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The World of Protozoa, Rotifera, Nematoda and Oligochaeta

Phallodrilus

Phallodrilus Pierantoni, 1902 (ref. ID; 5912, 6559, 6913)

Family Tubificidae (ref. ID; 5890, 5939): Subfamily Phallodrilinae Brinkhurst (ref. ID; 5945)

ref. ID; 5912

Phallodrilus is a predominately marine genus with nine species, only two of which are found in freshwater: P. hallae n. sp. and P. aquaedulcis Hrabe, 1960, which is know from phreatic waters in northwestern Germany (River Weser drainage system). (ref. ID; 5912)

Definition

Hair setae absent. Ventral setae of segment XI replaced by penial setae. Male pores paired near the penial setae. Spermathecal pores paired, placed laterally, or near the ventral setae, or in between. Vasa deferentia as long as, or longer than, the pear-shaped to cylindrical atria. Each atrium bears two discrete, petiolate prostate glands, one of which enters near the junction of the atrium and vas deferens. True penes absent. Spermatozeugmata rarely developed. Coelomocytes sparse to absent. (ref. ID; 5912)

Remarks

Cook (1969) included Aktedrilus monospermathecus Knollner, 1935 in Phallodrilus on the basis of their common possession of two prostate glands to each atrium; other characters of A. monospermathecus such as the unpaired mid-dorsal spermatheca, absence of penial setae, and presence of true penes, were considered to be significant only at the species level. Hrabe (1973), however, is of the opinion that the presence or absence of ture penes is a valid generic criterion and therefore he excludes A. monospermathecus from Phallodrilus. Furthermore, he described a new genus and species, Bacescuella pontica Hrabe, 1973, which also has two prostate glands to each atrium, possesses true penes, but has no (or only very short) vasa deferentia, and no spermathecae. Thus in this complex of small marine species Hrabe (1973) recognizes the two monospecific genera Aktedrilus and Bacescuella, and Phallodrilus with seven species. In the present account, Hrabe's restricted conception of Phallodrilus is adopted because the genus, now with nine species, forms a cohesive, easily defined taxon. Because some tubuficid genera exhibit various degrees of development of penes and penial setae, and because it seem unlikely that the dual prostate condition arose more than once within the family, we argue that Aktedrilus and Bacescuella are closely related to Phallodrilus; also, intermediate conditions may exist whose description would necessitate critical reexamination of the generic limits discussed above. (ref. ID; 5912)

Type species

Phallodrilus parthenopaeus Pierantoni, 1902 (ref. ID; 5912)

ref. ID; 6559

The genus Phallodrilus Pierantoni, 1902 is predominantly a marine taxon comprising about 70 species (Erseus 1990). The genus was first established by Pierantoni (1902) for a sublittoral marine tubificid oligochaete, P. parthenopaeus, from the Bay of Naples in Italy. It was not until 60 years later that another species of Phallodrilus, the meiobenthic P. aquaedulcis, was described by Hrabe (1960) from the River Weser in Germany. (ref. ID; 6559)
  1. Phallodrilus adriaticus Hrabe, 1971 (ref. ID; 5912)
  2. Phallodrilus albidus (ref. ID; 5882)
  3. Phallodrilus aquaedulcis Hrabe, 1960 (ref. ID; 5890, 5912, 6559)
  4. Phallodrilus biprostatus (Erseus and Baker, 1979) (ref. ID; 6602)
  5. Phallodrilus boeschi Erseus, 1984 (ref. ID; 6602)
  6. Phallodrilus caudatus (ref. ID; 5882)
  7. Phallodrilus christeri Davis, 1985 (ref. ID; 5890)
  8. Phallodrilus clavatus (ref. ID; 5882)
  9. Phallodrilus coeloprostatus Cook, 1969 (ref. ID; 5912, 6602)
  10. Phallodrilus cristolatus Erseus, 1983 (ref. ID; 5890)
  11. Phallodrilus crypticus (ref. ID; 5890 original paper, 6559)
  12. Phallodrilus cuspis Erseus & Dumnicka, 1988 (ref. ID; 6559) reported year? (ref. ID; 5890)
  13. Phallodrilus davisi Erseus, 1984 (ref. ID; 5890)
  14. Phallodrilus exiguus Erseus, 1979 (ref. ID; 5890)
  15. Phallodrilus extremus (ref. ID; 5882)
  16. Phallodrilus filithecatus (ref. ID; 5882)
  17. Phallodrilus geniculatus (ref. ID; 5882)
  18. Phallodrilus georgei Erseus, 1987 (ref. ID; 5890)
  19. Phallodrilus hallae Cook & Hiltunen, 1975 (ref. ID; 5912 original paper, 6559) reported author and year? (ref. ID; 5939)
  20. Phallodrilus inconspicuus Erseus, 1979 (ref. ID; 5890)
  21. Phallodrilus labouichensis Rodriguez & Giani, 1989 (ref. ID; 5890 original paper, 6559)
  22. Phallodrilus leukodermatus Giere (ref. ID; 5864)
  23. Phallodrilus lobatus Ersues, 1988 (ref. ID; 5890)
  24. Phallodrilus macmasterae Erseus, 1986 (ref. ID; 5890)
  25. Phallodrilus minutus Hrabe, 1973 (ref. ID; 5912)
  26. Phallodrilus monnioti Erseus, 1987 (ref. ID; 5890)
  27. Phallodrilus nidarosiensis Erseus, 1987 (ref. ID; 5890)
  28. Phallodrilus obscurus Cook, 1969 (ref. ID; 5890, 5912, 6602)
  29. Phallodrilus parthenopaeus Pierantoni, 1902 (ref. ID; 5912, 6618)
  30. Phallodrilus parviatriatus Cook, 1971 (ref. ID; 5912, 6602)
  31. Phallodrilus postspermathecatus Erseus, 1980 (ref. ID; 5890)
  32. Phallodrilus profundus Cook, 1970 (ref. ID; 5912)
  33. Phallodrilus prostatus (Knollner, 1985) (ref. ID; 5890) reported author and year? (ref. ID; 5892)
  34. Phallodrilus rectisetosus (ref. ID; 5882)
  35. Phallodrilus riparius Giani & Martinez-Ansemil, 1981 (ref. ID; 5890, 6559)
  36. Phallodrilus sabulosus Erseus, 1979 (ref. ID; 6602) reported author and year? (ref. ID; 5882)
  37. Phallodrilus stilus Erseus, 1986 (ref. ID; 5890)
  38. Phallodrilus simplidentatus Erseus, 1979, (ref. ID; 5890)
  39. Phallodrilus subterraneus Rodriguez & Giani, 1989 (ref. ID; 5890 original paper, 6559)
  40. Phallodrilus tempestatis Baker, 1981 (ref. ID; 5960 original paper) reported author and year? (ref. ID; 5882)
  41. Phallodrilus tenuissimus (ref. ID; 5882)
  42. Phallodrilus vanus Erseus, 1984 (ref. ID; 5890)
  43. Phallodrilus vulnus Erseus, 1983 (ref. ID; 5890)

