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The World of Protozoa, Rotifera, Nematoda and Oligochaeta

Lumbricillus

Lumbricillus Oersted, 1844 (ref. ID; 3692, 5967), Orsted, 1844 (ref. ID; 6913) or 1884 (ref ID; 1257, 5959)

Family Enchytraeidae (ref. ID; 1257, 1928, 7263)

Family Enchytraeidae: Subfamily Achaetinae (ref. ID; 5790)

ref. ID; 1923

Setae simple pointed and usually straight, spermathecae open between segments 4/5 or 3/4 and 4/5. Usually whitish in appearance and seldom more than 25 mm in length. Sigmoidal setae arranged in 4 bundles per segment. Esophagus gradually merging into intestine. Head pore between prostomium and segment 1; nephridia not plurilobed; peneal bulb as a rule without muscular strands, but covered by an investment of muscle. Testes deeply divided, forming a number of distinct lobes. (ref. ID; 1923)

ref ID; 5959

The genus is well characterized by the presence of large, lobed testes surrounded by well-developed seminal vesicles. In most other genera the testes and seminal vesicles are entire. (ref ID; 5959)

ref. ID; 5967

Remarks

Lumbricillus species are characterized by well-developed, lobed seminal vesicles; the absence of pharyngeal and oesophageal appendages and diverticula; and unnodulated setae which are most often sigmoid. A homogeneous generic diagnosis was made by Nielsen and Christensen (1959). There are only a few morphological characteristics useful for distinguishing the species of Lumbricillus. The key among these are the shape, size, and glandular attributes of the spermathecae which, in the marine species at least, will vary according to sexual maturity. In some Enchytraeidae the spermathecae are among the last of the reproductive organs to reach maturity (Welch 1914). Specimens in which developing ova are present as well as sperm in the ampulla of the spermatheca are usually specifically determinable. A complex of other morphological characteristics inculding setal shape and distribution, number of segments, relative dimensions of the sperm funnels and sperm funnel collars, and some details of the vas deferens add to the definition of species but because many of these vary between fairly broad limits they are not individually diagnostic. The penial bulb, type of Lumbricillus (Fig. 1B) is compact, penetrated by the vas deferens, and not penetrated, but covered, by a continuation of the longitudinal body wall musculature. (ref. ID; 5967)

Distribution

Cosmopolitan; the majority of species occur in littoral habitats but some also occur in limnic and rarely terrestrial environments (L. rivalis and L. pagenstecheri; see Nielsen and Christensen 1959). (ref. ID; 5967)

Type species

Lumbricillus lineatus (Muller, 1774) (ref. ID; 5967)
  1. Lumbricillus alaricus Shurova, 1974 (ref ID; 5959)
  2. Lumbricillus annulatus Eisen, 1904 (ref. ID; 5967)
  3. Lumbricillus arenarius (Michaelsen, 1889) (ref. ID; 6653)
  4. Lumbricillus belli Tynen, 1969 (ref ID; 5959, 5967)
  5. Lumbricillus buelowi (ref. ID; 5790)
  6. Lumbricillus charae (Tynen, 1970) (ref ID; 5959)
  7. Lumbricillus corallinae Shurova, 1977 (ref ID; 5959)
  8. Lumbricillus crassus (Claparede, 1862) (ref. ID; 1257)
  9. Lumbricillus crymodes (ref. ID; 5790)
  10. Lumbricillus curtu Coates, 1981 (ref ID; 5959 original paper, 5967)
  11. Lumbricillus dubius (ref. ID; 5790)
  12. Lumbricilus ebudensis (Claparede, 1862) (ref. ID; 1257)
  13. Lumbricillus eudioptus (ref. ID; 5790)
  14. Lumbricillus evansi Southern, 1909 (ref. ID; 1928 original paper)
  15. Lumbricillus fennicus Nurminen, 1964 (ref ID; 5959)
  16. Lumbricillus fossarum (Tauber, 1889) (ref. ID; 1257) reported year? (ref. ID; 1928)
  17. Lumbricillus franciscanus Eisen, 1904 (ref. ID; 5967)
  18. Lumbricillus franciscanus var. borealis Eisen, 1904 (ref. ID; 5967)
  19. Lumbricillus franciscanus var. unalaskae Eisen, 1904 (ref. ID; 5967)
  20. Lumbricillus helgolandicus (Michaelsen, 1934) (ref. ID; 1257, 5959)
  21. Lumbricillus ignotus Shurova, 1977 (ref ID; 5959)
  22. Lumbricillus kalatdlitus Nurminen, 1970 (ref. ID; 6913)
  23. Lumbricillus kamtschatkanus (Michaelsen, 1929) (ref. ID; 3692, 5959)
  24. Lumbricillus knoellneri (ref. ID; 5790, 6594)
  25. Lumbricillus kurilensis (ref ID; 5959)
  26. Lumbricillus lentus Shurova, 1978 (ref ID; 5959)
  27. Lumbricillus lineatus (O.F. Muller, 1771) (ref. ID; 3692) or 1774 (ref. ID; 1257, 5959, 5967, 6653, 7263) reported author and year? (ref. ID; 5863)
    Syn; Lumbricillus verrucosus Claparede, 1861 in Southern, 1909 (ref. ID; 1257)
  28. Lumbricillus litoreus (Hesse, 1893) (ref. ID; 1257) reported year? (ref. ID; 1928)
  29. Lumbricillus merriami Eisen, 1904 (ref. ID; 5967)
  30. Lumbricillus merriami var. elongatus Eisen, 1904 (ref. ID; 5967)
  31. Lumbricillus minimus (Cernosvitov, 1929) (ref. ID; 3692)
  32. Lumbricillus minutus (O.F. Muller, 1776) (ref. ID; 3692)
  33. Lumbricillus mirabilis Tynen, 1969 (ref. ID; 5967)
  34. Lumbricillus muscicolus (ref. ID; 5790)
  35. Lumbricillus niger Southern, 1909 (ref. ID; 1257, 1928 original paper)
  36. Lumbricillus orientalis (ref ID; 5959)
  37. Lumbricillus pagenstcheri (Ratzel, 1869) (ref. ID; 1257), pagenstecheri (Ratzel, 1869) (ref. ID; 5967, 6913) reported year? (ref. ID; 1928)
    Syn; Enchytraeus pagenstecheri Ratzel, 1869 (ref. ID; 5967)
  38. Lumbricillus pinquis (ref ID; 5959)
  39. Lumbricillus qualicumensis Tynen, 1969 (ref ID; 5959, 5967)
  40. Lumbricillus ritteri Eisen, 1904 (ref ID; 5959, 6913)
  41. Lumbricillus rivalis Levinsen, 1883 (ref. ID; 1257, 6913)
    Syn; Lumbricillus evansi Southern, 1909 (ref. ID; 1257); Lumbricillus subteraneus (Vejdovsky, 1889) (ref. ID; 1257)
  42. Lumbricillus rufulus (ref ID; 5959)
  43. Lumbricillus rupertensis Coates, 1981 (ref ID; 5959 original paper, 5967)
  44. Lumbricillus semifuscus Claparede, 1911 (ref. ID; 1257)
  45. Lumbricillus similis Shurova, 1977 (ref ID; 5959)
  46. Lumbricillus sterreri Lasserre and Erseus, 1976 (ref ID; 5959)
  47. Lumbricillus subterraneus (Vejdovsky) (ref. ID; 1928)
  48. Lumbricillus tsimpseanis Coates, 1981 (ref ID; 5959 original paper, 5967)
  49. Lumbricillus tuba Stephenson, 1911 (ref. ID; 1257, 5967)
  50. Lumbricillus verrucosus (Claparede) (ref. ID; 1928)
  51. Lumbricillus viridis Stephenson, 1911 (ref. ID; 1257)

