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The World of Protozoa, Rotifera, Nematoda and Oligochaeta

Tubificoides

Tubificoides Lastockin, 1937 (re-erected Holmquist, 1978) (ref. ID; 7240), Lastockin (Holmquist, 1978 partim) (ref. ID; 5945)

Family Tubificidae: Subfamily Tubificinae (ref. ID; 5857, 5945)

ref. ID; 5945

Vas deferentia enter atria opposite to large prostate glands, isolating distal cap-like portions of atria histologically different to rest. Main body of atria somewhat cylindrical, terminal setion again histologically distinct ending in penial structure bearing a penis sheath of varying form. Posterior dorsal setae, at least, frequently reduced to a simple-pointed form barely distinguishable from the short hair setae where these are present. Body wall variably papillate or at least with foreign matter coating. Other characteristics as for the subfamily. (ref. ID; 5945)

Type species

Tubificoides heterochaetus Lastockin, 1937 (ref. ID; 5945)

ref. ID; 7240

The genus Tubificoides Lastockin, 1937 was re-erected by Holmquist (1978) and expanded by Brinkhurst and Baker (1979) as part of the reorganization of the former polyphletic genus Peloscolex Leidy. The genus consists of a growing number of coastal and estuarine tubificids, most of which have much narrower geographic ranges than was originally believed. (ref. ID; 7240)
  1. Tubificoides aculeatus (Cook, 1974) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 7203)
    Syn; Peloscolex aculeatus Cook, 1974 (ref. ID; 5945)
  2. Tubificoides amplivasatus (Erseus, 1975) (ref. ID; 5945) reported author and year? (ref. ID; 7203)
    Syn; Peloscolex amplivasatus Erseus, 1975 (ref. ID; 5945)
  3. Tubificoides amplivastus (ref. ID; 5892)
  4. Tubificoides apectinatus (Brinkhurst, 1965) (ref. ID; 7240), (Brinkhurst, 1965) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 7203)
    Syn; Peloscolex apectinatus Br. Cook, 1969, 1971 (in part) (ref. ID; 7240); Peloscolex gabriellae apectinata Brinkhurst, 1965 (ref. ID; 7240); Peloscolex gabriellae apectinatus Brinkhurst, 1965 (ref. ID; 5945)
  5. Tubificoides bakeri Brinkhurst, 1985 (ref. ID; 7240 original paper)
  6. Tubificoides benedeni (Udekem, 1855) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 7203)
    See; Tubificoides benedii (ref. ID; 7248)
    Syn; Tubifex benedii Udekem, 1855 (ref. ID; 5945)
  7. Tubificoides benedii (d'Udekem, 1855) (ref. ID; 6593, 7248) reported year? (ref. ID; 6590)
    Syn; Edukemius benedii (Udekem, 1855) Holmquist, 1978, 1979 (ref. ID; 7248); Peloscolex benedeni (Udekem, 1855) Brinkhurst & Jamieson, 1971 (ref. ID; 7248) reported author and year? (ref. ID; 6593); Tubificoides benedeni (Udekem, 1855) (ref. ID; 7248)
  8. Tubificoides brevicoleus Baker, 1983 (ref. ID; 7203 original paper)
  9. Tubificoides brownae Brinkhurst & Baker, 1979 (ref. ID; 5945 original paper, 7248) reported author and year? (ref. ID; 7203)
    Syn; Tubificoides coatesae Brinkhurst & Baker, 1979 (ref. ID; 7248)
  10. Tubificoides coatesae Brinkhurst & Baker, 1979 (ref. ID; 5945 original paper)reported author and year? (ref. ID; 7203)
    See; Tubificoides brownae (ref. ID; 7248)
  11. Tubificoides crenacoleus Baker, 1983 (ref. ID; 7203 original paper, 7248)
  12. Tubificoides cuspisetosus Baker, 1983 (ref. ID; 7203 original paper)
  13. Tubificoides denouxi (Shirley & Loden, 1982) (ref. ID; 7203)
  14. Tubificoides diazi Brinkhurst & Baker, 1979 (ref. ID; 5945 original paper, 6602, 7248) reported author and year? (ref. ID; 7203)
    Syn; Tubificoides maureri Brinkhurst & Baker, 1979 (ref. ID; 7248)
  15. Tubificoides dukei (Cook, 1974) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 7203)
    Syn; Peloscolex dukei Cook, 1974 (ref. ID; 5945)
  16. Tubificoides euxinicus (Hrabe, 1966) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 7203)
    Syn; Peloscolex euxinicus Hrabe, 1966 (ref. ID; 5945)
  17. Tubificoides foliatus Baker, 1983 (ref. ID; 7203 original paper)
    See; Tubificoides wasselli (ref. ID; 7248)
    Syn; Peloscolex gabriellae Marcus, 1950 (ref. ID; 7203); Peloscolex gabriellae typica Brinkhurst & Cook, 1966 (ref. ID; 7203); Tubificoides gabriellae (Marcus) (ref. ID; 7203)
  18. Tubificoides fraseri Brinkhurst, 1986 (ref. ID; 7248 original paper)
    Syn; Tubificoides gabriellae (Marcus) Chapman et al., 1982 (ref. ID; 7248), (Marcus) Chapmann & Brinkhurst, 1980, 1981, 1984 (ref. ID; 7248)
  19. Tubificoides gabriellae (Marcus, 1950) n. comb. (ref. ID; 5945)
    See; Tubificoides foliatus (ref. ID; 7203)
    Syn; Peloscolex gabriellae Marcus, 1950 (ref. ID; 5945)
  20. Tubificoides heterochaetus Lastockin, 1937 (ref. ID; 5945) reported author and year? (ref. ID; 7203)
    Syn; Tubificoides swirenocowi Jaroschenko, 1948 (non T. heterochaetus (Mich.)) (ref. ID; 5945)
  21. Tubificoides heterochaetus (Michaelsen, 1926) n. comb. (ref. ID; 5945), (Michaelsen, 1926) emm. Baker, 1981 (ref. ID; 7248)
    Syn; Limnodrilus heterochaetus Michaelsen, 1926 (ref. ID; 5945, 7248); Peloscolex heterochaetus (Michaelsen, 1926) Brinkhurst, 1963 (ref. ID; 7248), (Michaelsen, 1926) Brinkhurst & Jamieson, 1971 (ref. ID; 7248), (Michaelsen, 1926) Cekanovskaja, 1962 (ref. ID; 7248), (Mich.) Devos, 1936 (ref. ID; 5945), (Michaelsen, 1926) de Vos, 1936 (ref. ID; 7248), (Michaelsen, 1926) Hrabe, 1964 (ref. ID; 7248)
  22. Tubificoides imajimai Brinkhurst, 1985 (ref. ID; 7240 original paper)
  23. Tubificoides insularis (Stephenson, 1922) Brinkhurst, 1985 (ref. ID; 7240 redescribed paper)
    Syn; Peloscolex benedeni (d'Udekem) in part: Brinkhurst 1965, 1971 (ref. ID; 7240)
  24. Tubificoides intermedius (Cook, 1969) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 7203)
    Syn; Peloscolex intermedius Cook, 1969 (ref. ID; 5945)
  25. Tubificoides kozloffi Baker, 1983 (ref. ID; 7203 original paper)
  26. Tubificoides longipenis (Brinkhurst, 1965) (ref. ID; 7248), (Brinkhurst, 1965) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 7203)
    Syn; Tubifex longipenis Brinkhurst, 1965 (ref. ID; 5945, 7248); Tubifex longipenis Cook, 1971 (ref. ID; 7248)
  27. Tubificoides maritimus (Hrabe, 1973) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 7203)
    Syn; Tubifex maritimus Hrabe, 1973 (ref. ID; 5945)
  28. Tubificoides maureri Brinkhurst & Baker, 1979 (ref. ID; 5945 original paper)
    See; Tubificoides diazi (ref. ID; 7248)
  29. Tubificoides motei Brinkhurst, 1986 (ref. ID; 7248 original paper)
  30. Tubificoides nerthoides (Brinkhurst, 1965) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 7203)
    Syn; Peloscolex gabriellae nerthoides Brinkhurst, 1965 (ref. ID; 5945)
  31. Tubificoides palacoleus Baker, 1983 (ref. ID; 7203 original paper)
  32. Tubificoides parapectinatus Brinkhurst, 1985 (ref. ID; 7240 original paper)
    Syn; Peloscolex apectinatus Brinkhurst and Simmons, 1968 (ref. ID; 7240); Peloscolex apectinatus Cook and Brinkhurst, 1975 (ref. ID; 7240)
  33. Tubificoides postcapillatus (Cook, 1974) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 7203)
    Syn; Tubifex postcapillatus Cook, 1974 (ref. ID; 5945)
  34. Tubificoides pseudapectinatus Brinkhurst, 1985 (ref. ID; 6602, 7240 original paper)
    Syn; Tubificoides apectinatus (Br.) (in part). Cook, 1969, 1971 (ref. ID; 7240)
  35. Tubificoides pseudogaster (Dahl, 1960) (ref. ID; 7248), (Dahl, 1960) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 6594, 7203)
    Syn; Isochaetides pseudogaster (Dahl) Hrabe, 1966 (ref. ID; 5945, 7248); Limnodrilus pseudogaster Dahl, 1960 (ref. ID; 5945, 7248); Tubifex pseudogaster (Dahl) Brinkhurst, 1962 (ref. ID; 5945, 7248)
  36. Tubificoides scoticus Brinkhurst, 1985 (ref. ID; 7240 original paper)
  37. Tubificoides swirencoides Brinkhurst, 1985 (ref. ID; 7240 original paper)
  38. Tubificoides swirencowi Jaroschenko, 1948 (ref. ID; 5945, 6618) reported author and year? (ref. ID; 7203)
  39. Tubificoides vestibulatus Erseus & Bonomi, 1987 (ref. ID; 6618)
  40. Tubificoides wasselli Brinkhurst & Baker, 1979 (ref. ID; 5945 original paper, 7248)
    Syn; Tubificoides foliatus Baker, 1983 (ref. ID; 7248)

Tubificoides apectinatus (Brinkhurst, 1965) (ref. ID; 7240), (Brinkhurst, 1965) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 7203)

Synonym

Peloscolex apectinatus Br. Cook, 1969, 1971 (in part) (ref. ID; 7240); Peloscolex gabriellae apectinata Brinkhurst, 1965 (ref. ID; 7240); Peloscolex gabriellae apectinatus Brinkhurst, 1965 (ref. ID; 5945)

Descriptions

Worms up to 5 mm long, up to 40 segments. Body wall with a thin layer of foregin matter from XV or XVI. Anterior dorsal setal bundles with 2-3 hair setae and 2-3 bifids with subequal teeth, the upper slightly the shorter, behind the clitellum there are 1 or 2 of each, the hairs are quite robust and the bifids are quite elongate distally, gently curved with very small teeth; posteriorly the setae are reduced to a single hair and bifid, the latter with very samll teeth. Anterior ventral setae 2-4 per bundle, with the upper teeth a little thinner and shorter than the lower, posteriorly the ventral setae become reduced to 1, with the upper tooth distinctly thinner than but only a little shorter than the lower. Spermathecae with voluminous ducts, a sperm trap, and elongate ampullae. Male ducts with a vas deferens at least three times the length of the atrium and thin, the atrium relatively short and thick. Penis sheath an elongate cylinder, often with a subterminal opening or recurved distal end. (ref. ID; 7240)

Remarks

Specimens from the Gorge, Victoria, B.C. (Brinkhurst 1979), have up to four hair setae and four bifids dorsally and as many as five bifids ventrally in anterior bundles. The penis sheaths are conical but the cuticule is only really thickened in a basal ring. (ref. ID; 5945)

The species as described in the original account has bifid dorsal setae in posterior bundles, but specimens from the Pacific coast of North America have simple pointed posterior dorsal setae. Not all of the Woods Hole material attributed to this species by Cook (1969, 1971) belongs here. Large number of specimens from the extensive Woods Hole collection were mounted under each cover slip, and so it is impossible to relate every specimen to a specific identification at this point without a great deal of effort, but slides said to include T. apectinatus contain two distinct species which can be discriminated now that the specific characteristics in the genus are better understood. The second species will be described below. The distinctions between this and other species in the swirencoi group with hair setae and bifid setae dorsally will be evaluated in the discussion. (ref. ID; 7240)

Habitat

Atlantic coast species, presumably subtidal, coastal. (ref. ID; 7240)

Distribution

Nova Scotia, Massachusetts. (ref. ID; 7240)

