and spermatozoa started to move in a half an hour. In females, the ovaries contained previtellogenic and vitellogenic oocytes of various diameters. Large oocytes occurred freely in the coelomic cavity. The smallest female with large oocytes in the coelomic cavity was about 10 mm long for 64 chaetigers. Paired intraepithelial seminal receptacles were present on the dorsal side of the female gametogenic chaetigers, posterior to the nototrochs, medial to the base of the branchia. The receptacles were oval chambers, about 20 x 25 µm in diameter, filled up with clusters of inactive spermatozoa. Females deposited up to 2500 eggs into 20 – 50 capsules which were joined to each other in a string. Each egg capsule was attached by two thin stalks to the inner wall of the burrow and contained from 10 to 60 eggs (Fig. 7 B, C). The eggs were spherical, 100 – 110 µm in diameter, filled with yolky globules. The majority of eggs in broods developed synchronously into larvae, but up to 10 % of the eggs did not undergo cleavage and were consumed by developing larvae. Synchronous beating of nototrochs on the dorsal side of the female produced a constant water flow inside the burrow and provided oxygen for the developing larvae. When no yolky globules were left in the digestive tract, the 3 - chaetiger larvae about 280 µm long hatched, escaped from the mother’s burrow and entered the plankton. Females brooding larvae in capsules had the next generation of vitellogenic oocytes up to 50 µm in diameter developing in the ovaries. Egg capsules in burrows occurred from February to September. Life history Males and females become mature when they have grown to about 55 and 65 chaetigers, respectively. Males produce spermatophores and pass them to females (Fig. 7 A). Females accumulate the sperm in seminal receptacles on the dorsal side of the gametogenic segments. The oocytes are steadily produced in ovaries and large oocytes are accumulated in the coelomic cavity within an extended period of reproduction. Females regularly lay broods which mostly contain eggs developing into larvae (Fig. 7 B, C). Fertilization is thought to occur in the egg capsule. Larvae develop 3 chaetigers in the capsule, then are released and continue development in the seawater, feeding on the plankton (Fig. 7 D). They are able to settle when they have grown to 16 – 18 chaetigers (Fig. 7 E). In the absence of suitable substrata, the larvae are able to postpone settlement and grow in the plankton until the 20 – 22 - chaetiger stage. Settlement is accompanied by gradual metamorphosis and loss of provisional larval features. The switch to a new mode of feeding after settlement is enabled by rapid elongation of the palps, modification of the prostomium, enlargement of the ventral peristomial lip and transformation of the lateral peristomial lips into the dorso-lateral ciliary folds. Newly settled individuals first construct a small silty tube on the shell surface (Fig. 7 F) and then begin boring into the shell. Boring is due to a combination of glandular secretion and abrasion, by which process the Ushaped burrow is permanently elongated in its middle part (Fig. 7 B). After settlement, larval pigmentation is totally reduced or retained only on 1 – 3 anterior chaetigers, and black and yellow pigments usually appear on palps in further development. The life span of the species is unknown but preliminary examination of the size structure of the population allows us to assume that adults are able to survive over 2 years in suitable conditions. Larval development of the species will be described elsewhere.	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF88FFA2057EFE998EED3635.taxon	discussion	Remarks Specimens from Brazil fit well the main characteristics of P. ecuadoriana originally described by Blake (1983) from Ecuador. Black bars on palps, incised prostomium, major falcate spines of chaetiger 5 with lateral tooth and subdistal longitudinal flange, and large, scoop-shaped pygidium are the diagnostic features of the species. Blake (1983: p. 258) reported the posterior chaetigers in the species “ with 4 – 5 long, thin capillaries and 3 – 4 short pointed spines ”; however, the paratypes of P. ecuadoriana (USNM 59920, 60549) were examined and notopodial slender capillaries were found gradually transforming