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Magalhães, Wagner F., Weidhase, Michael, Schulze, Anja, Bailey-Brock, Julie H. (2016): Taxonomic remarks on the genus Ctenodrilus (Annelida: Cirratulidae) including description of a new species from the Pacific Ocean. Zootaxa 4103 (4): 325-343, DOI: http://doi.org/10.11646/zootaxa.4103.4.2

Genus Ctenodrilus Clapar��de, 1863 Syn. Parthenope Schmidt, 1857 Type species: Parthenope serrata Schmidt, 1857. Diagnosis. Prostomium rounded, eyespots absent, nuchal organs shallow ciliary pits, postero-lateral, rounded; peristomium complete ring surrounding mouth; first segment complete chaetiger. Palps and branchiae absent. Body with few segments, up to 15. Chaetae thick and short, marginally smooth or serrated with variable number and size of teeth. Body with sparse cilia, concentrated on ventral region of anterior end. Digestive tract divided into three parts, anterior slender esophagus, inflated and pigmented stomach and intestine. A pair of nephridia present on peristomial region. Heart body enlarged dorsal vessel present through few anterior segments. Pygidium simple lobe, anal aperture usually dorsal. Asexual reproduction as paratomy, number of stolons variable; sexual reproduction as viviparity, protandrous or simultaneous hermaphrodites. ������continued on the next page ) Remarks. Ctenodrilus serratus (Schmidt, 1857) is currently considered as a valid species but type material has either been lost or never deposited. The identity of C. parvulus Scharff, 1887 and of the subspecies C. serratus limulicolus Sudzuki & Sekiguchi, 1972 is questionable (Wilfert 1973; Westheide et al. 2003; Dean & Blake 2014) and both species lack type material. Extensive comparisons have been made between C. serratus and C. parvulus. In addition to the anatomical and histological differences, the hooks in C. parvulus were illustrated as lacking distinct teeth (see Scharff 1887; Fig. 2). The subspecies C. serratus limulicolus has been considered as a synonym of C. serratus but it also differs in the alleged shape of nuchal organs and very distinct characteristics of the posterior end and pygidium (as seen in Fig. 14 d of Sudzuki & Sekiguchi 1972) having a ventral anus, cilia and distinct anal glands. C. paucidentatus was described as being distinct from C. serratus by the fewer number and shape of teeth on the multidentate hooks. The dentition and shape of hooks have been proved to be very variable among populations and even within an individual so the identity of C. paucidentatus is questionable based on the examination of type material. Morphology of C. serratus from the east coast of the United States and Caribbean is similar to that provided in Wilfert (1973) and Weidhase et al. (2016) (Fig. 1); further details in Remarks section below., Published as part of Magalh��es, Wagner F., Weidhase, Michael, Schulze, Anja & Bailey-Brock, Julie H., 2016, Taxonomic remarks on the genus Ctenodrilus (Annelida: Cirratulidae) including description of a new species from the Pacific Ocean, pp. 325-343 in Zootaxa 4103 (4) on pages 327-331, DOI: 10.11646/zootaxa.4103.4.2, http://zenodo.org/record/271202, {"references":["Claparede, E. (1863) Beobachtungen uber Anatomie und Entwicklungsgeschichte der wirbellosen Thiere an der Kuste von Normandie angestellt. Leipzig, W. Engelmann, 120 pp., 18 pls.","Schmidt, O. (1857) Zur Kenntnis der Turbellaria, Rhabdocoela und einiger anderer Wuermer des Mittelmeeres. Sitzungsberichte der Kaiserliche Akademie der Wissenschaften, Wien, Mathematisch-Naturwissenschaftliche Klasse, 23 (2), 347 - 366, pls. 1 - 5.","Scharff, R. (1887) On Ctenodrilus parvulus n. sp. Quarterly Journal of Microscopical Science, 27, 591 - 604.","Sudzuki, M. & Sekiguchi, K. (1972) Some remarks on five aberrant annelids from the culture water of Japanese horse-shoe crabs. The Science Reports of the Tokvo Kvoiku Daigaku Section B, 15, 39 - 56.","Wilfert, M. (1973) Ein Beitrag zur Morphologie, Biologie und systematischen Stellung des Polychaeten Ctenodrilus serratus. Helgolander wiss Meeresunters, 25, 332 - 346. http: // dx. doi. org / 10.1007 / BF 01611202","Westheide, W., Hass-Cordes, E., Krabusch, M. & Muller, M. C. M. (2003) Ctenodrilus serratus (Polychaeta: Ctenodrilidae) is a truly amphi-Atlantic meiofauna species - evidence from molecular data. Marine Biology, 142, 637 - 642.","Dean, H. K. & Blake, J. A. (2014) 7.15. 5. Ctenodrilidae Kennel, 1882. In: Schmidt-Rhaesa, A. & Walter de Gruyter, A. (Eds.), Handbook of Zoology, A Natural History of the Phyla of the Animal Kingdom. Berlin. [Database]","Weidhase, M., Bleidorn, C. & Simon, C. A. (2016) On the taxonomy and phylogeny of Ctenodrilus (Annelida: Cirratulidae) with a first report from South Africa. Marine Biodiversity, 1 - 10. [published online] http: // dx. doi. org / 10.1007 / s 12526 - 015 - 0355 - 3"]}