Examined materials

  • Holotype: United States National Museum (USNM) 32621, a whole-mounted specimen, East Pass, Halifax, Nova Scotia, June 1962, coll. T.W. Beak. (ref. ID; 7240)
  • Paratypes: USNM32622, 3 whole-mounted and 1 dissected specimen; American Museum of National Hisotry (AMNH) 3660, 3 whole-mounted and 2 dissected specimens; British Museum of National History (BMNH) 1964.15.1, 2 whole-mounted specimens; R.O. Brinkhurst collection, 4 whole-mounted specimens, all from the type locality. (ref. ID; 7240)
  • Other: Specimens from Cape Cod Bay, 1961-1962, det. Cook (1969, 1971). (ref. ID; 7240)

    Tubificoides bakeri Brinkhurst, 1985 (ref. ID; 7240 original paper)

    Descriptions

    Worms up to 7 mm long, up to 60 segments. A few median segments somewhat elongate, in one the posterior segments very short. Body wall with a variable covering of foreign matter from XIII. Anterior dorsal bundles with up to 4 hairs and 4 bifid setae with distinct teeth, divergent with the upper tooth thinner than but as long as the lower, only one of each in X, but beyond that 1-2 setae resembling the ventrals, no hair setae. Anterior ventral bundles with 3-4 setae each, the upper teeth thinner than the equally long lower teeth, but straight and diverging from the curved lower teeth, this difference becoming more pronounced posteriorly, where there are 1-2 quite stout curved setae per bundle. In one specimen the most posterior setae are thinner and straighter than the rest. The atrium is quite short, but other details of the reproductive system are not detectable a part from the penis sheaths, which are cylindrical with subterminal openings. None of the specimens had mated. (ref. ID; 7240)

    Remarks

    This a provisional description pending discovery of more material. Recent attempts to obtain such material failed. The species is distinguishable on the basis of the setal characters and penis sheath, and the atria are clearly those of a Tubificoides. (ref. ID; 7240)

    Habitat and distribution

    Pacific coast species. Known from the type locality, near Victoria, British Columbia, Canada and Los Angeles, California. (ref. ID; 7240)

    Examined material

  • Holotype: United States National Museum (USNM) 097294, a whole-mounted specimen, Mackenzie Bight, Saanich Inlet, near Victoria, British Columbia, Canada, 10 m, April 4, 1981, coll. H.R. Baker and K. Coates. (ref. ID; 7240)
  • Paratypes: USNM097295-8, four whole-mounted specimens from the type locality. (ref. ID; 7240)
  • Other: R.O. Brinkhurst collection, Palos Verdes Peninsula, Los Angeles, California, 61 m, 2 and 15 February 1983, 29 specimens, coll. T. Parker. (ref. ID; 7240)

    Tubificoides benedeni (Udekem, 1855) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 7203)

    Synonym

    Tubifex benedii Udekem, 1855 (ref. ID; 5945)

    Remarks

    Holmquist (1978) demonstrated that the male ducts of this heavily papillate species are basically of the form used here in defining this genus but used some fine differences to establish the new monotypic genus Edukemius that we see as a subgenus. Some small specimens attributed to this species that have short hair setae mentioned in the literature almost certainly belong to a distinct species. Both this and the very complex synonymy of this species are detailed in Brinkhurst and Jamieson (1971). All of the preceding species in the genus lack hair setae, whereas the following possess them, at least in post clitellar bundles, but they are characteristically short, usually bent. Like the above, they all possess very similar male efferent ducts and are recognizable as members of this genus. (ref. ID; 5945)

    Tubificoides benedii (d'Udekem, 1855) (ref. ID; 6593, 7248) reported year? (ref. ID; 6590)

    Synonym

    Edukemius benedii (Udekem, 1855) Holmquist, 1978, 1979 (ref. ID; 7248); Peloscolex benedeni (Udekem, 1855) Brinkhurst & Jamieson, 1971 (ref. ID; 7248) reported author and year? (ref. ID; 6593); Tubificoides benedeni (Udekem, 1855) (ref. ID; 7248)

    Descriptions

    Worms up to 55 mm long, 100 segments. Body wall closely papillate from II or at least from V or VI, but prostomium nonretractile. All setae with reduced or rudimentary upper teeth, 2-3 per bundle in II-III, thereafter 2 until XI, when only a single seta present in each bundle. Spermathecal and male pores in the ventral setal line, with raised lips around the pores. Vasa deferentia up to 37 µm wide, several times longer than the atria, which are up to 370 µm long by 70 µm wide at tne narrow point just above the penis sacs. Atria broader apically but otherwise typical in form and rearward curvature. Penis sheaths slightly conical with an inflated distal end, up to 100 µm long by 84 µm wide at the widest (basal) part. Spermathecae with ducts wide at the pore, then narrowing before entering voluminous ampullae via sperm traps. (ref. ID; 7248)

    Remarks

    Holmquist (1979) discussed the spelling of the name of this species, opting for the techinically correct original spelling of benedii rather than the widely known benedeni based on a later correction by W. Michaelsen. The species is abundant in North Atlantic shores on both sides of the ocean, where it is often found together with Clitellio arenarius (Muller). This is larger species than T. wasselli, which is also papillate but has a retractible prostomium like most papillate tubificids and very different setae with clearly bifid tips. The penis sheaths also differ in shape, and they are half the size in T. wasselli, but the other dimensions of the male duct are similar although the apical part of the atrium is wide in T. benedii. The early synonymy of the species is complex (see Brinkhurst and Jamieson 1971), but the specimens with hair setae in dorsal bundles included within this species in earlier accounts are now recognised as T. insularis (Stephenson, 1922) by Brinkhurst (1985). (ref. ID; 7248)

    Occurrence and ecology

    Marine oligochaeta. This species is regularly found in sulphide-rich sediments with extremely low oxygen tensions and can tolerate anaerobic conditions for several days. (ref. ID 6590)

    Material examined

    Over 200 specimens from all regions of Great Britain, many parts of Northern Europe (Sweden, Norway, Germany, Ireland), as well as New Brunswick, Canada, Connecticut and Maine, United States. (ref. ID; 7248)

    Tubificoides brevicoleus Baker, 1983 (ref. ID; 7203 original paper)

    Descriptions

    Length 10-20 mm, up to 105 segments, diameter at XI 550-860 µm (in fixed, slightly compressed whole-mounted sexually mature specimens). Prostomium as long as or slightly shorter than broad at peristomium, bluntly conical. Papillation usually present from 1/2III posteriad; papillation absent from 1/2X to 12/13 in sexually mature specimens. Clitellum over XI-XII. Anterior dorsal setal bundles with 2(3) bifid setae and (1)2 straight hair setae; bifid setae with upper tooth much thinner and shorter than lower. Posteriorly each dorsal bundle contains 1 bifid and 1 hair seta. Anterior ventral bundles with 2-3(4) bifid setae; posterior ventral bundles with 1 bifid seta. Upper teeth of bifid setae reduced posteriorly. Ventral setae absent in XI. Modified genital setae absent. Spermathecal pores paired in X, anterior and lateral to ventral setae of X; male pores paired posteriorly in XI in ventral setal line. (ref. ID; 7203)
  • Male genitalia: Sperm funnel large, vas deferens 29-43 µm wide with ciliated lumen and thick walls, may extend to XIV (up to 2500 µm long); joins atrium below swollen atrial head opposite entrance of the stalked prostate gland. Atrium cylindrical, histologically tripartite, with enlarged ental head and ectal penial bulb; external muscle layer of atrium thin (2.5-3.0 µm thick). Atrium (to base of penis sheath) 450-630 µm long, body of atrium 82-88 µm wide, narrowing basally to join penial bulb; swollen atrial head 100-120 µm wide. Atrium terminates in true penis with penis sheath; penial apparatus enclosed in short, muscular, penial sac. Penis sheath short and broad, approximately 30 µm long and 55 µm wide basally and apically, with a median narrowing approximately 42 µm wide. Spermathecae paired, each with bulbous base (100-115 µm high and 70-90 µm wide) narrowing abruptly to duct leading to large, thin-walled ampulla. Sperm present as elongate speramtozeugmata. Spermathecal ampullae may extend anteriad into IX or posteriad into XI. (ref. ID; 7203)

    Remarks

    Tubificoides brevicoleus appears related to T. denouxi (Shirley and Loden, 1982) and T. dukei (Cook, 1970) but is distinct in several respect. Tubificoides denouxi is papillate posteriorly but nonpapillate anteriorly; T. brevicoleus and T. dukei are papillate both anteriorly and posteriorly. The vasa deferentia of T. brevicoleus are very long (4-5 times as long as atria) while those of T. denouxi and T. dukei are much shorter (approximately 1 and 1+1/2 times length of atria, respectively). The forms of the penis sheaths are also distinct. Tubificoides denouxi has hair setae with slightly bifid tips (Shirley and Loden 1982), a feature that is unique within the genus. (ref. ID; 7203)

    Etymology

    From the Latin brevis, short; and coleus, sheath, scabbard; with a short (penis) sheath. (ref. ID; 7203)

    Habitat

    Intertidal sands with or without shell debris freshwater influence usually evident. (ref. ID; 7203)

    Type locality

    North tip of Luke Island, Fisher Channel, British Columbia. 52 degrees 6.6'N, 127 degrees 52.5'W. Low intertidal in Fucus sp. zone, shell sand, reducing below 2 cm; collected August 21, 1981 by H.R. Baker and K.A. Coates. (ref. ID; 7203)

    Type materials

  • Holotype: NMCIC (National Museums of Canada, National Museum of Natural Sciences Invertebrate Collection) 1982-1128, a mature, whole-mounted specimen. (ref. ID; 7203)
  • Paratypes: NMCIC1982-1129, -1130, two whole-mounted specimens from the type locality. USNM (United States National Museum, Smithsonian Institution) 079836-079837, two whole-mounted specimens from type locality. (ref. ID; 7203)
  • Other material examined

    Tubificoides brownae Brinkhurst & Baker, 1979 (ref. ID; 5945 original paper, 7248) reported author and year? (ref. ID; 7203)

    Synonym

    Tubificoides coatesae Brinkhurst & Baker, 1979 (ref. ID; 7248)

    Descriptions

    Length 20 mm; breadth 0.3 mm; 38 segments. Prostomium blunt, shorter than broad, clitellum 1/2X - 1/2XI (XI?). Body wall naked, sometimes with glandular (?) annulations just anterior to or in line with setae posteriorly. Segments elongating from VII to reach ~200 µm at XIV-XVI, and shortening posteriorly. Setae bifid, three or four (maximum of five) anteriorly with only one from X posteriorly, nodulus median; upper tooth thinner and shorter than lower anteriorly, posteriorly the lower tooth broadens. Setae increase in size to VII (45 µm) after which length about constant. Atria cylindrical with proximal enlarged head, distal penial bulb. Atria 280-430 µm long, penial bulb 66-71 µm long, 82-93 µm broad. Vasa deferentia slender (16 µm diameter) and about twice the length of the atria, joining subapically just below the head and opposite the prostate glands. Prostates stalked, about half the length of the atira and twice as broad. Penes with cuticular sheaths, short and broad (1.5 times as high as broad), with lateral openings. Spermathecae elongate (600 µm long) narrowing basally to enter separate exit ducts some 180-190 µm long. These ducts leave the main mass of the spermathecae laterally, creating sperm traps (?) and are enlarged (bulbous) between the sperm traps and the external pores. Spermathecal and penial pores ventral, each pair quite close together. Spermatozeugmata vermiform. (ref. ID; 5945)

    Body wall naked or covered with a thin layer of foreign matter. Three to five anterior setae with the upper teeth slightly shorter and thinner than the lower, single posterior setae with teeth quite divergent, the upper now distinctly thinner than the broad lower teeth, but more nearly equal in length. Spermathecae like those of other species in this assemblage, duct wide close to the pore, narrower towards the voluminous ampulla which it joins subapically. Atria 280-430 µm long, recurved posteriad, with vasa deferentia up to twice the length of the atria. Penis sheaths thin and often crumpled in whole mounts, but basically broadly conical (47-53 by 60-75 µm). Epithielium around the spermathecal and male pores glandular. Posterior segments may be elongate. (ref. ID; 7248)

    Remarks

    The somewhat shovel-shaped penis sheaths of this species are distinctive; the combination of other characters clearly separating this from other species. The elongate segments of the posterior body show slight development of the annulation seen in Tubificoides coatesae. (ref. ID; 5945)