into spine-like capillaries in posterior notopodia, the same as in the specimens from Brazil. Such gradually modified capillaries differ from true heavy spines which appear abruptly in posterior notopodia in addition to capillaries in some Polydora species (Blake 1979). Thus, the specimens from Brazil were identical to P. ecuadoriana and therefore referred to this species. The Brazilian worms exhibited significant variation in palp pigmentation; fine black lines and short prominent bars were present or pigmentation was totally absent. Besides specimens with the typical scoop-shaped pygidium, there were worms in Brazil which appeared surprisingly similar to P. ecuadoriana but had small disc-like pygidia. These worms are here referred to P. cf. haswelli. The two species occurred together in a shell in various proportions, thus one or another was usually more abundant in different areas. The differences between them are discussed below, in the Remarks section after the description of P. cf. haswelli. Some Polydora specimens boring into oyster shells from the Gulf coast of Mexico were also studied (ECOSUR SPIO- 75, 76). They were anterior fragments lacking a pygidium, thus a definitive identification could not be made. However, other features, such as black bars on the palps, caruncle extending to the end of chaetiger 3, and major falcate spines of chaetiger 5 with lateral tooth and subdistal longitudinal flange, appeared similar to those of P. ecuadoriana. These specimens were tentatively referred to P. ecuadoriana. With its boring mode of life and an unusual scoop-shaped pygidium, P. ecuadoriana is similar to Polydora pygidialis Blake & Woodwick, 1972 from California. The two differ, however, in that P. ecuadoriana has black bars on palps, an incised prostomium, caruncle extending posteriorly to the end of chaetiger 3, and major falcate spines with lateral tooth and subdistal flange, whereas P. pygidialis has plain palps, entire prostomium, caruncle to the end of chaetiger 2, and major spines with only a large lateral tooth.	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF88FFA2057EFE998EED3635.taxon	distribution	Distribution Ecuador; Gulf coast of Mexico; Brazil: Espírito Santo south to Santa Catarina.	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF9EFFA7057EFBDC8E9337A2.taxon	description	Polydora haswelli Blake & Kudenov (1978, pp. 259 – 260, Fig. 44).	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF9EFFA7057EFBDC8E9337A2.taxon	materials_examined	Material BRAZIL, Espírito Santo: Espírito Santo Bay: Ilha do Frade, 20 ° 17´S, 40 ° 17´W, intertidal, from shells of the oyster C. rhizophorae, coll. R. C. Nalesso and V. I. Radashevsky, 29 May 2003, MZSP 175 (12); 25 Feb 2004, MZSP 310 (5). São Paulo: Alcatrazes Is., 24 ° 06´S, 45 ° 42´W, 5 – 6 m, from live stony coral Mussismilia hispida (Verrill, 1808), coll. J. M. M. Nogueira, 4 Dec 1996, MZSP 173 (2), USNM 1022188 (2). Praia Fazenda, 23 ° 22´S, 44 ° 50.3´W, intertidal, sandy beach, from shell of a gastropod inhabited by hermit crab Dardanus insignis (de Saussure, 1858), 0 9 May 2001, coll. V. I. Radashevsky, MZSP (1 +). Praia São Francisco, 23 ° 45´S, 45 ° 25´W, intertidal, from shells of living oysters and gastropods, and from gastropods inhabited by hermit crabs, 16 Apr 2003, coll. V. I. Radashevsky, MZSP (1 +). Praia Araçá, 23 ° 48.8´S, 45 ° 24.2´W, sandy intertidal, coll. V. I. Radashevsky, from shell of oyster C. rhizophorae, 17 Apr 2003, MZSP (1 +). São Sebastião Island, Praia do Curral, 23 ° 51.5´S, 45 ° 25.9´W, 3 – 5 m, from empty shells of the gastropod Stramonita haemastoma inhabited by hermit crabs, 13 May 2004, coll. V. I. Radashevsky, (10 +, notes on live material, not fixed). São Sebastião, Praia do Saco Grande, 23 ° 49.7´S, 45 ° 25.5´W, 1 – 5 m, from shells of living saddle oyster Anomia ephippium Linnaeus, 1758 and gastropods Astraea olfersii (Philippi, 1846), Morula nodulosa (Adams, 1845), Pisania auritula (Link, 1807), Pisania pusio (Linnaeus, 1758), Siratus senegalensis (Gmelin, 1791), Stramonita haemastoma, Strombus pugilis, and Tegula viridula inhabited by hermit crabs Paguristes tortugae and Pagurus brevidactylus, 30 Jun 2004, coll. V. I. Radashevsky, MZSP 179 (50). Paraná: Paranaguá Bay: Ilha das Cobras, 25 ° 28.9´S, 48 ° 26´W, intertidal, from oyster shell, 23 Jul 1985, coll. P. C. Lana, CEM / UFPR 38 (2). Brasilia village of Ilha do Mel, 25 ° 33´S, 48 ° 19´W, 0.5 m, 25 Aug 2001, coll. V. I. Radashevsky, from shell of the gastropod Crepidula plana Say, 1822, MZSP 178 (1); from shells of gastropods Pugilina morio and Stramonita haemastoma inhabited by hermit crab Clibanarius vittatus, MZSP 176 (80). 