Palola Lizard Island Clade 1 (Fig. 2 A���C) Material examined. AM W. 43974, MI QLD 2344, sequenced; AM W. 44034, MI QLD 2352 (3, 1 sequenced and photographed); AM W. 44146, MI QLD 2356 (3, 1 sequenced and photographed); AM W. 44128, MI QLD 2358, sequenced; AM W. 44136, MI QLD 2358, sequenced; AM W. 44894, MI QLD 2401, sequenced; AM W. 44655, MI QLD 2398; AM W. 44656, MI QLD 2398; AM W. 44925, MI QLD 2390; AM W. 44202, MI QLD 2359; AM W. 43912, MI QLD 2335. Description. Specimens small and threadlike. Anterior fragments examined for all sequenced specimens, ranging in length from 7 to 18 mm and from 0.6 to 1.7 mm in width; with 38���78 chaetigers. Branchiae only observed in one specimen (AM W. 44034) from chaetiger 62 to the end of the fragment. No ventral eyespots. Head and body generally without pigmentation but with iridescent shine, or, if pigmentation is present light brown and restricted to prostomium and peristomium (Fig. 2 A). Mandibles usually protruding from mouth, thin and nearly transparent with serrated anterior margin (Fig. 2 B). Maxillae not examined. Antennae, palps and peristomial and parapodial cirri without pigment. Antennae and palps wrinkled, in preserved material, tapering and pointy tip. Median antenna reaches to chaetiger 2 or 3, lateral antennae reach to chaetiger 1 or 2 and palps reach to first or second peristomial ring. Tapering peristomial cirri reach forward to about 3 / 4 of the length of first peristomial ring. Eyes dark, oval or with ventral notch and nestled between lateral antennae and palps. Acicula brown. Other chaetae not examined. Dark pigment spots on dorsal side of posterior parapodia observed in two specimens (AM W. 44034; AM W. 44.136) (Fig. 2 C). Remarks. Although only distantly related in the phylogenetic tree and representing different species groups, Palola Lizard Island clade 1 is morphologically most similar to clade 5. Members of both clades are thin and threadlike, almost entirely lack pigmentation and have relatively thin and transparent mandibles. They can be distinguished by the relative length of the first and second peristomial rings: in clade 1, the first peristomial ring is 1.5 to 2 times as long as the second when viewed in the midlateral line where as in clade 5 they are more similar in length. Despite the morphological similarities, COI sequences are 26.9 % divergent (Kimura- 2 -Parameter model) between clades 1 and 5. Palola Lizard Island Clade 1 falls into species group A and is the sister group to a clade with an extremely wide geographic distribution throughout the tropical Eastern and Western Pacific, referred to as clade A 1 in Schulze (2006) and Schulze & Timm (2012). The average genetic distance to this clade is only 1.9 % and the two might represent the same species, although they are reciprocally monophyletic., Published as part of Schulze, Anja, 2015, Six genetically distinct clades of Palola (Eunicidae, Annelida) from Lizard Island, Great Barrier Reef, Australia, pp. 695-706 in Zootaxa 4019 (1) on pages 699-700, DOI: 10.11646/zootaxa.4019.1.23, http://zenodo.org/record/237471, {"references":["Schulze, A. (2006) Phylogeny and genetic diversity of palolo worms (Palola, Eunicidae, Polychaeta) from the tropical North Pacific and the Caribbean. Biological Bulletin, 210, 25 - 37. http: // dx. doi. org / 10.2307 / 4134534","Schulze, A. & Timm, L. E. (2012) Palolo and un: Distinct clades in the genus Palola (Eunicidae, Polychaeta). Marine Biodiversity, 42, 161 - 171. http: // dx. doi. org / 10.1007 / s 12526 - 011 - 0100 - 5"]}