    Reexamination of specimens from the type series of these taxa serves to emphasize the similarity between the east coast brownae and the west coast coatesae. The situation is made more complex by the reporting of T. brownae in San Francisco Bay by P.V. Dresler (personal communication), whose specimens certainly fit the description of this species, as do those formerly described as P. gabriellae from this area. Also, new material from Barnstable Harbour and Little Sippewisset Marsh, Massachussetts, identified by H.R. Baker as T. brownae, and the east coast material derived from C. Erseus collections have some traces of foregin matter on posterior segments, and elongate posterior segments are often visible. As the setae of these two sets seem identical, there remain no reliable characters separating them. The degree of lengthening of posterior segments may well be subject to variation simply due to killing and fixation method, and so differences reported here should be regarded with caution. The penis sheaths of dissected specimens in polyvinyl lactophenol from the type series appear bigger than those observed in the Canada balsam mounted material from Massachussets, which may be an artifact due to pressure exerted during mounting. The penis sheath in this material does appear to have a characteristics form with a suggestion of a projection on one side at least, but in many instances the small thimble to conical-shaped penis sheaths are crumpled as they are in most of these species. Specimens from the Waccamaw River, Winyah Bay, South Carolina, also appear to be identifiable as T. brownae. The worms have naked body walls, the segments are elongate posteriorly, and all the other characters seem appropriate. The European material is all clearly identifiable as T. brownae. The species is known from British Columbia to California, Massachussetts to South Carolina, England, and Saudi Arabia. (ref. ID; 7248)

    Etymology

    For Betsy Brown (College of Marine Sciences, Delaware), who collected the specimens. (ref. ID; 5945)

    Material examined

  • Brinkhurst collection: 3 specimens from the type series, Delaware; 7 from Barnstable Harbour, 1 June 1981 and 1 from Little Sippewisset Marsh, Massachusetts, 31 May 1981, coll. B. Day; 7 from the Waccamaw River, Winyah Bay, South Carolina, 5 November 1980, coll. K. Speer, Environmental Analysts of Florida Inc., at a salinity of 15 0/00; 1 from Eel Pond, Woods Hole, 24 June 1964, coll. R.O. Brinkhurt (identified as T. pseudogaster): 18 specimens from San Francisco Bay and Suisan Bay, California, 1961-1962, coll. R.A. Wagner (identified as Peloscolex gabriellae; Brinkhurst and Simmmons 1968); 9 specimens from the Medway, Kent, England, coll. B. Dives. (ref. ID; 7248)
  • Erseus collection: 3 specimens from the Marine Ecosystem Research Laboratory enclosures, Narragansett Bay, Rhode Island, 5 from Chesapeake Bay, Virginia, 5 from the Southern Bight, North Sea, and 5 from the Seven estuary, Great Britain, also from Coos Bay, Oregon, coll. D. Strehlow, and Saudi Arabia, material not seen (C. Erseus, personal communication). As Tubificoides coatesae: 7 specimens from the sample location and date as the type series, Chemainus, British Columbia, plus 1 other from the same location coll. C. Levings. (ref. ID; 7248)

    Tubificoides coatesae Brinkhurst & Baker, 1979 (ref. ID; 5945 original paper)reported author and year? (ref. ID; 7203)

    See

    Tubificoides brownae (ref. ID; 7248)

    Descriptions

    Dimensions 14 mm x 0.5-0.8 mm (at V), 1.05-1.15 mm (at XIII), 0.5-0.65 mm (at XVII), more than 46 segments. Prostomium rounded, conical, about as long as broad at base. Foreign matter adhering to body wall posteriorly in some specimens. Clitellum XI-1/2 XII. All setae bifid with upper teeth thinner and slightly shorter than or equal to lower teeth; upper teeth increase in length posteriorly while the lower teeth broaden and point away from the line of the shaft. Nodulus distal in anterior setae, becoming median posteriorly. Setae increase in size to VI or VII, remain this size to IX-X, then increased again to XIII-XIV after which length remains fairly constant. Setal length variable within bundles. Setae (two to five), usually three or four per bundle anteriorly, usually two in IX, one in X, zero in XI or one where present, posteriorly one seta. Body wall with glandular (?) rings in setal line posteriorly, segments elongate gradually to XX-XXI (600 µm long) than shorten again (300 µm by XXX), setal line three-quarters of the way from the anterior border of each segment. Atria elongate, tubular, with enlarged head end, 370-410 µm long, head 80 µm wide. Atria narrow medially and enlarge basally to join penial sacs. Penes with cuticular sheaths, conical to thimble shaped with distal openings, 47-53 µm wide at base, 60-75 µm long. Vasa deferentia 15-17 µm wide, longer than atria, which they join subapically opposite to the stalks of the prostate glands. Spermathecae elongate (450 µm) tubular, narrowing basally, with sperm traps. Spermatozeugmata vermiform. (ref. ID; 5945)

    Remarks

    If it were not for the elongate segments, observed in other species in the genus, this species would be largely inseparable from the T. pseudogaster assemblage on the basis of the other available characters. Only the form of the penis sheath gives another indication of its difference. Otherwise the whole group of T. gabriellae-pseudogaster-coatesae differ from the rest primarily in the form of the penis sheaths. (ref. ID; 5945)

    Etymology

    Named for K. Coates, the collector of this and much other British Columbia material. (ref. ID; 5945)

    Tubificoides crenacoleus Baker, 1983 (ref. ID; 7203 original paper, 7248)

    Descriptions

    Length 10-18 mm, 45-55 segments. Diameter at clitellum (in fixed, whole-mounted slightly compressed sexually mature specimens) 340-510 µm. Prostomium sharply conical, longer than broad at peristomium. Body wall naked. Clitellum present over 1/3X-XII. Anterior setal bundles with 3-5 (6) bifid setae, posteriorly 2-3 bifid setae per bundle; upper tooth slightly thinner and as long as or longer than the lower. Ventral setae absent in XI; modified genital setae absent. Spermathecal pores paired in X lateral to ventral setae in lateral line; male pores paired medially in XI, in ventral setal line. (ref. ID; 7203)
  • Male genitalia: Sperm funnel relatively large, upper lip slightly longer than lower, vas deferens 22-24 µm wide with ciliated lumen and thin walls, approximately 560 µm long; joins atrium subapically below swollen atrial head opposite to entrance of stalked prostate gland. Atrium cylindrical, histologically tripartite, with enlarged ental head and ectal penial bulb, external muscle layer of atrium 2.8-5.0 µm thick over body of atrium, very thin over enlarged head. Atrium (to base of penis sheath) 350-430 µm long, body of atrium 44-60 µm wide, widening slightly basally to join penial bulb; swollen atrial head 70-90 µm wide. Atrium terminates in true penis with penis sheath, penial apparatus enclosed in very muscular, deep penial sac. Penis sheath slightly cone-shaped, 34-40 µm wide basally, 67-100 µm long; with a distal blunt projection on the posterior edge, projection pointing directly away from sheath or slightly towards base of sheath. Penis sheath with distal opening. Spermathecae paired, each with bulbous base 45-65 µm wide narrowing to join large ampulla, sperm trap present. Muscle layer 2-4 µm thick over bulbous base, very thin or absent over ampulla. Spermatozeugmata long, vermiform; spermathecal ampullae often extended anteriorly into IX, posteriorly into XI or XII. (ref. ID; 7203)

    Length 10-18 mm, 45-55 segments. Body wall naked. Anterior setal bundles with 3-5 (6) setae per bundle, posteriorly 2-3, upper tooth slightly thinner than but as long as or longer than the lower. Spermathecal pores lateral to the ventral setae, male pores in ventral setal line. Vas deferens 22-24 µm wide, about 560 µm long; atrium 350-430 µm long, 44-60 µm wide, muscle layer 2.8-5.0 thick; penis sheath slightly cone shaped, opening distal to laterodistal, associated with a lateral projection of varying shape and position. Spermathecae with rather bulbous short ducts joining the pear-shaped ampullae subapically via sperm traps. (ref. ID; 7248)

    Remarks

    The form of the lateral projection of the penis sheath in T. crenacoleus is variable. While a lateral projection of some type was present in all penis sheaths examined the sheaths may very in appearance. This variation may be an artefact due to handling and fixation methods. T. crenacoleus appears to be related to T. diazi and T. longipenis. The penis sheaths of these three species are similar in that they possess a lateral projection; however, the shapes of the penis sheaths are distinct. (ref. ID; 7203)

    At the level of character analysis used here and by Baker (1983), this species now differs from T. pseudogaster primarily by the form of the penis sheath, which has a blunt projection on one edge (the posterior edge according to Baker 1983) that seems to provide a subterminal opening to the sheath in some instances. In others there appears to be a lateral lobe without any opening in the sheath. The sizes and proportion of parts of the male duct as judged from camera lucida drawings are similar in both species, as are the setae and the spermathecae, including their pore position. Apart from the type series (from the Coldville River, Beaufort Sea, Alaska) the species can be recognized among collection previously identified by the donors as T. pseudogaster. These extend the range of the species to the River Humber, Grimsby, Great Britain, and the Oslo Fjord, Norway. (ref. ID; 7248)

    Etymology

    From the Latin crena, notch or rounded projection; and the Latin coleus, sheath or scabbard; (penis sheath with a rounded projection.) (ref. ID; 7203)

    Habitat

    Subtidal habitats from 3 to 125 m. (ref. ID; 7203)

    Type locality

    At the mouth of the Coldville River, Beaufort Sea, Alaska. 70 degrees 30'00"N, 150 degrees 09'20"W. 3+1/2 m, collected August 25, 1976 by J.R. Cordwell. Fine sandy silt. (ref. ID; 7203)

    Type materials

  • Holotype: USNM (United States National Museum, Smithsonian Institution) 079844, a whole-mounted specimen. (ref. ID; 7203)
  • Paratypes: USNM079845-079846, two whole-mounted specimens from the type locality. NMCIC (National Museums of Canada, National Museum of Natural Sciences Invertebrate Collection) 1982-1141, -1142, two whole-mounted specimens from the type locality. (ref. ID; 7203)
  • Other material examined:

    Material examined

    Three specimens from the type series from the Beaufort Sea; one specimen, original material from the Baker collection from the Eskimo Lakes, Northwest Territories; 10 specimens, Beaufort Sea 70 degrees 30.54'N, 150 degrees 01.54'W, 25 August 1976, 14 August 1978, coll. H. Koch, J. Zehr; 53 specimens, Beaufort Sea, Tuktoyaktuk, Seakem Oceanography 1980; 2 specimens, labelled AWA Grimsby, H.R. Baker collection, clearly part of the River Humber survey of 1979-1980, coll. B.E. Barnett, Anglian Water Authority; 21 specimens, Oslo Fjord, Norway, coll. J. Gray; both European sets require confirmation. (ref. ID; 7248)

    Tubificoides cuspisetosus Baker, 1983 (ref. ID; 7203 original paper)

    Descriptions

    Length 10-14 mm, 39-67 segments. Width at clitellum (in fixed, whole-mounted, slightly compressed sexually mature specimens) 500-560 µm, in sectioned specimens 260-290 µm. Prostomium bluntly conical, may bear small papilla at tip, prostomium as long as or slightly shorter than broad at peristomium. Posteriorly body wall may have foreign material adhering. Anterior dorsal setal bundles (to VI or VII) with 2-4 short hair setae and 2-4 bifid setae; bifid setae with teeth of approximately equal length, teeth closely parallel. From VI(VII) posteriad dorsal setal bundles with 3-4 short hair setae and 3-4 sharply simple-pointed setae. Hair setae smooth (without serrations or lateral hairs). Anterior ventral setal bundles (to VI or VII) with 4-5 (6-7) bifid setae similar in shape to dorsal bifid setae, from VI(VII) posteriorly 3-4 (5) sharply simple-pointed setae per bundle. Ventral setae absent in XI, modified genital setae absent. Spermathecal pores paired in X, in ventral setal line just anterior to ventral setae, from VI(VII) posteriorly 3-4 (5) sharply simple-setal line. (ref. ID; 7203)
  • Male genitalia: Sperm funnel with lips of approximately equal length, vas deferens 18-22 µm wide with ciliated lumen and thin walls, approximately 1500 µm long; joins atrium subapically below swollen head opposite to entrance of stalked prostate gland. Atrium cylindrical, histologically tripartite, with enlarged ental head and ectal penial bulb, external muscle layer 1.5-4.5 µm thick over body of atrium, approximately 2 µm thick over enlarged head. Atrium (to base of penis sheath) 275-320 µm long, main body of atrium 47-63 µm wide, narrowing slightly basally to join penial bulb; swollen atrial head 54-62 µm wide. Atrium terminates in true penis with penis sheath, penial apparatus enclosed in short, muscular, penial sac. Penis sheath long and slender, distal tip may be flared slightly, sheath 75-100 µm long, 27-54 µm wide at base, 17-27 µm wide at distal tip. Spermathecae paired, each with slightly bulbous base (37-50 µm wide) and narrow duct (20-25 µm wide) leading to ampulla; sperm trap absent. Muscle layer thin. Spermatozeugmata long, vermiform; spermathecal ampullae may extend anteriad to IX or posteriad to XI. (ref. ID; 7203)