25 ° 28´S, 48 ° 27´W, 2 m, from shells of the cultured oyster C. rhizophorae, 13 Sep 2001, coll. V. I. Radashevsky, MZSP 174 (10). Santa Catarina, Florianópolis: Praia da Ponta de Sambaqui, 27 ° 28.5´S, 48 ° 33.7´W, 1 m, from shells of the cultured oyster C. gigas, coll. Y. M. B. Neptune, 15 Aug 2003, MZSP 177 (18).	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF9EFFA7057EFBDC8E9337A2.taxon	description	Adult morphology Up to 30 mm long and 1 mm wide at chaetiger 7 for 170 chaetigers. Body pale or light tan in life. Pigmentation on palps greatly variable: fine black lines or separate black bars present along longitudinal ciliated groove; occasionally pigmentation lacking. Pigment pattern and numbers of bars on palps not correlated with total number of chaetigers in an individual. Light yellow pigment often present along ciliated groove between black bars. Black pigment absent on body or black narrow strips present on either side of anterior part of prostomium, and indistinct paired patches present on dorsal side of peristomium and 1 – 3 anterior chaetigers. Small distinct yellow chromatophores present on dorso-lateral sides of posterior chaetigers in large individuals (Fig. 8 E). Yellow pigment on palps and posterior chaetigers absent in formaldehyde-fixed specimens. Prostomium incised anteriorly. Caruncle extending to end of chaetiger 3. Occipital antenna absent. Two pairs of black eyes usually present; occasionally one to three eyes present or eyes lacking. Palps as long as 10 – 20 chaetigers, with longitudinal groove lined with fine frontal cilia, latero-frontal motile compound cilia bordering groove, short lateral papillae with non-motile cirri arranged along either side of groove, and short compound non-motile cilia arising directly from palp surface and scattered on lateral and abfrontal palp surfaces. Chaetiger 1 with short capillaries in neuropodia and postchaetal lamellae in both rami; notochaetae absent. Posterior notopodia with only capillary chaetae. Chaetiger 5 greatly modified, almost twice as large as chaetigers 4 or 6, with 3 – 5 dorsal superior winged capillaries (Fig. 8 C), 5 – 8 major modified spines alternating with bilimbate-tipped companion chaetae and arranged in a slightly curved, diagonal row, and 5 – 8 winged ventral capillaries (Fig. 8 B); postchaetal lamellae absent. Dorsal superior and ventral capillaries shorter and fewer than those on chaetigers 4 and 6. Major spines falcate, with lateral accessory flange of variable size (Fig. 8 D). Hooded hooks in neuropodia from chaetiger 7, bidentate, with constriction on shaft (Fig. 8 A), up to 8 in a series. Branchiae from chaetiger 7, full-sized from chaetigers 9 – 10, distributed to middle or usually almost end of body, becoming much smaller on posterior chaetigers. Pygidium cup-shaped to disc-like, with dorsal gap to narrow incision, white due to numerous glandular cells; small notch often present on ventral side of pygidium, usually positioned asymmetrically (Fig. 8 E). Small knobs with short non-motile, probably sensory, cilia arranged along edge of pygidium, mainly along its dorso-lateral edge. Lateral ciliated organs as small pits between noto- and neuropodia on all chaetigers except 4 and 5. Glandular pouches from chaetiger 7, large in chaetigers 7 to 9, then greatly diminishing in size. Digestive tract without gizzard-like structure. Metanephridial segmental organs from chaetiger 7, opening to exterior laterally on anterior, sterile chaetigers and dorsally on gametogenic chaetigers; paired nephridia opening separately on all chaetigers. Habitat In Brazil, P. cf. haswelli was found intertidally and in shallow water in shells of live oysters Crassostrea brasiliana, C. gigas, C. rhizophorae and Anomia ephippium, the gastropod Crepidula plana, and empty shells of the gastropods Astraea olfersii, Pisania auritula, Pisania pusio, Pugilina morio, Siratus senegalensis, Stramonita haemastoma, Strombus pugilis, and Tegula viridula inhabited