Lumbricillus annulatus Eisen, 1904 (ref. ID; 5967)

Revised description

Medium to large, 12 to 24 mm, robust worms. Anterior segments very red due to blood colour, with 45-69 segments. Head pore 0/1. Setae robust anteriorly, slightly sigmoid, setae within a bundle more or less equal: preclitellar lateral (3-7), preclitellar ventral (5-10); postclitellar lateral (2-6), and postclitellar ventral (3-7, rarely 8 or 9). Brain concave posteriorly, lateral margins approximately parallel, length about 1.75 times width. Three pairs pharyngeal glands with ventral lobes, those of third pair large, almost concluding VII. At least two anterior pairs connected dorsally, third usually narrowly connected. Chloragocytes brownish, dense from VI, nucleate, very grandular. Lymphocytes nucleate, granular, extended spindle or oval shaped, with filamentous ends equal one-fifth to one-quarter seta length. Blood deep red or dark yellow in fixed specimens. Dorsal vessel arising 13/14 to mid-XV. Anteseptale of nephridia consisting only of funnel, postseptale large, lumpy, efferent duct posteroventral, terminal, or subterminal. Clitellum deep and dense, extending from one-half XI to XIII, rarely over the anterior to XIV. The following reproductive structures are laterally paired. Seminal vesicle regularly lobed in XI and X, occasionally extending into IX. Sperm funnel regular barrel shaped, length two to three times width; sperm collar wide, often convoluted. Vas deferens irregularly coiled in XII, rarely extending into XIII. Penial bulb large, oblong with thick muscular coat, length of bulb about one-third width segment XII. Ventral glands present in some of XIV to XX, increasing in size posteriad. Spermatheca simple, tubular with distinct ental and ectal ducts. Ectal duct length three to four times ampulla length, duct surrounded by small grandular cells decreasing in size entally, ectal orifice with a distinct, entally lobed rosette. Inner wall of ectal canal regularly folded into either a spiral ridge or a series of concentric rings (not determined which). Ampulla small, onion shaped, width just equal duct, connecting terminally to oesophagus at 5/6. (ref. ID; 5967)

Remarks

Shurova (1974) in an amended description of L. annulatus Eisen did not describe the pronounced ridging of the inner wall of the spermathecal ectal duct. Similar folding was observed (by K. Coates) infrequently in other Lumbricillus and Enchytraeus species; however, the relative length of the ectal duct and the consistent occurrence of regular ridging in mature individuals of L. annulatus makes the ridging of particular porminence for this species. Yamaguchi (1937) described Pachydrilus nipponicus from Akkeshi Bay, Japan. That species is scarcely distinguishable either from the original description of L. annulatus by Eisen (1904) or from the amended description of Shurova (1974). Japanese material of L. annulatus collected near Hakodate, Hokkaido in June 1960 has been examined. (ref. ID; 5967)

Lumbricillus belli Tynen, 1969 (ref ID; 5959, 5967)

Remarks

Lumbricillus belli may be distinguishable from other species of the genus on the basis of the description given by Tynen (1969). The existing illustrations and the original description of the species are however only adequate, not at all complete. Examination of longitudinally sectioned syntypes of L. belli Tynen yielded a little additional information. The clitellum, which extends over XII to XIII, is incomplete ventrally; the sperm duct is regularly coiled in XII; cutaneous glands are distributed in up to eight transverse rows per anterior segment; and the efferent ducts of the nephridia arise posteroventrally. It was not possible to confirm the presence of ventral glands on the ventral nerve cord in XV and XVI. Lumbricillus belli can be distinguished from L. qualicumensis, which Tynen (1969) found with L. belli, by size, setal distribution, length to diameter ratio of the sperm funnels, and relative width of the sperm funnel collar (Fig.6). Shurova (1977) has recently described three species, L. ignotus, L. similis, and L. corallinae, from the far eastern seas of the U.S.S.R. which are similar to both of L. belli and L. qualicumensis, particularly with regards to the morphologies of the spermathecae. A comparison of these species made by the senior author (1981) illustrated many interspecific similarities. Characteristic complexes of size, setal distribution and sperm funnel dimensions distinguish most of the species. (ref. ID; 5967)

Habitat

Upper intertidal, coarse sand, under seaweed. (ref. ID; 5967)

Lumbricillus curtus Coates, 1981 (ref ID; 5959 original paper, 5967)