    Remarks

    The change from bifid to simple-pointed setae in T. cuspisetosus is a gradual process in both dorsal and ventral setal bundles; the upper tooth becomes shorter and thinner over several segments until it disappears. The most anterior dorsal bundles usually contain only two hair setae and two bifid setae but by IV or V there are 3-4 of each type present in each bundles. Tubificoides cuspisetosus appears related to T. palacoleus sp. nov. which also has simple-pointed setae anteriorly. Details of the setation and male genitalia as well as differences in the shape of the penis sheaths serve to separate these species. Tubificoides swirencowi (Hrabe, 1964), T. amplivasatus (Erseus, 1975), T. apectinatus (Brinkhurst, 1965), and T. intermedius (Cook 1969) also possess hair and simple-pointed setae posteriorly but do not have simple-pointed setae anteriorly. This feature effectively separates these species from both T. cuspisetous and T. palacoleus. (ref. ID; 7203)

    Etymology

    From the Latin cuspis, pointed end of any object, and the Latin seta, seta; after the preponderance of sharply (simple) pointed setae. (ref. ID; 7203)

    Habitat

    Subtidal habitats 3-125 m depth. (ref. ID; 7203)

    Type locality

    Just off Oliktok Point, Beaufort Sea, Alaska. 70 degrees 30'54"N, 150 degrees 01'54"W. 3 m depth, salinity 28 ppt, collected September 14, 1978 by Dr. A.C. Broad, sediment muddy with much decaying tundra vegetation. (ref. ID; 7203)

    Type materials

  • Holotype: USNM (United States National Museum, Smithsonian Institution) 079839, a whole-mounted specimen. (ref. ID; 7203)
  • Paratypes: USNM 079840-079843, five whole-mounted specimens from the type locality. NMCIC (National Museums of Canada, National Museum of Natural Sciences Invertebrate Collection)1982-1133, -1134, four whole-mounted specimens from the type locality.
  • Other material examined:

    Tubificoides diazi Brinkhurst & Baker, 1979 (ref. ID; 5945 original paper, 6602, 7248) reported author and year? (ref. ID; 7203)

    Synonym

    Tubificoides maureri Brinkhurst & Baker, 1979 (ref. ID; 7248)

    Descriptions

    Length to 20 mm; breadth 0.2 mm: 73 segments. Prostomium rounded, about as long as breadth at the base, with small papilla on tip. Body wall may be encrusted with foreign matter. Clitellum XI-1/2 XII. Setae all bifid, two to six, usually three to five per bundle anteriorly in both dorsal and ventral bundles. Ventral setae reduced to two or three by segment VIII, dorsals one or two from XII. Segment X has one, two, or ventral setae, ventral setae of XI missing; dorsals of X-XI single or absent. Anterior setae with distal nodulus, upper teeth thinner and shorter than lower, posteriorly teeth equally long, the lower slightly broader. Setae small throughout (24-35 µm). Atria cylindrical (210-240 µm long) with proximal enlarged head 45-53 µm diameter and distal penial bulb (90-105 µm long by 70-80 µm). Vasa deferentia slender (16 µm diameter), three times as long as atria, with ciliated lumen, joining atria below apex opposite the prostate glands. Atria bent over posteriad. Penis sheaths four to five times as long as broad, with lateral openings. Just below, and on the same side as the lateral openings, lie large heavily cuticlarized rounded projections pointing towards base of penes. Spermathecae short (210 µm) thick (40 µm) cylinders with bulbous swellings on the innermost end (approximately 85 µm wide). Spermathecal and male pores open close together, ventrally and in setal line. (ref. ID; 5945)

    Length 20 mm, breadth 0.2-0.3 mm, up to 73 segments. Body wall naked or with a coating of foreign matter, posterior segments elongate. Anterior setal bundles with (2) 3-5 (6) setae per bundle, most in II-III, posteriorly (1) 2 (3), upper teeth thinner and shorter than the lower anteriorly, posteriorly lower teeth relatively broad but teeth about equally long. Spermathecal and male pore ventrolateral, in the line of the ventral setae. Atria short, up to 240 µm long, vasa deferentia 16-17 µm wide, from 3 to 6 times longer than the atria. Penis sheath curved, frequently bearing a spur developed to a variable extent, up to 140 µm long. Spermathecae with short ducts entering pear-shaped ampullae subapically via sperm traps. (ref. ID; 7248)

    Remarks

    The species resembles T. longipenis only in some details of the form of the penis sheaths, but is otherwise quite distinct. (ref. ID; 5945)

    The discovery of both the diazi and maureri forms (specimens with and without distinct spurs on the penis sheaths) in the collection from the type locality of diazi off New Jersey, and in other material from the same collector from off Virginia, led to the suspicion that these two species might prove to be synonymous. At first, the two seemed separable by the difference in the length of the penis sheaths (mean values of 95 vs. 125 µm for maureri and diazi), but then the Australian material was examined and that proved to have a mean length of the penis sheath (measured straight across the curved shape in the same way as before) of 110 µm. The complete ranges of measurements were 82-110, 96-140, and 90-131 µm, respectively, n varying from 16 to 19. Some specimens bear spurs on the penis sheath and some do not. In those with spurs, it would appear that the spur is a development of the upper lip of the ventrolateral opening of the penis sheath, and reference to the specimens in the type species show that this is the case rather than it being a separate structure on the body of the penis sheath as originally illustrated. There seems to be no reason to support the separation of these species further. The obvious difference between this species and both T. pseudogaster and T. crenacoleus is the overall form of the penis sheath. Otherhwise the two are very similar. The spermathecae are of the same form as that found in the rest of this group, with short ducts entering voluminous ampullae via sperm traps. There are often 6 thin setae per bundle anteriorly. The specimens from Virginia and Arcachon, France, seem quite typical of the diazi form, but the Oregon material is problematic in that the projections on the penis sheaths are far less obvious. In some of them the penis sheaths are shorter than usual, but these are unmated specimens (unlike the others) and the basal part of the penis may not be completely developed. The specimens from British Columbia are more typical, however, confirming the species for the Pacific. The single specimen from Kinneil, Firth of Forth, near Stirling, Scotland, strongly resembles this species. It has up to 5 setae per bundle anteriorly, reducing to 1 posteriorly, all with the upper tooth thinner and shorter than the lower. The penis sheaths are long and bent distally, without a rearward projection. The other single specimen from Barnstable Harbour resembles the maureri form. It has 3-4 anterior setae, usually 2 posteriorly, and the penis sheaths are long, curved, and open laterally rather than terminally. The species is known from the United States (Massachussets, Delaware, New Jersey, Virginia, Oregon), Canada (British Columbia), Scotland, France, and Australia (Victoria). (ref. ID; 7248)

    Etymology

    For R.J. Diaz (Virginia Institute of Marine Science), who collected the species. (ref. ID; 5945)

    Locality

    Marine species. (ref. ID; 6602)

    Material examined

    Brinkhurst and Erseus collections, 48 specimens from the type locality, offshore New Jersey to Virginia, 37 degrees 13.6' to 39 degrees 40'N, 72 degrees 55' to 75 degrees 60'W, 25-110 m, 1975-1976, coll. R.J. Diaz; 11 specimens, North of Cape Hatteras, North Carolina, 35 degrees 31' to 36 degrees 03'N, 75 degrees 10.6' to 75 degrees 13'W, 25-110 m, 1975 coll. D. Weston; 1 specimen from Barnstable Harbour, Massachusetts, June 1981, coll. B. Day; 8 specimens, intertidal zone, Coos Bay, Oregon, 1979, coll. D. Strehlow; 17 specimens, Boundary Bay, Vancouver, B.C., 49 degrees 01.36'N, 122 degrees 53.30' W, 15 November 1984, 11 m, coll. D.F. Moore; 5 whole-mounted and 2 sectioned specimens, Arcachon, France, Bassin d'Arcachon, Chenal de 1'Isle, plage d'Eyrac, low water low tide, Chenal de Courbery, March 1978, 5 m coll. C. Erseus; 1 specimen from Kinneil, Firth of Forth, Scotland, 56 degrees 01.58'N, 03 degrees 35.53'W, coll. B. Dicks; 20 specimens, Bancroft Bay, Gippslands Lakes, Victoria, Australia 30 May 1979, 6 m, coll. personnel of Marine Studies Group, Ministry for Conservation. (ref. ID; 7248)

    Tubificoides foliatus Baker, 1983 (ref. ID; 7203 original paper)

    See

    Tubificoides wasselli (ref. ID; 7248)

    Synonym

    Peloscolex gabriellae Marcus, 1950 (ref. ID; 7203); Peloscolex gabriellae typica Brinkhurst & Cook, 1966 (ref. ID; 7203); Tubificoides gabriellae (Marcus) (ref. ID; 7203)

    Descriptions

    Length 5.3-6.5 mm, 39-48 segments, width at XI 220-300 µm (in fixed, slightly compressed, sexually mature specimens). Prostomium bluntly conical with rounded tip, about as long as broad at peristomium, retractile. Body wall papillate from (III)IV posteriorly. Clitellum from 10/11 to 1/2XII. Anterior setal bundles with 2 bifid setae, upper tooth slightly shorter and thinner than lower, posteriorly 1 bifid seta per bundle, upper tooth reduced. Somatic setae absent in X and XI, modified genital setae absent. Gut without dilation in IX. From (IX, X) XIII posteriad 4-8 (10?) groups of raised sensory cells visible per segment, four of these sensory papillae are anterior to and slightly ventral to the setal bundles, the other sensory papillae being between the setal bundles; sensory papillae absent over clitellar region. Spermathecal and male pores paired in X and XI respectively, in the line of the (missing) ventral setae. (ref. ID; 7203)
  • Male genitalia: sperm funnel small, vas deferens 15-20 µm wide, 490-530 µm long with ciliated lumen and thin walls, joins atrium subapically below swollen atrial head opposite to entrance of stalked prostate gland. Atrium cylindrical, histologically tripartite, with enlarged ental head and ectal penial bulb, external muscle layer of atrium 1-2 µm thick over main body of atrium, may be slightly thicker over enlarged head. Atrium (to base of penis sheath) 195-230 µm long, swollen atrial head 45-50 µm wide; atrium 25-35 µm wide below swollen head narrowing medially to 15-20 µm, 25-30 µm wide just above penial bulb. Atrium terminates in true penis with penis sheath, penial apparatus enclosed in short, muscular, penial sac. Penis sheath conical, with slightly shovel-shaped tip, opening distal; penis sheath 25-35 µm wide basally, 40-48 µm long, cuticle of penis sheath thin. Spermathecae paired, each with a very short duct leading from the spermathecal pore to the slightly bulbous base; duct 13-14 µm high, 14-16 µm wide, spermathecal base 110-115 µm long, 35-50 µm wide basally, 22-28 µm wide at entrance into ampullae. Sperm traps present, spermatozeugmata vermiform. (ref. ID; 7203)

    Remarks

    Tubificoides foliatus is one of only two species in Tubificoides with a papillate body wall and only bifid somatic setae posteriad. Tubificoides benedeni and T. heterochaetus are also papillate (and lack hair setae) but have a least some simple-pointed setae posteriad. Details of the male genitalia and shape of the penis sheaths serve to separate T. foliatus from T. wasselli (Brinkhurst and Baker 1979). (ref. ID; 7203)

    Etymology

    From the Latin folium, leaf; after the leaf-shaped papillae of the cuticle. (ref. ID; 7203)

    Habitat

    Very low intertidal or subtidal muddy sands in areas of reduced salinities. (ref. ID; 7203)

    Type locality

    Limekiln Cove, Esquimalt Harbour, Esquimalt, British Columbia. 48 degrees 27.2'N, 123 degrees 27.6'W. Very silty shell debris - sand mixture with freshwater seepage over beach, midintertidal. Collected August 8, 1979 by H.R. Baker. (ref. ID; 7203)

    Type materials

  • Holotype: NMCIC (National Museums of Canada, National Museum of Natural Sciences Invertebrate Collection) 1982-1131, a whole-mounted sexually mature specimen. (ref. ID; 7203)
  • Paratypes: NMCIC1982-1132, two whole-mounted specimens from the type locality. USNM (United States National Museum, Smithsonian Institution) 079838, two whole-mounted specimens from the type locality. (ref. ID; 7203)
  • Other material examined:

    Tubificoides fraseri Brinkhurst, 1986 (ref. ID; 7248 original paper)