by hermit crabs Clibanarius vittatus, Paguristes tortugae and Pagurus brevidactylus, and also in live scleractinian corals. Up to 10 worms occurred per cm 2 of shell surface. Worm burrows were U-shaped with walls lined with fine silt. The burrows of P. cf. haswelli resemble those of P. ecuadoriana. The two species occurred together in shells in various proportions. Reproduction Polydora cf. haswelli is gonochoristic. Spermatids were interconnected in tetrads. Spermatozoa were introsperm with an elongated straight head about 1 µm in diameter, head + middlepiece 12 µm long, acrosome 1.5 µm, nucleus 6 µm, middlepiece 4.5 µm, and flagellum 45 µm long. Females deposited eggs about 100 µm in diameter into capsules which were joined to each other in a string and attached to the inner wall of the burrow by two stalks. Larvae developed inside the capsules until the 3 - chaetiger stage, when they hatched and continued development in the water column until the 16 – 18 - chaetiger stage, feeding on the plankton. Morphology of the capsules and early, encapsulated larvae resembled those of P. ecuadoriana. Larval development of the species will be described elsewhere.	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF9EFFA7057EFBDC8E9337A2.taxon	discussion	Remarks Worms identified herein as P. cf. haswelli resemble P. ecuadoriana in many characteristic features. The two species occur together in shells, have similarly variable pigmentation on palps and anterior chaetigers, an incised prostomium, caruncle extending posteriorly to the end of chaetiger 3, spermatids joined in tetrads, similar dimensions of oocytes and spermatozoa, and morphology of early larvae. The extreme forms of each species can easily be distinguished by the shape of pygidium: those referred to P. ecuadoriana have a large, scoop-shaped pygidium, while P. cf. haswelli individuals have a small, disc-like to cup-shaped pygidium. However, the morphology of the pygidium is variable in both species, including form intermediate between the cup- and scoop-shaped pygidia (Fig. 9 B). Other characteristics that might be diagnostic, such as the dentition of chaetiger 5 falcate spines (single lateral flange versus lateral tooth and small subterminal flange), and the presence of small yellow chromatophores on the posterior segments, are also variable, making delineation of the species ambiguous. Lateral additional structures on old falcate spines (situated in the anterior part of the row) were worn and indistinct, while on new spines (situated in the posterior part of the row) they appeared as one complete large flange or lateral tooth and small subterminal flange, depending on the view of observation and quality of the preparation. Yellow chromatophores were absent in some live worms with all the other characteristics corresponding to P. cf. haswelli. This variability raised a problem of conspecificity of worms with large, scoop-shaped and small, disc-like pygidia (see below doubts about identification as P. haswelli). Pelagic Polydora larvae caught in a creek entering Espírito Santo Bay in the state of Espírito Santo provided support for the idea of two sympatric species. Two kinds of larvae were found in the plankton and only P. ecuadoriana and P. cf. haswelli were present on the bottom, boring in oyster shells. The larvae appeared very similar in most diagnostic characteristics but differed unambiguously in the shape and position of melanophores on the lateral peristomial lips. It is plausible that two Polydora species co-occur in the area and they are able to interbreed on occasion, resulting in individuals with intermediate shape of the pygidium. Further molecular investigations are certainly needed to clarify this issue. Meanwhile worms with scoop-shaped and disc-like pygidia are distinguished taxonomically. Another problem to solve was the identification of the two species under discussion. The diagnostic features and taxonomic status of P. ecuadoriana are discussed above in this paper, while reasoning for P. cf. haswelli is provided here. Although both species demonstrate variable pigmentation (even absent in some individuals), they can be referred to a group of Polydora species having characteristic black bars on palps. Those species were reviewed by Williams & Radashevsky (1999: Table 1) and