Key to Palola clades from Lizard Island 1 Peristomial cirri reach at least to or beyond anterior margin of peristomium; fairly uniform brown pigmentation dorsally.............................................................................................. Palola clade 6 ��� Peristomial cirri not reaching to anterior margin of anterior peristomial ring; pigmentation, if present, faint or limited to poste- rior margins of segments................................................................................ 2 2. (1) Antennae and palps smooth and blunt-tipped..................................................... Palola clade 2 ��� Antennae wrinkled and with a pointy tip................................................................... 3 3. (2) Anterior margins of mandibles serrated.................................................................... 4 ��� Anterior margins of mandibles with a smooth edge........................................................... 5 4. (3) First peristomial ring 1.5 to 2 times as long as second peristomial ring in lateral mid-line.................. Palola clade 1 ��� First and second peristomial rings nearly equal in length in lateral midline.............................. Palola clade 5 5. (3) Lateral antennae about 2 / 3 the length of the median antenna......................................... Palola clade 4 ��� Lateral and median antennae more similar in length to each other..................................... Palola clade 3, Published as part of Schulze, Anja, 2015, Six genetically distinct clades of Palola (Eunicidae, Annelida) from Lizard Island, Great Barrier Reef, Australia, pp. 695-706 in Zootaxa 4019 (1) on page 703, DOI: 10.11646/zootaxa.4019.1.23, http://zenodo.org/record/237471

Schulze, Anja (2015): Six genetically distinct clades of Palola (Eunicidae, Annelida) from Lizard Island, Great Barrier Reef, Australia. Zootaxa 4019 (1): 695-706, DOI: http://dx.doi.org/10.11646/zootaxa.4019.1.23

Genus Palola Gray in Stair, 1847 Palola Gray in Stair, 1847: 1718.��� Fauchald 1992: 1179 ���1181. Type-species. Palola viridis Gray in Stair, 1847, by monotypy. Diagnosis. Eunicids with calcified, scoop-shaped mandibles. Three antennae (1 medial, 2 lateral) and one pair of palps arranged in a horseshoe shape on prostomium. Peristomium consisting of two rings, with peristomial cirri on second peristomial ring. Chaetal arrangement lacking subacicular hooks and pectinate chaetae along the entire body; branchiae simple, if present, usually from the mid-body region., Published as part of Schulze, Anja, 2015, Six genetically distinct clades of Palola (Eunicidae, Annelida) from Lizard Island, Great Barrier Reef, Australia, pp. 695-706 in Zootaxa 4019 (1) on page 697, DOI: 10.11646/zootaxa.4019.1.23, http://zenodo.org/record/237471, {"references":["Fauchald, K. (1992) Review of the types of Palola (Eunicidae: Polychaeta) Journal of Natural History, 26, 1177 - 1225. http: // dx. doi. org / 10.1080 / 00222939200770681"]}

Key to Palola clades from Lizard Island 1 Peristomial cirri reach at least to or beyond anterior margin of peristomium; fairly uniform brown pigmentation dorsally.............................................................................................. Palola clade 6 – Peristomial cirri not reaching to anterior margin of anterior peristomial ring; pigmentation, if present, faint or limited to poste- rior margins of segments................................................................................ 2 2. (1) Antennae and palps smooth and blunt-tipped..................................................... Palola clade 2 – Antennae wrinkled and with a pointy tip................................................................... 3 3. (2) Anterior margins of mandibles serrated.................................................................... 4 – Anterior margins of mandibles with a smooth edge........................................................... 5 4. (3) First peristomial ring 1.5 to 2 times as long as second peristomial ring in lateral mid-line.................. Palola clade 1 – First and second peristomial rings nearly equal in length in lateral midline.............................. Palola clade 5 5. (3) Lateral antennae about 2 / 3 the length of the median antenna......................................... Palola clade 4 – Lateral and median antennae more similar in length to each other..................................... Palola clade 3

Phascolion (Isomya) tuberculosum Th��el Phascolion tuberculosum Th��el, 1875: 15 ���16. Type locality: Koster, western Sweden, 25��� 35 m. Remarks: The single 25 mm representative of this species had been living in a gastropod shell and has a cluster of epizoans around the base of the introvert. The holdfast papillae are flattened discs some with a circular margin of pale hardened protein. There are sparsely scattered dark, large hooks. The two retractor muscles are of almost equal size and both originate very close to the posterior end of the trunk. Distribution: Common in the northeastern Atlantic, including the Azores, Mid��Atlantic Ridge, Bay of Biscay, and Scandinavian waters at bathyal depths (25���2700 m). The few specimens from Japan and New Zealand (as P. temporariae, Edmonds, 1976) from 93���300 m point to a low��density population in the western Pacific Ocean. This sample reinforces that previous report from New Zealand of what appears to be a low��density southern hemisphere population., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 11, DOI: 10.5281/zenodo.158002, {"references":["Theel, H. (1875) Etudes sur les gephyriens inermes des mers de la Scandinavie, du Spitzberg et du Groenland. Bihang till Kungliga Svenska Vetenskaps-Akademiens Handlingar, 3, 1 - 30.","Edmonds, S. J. (1976) Three sipunculan species (two new) from New Zealand. New Zealand Journal of Marine and Freshwater Research, 10, 217 - 224."]}