Phallodrilus adriaticus Hrabe, 1971 (ref. ID; 5912)

Descriptions

Marine species. Modified genital setae on segments X and XI, or XI only. All somatic setae bifurcate up to at least segment XVIII. Spermathecal epithelium of more or less uniform thickness. Up to seven penial setae per bundle as long as, or longer than, somatic setae. Spermathecal setae not modified. Penial setae two per bundle with sraight rounded ends. Atria erect with vas deferens junction median, near posterior prostae. Spermatozeugmata present. (ref. ID; 5912)

Phallodrilus albidus (ref. ID; 5882)

Descriptions

Marine species. (ref. ID; 5882)

Phallodrilus aquaedulcis Hrabe, 1960 (ref. ID; 5890, 5912, 6559)

Redescription (new material, Niagara River)

  • External features: Length 1.7-3.7 mm, mean 2.1 mm. Sixty percent of adult population 1.7-1.9 mm long. Width anteriorly (cleared, slightly compressed specimens) at segment II 104-156 µm, mean 128 µm. Width segment X, 156-229 µm, mean 190 µm, and width posteriorly 73-115 µm, mean 96 µm. Number of segments in mature individuals 29-48, mean 38. Prostomium rounded. Clitellum extending from 1/2 X anterior of chaetae to 1/2 XII posterior of chaetae. Somatic chaetae bifid, with upper tooh thinner and shorter than lower. Dorsal and ventral chaetae anteriorly 4-6 per bundle, normally 5, length 28-38 µm, mean 33 µm. Dorsal and ventral chaetae in postclitellar segments slightly shorter, with mean length 31 µm, usually 3 per bundle, with 2 per bundle in the terminal two to four segments. Penial chaetae thicker and distinct from somatic chaetae, bundles appear fanlike. Penial chaetae 5-7 per bundle, normally 6, longer than somatic chaetae, 28-40 µm, mean 36 µm. Width at thickest portion 1.5-2.5 µm. Distal end slightly deflected in some chaetae. Ventral chaetae of X not modified. Male pores located immediately lateral to penial chaetae, i.e., in line with ventral somatic chaetae, posteriorly in XI. Spermathecal pores in line with ventral somatic chaetae in most anterior part of X. (ref. ID; 6559)
  • Internal anatomy: Coelomocytes absent. Pharyngeal glands in IV-VI. Paired, coiled, very long nephridia present in VII, or VI-VII (at least?). Male genitalia paired. Vas deferens about 7-8 µm wide, appears somewhat shorter than atria, entering latter more or less apically. Atrium cylindrical, with 1-5 µm thick (i.e., highly variable) outer lining of muscles, and granulated and ciliated inner epithelium; ental end rounded, ectal end tapering and terminating in simple male pore on inconspicuous papilla. Atrium 60-90 µm long, 33-38 µm wide. Two roundish prostate glands present, anterior one attached to apical end of atrium, at junction with vas deferens, posterior one attached by short stalk to ectal end of atrium. Spermathecae consisting of short ducts, about as long (23-25 µm) as wide, and slender, somewhat club-shaped ampullae; ectal part of latter with distinctly thicker wall than ental part. Sperm mass (containing also secretion?) random, generally confined to inner, thin-walled part of spermathecal ampulla. (ref. ID; 6559)