Descriptions

Fixed specimens very small, 5 mm long, Holotype with 37 segments; other specimen with 34. Setae sigmoid, robust, bundles fan shaped; preclitellar lateral (2-5), preclitellar ventral (5-8); postclitellar lateral (3-6), and postclitellar ventral (3-7). Posterior setae with pronounced ectal "hook". Brain slightly indented posteriorly. Three pairs of pharyngeal glands present, all with ventral lobes, first or first and second pairs united broadly, third pair sometimes narrowly united, ventral lobes of third pair extending posteriad into VII, margins lobed. Chloragocytes nucleate, granular, small, present from V, dense from VII. Lymphocytes nucleate, small, ellipsoidal, granular, abundant. Dorsal vessel arising in XII, blood red in living specimen. Anteseptal part of nephridia consisting of funnel only, postseptale pear shaped, efferent duct varying from terminal to just subterminal, posteroventral. Clitellum very thin, extending over XII to XIII, no noticeable gland cell pattern. Reproductive structures laterally paired. Sperm funnel length 1.3 times width, collar regular, width just less than funnel. Vas deferencs relatively short, with only one or two loose loops in XII, entering penial bulb laterally near ectal edge of bulb. Penial bulb about same size as sperm funnel. Seminal vesicle with six or seven small, regular lobes, confined primarily to X. Spermatheca simple, attached to gut at one-half V. Ectal duct surrounded by fairly large, dense glands with a rosette of five or six low, rounded gland cells at ectal orifice. Ampulla round, thick walled ampullar cavity distinct, filled with sperm extending into ectal canal. Ventral nerve glands absent. (ref ID; 5959)

Remarks

With only one partially and two fully mature specimens available L. curtus sp. nov. is a minimally defined species. Nevertheless, it is distinctive within Lumbricillus because of the point of attachment of the spermatheca to the oesophagus; the extent and conformation of the septal glands; the small size of the sperm funnel and vas deferens; and the setal distribution. A table of comparison of L. curtus to similar Pacific species serves to show that it is justifiably designated a new species. Lumbricillus curtus differs from L. belli Tynen, 1969, L. similis Shurova, 1977, and L. corallinae Shurova, 1977 in the relative dimensions of the sperm funnel; those of the latter three species are considerably longer. Lumbricillus corallinae also is a significantly larger species than the other three. Lumbricillus qualicumensis Tynen, 1969, and L. ignotus Shurova, 1977 have narrow sperm collars (less than or equal to one-half of the width of the funnel) and long coiled vas deferens whereas L. curtus as a sperm collar about as wide as the funnel and a notably short vas deferens. Also characteristic for L. curtus is the anterior intraclitellar origin in XII of the dorsal blood vessel. Considerable overlap between the six species shown in Fig.9 in segment number and setal distribution reduce the taxonomic value of these characteristics. However, L. curtus and L. similis usually have more setae in ventral bundles than do the other four species. The number of segments and setal distributions tend to have limits which change as the number of specimens examined increases. Lumbricillus curtus is similar to the northern European L. semifuscus (Claparede, 1861) augm. (Stephenson, 1911) in possessing posteriorly enlarged ventral lobes on the third pair of septal glands. More significantly the relative dimensions and other details of the spermathecae serve to distinguish these two species. Because only three specimens of L. curtus with sperm in their spermathecae were obtained the ranges of variation of some characteristics are unknown. (ref ID; 5959)

Lumbricillus curtus (Fig.6) is distinctively small among Lumbricillus species found in British Columbia. The intersegmental (at half V) attachment of the spermatheca to the oesophagus and the small proportions of the sperm funnel and vas deferens are also characteristic (Cates 1981). (ref. ID; 5967)

Etymology

Curtus, Latin = short. (ref ID; 5959)

Habitat

Inlets of British Columbia, upper intertidal, usually in poorly sorted sediments. (ref ID; 5959)

Type locality

Neroutsos Inlet, British Columbia, 50 degrees 28.81'N, 127 degrees 34.10'W, uppper intertidal in poorly sorted, silty sediments. Collected June 19, 1978, by S.F. Cross. (ref ID; 5959)

Examined materials

  • Holotype: NMCIC1980-238, a mature, whole-mounted specimen, stained with borax carmine. (ref ID; 5959)
  • Other material: One mature specimen collected from Hastings Arm, British Columbia, 55 degrees 34.85'N, 129 degrees 48.30'W, June 22, 1977, by personnel of Dobrocky Seatech Ltd., Victoria, B.C., and a third specimen without eggs collected on Lord Island, Nuchatlitz Inlet, B.C., 49 degrees 46.75'N, 126 degrees 51.48'W, May 28, 1980, by H.R. Baker. (ref ID; 5959)

    Lumbricillus evansi Southern, 1909 (ref. ID; 1928 original paper)

    Diagnosis

    This species is most nearly related to L. subterraneus. The chief differences are: - L. subterraneus; epidermis smooth, without glands; corpuscles narrow; spermathecae spindle-shaped; male funnel with regular lip; freshwater species. L. evansi n. sp.; epidermis thickly covered with glands; corpuscles oval, frequently pointed; spermathecae roughly cylindrical, with constriction in the middle; male funnel with much enlarged and folded lip; marine littoral species. (ref. ID; 1928)

    Descriptions

    These worm are 10-14 mm long. The anterior bundles contain 6-9 setae. In each segment the epidermis is covered with numerous rows of small clear glands which alternate with fine lines. The brain is straight or slightly concave in front, deeply cut behind, where it is somewhat broader than in front. It is slightly longer than broad. There are two pairs of copulatory glands in the 13th and 14th segments, those in the 14th being the larger. This character is, however, very variable. The coelomic corpuscles are irregularly oval in shape, granular, and nucleated. In some cases the ends are drawn out into fine points. The intestine is covered with dark-brown glands. The girdle occupies segments 12 and 13. It is composed of small granular glands. The dorsal vessel rises in the 14th segment. There are three pairs of septal glands. The nephridia are formed of a small anteseptal, and a large broad, flat postseptal portion. The duct rises just behind the middle of the postseptal, and is about as long as this. Just behind the septum the nephridia are coloured brown. There are three "flames" in the nephridium, besides that in the funnel. The spermathecae are large and sac-shaped. They are constricted near the middle; and the base is surrounded by an enveloping glandular collar. The testes are of the usual shape and position. The male funnel varies greatly in shape and proportions according to its state of contraction. It varies from 6 to 10 times as long as broad; and the lip is thrown into large, conspicuous spreading folds. This remarkable character was very constant, and was found in specimens from widely distant localities, and easily servers to distinguish this from all other species. The duct is several times longer than the funnel. (ref. ID; 1928)