Radashevsky & Hsieh (2000: Table 1) but the Brazilian worms fit none of them. Both reviews, however, overlooked P. ecuadoriana and P. haswelli. The original description of the latter species referred ambiguously to “ additional pigment on palps ” (Blake & Kudenov 1978: p. 259) but James A. Blake clarified that " pigment spots occurred along the palps " (e-mail of 3 November 2003 to VIR). This clarification made P. neocaeca Williams & Radashevsky, 1999 (originally described from Rhode Island, U. S. A.) and P. haswelli Blake & Kudenov, 1978 (originally described from New South Wales, Australia) similar to each other and raised a problem of their identity. Both species are shell-borers with almost identical main characteristic features, including body and palp pigmentation and the shape of the pygidium. They differ in the shape of additional structures on major falcate spines of chaetiger 5, with large accessory tooth and smaller vertical flange above the tooth present in P. haswelli (Blake & Kudenov, 1978: fig. 44 C – E), and an obliquely curved flange present in P. neocaeca (Williams & Radashevsky 1999: fig. 1 D). These differences are not, however, unambiguous and should be studied more carefully since both kinds of spine lateral structures may vary. Sperm morphology might be informative for the species delineation and is used herein for a tentative identification of the Brazilian worms. Polydora neocaeca has spermatids joined in octads (Williams 2000) and spermatozoa with acrosome 0.9 ± 0.1 µm, nucleus 4.8 ± 0.4 µm, middlepiece 4.2 ± 0.4 µm (Williams & Radashevsky 1999; Williams 2000). Polydora cf. haswelli collected in Brazil have spermatids joined in tetrads, and the spermatozoa differ in measurements from those of P. neocaeca. The number of spermatids joined together by cytoplasmic bridges is species specific, 4 or 8 in examined Polydora species, and has been used for distinguishing between sibling species (Radashevsky & Pankova 2006). Information about spermatid aggregation and sperm morphology of P. haswelli from Australia is lacking. Pending more details on morphology of the Australian material, we refer to the Brazilian specimens as P. cf. haswelli. Remarkably, P. h a s w e l l i has not been reported outside of Australia, and P. ecuadoriana was recorded only from Ecuador. Further molecular investigations would clarify the identification of the Brazilian worms referred to these species.	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF9EFFA7057EFBDC8E9337A2.taxon	distribution	Distribution Australia: New South Wales; Brazil: Espírito Santo south to Santa Catarina.	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF9BFFA5057EFA7189083015.taxon	materials_examined	Adult morphology (based on material from Brazil) Specimens in poor condition. Up to 15 mm long and 0.6 mm wide at chaetiger 7 for 130 chaetigers. Black pigment diffused on palps along longitudinal ciliated grove, on sides of prostomium and on 1 – 4 anterior chaetigers; present only on prostomium or absent in some specimens. Largest individual with 130 chaetigers and with black bands on palps. Prostomium blunt to rounded anteriorly. Caruncle extending to middle of chaetiger 3. Occipital antenna absent. One to four eyes present or eyes absent. Chaetiger 1 with short capillaries in neuropodia and postchaetal lamellae in both rami; notochaetae absent. Posterior notopodia with only capillary chaetae. Chaetiger 5 greatly modified, with 2 – 3 dorsal superior capillaries, 4 – 6 major spines alternating with bilimbate-tipped companion chaetae, and ventral tuft of 4 – 6 capillaries; postchaetal lamellae absent. Dorsal superior and ventral capillaries shorter and fewer than those on chaetigers 4 and 6. Major spines falcate, with lateral tooth and narrow longitudinal flange located laterally on main fang above accessory tooth. Branchiae from chaetiger 7 to almost end of body, full-sized from chaetiger 10 – 11, greatly diminishing on posterior chaetigers. Hooks in neuropodia from chaetiger 7, bidentate, with constriction on shaft, up to 9 in a series. Pygidium small, cup-shaped to disc-like with dorsal incision. Gizzard-like structure in digestive tract not distinguished.	