    Remarks

    Only one specimen complete (24 segments). All post-copulatory. The specimen from cave Jivero-1 presents bifid penial setae, with upper tooth shorter and thinner than lower; the other two simple-pointed penial setae. 4-5 penial setae in the first specimen and 8 in the other two. Outer muscular lining of atria always very thin (less than 1 µm). (ref. ID; 5890)

    Modified genital setae on segments X and XI, or XI only. All somatic setae bifurcate up to at least segment XVIII. Spermathecal epithelium of more or less uniform thickness. Up to seven penial setae per bundle as long as, or longer than, somatic setae. Spermathecal setae not modified. Penial setae two to seven per bundle with hooked or curved ends. Aria procumbent, with vas deferens junction apical, near anterior prostate. Spermatozeugmata absent. Some anterior bundles with more than three setae. Three or more setae per bundle posteriorly, or, if only two per bundle, then some penial setae bifurcate. All dorsal setae bifurcate. Atria relatively small with very thin muscle walls. Porophores absent. Spermathecal pores in line with ventral setae. Four two six (rarely seven) unmodified (bifurcate) and simple-pointed penial setae present. Freshwater species. (ref. ID; 5912)

    Type locality

    Blumenthal, River Weser, Germany. (ref. ID; 6559)

    Type material

    Hrabe collection, examined by Erseus (1979). (ref. ID; 6559)

    New locality

    Tonawanda Channel, Niagara River, North America; sandy substrates, littoral zone to 17 m of water, fast-flowing water. (ref. ID; 6559)

    New material examined

    One specimen from Cave Mairulegorreta, at Zigoitia (Alava, Basque Country, Spain). 30T WN199 635, 880 m altitude. 08.VIII.1987. Fine to medium sand sediment. One specimen from Cave Jivero 1, at Ozana (Matienzo, Santander, Spain) 30T VN 537 948. 260 m altitude. 26.II.1984. Collection of Jos Notenboom, Zoologisch Museum of Amsterdam. One specimen from Cave of Labouiche, near Varnajoul (Foix, Ariege, France). 17.V.1985. Coarse sand sediment. Collected by V. Bouguenec & M. Gallissie. (ref. ID; 5890)

    New material: 2 stained, whole-mounted specimens in Canada balsam, 6 unstained, whole-mounted specimens in CMCP-10, and 1 sectioned specimen in Canada balsam deposited at the Royal Ontario Museum, Ontario, Canada (Catalogue Nos. ROMIZ I1433, ROMIZ I1435, and ROMIZ I1434, respectively); 20 whole-mounted specimens in CMCP-10 in the Farara collection and 10 whole-mounted specimens in CMCP-10 in the Erseus collection. (ref. ID; 6559)

    Phallodrilus biprostatus (Erseus and Baker, 1979) (ref. ID; 6602)

    Descriptions

    Marine species. (ref. ID; 6602)

    Phallodrilus boeschi Erseus, 1984 (ref. ID; 6602)

    Descriptions

    Marine species. (ref. ID; 6602)

    Phallodrilus caudatus (ref. ID; 5882)

    Descriptions

    Marine species. (ref. ID; 5882)

    Phallodrilus clavatus (ref. ID; 5882)

    Descriptions

    Marine species. (ref. ID; 5882)

    Phallodrilus coeloprostatus Cook, 1969 (ref. ID; 5912, 6602)

    Descriptions

    Modified genital setae on segments X and XI, or XI only. All somatic setae bifurcate up to at least segment XVIII. Spermathecal epithelium of more or less uniform thickness. Ten to 13 penial setae per bundle, shorter than somatic setae. Marine species. (ref. ID; 5912)

    Marine species. (ref. ID; 6602)

    Phallodrilus crypticus (ref. ID; 5890 original paper, 6559)