    Synonym

    Tubificoides gabriellae (Marcus) Chapman et al., 1982 (ref. ID; 7248), (Marcus) Chapmann & Brinkhurst, 1980, 1981, 1984 (ref. ID; 7248)

    Descriptions

    Worms up to 13 mm long, 60 segments. Body wall naked or with a fine covering of foreign matter, whole worms with a golden yellow tint posteriorly. Setae 2 (3) per bundle anteriorly, 1 posteriorly, the upper tooth thinner than the lower and shorter in anterior setae, shorter than or equal to the lower posteriorly. Spermathecal and male pores in the ventral setal line. Spermathecae always rudimentary, with broad ducts at the pore, narrowing before entering the ampullae, spermatozeugmata never observed. Male ducts with funnels with few if any sperm associated with them, vasa deferentia 13-18 µm wide, from 1.25 to 2.25 times the length of the atria, atria up to 260 µm long. Penis sheaths conical to cylindrical, about twice as long as wide, up to 52x30 µm. The atria are the usual shape for the genus, with prostate glands opposite the vasa deferentia and a bulbous cap of cells distal to these. The sperm sacs extend into XI at most, with few sperm morulae. The egg sacs occupy the posterior half of XI as well as XII. (ref. ID; 7248)

    Remarks

    This species possesses rudimentary spermathecae but does not seem reproduce by exchanging spermatozeugmata. The male funnels appear somewhat reduced from the usual condition, as they may have cilia in the mouth of the funnel, or sperm tails, but these do not stain as intensely as usual nor are they as abundant. The vasa deferentia are open and ciliated. The spermathecae may be sufficiently developed to have a lumen in the ampulla, but they are always small and empty. Extensive study of a population in the Fraser River, British Columbia was persued under the name Tubificoides gabriellae, the general name for what is now recognized as a series of papillate marine and estuarine worms. No mated specimens were ever found, and this led Baker (1983) to suggest that material from this and a related laboratory study was unavailable to him, when the actually meant that there were no mated specimens in the extensive material. However, in Baker's notes were rough descriptions of the material from Goldstream and the Australian material, and both putative species were described from unmated specimens. All of these specimens are derived from shallow estuarine locations, and two at least are associated with sewage and other sources of pollution. In the Fraser River, the species is capable of responding to seasonal fluctuations in intestitial salinity by rapidly producing very large populations, which would support the anatomical evidence for a parthenogenetic or self-fertilising mode of reproduction. The species clearly differs from the other nonpapillate species in respect to its reproductive behaviour and the effect this has on the reproductive anatomy, but it also has only 2 anterior and 1 posterior seta per bundle, and smaller atria and penes than all the others except T. motei. In the latter the atria are very short (190 µm) but while the penis sheaths are similar in size to those of T. fraseri, they are more often conical with subterminal openings, and there are more posterior setae. (ref. ID; 7248)

    Type materials

  • Holotype: USNM98836, a whole-mounted specimen, Fraser River, North Arm, Vancouver, British Columbia, Canada, January 1978, coll. P.M. Chapman. (ref. ID; 7248)
  • Paratypes: USNM98837-42, many whole-moutned specimens and specimens in fluid from the type locality. (ref. ID; 7248)
  • Other material examined: Many specimens, Fraser River, North Arm, 13 August 1973, coll. T.G. Northcote; Fraser River estuary, 13 September 1973, coll. C. Levings; 17 dissections and whole-mounted specimens, Goldstream estuary, near Victoria, British Columbia, 20 December 1978, coll. H.R. Baker; 1 specimen, Newport Bay, and 1, Mugu Lagoon, California, November 1979, coll. M.L. Quammen; 10 specimens, Prime Hook Creek, near junction with Broadkill River, Lewes, Delaware, 1979, coll. G.P. Burbank; 12 specimens, Werribee Sewage treatment farm, Port Philip Bay, Victoria, Australia, 38 degrees 00' S, 144 degrees 36' E, 23 November 1979, coll. J. Dorsey; various sites in Gippsland lakes area, Victoria, including Fraser Island, 1978-1979, coll. Marine Studies Group. (ref. ID; 7248)

    Tubificoides gabriellae (Marcus, 1950) n. comb. (ref. ID; 5945)

    See

    Tubificoides foliatus (ref. ID; 7203)

    Synonym

    Peloscolex gabriellae Marcus, 1950 (ref. ID; 5945)

    Descriptions

    Length 10-20 mm. Anterior bundles with up to six setae with more or less equal teeth. Penes with thin conical sheaths. Body wall variably papillate. (ref. ID; 5945)

    Tubificoides heterochaetus (Michaelsen, 1926) n. comb. (ref. ID; 5945), (Michaelsen, 1926) emm. Baker, 1981 (ref. ID; 7248)

    Synonym

    Limnodrilus heterochaetus Michaelsen, 1926 (ref. ID; 5945); Peloscolex heterochaetus (Michaelsen, 1926) Brinkhurst, 1963 (ref. ID; 7248), (Michaelsen, 1926) Brinkhurst & Jamieson, 1971 (ref. ID; 7248), (Michaelsen, 1926) Cekanovskaja, 1962 (ref. ID; 7248), (Mich.) Devos, 1936 (ref. ID; 5945), (Michaelsen, 1926) de Vos, 1936 (ref. ID; 7248), (Michaelsen, 1926) Hrabe, 1964 (ref. ID; 7248)

    Descriptions

    Length 7-9 mm. Up to five bifid setae anteriorly, reduced to one or two simple-pointed setae in post-clitellar bundles. Vasa deferentia short, atria cylindrical, penes reportedly without cuticular penis sheaths. Body wall papillate. (ref. ID; 5945)

    Worms up to 9 mm long. Body wall slightly papillate posteriorly, with foreign matter attached. Anterior setal bundles with upper teeth about as long and thick as the lower, posterior ventral and 1 of 2(3) dorsal setae simple-pointed, the other dorsal seta may have a very thin, rudimentary lower tooth. Spermathecal and male pores in the line of the ventral setae. Male funnels moderate in size, vas deferens 35.5 µm wide, about as long as atria, a little over 300 µm, prostates small, penis with thin cuticular sheaths, 46 µm by 36 µm, slightly cone-shaped. Spermathecae with ducts wide at the base, narrowing prior to entering the voluminous ampullae, with sperm traps. The species is known from Virginia to Louisiana, United States, and from the North Sea coast of Europe and the Danube estuary. (ref. ID; 7248)

    Remarks

    The species was previously known only from Europe but has been found in the tidal James River, Virginia by R.J. Diaz in salinities ranging from 0.1 to 18 0/00. Specimens have been donated to the National Museums of Canada, Museum of Natural Sciences (1979-15). No penis sheaths were observed. (ref. ID; 5945)

    The simple-pointed setae are rare among this group of species without hair setae, though simple-pointed setae of a different form are found in the dorsal bundles of half the species bearing hair setae (Brinkhurst 1985). The penis sheaths are very simple, only a little longer than broad and very thin in contrast to those of T. longipenis, which has simple-pointed posterior setae that differ in form and number to those observed here. (ref. ID; 7248)

    Material examined

    12 whole-mounted specimens, Cape Fear River, North Carolina, 28 Feb. 1980, 8 m, coll. B. Woodard, many whole-mounted specimens; James River, Virginia, 1976-1977, coll. R. Diaz; 2 whole-mounted specimens, Calcasicu Lake, Louisiana, 16 January 1976 and 9 June 1976, coll. M.S. Loden. (ref. ID; 7248)

    Tubificoides imajimai Brinkhurst, 1985 (ref. ID; 7240 original paper)

    Descriptions

    One specimen about 5 mm long, 47 segments. Median segments elongate. Body wall with a thin covering of foregin material from XI to XV, surface with ridges and furrows but no papillae. Anterior bundles with 3 hair setae and 3 bifid setae with long parallel teeth, behind the clitellum reduced to 2 of each, in XV or XVI 1 bifid and 1 simple pointed seta in one specimen, both setae simple pointed beyond that. Anterior ventral setae 3-4 per bundle with the upper tooth very slightly thinner and shorter than the lower or teeth equal, posteriorly 2 setae with obvious, quite elongate, teeth, setae long beyond the nodulus. Vas deferens 3-4 times the length of the atrium and relatively broad; atrium short, strongly curved, terminating in an elongate penis with a penis sheath in the form of a modified cone. Spermathecal pores in the setal line; spermatheca with a large vestibule behind the pore and an elongate sac shaped ampulla with a sperm trap at the union of the two. (ref. ID; 7240)

    Remarks

    While there are only two specimens of this species available, the material is deemed sufficient to permit a name to be attached to this oriental species. (ref. ID; 7240)

    Habitat and distribution

    Pacific coast species. Knowledge limited to the type location, Japan. (ref. ID; 7240)

    Examined material

  • Holotype: United States National Museum (USNM) 097299, a whole-mounted specimen, Nagaura Bay, Yokosuka, Kanagawa Prefecture, Japan, 12.5 m, October 1977, coll. M. Imajima. (ref. ID; 7240)
  • Paratype: USNM097300, a whole-mounted specimen from the type locality. (ref. ID; 7240)
  • Other: 5 specimens in fluid (not examined) Japanese National Science Museum, Tokyo (NSMT)-An-101. (ref. ID; 7240)

    Tubificoides insularis (Stephenson, 1922) Brinkhurst, 1985 (ref. ID; 7240 redescribed paper)

    Synonym

    Peloscolex benedeni (d'Udekem) in part: Brinkhurst 1965, 1971 (ref. ID; 7240)

    Descriptions

    Worms up to 30 mm long, up to 150 segments. Body wall papillate from between III and VI rearwards. Anterior dorsal bundles with 1-3 (4) short bent hair setae and the same number of bifid setae, each with the upper tooth somewhat shorter and thinner than the lower. Behind the clitellum, the dorsal bundles contain 1 hair and 1 bifid setae. anterior ventral setal bundles with 3-4 bifid setae with the upper tooth distinctly shorter and thinner than the lower, falling to 1 seta posterior, with an exceedingly thin upper tooth. Spermathecal and penial openings close to the ventral setal lines in X and XI. Spermathecae globular, with distinct muscular ducts about as long as ampullae, only a slight distension of the duct close to the pore and sperm trap. Male ducts (paired) with a very long vas deferens coiled in the sperm sacs as far back as XVII, entering the elongate atrium opposite the prostate gland. Atria exceptionally long and thin. The penis sheath, set in a deep penial sac, is sharply conical but bears a distended ectal end. Spermatozeugmata unusually short and broad. (ref. ID; 7240)

    Remarks

    The type material consists of serially sectioned specimens which are hard to scan because of frequent changes in the orientation and sequence of the sections. There is no doubt, however, that they and the new specimens collected by H.R. Baker are the same species when these are compared with the original description. The species is large compared with many species in the genus, but is similar in general appearance to the common intertidal species T. benedii, (= T. benedeni). As it is usually found with T. benedii, care should be taken to ensure that the two species are recognized in future. The hair setae now provide an obvious distinction, and there are also more bifid setae in both dorsal and ventral bundles than in T. benedii. The setae also differ in shape, but while the penis sheaths are similar, the atria of T. insularis are much longer than those of T. benedii. The latter will be redescribed in a subsequent review of species lacking hair setae. In the original account, Stephenson (1922) stated that there were no papillae on the body wall behind the clitellum in contrast to T. benedii, but he suggested that this was known to be a variable character. Papillae are usually present throughout the body in the new material, with the exception of a variable number of anterior segments that can be retracted, as in so many tubificids with armoured body walls. Stephenson also noted the presence of simple pointed setae in both dorsal and ventral bundles. In the new material the dorsal setae seem to be always bifid even if the teeth are very small in posterior setae, but they appear simple pointed in lateral view. Some ventral setae have the upper tooth every much reduced and sometimes this is worn or broken off, but in general the setae do not become transformed into simple-pointed setae posteriorly as they do in some species in this genus. Stephenson also commented on the reduced number of spermatozeugmata in his material, but this is clearly a question of the reproductive state of the individual; he stated that the spermatozeugmata were exceptionally thick. (ref. ID; 7240)

    Habitat

    Atlantic coast species, muddy intertidal and estuarine situations. (ref. ID; 7240)

    Distribution

    New Brunswick, Maine, and Great Britain. (ref. ID; 7240)

    Syntype

    British Museum of Natural History (BMNH) 1933.5.25.1218-1223, serial sections of three specimens, Ascog, Bute, Scotland. (ref. ID; 7240)