    Etymology

    This species is named after Mr. W. Evans, of Edinburgh, who collected my species for me. (ref. ID; 1928)

    Lumbricillus fossarum (Tauber, 1889) (ref. ID; 1257) reported year? (ref. ID; 1928)

    Descriptions

    This species was very briefly described without figures by Tauber and Levinsen. Ude (tom. cit.) gave a fuller description, and figured the spermatheca, nephridium, and copulatory glands. The cuticle in each segment bears several rows of clear glands. The brain is slightly concave before and behind, and is broader at the back than the front. The anterior ventral bundles contain 6-8, rarely 9 setae. The coelomic corpuscles are oval or pear-shaped, granular and nucleated. The clitellum is very prominent, and occupies segment 12 and half 13. The dorsal vessel rises in the 13th segment (Ude says between the 14th and 15th segments). (ref. ID; 1928)

    Lumbricillus franciscanus Eisen, 1904 (ref. ID; 5967)

    Remarks

    The type material of L. franciscanus was deposited at CASF and subsequently destroyed in 1906 (personal communication, D. Chivers). Lumbricillus fanciscanus is not described from an intertidal or marine habitat but other forms of the species, L. f. var. borealis and L. f. var. unalaskae, subsequently described from Alaska may have been found in intertidal habitats. Stephenson (1922) thought that L. aegialites Stephenson, subsequently synonymized with L. pagenstecheri sensu Nielsen and Christensen (Nurminen 1965), was not unlike L. franciscanus; however, on the basis of the existing descriptive material, it seems that the glandular arrangements around the spermathecal ectal ducts of the two species are distinctively different. Lumbricillus fransicanus is not known to have a distinct rosette of glands at the ectal orifice. (ref. ID; 5967)

    Habitat

    Moist soil of river bank. (ref. ID; 5967)

    Type locality

    The original material was collected from the banks of the Santa Clara River, California. (ref. ID; 5967)

    Lumbricillus franciscanus var. borealis Eisen, 1904 (ref. ID; 5967)

    Remarks

    The type material could not be located. It was not at UWMS as indicated by Reynolds and Cook (1976). The description of this taxon as a form of the species L. fraciscanus was based on two mature specimens. It is not determinable from the scanty morphological details in Eisen's (1904) description whether this is a taxon of the specific, subspecific, or infrasubspecific level in accordance with International Code of Zoological Nomenclature regulations. The location and habitat information that Eisen (1904, p.89) gave for L. franciscanus var. borealis was not complete; however, because the greatest diversity of Lumbricillus species is found in intertidal habitats it is not unreasonable to suppose that these specimens were found there also. (ref. ID; 5967)

    Type locality

    The original material was collected on St. Paul Island, Pribilof group, Alaska. (ref. ID; 5967)

    Lumbricillus franciscanus var. unalaskae Eisen, 1904 (ref. ID; 5967)

    Remarks

    Type material were not located at UWMS. Lumbricillus franciscanus var. unalaskae Eisen is also considered a taxon of, at present, indeterminate status. (ref. ID; 5967)

    Type locality

    The original materials were collected at Unalaska, Unalaska Island, Aleutian group, Alaska. (ref. ID; 5967)

    Lumbricillus knoellneri (ref. ID; 5790, 6594)

    Descriptions

    Marine species. (ref. ID; 6594)

    Lumbricillus lineatus (O.F. Muller, 1771) (ref. ID; 3692) or 1774 (ref. ID; 1257, 5959, 5967, 6653, 7263) reported author and year? (ref. ID; 5863)

    Synonym

    Lumbricillus verrucosus Claparede, 1861 in Southern, 1909 (ref. ID; 1257)

    Descriptions

    Polyploidism. (ref. ID; 7263)

    Remarks

    Lumbricillus rupertensis Coates (Fig.5) has a similar spermathecal morphology and setal distribution to L. lineatus but is clearly distinct with regards to its very short sperm funnels. (ref. ID; 5967)

    Habitat

    Salt marshes, upper to lower littoral zone, driftline algal deposits; sand, gravel, mud under rocks; near domestic and industrial marine outfalls. In British Columbia it was collected in greatest abundance near marine pulp mills. (ref. ID; 5967)

    Lumbricillus litoreus (Hesse, 1893) (ref. ID; 1257) reported year? (ref. ID; 1928)

    Diagnosis

    The only differences between this species and L. lineatus appear to be (1) number of setae in a bundle; (2) The structure of the copulatory glands; (3) the nature of the glands at the spermathecal pore. None of these differences seem of great importance; and it is doubtful whether there is sufficient justification for keeping the two species separate. (ref. ID; 1928)

    Descriptions

    This specimens found in soil and in brackish water. (ref. ID; 1928)

    Lumbricillus merriami Eisen, 1904 (ref. ID; 5967)

    Remarks

    Lumbricillus merriami as described is similar in many morphological characters to both L. annulatus Eisen and L. nipponicus Yamaguchi (Fig.7). The relative dimensions of the spermatheca of L. merriami are different than those of the above-mentioned species, with the ampulla being somewhat longer relative to the length of the ectal duct. The vas deferens of L. merriami is shorter than that of L. annulatus. The L:D of the sperm funnel is 3:1 for L. merriami, usually slightly longer than the 2-3:1 found in L. annulatus. Like L. annulatus the anterior part of the postseptale of the nephridia are described (Eisen 1904) as covered by wartlike projections. With respect to habitat, the three species are similar. The specific status of L. merriami is considered to be unstable and indeterminate at present. Investigation of new material is desirable. (ref. ID; 5967)

    Habitat

    Intertidal under decaying seaweeds. (ref. ID; 5967)

    Lumbricillus merriami var. elongatus Eisen, 1904 (ref. ID; 5967)