    Descriptions

    1 specimen not complete, posterior end missing (25 segments). Diameter at V (slightly compressed) 89.6 µm, at XI 106.4 µm. Non-prominent, inconspicuous clitellum. Prostomium conic, 34.0 µm long and 27.2 µm wide. Secondary annulations from II to VII, but never very distinct. Somatic setae bifid, with upper tooth shorter and thinner than lower. Dorsal setae present at XI. 3 (rarely 4) setae per bundle anteriorly, and 2 setae from IX backwards. Ventral setae of XI modified into penial setae, bifid 7-8 per bundle; 21.4-22.8 µm long and about 0.7 µm thick, fan-wise arranged; convergent towards the male pore. One pair of male pores in line with ventral setae at XI, located in the posterior third of segment, at the top of a projecting porophore. One pair spermathecal pores in most anterior part of X, in line of ventral setae. Pharyngeal glands present in IV-VI (scarcely over the posterior part of III). Male genitalia paired. Vas deferens shorter than atrium, about 42 µm long and 4 µm in diameter. Atrium spindle-shaped and curved, about 107 µm long and 26.4 µm of maximum diameter, with a very thick muscular layer (up to 7.14 µm), and inconspicuous epidermal layer. Atrial lumen is filled by spermatozoids. Ectally atrium constricts before entering atrial bulb which is a heavily muscular structure at the male pore, possibly acting as a pseudopenis. Sometimes atrial internal epithelium is projecting towards the exterior through the pore. Spermathecae elongate, 51-54.4 µm long and 17.9 µm maximum diameter. One ampulla into X, the other extending over IX. Spermathecal wall with distinct muscular outer lining (about 2.0 µm thick). Duck short, 21.4 µm long and 14.3 µm diameter. A sperma bundle fills the spermathecal lumen. (ref. ID; 5890)

    Remarks

    This species is characterized by fan-shaped bifid penial setae, thick muscular atria terminating in a bulb at the male pores and the asymmetrical position of spermathecal ampulla in IX and X. P. crypticus n. sp. is a member of the group of Phallodrilus species that possess numerous penial setae (not club-shaped), and spermathecal pores on the line of ventral setae. In this complex, only P. aquaedulcis Hrabe, 1960, P. obscurus Cook, 1969, P. riparius Giani & Martinez-Ansemil, 1981 and P. prostatus (Knollner, 1985) present the vas deferens shorter than the atrium, a character shared also by P. crypticus n. sp. This species is clearly distinguished from the precedent species by the combination of characters mentioned above. Bifid condition of penial setae is not very common, being known only in P. aquaedulcis, P. vulnus Erseus, 1983, P. cristolatus Erseus, 1983, P. vanus Erseus, 1984 and P. christeri Davis, 1985. P. crypticus n. sp. is also closely related to the recently described P. monnioti Erseus, 1987, from a subtidal locality in Corsica (France), that also shares an atrial bulb at the male pore, a very peculiar feature in the genus, and a very thick atrial musculature. They are, however, distinguished by the different position of the spermathecal pores (lateral in P. monnioti) and the shape and number of penial setae (simple-pointed and 15-20 per bundle in P. monnioti). The asymmetrical disposition of the two spermathecal ampullae was only known in P. tempestatis Baker, 1981. This species has been found in mid intertidal sediment, influenced by a small trickle of freshwater (Baker 1981). (ref. ID; 5890)

    Etymology

    The name crypticus (Latin 'crypta' for cave) refers to the habitat of this species. (ref. ID; 5890)

    Type locality

    Cave del Orillon at Ozana (Matienzo, Santander, Spain). Collected by Jos Notenboom, 26.II.1984. Coordinates 30TVN 543 947, 275 m altitude. (ref. ID; 5890)

    Material examined

    1 specimen, stained in haematoxylin, dissected and mounted in Canada balsam. Collection of Jos Notenboom, Zoologisch Museum of Amsterdam. (ref. ID; 5890)

    Phallodrilus cuspis Erseus & Dumnicka, 1988 (ref. ID; 6559) reported year? (ref. ID; 5890)

    Descriptions

    Freshwater species. (ref. ID; 6559)

    Phallodrilus extremus (ref. ID; 5882)

    Descriptions

    Marine species. (ref. ID; 5882)

    Phallodrilus filithecatus (ref. ID; 5882)

    Descriptions

    Marine species. (ref. ID; 5882)

    Phallodrilus geniculatus (ref. ID; 5882)

    Descriptions

    Marine species. (ref. ID; 5882)

    Phallodrilus hallae Cook & Hiltunen, 1975 (ref. ID; 5912 original paper, 6559) reported author and year? (ref. ID; 5939)