Themiste (Lagenopsis) minor huttoni (Benham) Phascolosoma huttoni Benham, 1903: 177 ���184. Type locality: New Zealand. Remarks: These 11 worms with smooth dark sausage��shaped trunks measure 3���20 mm long. The short introverts bear scattered dark hooks and the collar region at the base of the branched tentacles is pigmented. The orientation of these hooks is generally towards the posterior end but some small deviation from this can be found (not to be confused with T. variospinosa from Australia where they point in all directions). Internally the single pair of strong retractor muscles originate near the middle of the trunk and digitiform contractile vessel villi are well developed in the larger worms. Distribution: Australia, New Zealand (including Chatham Islands), commonly in intertidal, hard substrata. This material is therefore not surprising except for the greater depths., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on pages 12-13, DOI: 10.5281/zenodo.158002, {"references":["Benham, W. B. (1903) The sipunculids of New Zealand. Transactions of the Royal Society of New Zealand, 36, 172 - 189."]}

Aspidosiphon Diesing Diagnosis: Introvert usually longer than trunk. Recurved hooks in numerous rings (absent in three species, scattered in two). Trunk with anal shield composed of hardened units that may be inconspicuous. Introvert protrudes from ventral margin of shield. Body wall either with continuous longitudinal muscle layer or with longitudinal muscle layer separated into anastomosing, sometimes ill��defined bundles. Small tentacles enclose dorsal nuchal organ but not mouth. Contractile vessel without villi. Two introvert retractor muscles may be almost completely fused. Spindle muscle attaches posteriorly. Two nephridia. One species may exceed 100 mm, but most are less than 40 mm long. Commonly inhabiting some sort of shelter, e.g. holes in coral or soft rock, mollusc shells, or arenaceous foraminiferan tests., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 14, DOI: 10.5281/zenodo.158002

Phascolosoma Leuckart Diagnosis: Introvert variable in length, often longer than or equal to trunk, with numerous rings of recurved hooks. Fewer than 30 small tentacles in crescent around nuchal organ (peripheral tentacles lacking). Contractile vessel without true villi. Four introvert retractor muscles, lateral pairs sometimes partially fused. Two unilobed nephridia. Body wall musculature in separate bands., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 13, DOI: 10.5281/zenodo.158002

Phascolion (Phascolion) strombus (Montagu) Sipunculus strombus Montagu, 1804: 74 ���76. Type locality: Atlantic coast of northern Europe. Remarks: The 10 specimens are 9���20 mm long and bear well��developed tentacles and hooks, and in most cases distinct U�� or V��shaped holdfast papillae towards the posterior end of the trunk. Edmonds (1980) erected a new species from Australia that he called Phascolion cronullae, but Cutler & Cutler (1985) demoted this to the status of a subspecies of P. strombus, and this was maintained in Cutler (1994). The distinction between these two geographically isolated populations was based on: (a) Relative size of the two retractor muscles and their point of origin. In the nominate form the ventral is much smaller, commonly 10���20 % the diameter of the dorsal, but may be up to 35 % of the size of the dorsal and have their origins at different anterior/posterior levels. In the Australian form they are more similar in size with the ventral 50���75 % the size of the dorsal, both originating at nearly the same level near the posterior end of the trunk. (b) Anterior papillae around the base of the introvert. In both populations there is a significant array of large dark mammilate or columnar papillae. The nominate form is alleged to have a single nipple��like tip, while the Australian form has 1���4 tips, around 25 % having more than one. We now conclude that these character states are not taxonomically useful or fixed and that there is more variation within this widespread and eurytopic species than previously acknowledged. In this sample, the ventral retractors ranged from 20��� 66 % of the size of the dorsal muscle and a few multi��tipped papillae could be found on most, but not all, of these. Since many had been removed from gastropod shells, the posterior end of the trunk was not always present, making precise observations about muscle origins impossible. Distribution: Very common and eurytopic in the North Atlantic and Arctic oceans, most numerous between 200 and 2000 m. It is also found in deep water in the Caribbean, plus scattered records from the Mediterranean, Red Sea, Gulf of Aden, Madagascar, and South Africa; two Antarctic records; and records from Argentina and Chile. Other Pacific Ocean records come from the South Pacific, New Zealand, and Japan. Remarkably, it is known from depths of 1���4030 m. The only previous record from New Zealand was that of Edmonds (1976) when he described P. t o r t u m, later synonomised with P. strombus. A few Antarctic reports have been called into question by Cutler et al., (2001). Thus, these records provide more evidence that this species, while morphologically quite plastic, is present near New Zealand., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 10, DOI: 10.5281/zenodo.158002, {"references":["Montagu, G. (1804) Description of several marine animals found on the south coast of Devonshire. Transactions of the Linnean Society of London, 7, 61 - 85.","Cutler, E. B. & Cutler, N. J. (1985) A revision of the genera Phascolion Theel and Onchnesoma Koren and Danielssen (Sipuncula). Proceedings of the Biological Society of Washington, 98, 809 - 850.","Cutler, E. B. (1994) The Sipuncula, their systematics, biology and evolution. Cornell University Press, Ithaca. 480 pp.","Edmonds, S. J. (1976) Three sipunculan species (two new) from New Zealand. New Zealand Journal of Marine and Freshwater Research, 10, 217 - 224.","Cutler, E. B., Dean, H. K. & Saiz-Salinas, J. I. (2001) Sipuncula from Antarctic Waters. Proceedings of the Biological Society of Washington, 114, 861 - 880."]}