    Remarks

    The type specimens of L. merriami var. elongatus deposited at CASF were destroyed in 1906 (personal communication, D. Chivers). The sperm funnels of L. merriami var. elongatus are of notably different relative dimensions than those of these species, "globular", and three times longer than wide, respectively; the setal distribution is quite different from that of the species; and the relative dimensions of the spermathecae are different. If it had been indicated by Eisen that the specimens of L. merriami and var. elongatus were at the same level of sexual maturity then it would be justified to consider these taxa as two distinct species. In the existing circumstance it is not possible to resolve these two taxa and L. merriami var. elongatus is of indeterminate status. (ref. ID; 5967)

    Habitat

    Intertidal, under seaweed (with specific form). (ref. ID; 5967)

    Type locality

    The original material was collected at Metlakatla, Alaska, On June 4, 1899. (ref. ID; 5967)

    Lumbricillus mirabilis Tynen, 1969 (ref. ID; 5967)

    Remarks

    Examination of the syntype of Lumbricillus mirabilis Tynen led to some amendments and additions to the original taxon description (Tynen 1969). The posterior setae of the type specimen are slightly sigmoid, the posterior margin of the brain is slightly concave and the nephridial efferent ducts may be terminal or just subterminal. The clitellum covers segments XII to half XIV, not XI to half XIV. Five eggs are present, extending posteriad to XVI; the lobes of the seminal vesicle extend through all of X-VIII. A note of particular importance is that the spermathecal structure of the syntype was not well represented by Tynen's illustration (1969, Fig.9). The spermathecal structure in the syntype is as shown in Fig.6 (a tracing made with a drawing tube). Examination of numerous mature specimens from recent collections allowed further additions and served to confirm amendments based on the type specimen. One change not indicated in Fig.6 was allowance for a high degree of variability in dorsal fusion of the pharyngeal gland pairs. Shurova (1974) also proposed similar amendments to the original taxon description (Fig.6). The spermathecae as described and illustrated by Shurova (1974, Fig.3b) are very similar to what was observed in both the type and the new British Columbia material; however, the distribution of sperm in the ampullar cavity is more diffuse than found in British Columbia specimens. Shurva (1974) did not record the wide, convoluted sperm funnel collar that is characteristics for sexually mature L. mirabilis found in British Columbia. The extent of morphological similarity between L. mirabilis and L. pagenstecheri is shown in Fig.6. The most significant morphological difference found in mature, fixed specimens of the two species was in the relative dimensions of the spermathecae. Lumbricillus pagenstecheri has a longer ectal duct and a denser envelope of glandular cells. Chloragogen colour, dicussed by Tynen (1969), is not a consistent characteristics from specimen to specimen and is probably nutritionally and physiologically dependent. (ref. ID; 5967)

    Habitat

    From 1.8 to 5 m above chart datum, in coarse sand, gravel, shell deposits; frequently associated with driftline algae. (ref. ID; 5967)

    Lumbricillus niger Southern, 1909 (ref. ID; 1257, 1928 original paper)

    Descriptions

    This species is at once distinguished by its dark appearance. To the naked eye it appears quite black. This is due to the presence of very dark brown pigment in the cells which cover the gut. They are small, with granular contents, and there is a small, clear space in the centre of each. In the anterior segments these cells are absent, and the head of the worm is of the normal pink colour. They begin sparsely in the 4th segment; and from the 8th segment onwards they surround the gut and the dorsal vessel. The worms are 10-15 mm long. The anterior ventral bundles contain 5-7 setae, the lateral ones 4-6. The setae are not so curved, nor are they arranged in such a fan-shaped manner, as is usual in this genus. The epidermis of each segment is composed of several rings formed by lines of fine dotted glands; and each ring has several rows of clear, oval glands. Contrary to the usual rule, these rings are even more prominent in the posterior than in the anterior segments. The clitellum occupies segment 12. It is formed of rows of very small granular glands. The prostomium and 1st segment are covered with small papillae, probably sensory in function. The head-pore is situated between the prostomium and the 1st segment. It is small and round. The brain is somewhat longer than broad. It is deeply emarginate behind, and straight or slightly concave in front, and the sides are almost parallel. The coelomic corpuscles are very thin and fragile in appearance. Their contents are faintly granular, with a clear spot in the middle. They are so thin as to be bent into folds as they flow about in the coelomic fluid. They are of various shapes, and resembles those of Marionina arenaria, which Michaelsen has figured. In some of them the ends are drawn out into fine points. In others one side is rounded, and the other drawn out into fine points. In others one side is round, and the other drawn out into a number of fine pseudopod-like processes, which may be branched. There are three pairs of septal glands in the 4th-6th segments. The nephridia are composed of a long, slender anteseptal, and a large, flat postseptal, which passes gradually into a long duct. The dorsal vessel rises between the 13th and 14th segments. The copulatory glands are very small, and seem to occur only in the 14th segment. The spermathecae are in the normal position, and communicate with the oesophagus by a narrow duct. The ampulla is oval in shape, and is shorter than the narrow duct. The latter is surrounded at the pore by a massive collar of glands. The testes are lobed, and occupy the 10th segment. The sperm-funnels are very variable in shape; and the relative proportion of length to breadth varies from 4-7 according to the tension on the organ. The lip is prominent and usually slightly folded. The duct is long and coiled, and ends in a large prostate. This species is chiefly characterized by its dark colour and the structure of its nephridia, spermathecae, and coelomic corpuscles. (ref. ID; 1928)

    Lumbricillus pagenstcheri (Ratzel, 1869) (ref. ID; 1257), pagenstecheri (Ratzel, 1869) (ref. ID; 5967, 6913) reported year? (ref. ID; 1928)

    Synonym

    Enchytraeus pagenstecheri Ratzel, 1869 (ref. ID; 5967)

    Descriptions

    This species occurs commonly in manure and garden soil, and also in brackish water near the sea. In specimens from England the nephridia were peculiar in having no differentiated duct, the postseptal continuing of the same diameter up to the external pore. The size is 8-10 mm. Setae 4-7. Only two pairs of commissural vessels enter the two anterior loops of the ventral vessel, in front of the junction of the latter, and not three as Vejdovsky figures. Ude separated L. henkingi form this species on account of the structure of the copulatory glands, all other characters being approximately in agreement. There seems nothing in the figures given to justify this proceeding, especially as the copulatory glands vary considerably in the same species. (ref. ID; 1928)