    Descriptions

    About 2.8 to 3.7 mm long; 0.115 to 0.170 mm diameter anteriorly, 0.15 to 0.18 mm at the clitellum, and 0.10 to 0.12 mm posteriorly. Twenty-eight to 34 segments (usually 33). Prostomium rounded, as long as, or somewhat shorter than, it is wide at the peristomium junction. Clitellum weakly developed around the spermathecal pores and on segments XI and 1/2 XII. Dorsal and ventral setae similar in number, size and shape up to segment XIX, each bundle containing three to five bifurcate setae, with the upper tooth shorter and thinner than the lower, 35 to 50 µm long. Between segments XX and XXII upper teeth of dorsal setae become progressively smaller and setal length increases; from segment XXIII to terminal segment, dorsal setae single-pointed, 50 to 65 µm long. Ventral setae bifurcate, similar to anterior setae, from segment XX to XXIX but simple-pointed on segment XXX to terminal segment. Ventral setae of segment XI modified into penial bundles each containing three to six single-pointed, distally hooked penial setae 60 to 65 µm long (small persentage of penials examined possessed a very small rounded elevation on the outside terminal curve that probably represents a rudimentary upper tooth). Paired spermathecal pores located on the lateral line of segment X, near intersegmental furrow IX/X. Paired male pores on summits of paired conical porophores located just lateral to, and in front of, the penial setae. Coelomocytes absent. Pharyngeal glands present in segment IV and V. Male genital system (all structures paired): small male funnel opens into a short vas deferens 4.5 to 7 µm diameter, which merges into the atrium. Each atrium spindle-shaped to cylindrical, 28 to 33 µm maximum external diameter, about 150 to 200 µm total length; long axis of atria directed anteriorly. Two prostate glands join each atrium, one with a long discrete prostatic duct joins posteriorly and laterally 35 to 50 µm from the external male pore, and one with indistinct ducts joins anteriorly and medially in the region of the vas deferens - atrial junction. Atrium consists of a narrow lumen, inner lining cells 5 to 7 µm thick, and a robust outer muscle layer (with a characteristics reticulated appearance in fixed specimens) 8 to 12 µm thick; muscle layer extends for some distance along the vas deferens and along the posterior prostatic duct. True penes absent: atrium terminates simply on the apex of a conical non-retractile porophore directed posteriorly. Penial setae lie more or less horizontally, median to the atrium and posterior prostatic duct, and terminate in a deep fold in the body wall located immediately behind the porophore; a set of enlarged setal retractor muscles apparently maintains this ofld, and causes the penial setae to lie parallel to the long axis of the body. Paired cylindrical spermatheca with short indistinct ducts open in the lateral line near the anterior margin of segment X: spermathecal ampulla 70 to 110 µm long, 23 µm maximum external diameter; duct about 18 µm long. Each spermatheca consists of an inner layer of lining cells (very thin at the distal end of the ampulla and thickened in the middle) and an outer muscle layer 3 to 6 µm thick. Sperm in spermathecae in random masses. (ref. ID; 5912)

    Etymology

    Species named for Trudy Hall, a patient and charming lady. (ref. ID; 5912)

    Type deposition

  • Holotype: Royal Ontario Museum, Toronto (ROM) I-42: Lake Superior 46 degrees 49'06"N, 85 degrees 57'30"W; depth 188 m; collected by B. Riddell, 1 June 1973. (ref. ID; 5912)
  • Paratypes ROM I-43: 47 degrees 04'48"N, 85 degrees 49'48"W; depth 98 m; coll. B. Riddell, 1 June 1973. ROM I-44: 47 degrees 58'12"N, 89 degrees 08'48"W; depth 168 m; coll. B. Riddell, 8 May 1973. ROM I-45: 47 degrees 59'06"N, 87 degrees 39'30"W; depth 241 m; coll. B. Riddell, 11 May 1973. (ref. ID; 5912)
  • National Museum, Ottawa (NMNS) 4404: 48 degrees 10'00"N, 87 degrees 08'06"W; depth 197 m coll. B. Riddell, 12 May 1973. NMNS 4405: 48 degrees 36'54"N, 87 degrees 48'00"W; dpeth 150 m coll. B. Riddell, 10 May 1973. (ref. ID; 5912)
  • Smithsonian Institution, Washington (US-NM) 51624: data as for NMNS4405. USNM51625: 47 degrees 16'12"N, 86 degrees 52'06"W; depth 269 m; coll. B. Riddell, 15 May 1974. (ref. ID; 5912)

    Phallodrilus labouichensis Rodriguez & Giani, 1989 (ref. ID; 5890 original paper, 6559)

    Descriptions

    One specimen complete with 28 segments, one specimen with posterior end missing. Diameter at VII (slightly compressed) 107 µm, at XI 128 µm. Clitellum extending over XI-XIII, not very conspicuous. Prostomium rounded, 30-42.5 µm long, and 39-59 µm wide. Secondary annulations from II to IV, but distinct only in II. Somatic setae bifid, with upper tooth shorter than lower. Ventral setae present on X and XI, but not modified into genital setae. 3 setae per bundle in segment II and 4 setae in all other preclitellar segments, 2-3 setae in postclitellar bundles. The length of setae varies between 20-27 µm. One pair male pores in line with ventral setae at XI, located in about the middle of segment, at the top of a projecting porophore. One pair spermathecal pores in the anterior part of X, near septum IX/X, in line of ventral setae. Pharyngeal glands in III-IV. Chloragogen cells from VI onwards. The gut presents a dorsal appendage without lumen at the intersegment VI/VII, extending over VII, and a stomachal dilatation at VIII. Coelomocytes small and scare. Male genitalia paired. Seminal vesicle anteriorly extended into IX. Ovisac in X and XI. Vas deferens shorter than atrium, entering apically. Atrium tubular, about 87 µm long and 7.5 µm diameter. One prostate gland attached to the ental part of the atrium, near the junction with vas deferens and the other in the ectal part. Atria opening to exterior through a simple pore, where it can be protruded, without forming a true penial apparatus but a protrusible pseudopenis. Spermathecae each with a short and muscular duct, 14.3 µm and an elongated ampulla vertical and perpendicular to the main axis of the body. Sperm filled the ampulla without a particular ordination. (ref. ID; 5890)