Aspidosiphon (Akrikos) cf. zinni Cutler Aspidosiphon zinni Cutler, 1969: 209 ���211. Type locality: East Coast of the U.S., 39.43 �� N, 70.6 �� W, 2496 m. Remarks: The two 5 mm transparent thin worms have continuous longitudinal musculature and a pale, ungrooved anal shield made up of fine uniform granular units. The introvert is shorter than the trunk and has scattered, small, pale hooks and no observable tentacles. Distribution: Common in the northern Atlantic Ocean at depths of 1100���4400 m, plus two records from around 9 �� S off the Congo River. In the Indian Ocean there is one report from the Mozambique Channel and one from the Reunion Islands. The worms frequently live in arenaceous foraminiferan tests. The present specimens are thousands of kilometres from any previously known populations of A. zinni. Since they are very small deep��water animals it is conceivable that the species has been overlooked by previous collectors Before accepting the present material as unequivocally this species, we prefer to have additional specimens for confirmation., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 15, DOI: 10.5281/zenodo.158002, {"references":["Cutler, E. B. (1969) New species of Sipuncula from the western North Atlantic. Proceedings of the Biological Society of Washington, 82, 209 - 218."]}

Themiste (Lagenopsis) dehamata (Kesteven) Dendrostoma dehamatum Kesteven, 1903: 69 ���73. Type locality: Port Jackson, New South Wales, 1��� 2 m. Remarks: Seventeen worms are assigned to this species, having trunks 10���80 mm long and neither introvert hooks nor pigment around the collar region just behind the tentacles. Several have a reddish tint and some exhibit a reticulate or annulate pattern of fine wrinkles in the thick skin. The digitiform contractile vessel villi were not well preserved or underdeveloped in a few specimens. The strong pair of introvert retractors originate near the middle of the trunk. Worms collected in December (Stn T 524) had mature eggs in their body cavity. Distribution: New South Wales and southern Australia, in intertidal unconsolidated sediments. Unpublished records of 12 specimens from 10���40 m mud/shell/gravel habitats in Omaha Bay, New Zealand (R. Taylor, pers.comm.) supplement these from 16���110 m to confirm its presence around New Zealand., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 12, DOI: 10.5281/zenodo.158002, {"references":["Kesteven, H. L. (1903) A new species of Dendrostoma. Records of the Australian Museum, 5, 69 - 73."]}

Cutler, Edward B., Schulze, Anja, Dean, Harlan K. (2004): Zealand species. Zootaxa 525: 1-19, DOI: 10.5281/zenodo.158002

Phascolion (Isomya) cf. hedraeum Selenka & de Man Phascolion hedraeum Selenka & de Man in Selenka et al., 1883: 49 ���50. Type locality: Rio de Janeiro, 7 m. Remarks: The six 2���12 mm worms had been living in gastropod shells. The holdfast papillae are circular, with only their anterior margins showing pale hardened protein, which may be dark in a few cases. Scattered pale hooks can be seen. The two retractor muscles are of almost equal size, but the smaller one may be only 65���70 % the diameter of the larger, both having their origins at the same level very close to the posterior end of the trunk. The similarities to P. tuberculosum are significant and in some ways these small specimens did not fit perfectly into either taxon (thus the cf. designation), but most closely resemble P. hedraeum. Distribution: Southern Atlantic reports give latitudes down to 74 �� S. Some records are from the southern Pacific, near Australia, and also off Japan. They live at shelf and slope depths, rarely over 800 m, in gastropod or scaphopod shells. The present collection suggests the possible existence of a low��density bathyal population in this part of the southern hemisphere., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 11, DOI: 10.5281/zenodo.158002, {"references":["Selenka, E., de Man, J. G. & Bulow, C. (1883) Die Sipunculiden, eine systematische Monographie. Semper Reisen im Archipel der Philippinen 2, 4, 1 - 131."]}