    Lumbricillus qualicumensis Tynen, 1969 (ref ID; 5959, 5967)

    Revised description

    Small, yellow specimens, 7 to 10 m long, with 30 to 42 segments. Cutaneous glands in approximately 6 transverse rows per segment. Setae sigmoid, without nodulus: preclitellar lateral (2-6), preclitellar ventral (2-6), postclitellar lateral (1-5), and postclitellar ventral (2-5). Brain distinctly concave posteriorly, maximum length approximately 1.5 times width. Three pairs of pharyngeal glands present, all with ventral lobes, all united dorsally. Lymphocytes nucleate, grandular, spindle shaped, with attenuated tips, one-half to three quarters setal length. Chloragocytes large, colourless or pale gold, sparse from V, dense from VII, one-half setal length, nucleate. Blood faint yellow to pink. Dorsal vessel usually originating at 13/14, ranging from one-half XII to one-half XIV. Anteseptal part of nephridium consisting of funnel only, postseptale, oval to egg shaped, ectal duct posteroventral, terminal or subterminal. Cells of clitellum deep, dense scattered, extending from XII to XIII, ventrally also over posterior XI. Reproductive structures are laterally paired. Seminal vesicle small, three to six lobes in XI and X. Sperm funnel length about 1.2 to 1.5 times width, collar constricted. Vas deferens long and narrow, regularly coiled in XII. Penial bulb more or less sphaerical, height of bulb one-quarter diameter XII. Small ventral bodies usually present in XIV and XV. Spermatheca (Fig.6) with distinct ental and ectal ducts. Ectal duct length 2 to 2.5 times ampulla length, covered with small glandular cells; with distinct, slightly lobed rosette around orifice. Ampulla round, with regular, central ring of sperm, connecting via terminal ental canal with oesophagus. (ref. ID; 5967)

    Remarks

    The original description of L. qualicumensis Tynen gave no details of lymphocytes and chloragocytes, of brain shape or dimensions, of nephridia, or of the morphology of the pharyngeal glands. No type was designated for this taxon (Tynen 1969, 1970), nor could any type material be located with other material of Tynen's (1969) Vancouver Island species held in NMCIC. In some details the specimen identified as L. qualicumensis did not agree with the original description. These differences are recorded in Fig.6; they include a more anterior dorsal blood vessel origin in the recent material, greater variability in segment number range and setal distribution, and the presence of small ventral bodies in XIV and XV of mature worms. Lumbricillus ignotus Shurova, 1977 is similar to L. qualicumensis. Minor differences between these taxa (Fig.6) exist in segment number ranges and setal distributions. The origin of the dorsal blood vessel for L. ignotus is like L. qualicumensis as described by Tynen (1969). The presence of ventral glands in XIV and XV of L. ignotus is like the condition of the recent material of L. qualicumensis. The clitellum of the British Columbia taxon is not as extensive as described for L. ignotus (Shurova, 1977) but for the two species the described morphologies of the spermathecae and sperm funnels are the same. The few preceeding remarks are made to show that the specimens identified as L. qualicumensis were neither exactly the same as the taxon described by Tynen (1969) nor as L. ignotus and the significant morphological differences probably do not exist between L. ignotus and L. qualicumensis. Because the specimens of L. qualicumensis were collected in the vicinity of the reported type locality of that species and in spite of resevations on account of the limited detail of the original description and absence of type material, they were identified as L. qualicumensis and the taxon description was slightly amended as above and in Fig.6. It is not advisable to propose the synonymy of L. qualicumensis to L. ignotus without examination of the type material of the latter. Lumbricillus qualicumensis is distinguished from L. belli Tynen, 1969 and L. pagenstecheri (Fig.6) on the basis of relative dimensions of the sperm funnel and morphology of the sperm funnel collar. Lumbricillus pagenstecheri (Fig.6) is also a noticeably longer and more robust worm, with straight or only slightly sigmoid setae. (ref. ID; 5967)

    Habitat

    Middle to high intertidal; in sandy substrates, frequently associated with wash and (or) attached algae. (ref. ID; 5967)

    Lumbricillus rupertensis Coates, 1981 (ref ID; 5959 original paper, 5967)

    Descriptions

    Small intertidal species, size (fixed) 8 to 10 mm, with 33 to 41 segments. Cutaneous glands in obvious rows anteriorly. Setae sigmoid, without nodulus. Distribution of setae: preclitellar lateral (3-5), preclitellar ventral (4-6); postclitellar lateral (2-4), and postcilitelar ventral (3-6). Brains concave posteriorly, lateral margins converge anteriad length approximately twice width. Three pairs of pharyngeal glands present, all with ventral lobes, all united dorsally, third pair only narrowly. Chloragocytes present from VI, golden, large and dense posterior to clitellum. Lymphocytes elliptical, nucleate, abundant, length one-quarter to one-third setal length. Dorsal vessel arising in posterior of XIII or XIV. Blood yellow in fixed specimens. Anteseptal parts of nephridia consisting of funnel only, efferent duct posteroventral. Clitellum extending over XII and XIII, not covering posterior to XIII, cells tall, granular, less dense ventrally, distribution irregular. The following reproductive organs are laterally paired. Seminal vesicle regularly lobed, about six lobes, extending anteriad to IX. Sperm funnel tapered to duct end, length about 1.5 times width, approximately one-half body width, collar equal to funnel width, regular. Vas deferens tightly coiled in XII. Penial bulb small, spherical. Spermatheca of mature specimens with a single medioventral ampullar expansion, ampulla attached to gut at dorsal apex of expansion. Ental and ectal ducts not distinct, no distinct ampullar cavity. Spermatozoa embedded in the thick walls of the ampulla proper and of the expansion. Lobed, medium-sized rosette of fused glands at ectal pore, no glands along ectal duct. One or two developing eggs present at one time. Small ventral bodies on nerve cord in XIII to XV. (ref ID; 5959)