    Other material examined

    United States National Museum (USNM) 32600 (from 26403), one whole-mounted dissected specimen indentified as "Peloscolex benedeni with hairs" (R.O. Brinkhurst), Grand Manan, New Brunswick; 32573, one specimen as above, Sheepscot River, Maine, both coll. M. Pettibone. H.R. Baker collection. All specimens coll, et det. H.R. Baker, 1981: Sherkin Island, Co. Cork, 12 locations, September 7-20 (USNM97311-13); Doctors Bay, south of Kircubbin, Co. Down, three localities, October 5; near Biological Station and at Mweenish Island, Mulroa Point, Galway Bay, September 26-28; south of Penmon Point, Gwynedd, North Wales, October 25; Tay Estuary and St. Andrews East Sands, Scotland, October 18-19; Filey Brigg, Hull, England, October 28. 52 whole-mounted or dissected specimens. (ref. ID; 7240)

    Tubificoides intermedius (Cook, 1969) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 7203)

    Synonym

    Peloscolex intermedius Cook, 1969 (ref. ID; 5945)

    Remarks

    The male ducts of the type specimen of this species are like those of the rest of the genus, not as illustrated by Cook (1969) with the vas deferens joining the atria apicallly. The species was recorded from Cape Cod Bay, 36.5-42.6 m (type locality). (ref. ID; 5945)

    Tubificoides kozloffi Baker, 1983 (ref. ID; 7203 original paper)

    Descriptions

    Length 5-8 mm, 50-62 segments, width at clitellum 270-400 µm (in fixed, slightly compressed, sexually mature specimens). Prostomium conical with rounded tip, shorter than broad at peristomium. Body wall naked. Clitellum over XI-XII. Anterior, dorsal, setal bundles with 2-3 pectinate setae and 1-2 long straight hair setae, pectinate setae with long, almost parallel teeth, upper tooth slightly shorter than lower, intermediate teeth very thin; posterior dorsal setal bundles with one bifid seta (not pectinate, upper tooth shorter and thinner than lower) and one straight hair seta. Anterior ventral setal bundles with (1) 2 (3) bifid setae per bundle, from (VIII)IX posteriorly 1 seta per bundle; all ventral setae with upper tooth shorter and thinner than lower, posteriorly upper tooth much thinner. Ventral sometimes absent in X, absent in XI, modified genital setae absent. Spermathecal pores paired in X in line with but anterior to ventral setae of X; male pores paired posteriorly in XI in ventral setal line. (ref. ID; 7203)
  • Male genitalia: Sperm funnel relatively large, upper lip slightly longer than lower. Vas deferens 12.5-15 µm wide, approximately 1000 µm long with ciliated lumen and thin walls; joins atrium subapically below swollen atrial head opposite to entrance of stalked prostate gland. Atrium cylindrical, histologically tripartite, with enlarged ental head and ectal penial bulb, external muscle layer of atrium thin (approximately 1 µm thick). Atrium (to base of penis sheath) 210-270 µm long, body of atrium 24-31 µm wide, narrowing slightly at penial bulb; swollen atrial head 50-75 µm wide. Atrium terminates in true penis with penis sheath, penial apparatus enclosed in short, muscular, penial sac. Penis sheath cylindrical, short and broad with basal and distal wideing, 40-52 µm long, 24-30 wide basally. Spermathecae paired, each with slightly bulbous base 23-31 µm wide and narrow duct 18-24 µm wide, duct widening to join large thin-walled ampulla; bulbous base and duct 100-115 µm long. Sperm present as short, thick spermatozeugmata. (ref. ID; 7203)

    Remarks

    Details of setation and the male genitalia, especially the form of the penis sheath, serve to distinguish T. kozloffi from the apparently related T. aculeatus (Cook, 1970) and T. nerthoides (Brinkhurst 1965; Cook and Brinkhurst 1973; Brinkhurst and Baker 1979). It should be noted here that issue of the Transactions of the American Microscopical Society in which the descriptions of T. aculeatus (Cook) and T. dukei (Cook) were published (Vol.88, No.4) was actually published on January 28, 1970, not in 1969 as was the rest of the volume; the species are therefore T. aculeatus (Cook, 1970) and T. dukei (Cook, 1970). (ref. ID; 7203)

    Etymology

    Named after Dr. Eugene N. Kozloff, who has worked with the marine invertebrate fauna of the Pacific coast for many years. (ref. ID; 7203)

    Habitat

    Intertidal muddy marine sands with some freshwater influence. (ref. ID; 7203)

    Type locality

    Just north of the Friday Harbour Marine Laboratory, San Juan Island, Washington; on Point Caution nearly opposite to Pt. George, Shaw Island. 48 degrees 33.6'N, 123 degrees 00.7'W. In a wide crevice in rock at the high tide line, small freshwater trickle present, very muddy sand between rock chips, sand under two inches of dry rock chips; approximately one foot above the top of the Fucus zone. Collected July 15, 1982 by H.R. Baker. (ref. ID; 7203)

    Type materials

  • Holotype: USNM (United States National Museum, Smithsonian Institution) 079833, a mature, whole-mounted specimen. (ref. ID; 7203)
  • Paratypes: USNM079834-079835, two whole-mounted mature specimens from the type locality. NMCIC (National Museums of Canada, National Museum of Natural Sciences Invertebrate Collection) 1982-1126, -1127, two whole-mounted mature specimens from the type locality. (ref. ID; 7203)
  • Other material examined:

    Tubificoides longipenis (Brinkhurst, 1965) (ref. ID; 7248), (Brinkhurst, 1965) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 7203)

    Synonym

    Tubifex longipenis Brinkhurst, 1965 (ref. ID; 5945, 7248); Tubifex longipenis Cook, 1971 (ref. ID; 7248)

    Descriptions

    Worms up to 30 mm and 108 segments. Body wall presumed naked (not cited in literature descriptions, but species formerly classified as a Tubifex). Dorsal and ventral setae similar in number, size, and shape, anteriorly 2-4 broad bifid setae with the upper teeth shorter than the lower, from XII only a single simple-pointed seta per "bundle". Spermathecal and male pores in the line of the ventral setae. Vasa deferentia up to 23 µm wide, very long and coiled, atria up to 510 µm long. Penis sheaths; 320-450 µm long by 34-73 µm wide, with a cuticular hook some 30 µm long near the distal end. (ref. ID; 7248)

    Remarks

    The species was described from a single specimen with highly characteristic penis sheaths from Georgetown, Maine. Cook (1971) redescribed the species from additional specimens from Cape Cod Bay. A brief abstract of the description follows: 25-30 mm. Two to four broad bifid setae anteriorly, upper teeth shorter than the lower, from XII a single simple-pointed seta. Penes thickly cuticularized, each with a hook near its distal end. Non-papillate body wall. (ref. ID; 5945)

    This species has elongate penis sheath, but in this instance there are only 2-4 anterior bifid setae and the single posterior setae are simple pointed. The species is known from Maine and Massachussetts. No new material is to hand. (ref. ID; 7248)

    Tubificoides maritimus (Hrabe, 1973) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 7203)

    Synonym

    Tubifex maritimus Hrabe, 1973 (ref. ID; 5945)

    Remarks

    The type specimens of this species lack the anterior end, but as there are hair setae and bifids from XV it is possible to assume that they may be present anteriorly, although postcapillatus has hair setae from XIII on that are missing anteriorly. The penis sheaths in this Black Sea species are cylindrical, and the illustration of the male duct places the species in this genus. Less certain is the position of Peloscolex debilis Finogenova, 1972, which has short naked hair setae but pectinate setae anteriorly and has no cuticular penis sheaths. It may be assigned to this genus once a clear illustration of the form to the atrium is available. (ref. ID; 5945)

    Tubificoides maureri Brinkhurst & Baker, 1979 (ref. ID; 5945 original paper)

    See

    Tubificoides diazi (ref. ID; 7248)

    Descriptions

    Length to 20 mm; breadth 0.3 mm; 31 segments. Prostomium conical, shorter than broad at the base Body wall naked. Clitellum 1/2 XI-XII. Segments elongating from VII to ~250 µm at XXII and shortening posteriorly. Setae bifid, four to five per bundle anteriorly, one or two posteriorly, no setae on XI. Setal line two-thirds of the way from the anterior broader of each segment. Nodulus distal. Upper teeth of anterior setae thinner and shorter than lower, posteriorly lower teeth broaden. Setae increase in length to V (36-41 µm) after which the length is constant. Atria elongate cylinders (120-240 µm long) widening slightly proximally. Vasa deferentia three to six times length of atira, 17 µm diameter, enter atria subapically opposite large stalked prostates. Atria terminate in penes with elongate, slightly curved cylindrical sheaths with laterodistal openings. Spermathecae elongate with median broadening (200-220 µm long, 55 µm wide in the middle) with short ducts leading to spermathecal pores. Spermathecal and penial pores ventrolateral, in setal line. This species also has elongate penis sheaths, but they are not so elaborate as those in the preceding species. The vasa deferentia area relatively elongate. (ref. ID; 5945)

    Etymology

    For Don Maurer, student of the benthos at the College of Marine Sciences, Delaware. (ref. ID; 5945)

    Tubificoides motei Brinkhurst, 1986 (ref. ID; 7248 original paper)

    Descriptions

    Length 5-6 mm, width 0.25 mm (preserved and whole-mounted), up to 40 segments. Body wall naked anteriorly, grooved and ridged from just behind the clitellum posteriad, segments quite clearly elongate, becoming so quite rapidly between XV and XVII, but becoming shorter again posteriorly. Anterior setal bundles with 2-4, mostly 3, setae per bundle, both teeth quite short, but the upper usually shorter and thinner than the lower. Posteriorly 2, rarely 3, rather similar setae per bundle. Spermathecal pores lateral, male pores ventrolateral in line with the ventral setae. Spermathecae with distinct ducts entering the ampullae via sperm traps, the ampullae spherical to ovoid. Male ducts each with a vas deferens about as long as the atrium, 10 µm wide, atria 190 by 30 µm. Penis sheaths in this species are broadly conical, tapering to a narrow distal end, possibly with an oblique opening (50x26 µm). (ref. ID; 7248)

    Remarks

    The number of posterior setae, the form of the penis sheath and atrium, and the distinctly modified body wall of the posterior segments serve to distinguish this species from T. brownae. The Sechelt Inlet specimens do appear to be the same species, even those the zoogeographic distribution indicated by the inclusion of this material in the same species is at first surprising. Comparison with the distribution patterns reported elsewhere in this study suggests that many species are quite widely distributed, however. C. Erseus (personal communication) agrees that the Sechelt specimens do conform with the rest. (ref. ID; 7248)

    Type locality

    Southwest Florida continental shelf at 26 degrees 16.47'N, 82 degrees 38.21'W, November 1980, 26 m, coll. for the Mote Marine Laboratory by M.R. Milligan. (ref. ID; 7248)

    Type materials

  • Holotype: USNM98843, a whole-mounted specimen. (ref. ID; 7248)
  • Paratypes: USNM98844-98851, 14 whole-mounted specimens on 8 slides; More Marine Laboratory collection, 12 whole-mounted specimens on 5 slides; Brinkhurst collection, 10 specimens on 5 slides, 24 degrees 47.49' to 26 degrees 16.47' N, 82 degrees 13.14' to 83 degrees 12.49'W, November 1980. (ref. ID; 7248)
  • Other material examined:

    Tubificoides nerthoides (Brinkhurst, 1965) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 7203)

    Synonym

    Peloscolex gabriellae nerthoides Brinkhurst, 1965 (ref. ID; 5945)

    Remarks

    The penis sheaths of this species can now be illustrated properly, and prove to bear at thickened flange on one side. The setae are illustrated here as they have not been shown in detal elsewhere. (ref. ID; 5945)

    Tubificoides palacoleus Baker, 1983 (ref. ID; 7203 original paper)

    Descriptions

    Length to 6 mm, at least 34 segments (all specimens incomplete), width at XI 340 µm (in fixed, slightly compressed, nearly mature specimen). Prostomium bluntly conical, as long as or slightly shorter than broad at peristomium. Body wall naked. Clitellum not observed. Anterior dorsal setal bundles (to VIII-X) with (1) 2-3 bifid setae and (1) 2-3 straight hair setae; bifid setae with teeth of approximately equal length, teeth almost parallel, hair setae smooth. From VIII to X posteriad dorsal setal bundles with 1 hair seta nd 1 sharply simple-pointed seta. Anterior ventral setal bundles with 2-3 bifid seta, upper tooth shorter than lower, upper tooth reduced posteriorly; from IX to X posteriorly 1-2 sharply simple-pointed setae per bundle. Ventral setae absent in XI, modified genital setae absent. Spermathecal pores paired in X in ventral setal line just anterior to the ventral setae of X; male pores paired medially in XI, in ventral setal line. (ref. ID; 7203)
  • Male genitalia: sperm funnel small, vas deferens 18-21 µm wide with ciliated lumen, approximately 800 µm long; joins atrium subapically below swollen atrial head opposite to entrance of stalked prostate gland. Atrium cylindrical, histologically tripartite, with enlarged ental head and ectal penial bulb, external muscle layer 1.0-1.5 µm thick over atrium. Atrium (to base of penis sheath) approximately 410 µm long, main body of atrium 40-43 µm wide, narrowing slightly basally to join penial bulb; swollen atrial head approximately 70 µm wide. Atrium terminates in true penis with penis sheath, penial apparatus enclosed in short, muscular, penial sac. Penis sheath short and broad, narrowing before the flared tip, 63-68 µm long, 45-47 µm wide at base, median narrowing 27-30 µm wide, flared tip 45-50 µm wide, narrowing apically. Spermathecae paired, each with bulbous base (approximately 65-75 µm high, 50-60 µm wide) and narrow duct (approximately 22 µm wide, 28 µm long) leading to ampulla; sperm trap apparently absent. Spermatozeugmata not observed. (ref. ID; 7203)