Themiste Gray Diagnosis: Introvert shorter than trunk. Body wall with continuous muscle layers. Tentacles basically surrounding mouth but extending along branching stem like outgrowths as the animal grows. With or without hooks. Two large introvert retractor muscles; contractile vessel with distinct villi or tubules. Spindle muscle adheres closely to rectum and is not attached posteriorly. Two nephridia. Animals small to large (adults 4���400 mm long)., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 12, DOI: 10.5281/zenodo.158002

Nephasoma diaphanes (Gerould) Phascolosoma diaphanes Gerould, 1913: 395. Type locality: Off eastern seaboard of the U.S., Cape Cod to New Jersey, 90���2400 m. Remarks: These 99 worms are commonly 2���10 mm long with a few large ones in the 15���20 mm range, and one 25 mm giant. They are slender, transparent, have small pale scattered hooks, and reduced lobe��like tentacles. The papillae are usually inconspicuous, but in some individuals those towards the posterior end may be quite distinct and dark reddish��brown. It is not uncommon to find this species living inside arenaceous foraminiferan tests or vacated polychaete tubes. A parasitic nematode was living within the body cavity of a worm from Stn S 201. Distribution: Cosmopolitan from 82 �� N to 78 �� S in cold water, mostly from bathyal and abyssal depths, down to 5300 m. Edmonds (1976) reported the species from off New Zealand in 600���660 m at 46 �� S, 171 �� E. These records strengthen the assertion that this is a truly cosmopolitan cold��water species., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on pages 7-8, DOI: 10.5281/zenodo.158002, {"references":["Gerould, J. H. (1913) The sipunculids of the eastern coast of North America. Proceedings of the United States National Museum, 44, 373 - 437.","Edmonds, S. J. (1976) Three sipunculan species (two new) from New Zealand. New Zealand Journal of Marine and Freshwater Research, 10, 217 - 224."]}

Phascolion (Montuga) pacificum Murina Phascolion pacificum Murina, 1957: 1777 ���1781 Type locality: Northwest Pacific, 5100���6900 m. Remarks: Most of the 146 worms had been living in gastropod shells (some in scaphopod shells) and many had been damaged upon removal. Thus, body lengths are difficult to measure. They are all small, ranging from 2���10 mm, and have some limited array of V�� or U��shaped holdfast papillae on the posterior half of the trunk. These may be weakly developed, especially in the smaller worms. The introvert retractor muscles appear as a single column until very near the posterior end where they divide into a pair of distinct roots. Distribution. This bathyal and abyssal species (300���6900 m) is widespread at high latitudes in the northwestern and southwestern Pacific, the northeastern, southern and Antarctic Atlantic, and the subantarctic Indian Oceans, including northern Australia. While there are recorded populations at some distance to the west and southeast of New Zealand (Cutler, 1977 a; Cutler et al., 2001), this material does suggest a more continuous, albeit not dense distribution., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 9, DOI: 10.5281/zenodo.158002, {"references":["Murina, V. V. (1957) Sipunculids collected on the first trip of the complex Antarctic expedition on the Ob' in 1956. Zoologicheskii Zhurnal, 36, 992 - 998.","Cutler, E. B., Dean, H. K. & Saiz-Salinas, J. I. (2001) Sipuncula from Antarctic Waters. Proceedings of the Biological Society of Washington, 114, 861 - 880."]}

Phascolion Th��el Diagnosis: Introvert 0.5���4 times trunk length, with or without hooks. Trunk usually with modified holdfast papillae. Body wall with continuous muscle layers. Tentacles arranged around mouth. Introvert retractor muscle system modified by fusion of dorsal and ventral pairs; relative size and degree of fusion define the subgenera. Contractile vessel generally without villi. Gut coiling generally loose, without axial spindle muscle. One nephridium (usually right). Small to medium��sized worms (, Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 8, DOI: 10.5281/zenodo.158002