    Remarks

    Lumbricillus rupertensis sp. nov. is morphologically similar to several other Lumbricillus species including L. lineatus (Muller, 1774), L. fennicus Nurminen, 1964, and L. alaricus Shurova, 1974. The sperm funnel ratio of L. rupertensis is like that of L. fennicus and L. alaricus; however, among these three species, L. fennicus has characteristically divided or lobed sperm funnel. The general morphology of the spermathecae of L. rupertensis, an indistinct ampullar cavity and only a crown of lobed glands around the ectal pore, is similar to L. lineatus. L. fennicus, and L. alaricus. Lumbricillus lineatus is characterized with regard to its spermathecal morphology by distinct ental and ectal ducts and an ampullar constriction dividing it into an ectal and an ental part. The detailed morphology of the spermathecal ampulla of L. rupertensis, similar to the ampullae of L. sterreri Lasserre and Erseus, 1976 and L. charae (Tynen, 1970), i.e., a globular expansion of the ampulla medial to the junction with the oesophagus, is unique among the morphologically similar species shown Fig.12. (ref ID; 5959)

    Lumbricillus rupertensis bears many morphological similarities to L. lineatus (Fig.5), also recorded from British Columbia, but these species can be distinguished by their distinctly different sperm funnel morphologies as well as by the details of their respective spermathecal ampullae (Coates 1981). The ampulla of L. lineatus is divided by a constriction into distinct ectal and ental parts whereas mature specimens of L. rupertensis have a median globular expansion at the ental apex of the ampulla. (ref. ID; 5967)

    Etymology

    From Prince Rupert, British Columbia. (ref ID; 5959)

    Habitat

    Upper intertidal in silt and woodchips. (ref ID; 5959)

    Type locality

    Near Prince Rupert, British Columbia, 54 degrees 13.58'N, 130 degrees 18.02'W, intertidal. Collected August 22 to 24, 1974, by personnel of P.E.I., Vancouver, B.C. (Levings et al. 1975). (ref ID; 5959)

    Examined materials

  • Holotype: NMCIC1980-236, a mature stained specimen mounted in Canada balsam. (ref. ID; 5959)
  • Paratype: NMCIC1980-237, a whole mounted specimen, location as for Holotype. (ref ID; 5959)
  • Other material: USNM60348 and 60349, two stained whole mounts. Author's collection four whole-mounted specimens and one whole, stained specimen in 70% ethanol. All collected intertidally in the immediate vicinity of the type locality (Levings et al. 1975). (ref ID; 5959)

    Lumbricillus subterraneus (Vejdovsky) (ref. ID; 1928)

    Descriptions

    This species found sewage works, under-ground waters, stream (excessively contaminated with trade effluents). The worm are 12-18 mm long. The anterior ventral bundles contain 5-7 setae. The cuticle is smooth and without glands. The spermathecae are spindle-shaped, and without sharply defined duct, and are surrounded at the base with prominent glands. The dorsal vessel rises in the 14th or 15th segment. The copulatory glands vary greatly. Sometimes large glands occur in the 13th and 14th segments; sometimes they are small, or quite absent. The brain, nephridia, and genital organs agree with the description of Vejdovsky. (ref. ID; 1928)

    Lumbricillus tsimpseanis Coates, 1981 (ref ID; 5959 original paper, 5967)

    Descriptions

    Mature fixed specimens 8 to 10 mm long, with 30 to 34 segments. (Mature specimens from Ocean Falls had only 27 and 25 segments but in all other respects fit this description which is based on the specimens found at Prince Rupert.) Setae sigmoid, bundles fan-shaped; preclitellar lateral (3-4, rarely 2 or 5), preclitellar ventral (3-6), postclitellar lateral (2-5), postclitellar ventral (3-6, rarey 1, 2, or 7). Brain indented posteriorly, narrowing gradually anteriad, length 1.5 to 2 times maximum width. Two anterior paires (4/5 and 5/6) pharyngeal (= septal) glands united dorsally, third pair narrowly or not united; dorsal lobes of posterior pair (6/7) extending posteriad into VII, lobed along posterior edges; ventral lobes present for all pairs. Few chloragocytes present in V, dense from VI, tall, yellowish. Lymphocytes small, oval to elongate, nucleate, abundant, largest cells one-third to one-half length of setae. Dorsal vessel arising in XII or XIII, blood yellow in fixed specimens, light red in living specimens. Anteseptal parts of nephridia short, wide, consisting of funnel only, postseptale large, with cellular interstitial tissue, efferent duct wide, terminal. Clitellum extending from posterior XI to anterior XIV. Male and female reproductive structures are laterally paired. Seminal vesicle extending through XI to IX, with five to nine regular lobes. Sperm funnel cylindrical, narrowing ectally, usually bent; length 1.5 to 2.5 times width, length subequal to body width. Sperm collar width just less than funnel width, not convoluted. Vas deferens long, narrow, thin walled, arising postero-laterally from sperm funnel, coiled tightly in XII. Penial bulb of medium size with muscular coat. Spermatheca simple, ampulla large, onion shaped, with relatively thick walls, connected to gut at 4/5 by short ental duct; ectal duct as long as or longer than ampulla, of almost constant width, canal widening slightly at ampulla. Sperm in irregular ring-shaped swirl in ampulla. Ectal duct with some all gland cells; with distinct, small rosette around the ectal orifice. Ventral bodies on nerve cord absent. (ref ID; 5959)