    Remarks

    In addition to P. labouichensis n. sp., there are five Phallodrilus species lacking penial setae; P. simplidentatus Erseus, 1979, P. inconspicuus Erseus, 1979, P. davisi Erseus, 1984, P. nidarosiensis Erseus, 1987, P. georgei Erseus, 1987. All five species present a series of characters that clearly distinguish them from P. labouichensis n. sp. P. simplidentatus presents simple-pointed setae in the posterior region of the body and a peculiar position of spermathecae in the middle of segment X. P. inconspicuus shows bipartite spermathecae, well developed prostate glands and vas deferens longer than atrium. P. davisi is unique in this group of species presenting a well-developed, true penis. P. georgei has a peculiar atrial structure and a bipartite spermathecal ampulla. Finally P. labouichensis n. sp. is related to P. nidarosiensis by the presence of male protuberances or porophores in segment XI. Both species may be distinguished, however, by the position of spermathecal pores (lateral in nidarosiensis) and structure of spermathecae (ampulla about the same length than duct in nidarosiensis) and atria ('somewhat comma-shaped', Erseus 1987). P. labouichensis n. sp. is also well characterized by the structure of its alimentary tract. A similar one is known only in P. vanus Erseus, 1984, in which it occurs as a change of gut structure in segments VIII and IX. (ref. ID; 5890)

    Etymology

    The species is named after the undergroud stream of Labouiche. (ref. ID; 5890)

    Type locality

    Underground stream of Labouiche, near Vernajoul. 5 km northeast of Foix, Ariege, France. Collected by V. Bougenec and M. Galissie, 17.V.1985. (ref. ID; 5890)

    Material examined

    1 specimen dissected, stained in haemtoxylin and mounted in Canada balsam. 1 whole-mounted specimen in Canada balsam. Collection of V. Bougenec and M. Galissie, Universite Paul Sabatier, Toulouse, France. (ref. ID; 5890)

    Phallodrilus leukodermatus Giere (ref. ID; 5864)

    Descriptions

    The outstanding morphological features of P. leukodermatus are the absence of the gut, mouth and anus, two well-developed blood vessels and the nerve cord transversing the spacious but septate coelomic cavity, and complete reduction of nephridia. Fluffy call digitations emerging from the vascular endothelium are best shown in cross sections. These structures more or less fill the coelom and contain an unusually 'active' cytoplasm with numerous mitochondria, Golgi-apparatus and a well-developed rough endoplasmic reticulum. In the thick body wall of the worms, a tenuous outer ring of circular muscle fibres surrounds a solid inner layer of longitudinal muscle cells. Embedded in this are conical portions of musculature, a feature quite unusual in annelids. They contain many mitochondria. A muscular sheath also encloses dorsally the thick nerve cord. The integument covering the long, slender, much annulated body of P. leukodermatus (ca. 20-30 mm long, ca. 0.15 mm wide) is a soild, conspicuous 'mantle'. Its surface, often folded into numerous tiny irregular ridges, is covered by countless epicuticular projections (nomenclature according to Richards 1977) which increase the surface area of the worm by approximately ten times. They emerge from digitated ramifications of the numerous long and irregular microvilli which cross the orthogonal grid of cuticular fibres and the loose matrix of the epicuticle. (ref. ID; 5864)

    Phallodrilus minutus Hrabe, 1973 (ref. ID; 5912)

    Descriptions

    Modified genital setae on segments X and XI, or XI only. All somatic setae bifurcate up to at least segment XVIII. Spermathecal epithelium of more or less uniform thickness. Up to seven penial setae per bundle as long as, or longer than, somatic setae. Spermathecal setae not modified. Penial setae two to seven per bundle with hooked or curved ends. Aria procumbent, with vas deferens junction apical, near anterior prostate. Spermatozeugmata absent. Three setae pe bundle anteriorly, two per bundle posteriorly. Five simple-pointed penial setae per bundle. Marine species. (ref. ID; 5912)

    Phallodrilus obscurus Cook, 1969 (ref. ID; 5890, 5912, 6602)

    Descriptions

    Modified genital setae on segments X and XI, or XI only. All somatic setae bifurcate up to at least segment XVIII. Spermathecal epithelium of more or less uniform thickness. Up to seven penial setae per bundle as long as, or longer than, somatic setae. Spermathecal setae not modified. Penial setae two to seven per bundle with hooked or curved ends. Aria procumbent, with vas deferens junction apical, near anterior prostate. Spermatozeugmata absent. Some anterior bundles with more than three setae. Three or more setae per bundle posteriorly, or, if only two per bundle, then some penial setae bifurcate. All dorsal setae bifurcate. Atria relatively small with very thin muscle walls. Porophores absent. Spermathecal pores in line with ventral setae. Two or three (rarely four) simple-pointed penial setae present. Marine species. (ref. ID; 5912)