Golfingia margaritacea (Sars) Sipunculus margaritaceus Sars, 1851: 196 ���197. Type locality: Norway. Remarks: The eight worms assigned to this species are 2���187 mm long and some are damaged or partly decayed. The presence of well��developed tentacles and the absence of hooks, distinct papillae, or other external features characterise this species. Distribution: Widespread in the Atlantic, Arctic, and Antarctic oceans (80 �� N to 78 �� S). In the Pacific Ocean from higher latitudes (> 30 �� N and S) and reports from lower latitudes are from depths> 2000 m. Indian Ocean records are limited to subantarctic latitudes. Reported depth ranges are 1���5300 m, but most have been collected from depths less than 400 m. Thus these new records supplement previous reports from Australian (Edmonds, 1980 as G. m. adelaidensis) and New Zealand waters (Edmonds, 1960 as G. cantabriensis), and numerous records from the nearby Antarctic Ocean (Cutler et al., 2001)., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on pages 6-7, DOI: 10.5281/zenodo.158002, {"references":["Sars, M. (1851) Beretning om en i Sommeren 1849 foretagen zoologisk Reise i Lofoten og Finmarken. Nytt Magasin for Naturvidenskapene, 6, 121 - 125, 190 - 211.","Edmonds, S. J. (1960) Sipunculids from New Zealand and the Chatham Islands. New Zealand Department of Scientific and Industrial Research Bulletin 139, 159 - 167.","Cutler, E. B., Dean, H. K. & Saiz-Salinas, J. I. (2001) Sipuncula from Antarctic Waters. Proceedings of the Biological Society of Washington, 114, 861 - 880."]}

Aspidosiphon muelleri Diesing Aspidosiphon muelleri Diesing, 1851: 68. Type locality: Mediterranean Sea. Remarks: The single representative of this species was removed from a piece of calcareous polychaete tube and is only 5 mm long. The anal shield is not dark but shows the square and rectangular units divided by grooves typical for this species. The caudal shield with radiating grooves is clearly developed and many rings of hooks can be seen. Distribution: Common in the northeastern Atlantic from Norway to West Africa (48 ��� 10 �� N). It extends through the Mediterranean into the Red Sea and the Gulf of Aden along the coast of East Africa to Madagascar and South Africa. Also recorded from Sri Lanka, and there are sparse reports from central Japan through Thailand, Vietnam, Indonesia, and down to Australia, New Guinea, and the Kermadec Islands. It is not found in most of the Pacific Ocean; one record from Juan Fernandez Island off Chile and one from southern Brazil are the only reports from near the American continents. The species inhabits shelf depths (5���400 m) throughout most of its range, but there are a few as deep as 2900 m. It is most often found in discarded mollusc shells. Some from shallow, warm water live in the bases of solitary corals that have overgrown small gastropods, forming a commensal relationship. This is a new record for New Zealand, but only a modest extension of its known range., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 14, DOI: 10.5281/zenodo.158002, {"references":["Diesing, K. M. (1851) Systema Helminthum. Braumiller, Vindobonae."]}

Nephasoma Pergament Diagnosis: Introvert about equal to or shorter than trunk. Hooks, when present, usually scattered. Body wall with continuous muscle layers. Tentacles arranged around the mouth but may be reduced in both size and number. Two introvert retractor muscles, often partially fused. Contractile vessel without villi. Spindle muscle not attached posteriorly. Two nephridia. Species generally small to medium sized (trunk, Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 7, DOI: 10.5281/zenodo.158002

Phascolosoma annulatum Hutton Phascolosoma annulatum Hutton, 1879: 278. Type locality: Cape Campbell, New Zealand. Remarks: The 74 worms have trunk lengths of 5���45 mm, mostly in the 15���30 mm range. The trunk is light brown with many scattered dark papillae These papillae often appear to have polygonal (not circular) bases and each one is covered by small dark polygonal platelets that spread over the inter��papillary area in a distinctive manner. Less than 25 complete rings plus several incomplete patches of dark hooks circle the tip of the introvert that is longer than the trunk. Distinct short tentacles can be seen forming a crescent dorsal to the nuchal organ. The two pairs of introvert retractor muscles may appear to be fused into a single column for some distance when the introvert is withdrawn. The coelom of one worm collected in May (Stn B 230) was full of eggs. As most members of this genus live in some sort of protective hole or crevice it was interesting but not surprising to find one living in a serpulid polychaete tube. Distribution: Southern Australia, New Zealand, Campbell Island, in shallow cool water. This material reinforces previous records from New Zealand (Edmonds, 1960; Taylor, 1991)., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 13, DOI: 10.5281/zenodo.158002, {"references":["Hutton, W. K. (1879) Additions to the list of New Zealand worms. Transactions of the Royal Society of New Zealand, 12, 277 - 278.","Edmonds, S. J. (1960) Sipunculids from New Zealand and the Chatham Islands. New Zealand Department of Scientific and Industrial Research Bulletin 139, 159 - 167.","Taylor, P. D. (1991) Observations on symbiotic associations of bryozoans and hermit crabs from the Otago shelf of New Zealand. In: Bigey, F. P. (Ed.) Bryozoaires Actuels et Fossiles: Bryozoa Living and Fossil. Bulletin de la Societe des Sciences Naturelles de l'Ouest de la France, Memoire HS. 487 - 495."]}