    Remarks

    Although only three nearly mature (precopulatory) specimens T. palacoleus were available to describe the genitalia, the form of the atria and penis sheaths, combined with the distinctive setation of all eight specimens, allow this taxon to be described. The setation of T. palacoleus resembles that of T. cuspisetosus, and to a lesser degree, that of T. swirencowi and T. amplivasatus but the number of setae, and the segment where the bifid setae turn simple-pointed is quite different in these species (see Remarks under T. cuspisetosus). The shape of the penis sheath of T. palacoleus is also obviously different to that of T. cuspisetosus. Tubificoides wasselli, a papillate species with only bifid setae, also has a somewhat shovel-shaped penis sheath but the apical flare is not nearly as developed. (ref. ID; 7203)

    Etymology

    From the Latin pala, shovel-shaped and the Latin coleus, sheath or scabbard; with a shovel-shaped (penis) sheath. (ref. ID; 7203)

    Habitat

    Subtidal at 48 m. (ref. ID; 7203)

    Type locality

    Frobisher Bay, North West Territories. 63 degrees 18.6'N, 68 degrees 06.2'W. 48 m, collected July 28, 1967 by Dr. J.W. Wacasey. (ref. ID; 7203)

    Type materials

  • Holotype: NMCIC (National Museums of Canada, National Museum of Natural Sciences Invertebrate Collection) 1982-1135, a whole-mounted nearly mature (precopulatory) specimen. (ref. ID; 7203)
  • Paratypes: NMCIC1982-1136 to -1140, one dissected and six whole-mounted precopulatory specimens from the type locality. (ref. ID; 7203)

    Tubificoides parapectinatus Brinkhurst, 1985 (ref. ID; 7240 original paper)

    Synonym

    Peloscolex apectinatus Brinkhurst and Simmons, 1968 (ref. ID; 7240); Peloscolex apectinatus Cook and Brinkhurst, 1975 (ref. ID; 7240)

    Descriptions

    Worms up to 20 mm long, 75 segments. Posterior segments markedly elongate. Body wall with a thin covering of foregin matter from XVI, and ridged. Anterior dorsal setal bundles with 2-3 hair setae and 2-4 bifid setae, the latter with fairly long, divergent teeth, from XII the bifid setae are replaced by simple pointed setae, and while the number remains initially at 2-3 of each it falls posteriorly to one of each. Anterior ventral setae 2-4 or up to 7 per bundle, with broad subequal teeth, posteriorly reducing to 2-3 and eventually to 1 in which the upper tooth becomes distinctly longer and thinner than the lower. Spermathecae with rounded vestibule behind the pore, leading to a narrow duct that enters a spherical ampulla basally, no obvious sperm trap. Male duct with vas deferens (width 30 µm) twice the length of the atria (0.9 versus 0.45 mm in one specimen), atrium moderately long terminating in a bluntly conical penis with thin distal end which protrudes well beyond the collarlike basal penis sheath when everted. Spermatozeugmata broad. (ref. ID; 7240)

    Remarks

    This species was originally identified as T. apectinatus but the detailed form the setae and penis sheaths differ. (ref. ID; 7240)

    Habitat

    Shallow estuarine situations. (ref. ID; 7240)

    Distribution

    Pacific coast species. Victoria, British Columbia and San Francisco Bay, California. (ref. ID; 7240)

    Examined materials

  • Holotype: United States National Museum (USNM) 097301, a whole-mounted specimen, The Gorge, Victoria Harbour, British Columbia, June 7, 1982, coll. K. Sendall and G. Hawkins. (ref. ID; 7240)
  • Paratypes: USNM097302-5, two whole mounted specimens on one slide; R.O. Brinkhurst collection. 3 dissected specimens, all from the type locality. (ref. ID; 7240)
  • Other: R.O. Brinkhurst collection, 4 whole-mounted mature specimens, several immature specimens, San Francisco Bay (Brinkhurst and Simmons 1968). (ref. ID; 7240)

    Tubificoides pseudapectinatus Brinkhurst, 1985 (ref. ID; 6602, 7240 original paper)

    Synonym

    Tubificoides apectinatus (Br.) (in part). Cook, 1969, 1971 (ref. ID; 7240)

    Descriptions

    Worms up to 20 mm long, up to 52 segments. Body wall very little modified, slightly furrowed from V and with very little foreign material. Anterior dorsal setal bundles with 3 bifid setae with the teeth quite long in II but rapidly changing to setae with the upper teeth shorter and thinner than the lower, teeth diverging, accompanied by 3 fine hair setae. Behind the clitellum the dorsal bundles have a single curved bifid seta with the upper tooth distinctly thinner and shorter than the lower, accompanied by exceedingly fine, single hair setae which are frequently missing (or broken off?). Anterior ventral setae 3 per bundle, with the upper tooth thinner but only slightly shorter than the upper (unless broken), posteriorly 2 per bundle, sigmoid, with thin, short upper teeth, ventral and dorsal setae essentially the same shape. Atria elongate, thin walled. Penis sheath simple thimble shaped, with subterminal opening. Spermatozeugmata broadly leaf shaped, spermathecae spherical with simple ducts. (ref. ID; 7240)

    Remarks

    This species is immediately separable from T. apectinatus in regard to both size and the postclitellar dorsal setae. The hair setae of posterior bundles are normal hair setae in T. apectinatus, but they are very short and thin or absent in T. pseudapectinatus. Also, in T. apectinatus the posterior ventral setae appear longer and straighter beyond the nodulus, and project further from the body wall than in the other species. The spermathecae of the two species clearly differ. The atria of T. pseudapectinatus are clearly elongate and rather thin walled, but further details need to be established from fresh material. The clearly tubificine nature of the setae, penis sheaths, and spermatozeugmata strongly suggest that this species be located in Tubificoides, the only predominantly salt water genus in the subfamily. (ref. ID; 7240)

    Habitat

    Marine species. (ref. ID; 6602)

    Atlantic coast species, shallow marine sites. (ref. ID; 7240)

    Distribution

    Cape Cod Bay. (ref. ID; 7240)

    Examined materials

  • Holotype: United States National Museum (USNM) 097306, a whole-mounted specimen, Cape Cod Bay, 1961-1962, station 1730 E3. (ref. ID; 7240)
  • Paratypes: USNM097307-10, many whole-mounted specimens from the type locality. (ref. ID; 7240)
  • Other material available: Cook collection. Specimens in fluid mixed with other species, Gray Museum, Marine Biological Laboratory, Wood Hole, Massachusetts. (ref. ID; 7240)

    Tubificoides pseudogaster (Dahl, 1960) (ref. ID; 7248), (Dahl, 1960) n. comb. (ref. ID; 5945) reported author and year? (ref. ID; 6594, 7203)

    Synonym

    Isochaetides pseudogaster (Dahl) Hrabe, 1966 (ref. ID; 5945, 7248); Limnodrilus pseudogaster Dahl, 1960 (ref. ID; 5945, 7248); Tubifex pseudogaster (Dahl) Brinkhurst, 1962 (ref. ID; 5945, 7248)

    Descriptions

    Marine species. (ref. ID; 6594)

    Worms up to 14 m long, 50 or more segments. Body wall naked, or with fine coating of foreign matter. Segments elongate posteriorly, with glandular rings in the setal line two-thirds the way along the segment. Anterior setal bundles with 3-5 (6) setae per bundle, bifid with the upper teeth as long as or somewhat longer than the lower, posteriorly (1) 2 (3) per bundle with the upper teeth thinner and longer than the lower. Spermathecal pores lateral, male pores ventrolateral in the line of the ventral setae. Spermathecal ducts short and broad, entering pear-shaped ampulla subterminally through sperm traps. Dimensions for the male duct are given as extreme range followed by mean values and standard error of the means of 38 specimens: atrium length 250-430 µm (289+/-7); width below head 36-85 µm (55+/-2); vas length 510-1390 µm (838+/-46); vas width 14.3-27.1 µm (19.9+/-0.5); penis length 47-94 µm (72+/-1.7); thickness of atrial muscles 1.6-4.3 µm (2.5+/-0.1). This considerable variation observed within the set of 38 specimens can be readily expanded by reference of more material as evidenced by the earlier published values for the length of the penis sheath. The penis sheath is a simple cylinder. (ref. ID; 7248)

    Remarks

    The original material of this species was redescribed by Baker (1979). The material consisted of unmated specimens with partially developed spermathecae. Examination of material from a wide variety of locations makes is possible to state that the spermathecal pores are lateral in position, that they enter short, broad ducts that narrow before they open into extensive pear-shaped ampullae via the so-called traps, and that the spermatozeugmata vary from a somewhat vermiform to a tadpole-like form. The spermathecae may be enlarged to occupy parts of segments IX-XI. While it is important to establish the form of these structures for the species, the result is disappointing in that all other members of the genus that share the same setal form and lack of papillation have spermathecae of this pattern, though the position of the pores may differ. None of the species in this group appear to have the more spherical form of spermatheca exhibited by some of those species in the genus with hair setae (Brinkhurst 1985). In describing T. coatesae, Brinkhurst and Baker (1979) introduced a new character, the distinctive elongate posterior segments of that species, with the setae set well towards the posterior end of the segment. The setae are associated with a ring of supposed glandular tissue. Careful examination of European material of T. pseudogaster reveals the presence of elongate segments in that species too. Only those segments fixed in the relaxd state show this characteristic, of course, and the glandular rings are only clearly visible in a few preparations that have taken up the stain in a particular way. This character appeared to be so distinctive for T. coatesae because none of the British Columbia specimens of T. pseudogaster in the collection show this, though they appear to resemble the European members of the species in every other respect. The setae vary somewhat in shape in the various sets of material. The upper tooth is always thinner than the lower, and it usually becomes longer relative to the lower tooth posteriorly, but the relative lengths of the upper and lower teeth do vary somewhat. Considerable intraspecific variation in setal form in species with bifid setae was clearly established long ago by Kennedy (1969) in Limnodrilus, another genus in this subfamily. The number of setae is more consistent, varying from 3 to 5 or 6 anteriorly, and reduced to (1) 2 (3) in posterior bundles. The body wall is usually naked in these worms, but these may be a fine coating of foreign matter adherent the body wall, usually posteriorly but sometimes anteriorly (as in specimens from the Clyde River, Scotland). The atria are quite consistently short and broad, frequently curved over the penis sacs. The vasa deferentia, where visible, are clearly over twice the length of the atria and are usually half the width of the atrium at the narrow point below the union with the vas. Length of the penis shows variation due to the degree of maturation, unmated specimens having smaller male ducts than mated specimens. Squashing material that has been dissected out in polyvinyl lactophenol appears to increase the apparent dimensions of both atria and penis sheaths in some, but not all, such material. The form of the penis sheath a traditionally significant freature of tubifid anatomy, seems now to be consistent. Most of the material in the Baker collection is identifiable as pseudogaster as suggested by the donors, but among the material from Hull, England and Oslo, Norway, were specimens of T. crenacoleus. In addition, I found one specimen of pseudogaster among the type series of crenacoleus from the Beaufort Sea. The penis sheaths of pseudogaster are cylindrical, the feature that caused the original author to classify this species in the genus Limnodrilus. The penes of crenacoleus have a distal expansion, those of diazi are curved and open laterally, and in other species the penes are conical to thimble shaped. Specimens from Eel Pind, Wood Hole, reported by Brinkhurst (1965) as probable, Tubifex pseudogaster, the first possible record of the species of North America, differ from this description of the species. There are only 2-3 setae per bundle anteriorly, dropping to single setae from VIII-IX; the anterior setae have the upper tooth markedly shorter than the lower. The penis sheaths are shorter than those of pseudogaster (40-47 µm) and are somewhat conical. As this is older material mounted in polyvinyl lactophenol, other details are unavailable for study, but the specimen may belong to T. brownae (q.v.). (ref. ID; 7248)