    Marine species. (ref. ID; 6602)

    Phallodrilus parthenopaeus Pierantoni, 1902 (ref. ID; 5912, 6618)

    Descriptions

    Modified genital setae on segments X and XI, or XI only. All somatic setae bifurcate up to at least segment XVIII. Spermathecal epithelium of more or less uniform thickness. Up to seven penial setae per bundle as long as, or longer than, somatic setae. Spermathecal setae modified with upper teeth 2.5 times longer than the lower. Marine species. (ref. ID; 5912)

    Marine species. (ref. ID; 6618)

    Phallodrilus parviatriatus Cook, 1971 (ref. ID; 5912, 6602)

    Descriptions

    Modified genital setae on segments X and XI, or XI only. Somatic setae bifurcate up to about segment IX; dorsal and ventral setae simple-pointed from about segment X. Spermathecal epithelium thickened at regular intervals, imparting appearance of a chain of closely joined spheres internally. Marine species. (ref. ID; 5912)

    Marine species. (ref. ID; 6602)

    Phallodrilus profundus Cook, 1970 (ref. ID; 5912)

    Descriptions

    Two modified genital setae in each ventral bundle of segments IX, X, and XI. Marine; deep-sea species. (ref. ID; 5912)

    Phallodrilus rectisetosus (ref. ID; 5882)

    Descriptions

    Marine species. (ref. ID; 5882)

    Phallodrilus riparius Giani & Martinez-Ansemil, 1981 (ref. ID; 5890, 6559)

    Descriptions

    Freshwater species. (ref. ID; 6559)

    Phallodrilus sabulosus Erseus, 1979 (ref. ID; 6602) reported author and year? (ref. ID; 5882)

    Descriptions

    Marine species. (ref. ID; 5882, 6602)

    Phallodrilus subterraneus Rodriguez & Giani, 1989 (ref. ID; 5890 original paper, 6559)

    Descriptions

    No complete specimen (more than 16 segments, posterior end missing). Diameter in slightly compressed whole-mounted specimen at segment XI 127.5 µm. Prostomium rounded, longer than wide (length: 50 µm, width: 39.1 µm). Secondary annulations from VIII onewards, 5-7 on each segment. Somatic setae all bifid, upper tooth shorter and much thinner than lower. 2 to 5 setae per bundle present anteriorly, 31.5-35.7 µm long, and 3 posteriorly. Ventral setae of XI modified into penial setae, one on each pore, simple-pointed and slightly sigmoid (1.5 µm thick). These setae have a perpendicular disposition to the main axis of the body. Male pores located on prominent porophores near the mid-ventral line. One pair of spermathecal pores located ventrally in XII, near the intersegmental septum XI/XII. Pharyngeal glands in V-VI, dorsal or lateral to the alimentary tract. Male genitalia paired. Vas deferens very coiled, in the anterior part of segment XI, shorter than atrium, entering apically. Atrium divided into two parts: an ental enlarged one, up to 25 µm diameter, and covered by a very thick muscular layer (up to 6.4 µm), with a ciliated and conspicuous epithelium layer (5 µm thick). The ectal part is thinner (down to 12.9 µm) terminating in a pseudopenis. Two non-pedunculate prostates attached to the atrium, ental one very large, near the junction to the vas deferens, and the second one small, near the pore. One pair spermathecae in XII, with elongated ampulla of about 57.1 µm long and up to 21.4 µm wide, and a short duct 14.3 µm long and 9.3 µm diameter. Sperm in an elongated bundle into the ampulla. Spermathecal wall with a 2.8 µm thick muscular layer. (ref. ID; 5890)

    Remarks

    Phallodrilus subterraneus n. sp. is closely related to P. exiguus Erseus, 1979, P. lobatus Ersues, 1988, P. stilus Erseus, 1986 and P. macmasterae Erseus, 1986. These species have one simple-pointed penial seta on each male pore, and spermathecal setae are not modified. However, the new species may be distinguished from them by the general structure of the male duct: vas deferens short in relation to the length of the atrium, presence of a pseudopenis, atrium with a very thick muscular layer and divided into two parts (ciliated and non-ciliated). P. lobatus is well separated from the species quoted above by the presence of an unpaired mid-ventral crypt between male pores, and also a papillae area around the spermathecal pores. P. stilus is distinguished by peculiar 'flaplike male protuberances' in XI and tubular atria. The dorsal and lateral position of spermathecae in P. exiguus and P. macmasterae, respectively, distinguish these species from Phallodrilus subterraneus n. sp.. All four species lack pseudopenes, the atria taper directly into a simple opening, and the atrial muscular layer is much thinner. Phallodrilus subterraneus n. sp. is also related to P. postspermathecatus Erseus, 1980, by the position of spermathecae in the postatrial segment. However, this species is clearly separate from P. subterraneus n. sp. by the characteristic spermathecal walls, filled by granules of secretion; and its length, passing over segment XIII. P. postspermathecatus presents also a pseudopenis, but the atrium is shorter than the vas deferens and has a thin musculature. Finally, it should be noted that an additional species of Phallodrilus (P. cuspis Erseus & Dumnicka, in print) with a single penial setae, muscular atria and spermathecae located in segment XII was recently found in some wells in Italy (Erseus & Dumnicka, in print and C. Erseus, pers. comm.). P. subterraneus n. sp. and P. cuspis are very closely related. They differ mainly by the shape of the spermathecae and by the size of the penial setae; these setae are much thinner and slender in P. subterraneus than in P. cuspis. (ref. ID; 5890)