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF9BFFA5057EFA7189083015.taxon	discussion	Remarks	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF9BFFA5057EFA7189083015.taxon	distribution	Distribution Pacific Mexico; Chile;? Brazil: São Paulo.	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF96FFAF057EFE9988DC3447.taxon	materials_examined	Material Paraná, Paranaguá Bay, 25 ° 28´S, 48 ° 27´W, 2 m, from shells of the oyster C. rhizophorae, 13 Sep 2001, coll. V. I. Radashevsky, MZSP 182 (holotype). Santa Catarina, Florianópolis, Praia da Ponta de Sambaqui, 27 ° 28.5´S, 48 ° 33.7´W, 1 m, from shells of the cultured oyster C. gigas, coll. Y. M. B. Neptune, 25 Apr 2003, MZSP 181 (1 paratype).	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF96FFAF057EFE9988DC3447.taxon	description	Adult morphology Holotype complete male, with 103 chaetigers; paratype anterior fragment of female. Prostomium rounded anteriorly. One pair of black eyes present in holotype; eyes absent in paratype. Caruncle extending to middle of chaetiger 3. Occipital antenna absent. Black pigment diffused on dorso-lateral sides of peristomium, in front of palp bases, and on dorsal side of four anterior chaetigers; segmental patches of pigment larger on anterior chaetigers (Fig. 10 A). Narrow black line present along longitudinal ciliated groove on palps. Chaetiger 1 with short capillaries in neuropodia and small postchaetal lamellae in both rami, notochaetae absent. Posterior notopodia with packets of needle-like spines besides capillaries. Needles not protruding beyond body surface, beginning from chaetiger 27 in holotype. Chaetiger 5 greatly modified, with 4 dorsal superior winged capillaries (Fig. 10 C), 6 major modified spines alternating with bilimbate-tipped companion chaetae and arranged in a slightly curved, diagonal row (Fig. 10 D), and 6 winged ventral capillaries (Fig. 10 B); postchaetal lamellae absent. Dorsal superior and ventral capillaries shorter and fewer than those on chaetigers 4 and 6. Major spines falcate, with small lateral accessory tooth. Hooded hooks in neuropodia from chaetiger 7, bidentate, with constriction on shaft. Branchiae on chaetigers 7 – 90, full-sized from chaetiger 11. Nototrochs from chaetiger 7 onwards (Fig. 10 A). Pygidium small, cup-shaped. Holotype male, with sperm first present in chaetiger 27. Habitat Polydora carinhosa bores in shells of the oysters C. rhizophorae and C. gigas. Reproduction Polydora carinhosa is gonochoristic. Females deposit eggs into capsules which are joined to each other in a string. Each egg capsule is attached by two thin stalks to the inner wall of the burrow. Larvae develop inside the capsules until about the 14 - chaetiger stage. The kind of lecithotrophy (endo- or exolecithotrophy) in P. carinhosa is unknown since only two broods with 10 - and 13 – 14 - chaetiger larvae were found. Larval development Ten-chaetiger larvae (Fig. 11 A) about 650 µm long, with three pairs of black eyes including two pairs of lateral eyes and one pair of median eyes; lateral eyes positioned close to each other and obscured by ramified melanophores positioned above them. Prostomium and peristomium weakly demarcated. A group of short non-motile apical cilia present on frontal edge of prostomium. Lateral lips of peristomium well developed, forming a voluminous vestibulum lined with short fine cilia. Vestibular ciliation running posteriorly over low ventral peristomial lip as a triangular neurotroch. One pair of small ciliated cells positioned on either side of neurotroch in the middle of chaetiger 1. Prototroch formed by two bands of long cilia running along lateral sides of peristomium, and two shorter bands of shorter cilia running across lateral lips of peristomium. Short compound cilia positioned along outer edges of vestibulum; these cirri probably sensory, moving slower than simple cilia. Nototrochs from chaetiger 3 onwards. Grasping cilia on either side of each nototroch, beating perpendicular to body axis and holding long bristles along the dorsum when larva is swimming in the capsule. Gastrotrochs on chaetigers 3, 5, 7, and 10. Telotroch interrupted middorsally, forming a gap where long bristles are held when larva is swimming in the capsule. Fine grains of yellow pigment dispersed on anterior part of prostomium, on ventral side of posterior chaetigers and on pygidium. A pair of ramified melanophores present on prostomium between median and