Nephasoma cutleri (Murina) Golfingia cutleri Murina, 1975: 1087 ���1088. Type locality: Pacific Ocean, 12 �� S, 178 �� E, 2614 m. Remarks: The two very slender, thread��like, complete specimens, have trunks 11���12 mm by 0.5 mm with introverts 40���60 % of the trunk length. There is a third specimen but most of its trunk is missing. Small, pale, scattered hooks and a slight constriction at the base of the introvert can be seen, but no other external features. The retractor muscles originate about 30 % of the distance towards the posterior end of the trunk and the intestine appears as a loose but regular double helix. Distribution: Scattered reports from the Pacific Ocean at abyssal depths (2600���4600 m), and the Comoro Islands and Sri Lanka in the Indian Ocean from 100���3700 m. These three worms constitute a significant range extension, and suggest a more uniform but sparse distribution throughout the deep water of the Indo��Pacific region., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 7, DOI: 10.5281/zenodo.158002, {"references":["Murina, V. V. (1975). New taxa of the genus Golfingia. Zoologicheskii Zhurnal, 54, 1085 - 1089."]}

Nephasoma pellucidum (Keferstein) Phascolosoma pellucidum Keferstein, 1865: 433. Type locality: St Thomas, Virgin Islands, 4 m. Remarks: The eight representatives of this species in the collections have sausageshaped trunks 10���50 mm long that are uniformly covered with distinct papillae which were usually darker than the underlying light brown body. Scattered pale hooks can usually be seen and the single pair of retractor muscles originate in the middle third of the trunk. Some had ruptured body walls or the internal organs were poorly preserved. Two earlier reports from the Galathea expeditions (Cutler, 1977 a: 143; 1977 b: 152) included hesitant and tentative identifications of a few worms from New Zealand waters at 30 m and 660 m that were not well preserved and/or did not exhibit the normal array of hooks and papillae. Therefore, these were intentionally omitted from later works (Cutler in press; Cutler & Saiz, in press). Distribution: Generally a shallow��water species, with a few bathyal records, from the western Atlantic and Caribbean south to Brazil. In the South Pacific and Indian oceans from Indonesia and Australia, southern Japan, and one record each from Cape Province and India. These records confirm the presence of this species in the New Zealand region., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 8, DOI: 10.5281/zenodo.158002, {"references":["Keferstein, W. (1865) Beitrage zur anatomischen und systematischen Kenntniss der Sipunculiden. Zeitschrift fur wissenschaftliche Zoologie, 15, 404 - 445."]}

Golfingia Lankester Diagnosis: Introvert about equal to or shorter than trunk. Hooks, when present, usually scattered. Body wall with continuous muscle layers. Tentacles arranged around mouth. Four introvert retractor muscles. Contractile vessel without villi. Spindle muscle not attached posteriorly. Two nephridia. Worms small to medium sized., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 6, DOI: 10.5281/zenodo.158002

Phascolion (Montuga) lutense Selenka Phascolion lutense Selenka, 1885: 16 ���17. Type locality: South Pacific, 3500 m. Remarks: The 134 specimens in the collection were living within the unique but typical cylindrical gray clay/mucus tubes of their own construction, 8���26 mm long. Other than the dense array of small gray papillae surrounding the base of the introvert there are no other epidermal ornamentations on the light gray bodies. Internally the strong introvert retractor muscles are fused into a single column until very near the posterior end where they bifurcate for a short distance. Distribution: A cold��water species (1000���6860 m), unknown from lower latitudes, even in deep water. Widely collected in the Southern Hemisphere in all the worlds oceans, including the Antarctic. In northern waters it is recorded from the northwestern Pacific and the northeastern Atlantic. These are the first records from New Zealand waters, but are only a modest range extension of the large nearby Antarctic population (Cutler et al., 2001)., Published as part of Cutler, Edward B., Schulze, Anja & Dean, Harlan K., 2004, Zealand species, pp. 1-19 in Zootaxa 525 on page 9, DOI: 10.5281/zenodo.158002, {"references":["Selenka, E. (1885) Report on the Gephyrea collected by H. M. S. Challenger during 1873 - 76. Report on the Scientific Results of the Voyage of H. M. S. Challenger, Zoology, 13, 1 - 25.","Cutler, E. B., Dean, H. K. & Saiz-Salinas, J. I. (2001) Sipuncula from Antarctic Waters. Proceedings of the Biological Society of Washington, 114, 861 - 880."]}

Southward, Eve C., Schulze, Anja, Tunnicliffe, Verena (2002): Vestimentiferans (Pogonophora) in the PaciŽc and Indian Oceans: a new genus from Lihir Island (Papua New Guinea) and the Java Trench, with the Žrst report of Arcovestia ivanovi from the North Fiji Basin. Journal of Natural History 36 (10): 1179-1197, DOI: 10.1080/00222930110040402, URL: http://www.tandfonline.com/doi/abs/10.1080/00222930110040402