    Etymology

    The name subterraneus (Latin form 'underground') alludes to the habitat of this species. (ref. ID; 5890)

    Type locality

    Cave Jivero 1 at Ozana (Matienzo, Santander, Spain). Coordinates: 30TVN 537 948, 260 m altitude. 26.II.84. (Reference from the Trans. British Cave Res. Ass., 8(2), 1981). Cave with a river that flows all year. (ref. ID; 5890)

    Material examined

    1 specimen completely mature, stained in haematoxylin, dissected and mounted in Canada balsam. Collection of Jos Notenboom, Zoologisch Museum of Amsterdam. (ref. ID; 5890)

    Phallodrilus tempestatis Baker, 1981 (ref. ID; 5960 original paper) reported author and year? (ref. ID; 5882)

    Descriptions

    Marine species. (ref. ID; 5882)

    Length 10 mm to 14 mm, width anteriorly and posteriorly 270 µm to 400 µm, width at X 350 µm to 580 µm ( infixed, slightly compressed, sexually mature specimens), up to 52 segments. Prostomium bluntly conical, about as long as broad at peristomium. Clitellum extending over 1.2X-1/2XII. Somatic setae all bifid, three or four per bundle anteriorly, two per bundle posteriorly; upper tooth shorter and thinner than lower. Ventral setae of X unmodified, ventral setae of XI modified into penial bundles each of which contains three to five (usually four) slender, slightly sigmoid, ectally hooked penial setae. Penial setae (ps) maximally 5 µm thick, 90 µm to 130 µm long, enclosed in muscular sac (ms). Male pores open slightly posteriorly in XI in setal line on small papillae; pores open laterally penial setae. Spermathecal pores at 9/10 just lateral to setal line. Pharyngeal glands extend into V. Normal alimentary canal present. Male genitalia (all structures paired): sperm funnel (sf) small with fleshy lips, vas deferens (vd) from 25-28 µm wide with thick walls and ciliated lumen, opens apically into atrium; vas deferens three to six times as long as atrium. Atrium (a) elongate, pear shaped, composed of large irregular cells, atrium from 100 µm to 140 µm long, 39 µm to 46 µm wide; atrial muscle layer very thin; atrial lumen ciliated; no penis formed. Two stalked prostate glands (pr 1,2) present, anterior one with very wide base; anterior prostate (pr 1) opens into median portion of atrium, posterior prostate (pr 2) opens into atrium opposite and slightly basal to the anterior prostate. Spermathecae (s) with short, thick ducts, ducts with basal swelling, ampullae with thin walls; ampulla of open spermatheca often extended into IX, rarely not XI. Spermathecal pores elongate laterally. Sperm present in loose bundles. (ref. ID; 5960)

    Remarks

    This species is of a rather uniform width throughout the length of the body; the clitellar region is not extremely swollen. The histological differences between the vasa deferentia (highly organized small cells) and atria (large, rather amorphous irregular cells) are very marked with the border between the two cell types being very distinct. (ref. ID; 5960)

    Etymology

    This species is named in memory of the storm ("tempestas," Latin) which forced my sampling expedition to seek shelter on McInnes Island and which resulted in the collection of the type material of this species. (ref. ID; 5960)

    Habitat

    Known only from intertidal sediments (usually coarse sands or shell sands with some fine silts present), usually with some degree of freshwater influence. (ref. ID; 5960)

    Type locality

    Southeast side of McInnes Island, near Milbanke Sound, British Columbia, 52 degrees 15.7'N, 128 degrees 43.3'W, collected in midintertidal in a crevice in rock. Sediment: shell sand under rocks with a small amount of organic material present. Small trickle of fresh water over sediment. Collected by H.R. Baker, May 19, 1980. (ref. ID; 5960)

    Examined material

  • Holotype: NMCIC1981-19, a whole mounted specimen. (ref. ID; 5960)
  • Paratypes: NMCIC1981-20a, b, c; NMCIC1981-21a, b, five whole-mounted specimens from the type locality. (ref. ID; 5960)

    Phallodrilus tenuissimus (ref. ID; 5882)

    Descriptions

    Marine species. (ref. ID; 5882)