    Material examined

    The type series from Denmark, together with material from the Isle of Man, Grimsby, Scapa Flow, and the Forth and Clyde rivers, Great Britain; the Schleifjord, West Germany; Schelphook, Netherlands; Oslo Fjord, Norway; and Sweden. North American material from Friday Harbor, Washington State; Burrard Inlet and Victoria, British Columbia; and Beaufort Sea, Alaska. (ref. ID; 7248)

    Tubificoides scoticus Brinkhurst, 1985 (ref. ID; 7240 original paper)

    Descriptions

    Specimens mostly incomplete, one 9 mm long, 50 segments. Median segments elongate. Body wall variable from naked to coated in foreign matter or slightly papillate posteriorly. Dorsal bundles anteriorly with hair and bifid setae, 2-3 (4) of each, the bifids with broad lance-shaped teeth, the upper shorter than the lower, sometimes bifurcate; from VIII the same number of hairs and blunt simple pointed setae. Anterior ventral setae 2-5 per bundle, bifid like the dorsals, posteriorly 3-4 per bundle, probably simple pointed but setae unbroken. Vas deferens length not established, but apparently about 2-3 times as long as the atrium. Atrium moderately long, terminating in an elongate penis in along cylindrical sheath. The spermatheca has a fairly elongate duct, distinct sperm traps and ovoid ampulla. The spermatozeugmata is quite elongate, broadest medially. (ref. ID; 7240)

    Remarks

    The setae of this species are unique, the closest parallel being those of T. nerthoides, which are pectinate. The other species with elongate, unornamented penes like those of the new species is T. cuspisetosus Baker in which the setae differ in shape except for the presence of simple pointed posterior ventral setae, and the spermatheca is elongate without a sperm trap. (ref. ID; 7240)

    Habitat and distribution

    Atlantic coast species. Sandy mud in shallow estuarine habitat, Kinneil, Firth of Forth. (ref. ID; 7240)

    Examined materials

  • Holotype: British Museum of Natural History (BMNH) 1984.15.1, a whole-mounted specimen, Firth of Forth, Kinneil, 10 m sandy mud, coll. B. Dicks, 1979, (56 degrees 2'23"N, 3 degrees 37'13"W). (ref. ID; 7240)
  • Paratypes: BMNH1984.15.2-11, 10 whole-mounted specimens from the type locality. R.O. Brinkhurst collection 18 whole-mounted specimens from the type location. (ref. ID; 7240)

    Tubificoides swirencoides Brinkhurst, 1985 (ref. ID; 7240 original paper)

    Descriptions

    Worms up to 10 mm long, up to 75 segments. Body wall closely papillate from VII or at least behind the clitellum, small posterior naked projection of undeveloped, narrow segments present. Dorsal setae (1) 2-3 (4) hairs and bifid per bundle, the bifids with two closely applied, usually short teeth, those of V-VII may bear short supernumary teeth above the upper teeth; from X on 1-2 hairs and 1-2 simple pointed setae per bundle. Anterior ventral setae 2-3 per bundle, at first with short, broad, teeth, the upper slightly longer than the lower, by VI the upper teeth clearly longer and thinner than the lower, posteriorly 1-2 per bundle with upper teeth thinner than but only a little longer than the lower, setae quite long and straight beyond the nodulus. Male ducts with quite short atrium, with very wide vas deferens at least 2-3 times the length of the atrium, penis sheath mostly twice as long as broad, cylindrical. Spermatheca with short, broad duct, somewhat special ampulla twice the length of the duct. Spermatozeugmata elongate, slender. (ref. ID; 7240)

    Remarks

    This species is similar to T. swirencoi and T. intermedius, but differs from the former in having longer penis, wider vas deferens, longer vas deferens (that of swirenoci being 1-1.5 times the atrium length, not 2.5 as in Cook (1969)). The penis sheath in T. intermedius is longer than that of the new species and the vas deferens is again narrower (15 µm versus 3-0 µm) and the body is less papillate in intermedius. There is some sign of a glandular thickening of the body wall around the spermathecae and male pores as reported in T. swirencoi by Hrabe (1964). (ref. ID; 7240)

    Habitat and distribution

    Atrantic coast species. Anoxic mud in the River Humber, around Grimsby, intertidal to a depth of 1-5 m. (ref. ID; 7240)

    Examined materials

  • Holotype: British Museum of Natural History (BMNH) 1984.15.12, a whole-mounted specimen, R. Humber, Immingham, South Killingholme Haven, intertidal, 53 degrees 35'12"N, 0 degrees 2'48"W, England, coll. B. Barnett, 16 June, 1981 (ref. ID; 7240)
  • Paratypes: BMNH1984.15.13-21, 9 whole-mounted specimens from the type locality. R.O. Brinkhurst collection. 15 whole-mounted specimens from the type locality; plus Grimsby Roads, 19 November 1979, same map reference. (ref. ID; 7240)
  • Other: Available from A.W.A., Marine Laboratory, Elsham Wolds. (ref. ID; 7240)

    Tubificoides swirencowi Jaroschenko, 1948 (ref. ID; 5945, 6618) reported author and year? (ref. ID; 7203)

    Descriptions

    Marine species. (ref. ID; 6618)

    Tubificoides vestibulatus Erseus & Bonomi, 1987 (ref. ID; 6618)

    Descriptions

    Marine species. (ref. ID; 6618)

    Tubificoides wasselli Brinkhurst & Baker, 1979 (ref. ID; 5945 original paper, 7248)

    Synonym

    Tubificoides foliatus Baker, 1983 (ref. ID; 7248)

    Descriptions

    Length 8 mm; breadth 0.3 mm; 51 segments maximum. Body wall closely papillate. Bifid setae two per bundle anteriorly, upper tooth thinner than but as long as or shorter than the lower, nodulus median. Posteriorly setae single, teeth about the same size. Setae increase in length to VI (55 µm) after which length remains fairly constant. Atria long (300-380 µm) slightly curved cylinders with enlarged proximal end 1.8-2.0 times the diameter of the distal part. Atrial walls 4-8 µm thick. Vasa deferentia elongate, thin (3-5 times length of atria, 19-22 µm wide) joining atria subapically opposite the entry of the prostate glands. Atria terminate in penes with cuticular penis sheaths elongated thimble shaped, about twice as long as breadth at the base with distal opening lateral. Spermatheca elongate with long ducts (600-680 µm long total), sperm traps present. Spermatozeugmata elongate. The form of the penis sheath and the number and form of the setae are characteristic. The spermathecal ducts join the narrowest part of the ampullae just below their tips in what we term a sperm trap. (ref. ID; 5945)

    Length to 11 mm, breadth to 0.3 mm, up to 50 segments. Prostomium and I retractile, body wall closely papillate, especially behind the clitellum. Setae 2, rarely 3 anteriorly, with upper teeth shorter and thinner than the lower, from between VI and VIII reduced to 1 seta with thick, recurved lower tooth. Spermathecal and male pores in place of the missing ventral setae of X and XI, respectively. Atria appear relatively elongate, sometimes extending in the sperm sacs to XII, up to 380 µm long, vasa deferentia up to 20 µm thick, 2-3 (5) times as long as atria. Penes with penis sheaths conical, lateral opening, 2-3 times longer than broad (up to 50x30 µm). Spermathecae with large ovoid ampullae, spermathecal ducts distinct, wide at pore then narrowing of enter pear-shaped ampullae subapically via sperm traps. Spermatozeugmata vary from broad to elongate. (ref. ID; 7248)

    Remarks

    The type series of this species was restricted to 8 specimens, all dissected or whole mounted in polyvinyl lactophenol. Despite that, the original description is fairly complete. Reexamination of four of these species suggests that prostomium is retractible and shows that the posterior segments may be elongate. Rings of enlarged sensory papillae protrude through the papillate body wall just in front of the setal line in two of the four specimens. The difference between these specimens is probably due to the state of these animals when killed, as the sensory papillae only ever seem to be visible in a proportion of specimens of species with papillate body walls. These sensory structures might be expected to be retractile. When everted, they do create wide nodes where the body width is greater than elsewhere. The setae enlarge gradually between VI and VIII, where a single seta replaces the usually paired anterior setae. Some posterior setae are quite sigmoid, with short proximal ends and large lower teeth. In one specimen the vas deferens is only 12 µm wide, and is 3 times the length of the atrium which, at about 400 µm (stated to be 300-380 µm for the species in the original account) excluding the penis sac, seems to be stretched by the dissection process. The range observed in other material is 220-280 µm. The original description cites the vas deferens as being 19-22 µm wide; it is up to 20 µm wide in Canada balsam mounted material. The penis sheaths are clearly visible in only one of the paratypes examined, and these are squashed and enlarged in a way that seems typical for material mounted in polyvinyl lactophenol. The retractile anterior end is confirmed in other specimens. The description of T. foliatus suggests few differences between it and T. wasselli. This west coast species included the papillate specimens of P. gabriellae sensu Brinkhurst reported from British Columbia and California, as noted in the original description. (The nonpapillate forms are attributable to T. coatease, now synonymised with T. brownae as noted above, plus the other nonpapillate Tubificoides species reported here for the northeast Pacific coast). In describing T. foliatus, Baker (1983) stated that "details of the male genitalia and shape of the penis sheaths serve to separate T. foliatus from T. wasselli". The illustration shows that penis sheaths to be quite similar in shape, in fact, and the table lists the sizes of the male duct of foliatus to be just about half those for wasselli for all features apart from the width of the vas deferens. Two of the three specimens from Cape Hatteras have atria and penes in the range measured for T. foliatus. The specimens do have strongly curved, short posterior setae with very large lower teeth but are otherwise similar to specimens of T. foliatus. The specimens from the Gulf coast of Florida have similar atrial and penial dimensions, but one has atria 280 µm long (with a penis sheath of 48x30 µm), which is intermediate in length between those of wasselli and foliatus, as is the penis sheath on another (51x39 µm). In some of these the atria enter the sperm sacs in XII. The relatively large size of the penis sheaths from the dissected and flattened specimen in polyvinyl lactophenol from the type series is considered to be an artifact, as in the type series of T. brownae. These two species were described from material collected and mounted under classroom conditions, and they were not treated with the same care as subsequent material mounted in Canada balsam. The specimens from the Weeki Wachi River, Florida, had been mounted in Ammans lactophenol before being remounted in Canada balsam, and few details could be discerned. They do conform to the general description of T. wasselli. The posterior chaetae have very large lower teeth but they are quite long and usually straight distally. The species is now known from Delaware to Florida, British Columbia to California. (ref. ID; 7248)

    Etymology

    For Jim Wassell who collected the material. (ref. ID; 5945)

    Material examined

    Type series, 4 specimens on 3 slides, Indian River Bay, Delaware, National Museum of Canada, Museum of Natural Sciences 979-0032; Brinkhurst collection, 20 specimens on 10 slides, The Gorge, Victoria, British Columbia, May 1982, coll. K. Sendall; 2 specimens, The Groge, Victoria, 1978, coll. W. Carolsfeld; 25 specimens Boundary Bay, Vancouver, 15 November 1984, 1.5-11 m, coll. D.F. Moore; 4 specimens on 2 slides, Point Richmond, California, 15 July 1957, coll. M.L. Jones; 10 specimens on 10 slides, San Francisco ay, Suisun Bay, California, 1961-1962, coll. R.A. Wagner (as "papillate P. gabriellae" in Brinkhurst and Simmons 1968); 10 specimens on 1 slide, James River, Virginia, 1983, coll. R.J. Diaz. Erseus collection, 3 specimens on 3 slides, off Cape Hatteras, North Carolina, 35 degrees 00.5' to 35 degrees 01.1'N, 75 degrees 33.5' to 75 degrees 34.0'W, November 1977, 33-40 m, coll. D. Weston; 4 specimens, Biscayne Bay, Miami, Florida, 25 degrees 35.6'N, 80 degrees 14.2'W, November 1977, 2-3 m; 9 specimens, Hutchinson Island, Florida, 27 degrees 20.8' to 27 degrees 22.6'N, 80 degrees 12.9' to 80 degrees 14.1'W, 1973, 6-11.5 m. Mote Marine Laboratory collection, 8 specimens on 7 slides, offshore from Charlotte Harbor, Florida to the Dry Tortugas, 24 degrees 47.7'N, 83 degrees 13.0'W to 26 degrees 45.50'N, 82 degrees 45.18'W at depths between 25 and 60 m, November 1980, May 1981, and Weeki Wachi River, Florida, August 1984, coll. M. Milligan. (ref. ID; 7248)