lateral eyes. A pair of small melanophores on ventral side of lateral peristomial lips, just on prototroch level. Black pigment present laterally between chaetigers 1 and 2. Distinct transverse paired melanophores on dorsal side of chaetigers 3 – 6, in front of nototrochs; from chaetiger 7 those melanophores becoming stellar, ramified. Paired black pigmentation present on dorsal sides of pygidium. Larval serrated bristles in all notopodia, those on chaetiger 1 longest. Voluminous vestibulum opening posteriorly into short esophagus which extends to end of chaetiger 2. Buccal bulb absent. A muscular sphincter present between esophagus and stomach. The stomach narrowing posteriorly and weakly separated from the hindgut. Wall of the stomach containing numerous oil drops; its inner surface lined with fine cilia. Two pairs of protonephridia in chaetigers 1 and 2. Fully developed, ready to hatch larvae 800 – 850 µm long for 13 – 14 chaetigers. Fine granules of yellow pigment dispersed on anterior part of prostomium, on pygidium, and on ventral side from chaetiger 6 (Fig. 12 C). Two large black patches on dorsal side of pygidium, along edge of gap; no middorsal melanophore. Short single motile cilia on tips of postchaetal lamellae. Numerous non-motile sensory cilia on frontal edge of prostomium, on palps and on posterior edge of pygidium. One pair of banana-shaped cells with striated contents positioned posterior to median eyes; large fusiform cells with striated contents positioned inside anterior end of prostomium and in palps; elongated cells of irregular shape present on dorsal and ventral sides of chaetigers, and inside pygidium. Gastrotrochs on chaetigers 3, 5, 7, 10, and 13. Those on chaetigers 3 and 5 composed of two lateral ciliated cells, those on other chaetigers with five ciliated cells. Chaetiger 5 with 2 dorsal superior capillaries (Fig. 11 E), 4 dorsal modified chaetae (Fig. 11 F – H) and 3 ventral capillaries (Fig. 11 D). Modified chaetae including first two provisional spines and posterior two chaetae of quasi-adult type. Provisional spines including one heavy falcate spine with longitudinal groove (Fig. 11 G), and one thinner, awl-like spine with sigmoid distal end (Fig. 11 F); chaetae of quasi-adult type including heavy falcate spines with two lateral teeth, and bilimbate-tipped companion chaetae (Fig. 11 H). Hooded hooks in neuropodia from chaetiger 7, 2 – 3 in a series, accompanied by 2 – 3 winged capillaries (Fig. 11 B, C). Lateral organs as small pits 3 – 5 µm in diameter with stiff, non-motile cilia 10 – 15 µm long between noto- and neuropodia on all chaetigers. Glandular pouches in chaetigers 6 – 11, large in anterior chaetigers and gradually diminishing in size posteriorly, each composed of 1 – 2 large secretory cells enveloped by thin common membrane but opening to the exterior separately. Circulatory system developed and functional. Protonephridia in chaetigers 1 and 2. Metanephridia from chaetiger 7 onwards. Settlement and metamorphosis Larvae of P. carinhosa underwent gradual metamorphosis and loss of provisional larval features inside the egg capsules when they grown to 900 µm long for 14 chaetigers. The largest larva without bristles in notopodia was 1035 µm long for 14 chaetigers. The 14 - chaetiger larvae hatched and settled after a short planktonic stage (Fig. 13). Adult mode of feeding after settlement was enabled by rapid elongation of the palps, modification of the prostomium, enlargement of the ventral peristomial lip and transformation of the lateral peristomial lips into dorso-lateral ciliary folds. In further development, the prostomium became separated from the peristomium; the caruncle and nuchal ciliated bands elongated posteriorly over chaetiger 3, and nototrochs were lost on anterior chaetigers.	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF96FFAF057EFE9988DC3447.taxon	discussion	Remarks	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
F72C87EBFF96FFAF057EFE9988DC3447.taxon	distribution	Distribution Brazil: Paraná south to Santa Catarina.	en	Radashevsky, Vasily I., Lana, Paulo C., Nalesso, Rosebel C. (2006): Morphology and biology of Polydora species (Polychaeta: Spionidae) boring into oyster shells in South America, with the description of a new species. Zootaxa 1353: 1-37, DOI: 10.5281/zenodo.174538