    Remarks

    One specimen identifiable as Lumbricillus tsimpseanis with three spermathecae, a pair in V and one in VI, was found in Neroutsos Inlet. Lumbricillus tsimpseanis sp. nov. is morphologically similar to two marine intertidal Lumbricillus species described by Michaelsen, L. helgolandicus (1927 and 1934) and L. kamtschatkanus (1929). Some taxonomically important characteristics of these species are shown in Fig. 4. Lumbricillus tsimpseanis differs from L. helgolandicus, from all accounts, by having fewer segments and being substantially smaller, 8 mm rather than 14 mm. Lumbricillus helgolandicus was originally described as having a sperm funnel approximately 12 times as long as wide (Michaelsen 1934; "Samentrichter vielfach so lang wie dick, bie einem naher untersuchten Stuck ungefahr 12 mal so lang wie in der Mitte dick ..." p.138). Nielsen and Christensen (1959) redescribed L. helgolandicus as having sperm funnel only two to three times longer than wide and with sigmoid setae rather than straight setae as were observed by Michaelsen (1934). Von Bulow (1957) recorded L. helogolandicus from Kimbrischen Halbinsel but did not report discrepancies in the sperm funnel dimensions or setal shapes between her specimens and those originally described by Michaelsen. However, in the complete description of L. helgolandicus, Michaelsen (1934) suggested that his species was synonymous with L. pagenstecheri sensu Stephensen 1922 (non Ratzel) even though the latter was described with male funnels only three to five times as long as thick. Specimens of L. tsimpseanis and L. helgolandicus sensu Nielsen and Christensen (1959) have sperm funnel dimensions as described for L. kamtschatkanus. Nurminen (1973) pointed out the possibility of synonymy between L. kamtschatkanus and L. helgolandicus sensu Nielsen and Christensen. Lumbricillus kamtschatkanus was described without illustrations so that the exact shape of the spermatheca is not determinable. It is described, like those of L. helgolandicus and L. tsimpseanis, with long, thin ectal duct and an inflated onion-shaped ampulla. The duct is described as naked like L. helgolandicus, unlike L. tsimpseanis; however, the rosette at the ectal orifice is small as for L. tsimpseanis. It was not possible to locate any type materials of L. kamtschatkanus nor has the species been recorded recently from the far-eastern seas of the U.S.S.R. (Shurova 1974, 1977). Lumbricillus lentus Shurova, 1978, which is described as being like L. ritteri Eisen, 1904 (= L. pagenstecheri (Ratzel) (personal observation)), but is lacking the covering of glandular cells around the spermathecal ectal duct, was found the intertidal of the eastern coast of Kamchatka and may be synonymous with L. kamtschatkanus. (Michaelsen (1929) also comments that L. kamtschatkanus is like L. pagenstecheri without glands along the ectal duct.) Shurva (1974, 1977) also has described several northern Pacific Lumbricillus species with glands along the spermathecal ectal duct. These include L. corallinae, L. ignotus, L. pinquis, L. similis, L. kurilensis, L. orientalis, and L. rufulus from the far-eastern seas of the U.S.S.R. These all differ from L. tsimpseanis in the density of their glandular coverings along the spermathecal ectal duct; in their greater size and numbers of segments; in setal distributions; in sperm funnel dimensions; and in the details of their spermathecal morphologies. (ref ID; 5959)

    Lumbricillus tsimpseanis (Fig.5) is distinguished from other Lumbricillus species found in British Columbia by the distribution of sperm in the spermathecal ampulla and by the arrangement of glands along the ectal duct of the spermatheca, a small rosette and a few separate gland cells surrounding the ectal half of the duct. (ref. ID; 5967)

    Etymology

    From Tsimpsean Peninsula where the type specimens were collected. (ref ID; 5959)

    Habitat

    Upper intertidal, estuarine, in filamentous algae (Enteromorpha?) either washed up or attached on logs and rocks. (ref ID; 5959)

    Type locality

    Near Prince Rupert, British Columbia, 54 degrees 14.55'N, 130 degrees 16.26'W; upper intertidal-estuarine. Collected August 22 to 24, 1974, by personnel of the Pacific Environment Institute (P.E.I.), Vancouver, B.C. (Levings et al. 1975). (ref ID; 5959)

    Examined materials

  • Holotype: NMCIC1980-2100, one mature specimen, with eggs, stained with borax carmine, mounted whole in Canada balsam. (ref ID; 5959)
  • Paratypes: USNM60347, one specimen from Prince Rupert, 54 degrees 12.89'N, 130 degrees 17.22'W, not fully mature, collected by the author; and USNM62421, a mature specimen with eggs from the type locality, NMCIC1980-235, from the same location and collected at the same time as USNM60347, also less than fully mature; and NMCIC1980-2101a to NMCIC1980-2101d, four mature specimens collected with the Holotype. All paratypes are whole mounted. (ref ID; 5959)
  • Other material: Five mature specimens from the type location, collected May 12, 1980, by the author; Neroutsos Inlet, Vancouver Island, 50 degrees 23.87'N, 127 degrees 28.42'W. 12 specimens, collected June 13, 1976, and several other specimens collected at the same time in the same general area; Neroutsos Inlet, 50 degrees 20.52'N, 127 degrees 26.30'W, 5 specimens collected August 26-27, 1974, by P.E.I. personnel; Ocean Falls, B.C. 52 degrees 21.53'N, 123 degrees 43.12'W, two specimens of collected February 26, 1977; and one specimen from the Brooks Peninsula, Vancouver Island, 50 degrees 12.02'N, 128 degrees 47.95'W, collected July 24, 1976. These specimens are held in the author's collection. (ref ID; 5959)

    Lumbricillus tuba Stephenson, 1911 (ref. ID; 1257, 5967)

    Remarks

    The segment number of the British Columbia specimens is more variable and on average higher than that indicated either by Stephenson (1911), 35 to 39, or by Nielsen and Christensen (1959), 33 to 41, for L. tuba. The presence and extent of ventral glands is variable; none mentioned by Nielsen and Christensen (1959); small, in segments XIII to XVI recorded by Stephenson (1911); and in the British Columbia specimens small, in XIV and XV. Lumbricillus tuba differs most significantly from other Lumbricillus species described from the British Columbia coast in that it has a long, distinct, naked spermathecal ectal duct. Lumbricillus kamtschatkanus (Michaelsen, 1929) is apparently a similar species; however, L. kamtschatkanus is not indicated as having the same distinctive spermathecal shape as L. tuba. Michaelsen (1929) described the spermatheca of L. kamtschatkanus "...Ampulle dick-zwiebel-forming... Ausfuhrgang scharf von der Ampulle abgesetzt, fast doppelt so lang, wening mehr als halb so dick..." (p.317). The dimensions of the sperm funnel of L. kamtschatkanus are also different than those of L. tuba. Nurminen (1973), in the respect of the spermatheca, found L. kamtschatkanus to be similar to L. helgolandicus (Michaelsen, 1927) augm. (Michaelsen, 1934) aff. Nielsen and Christensen, 1959. (ref. ID; 5967)

    Habitat

    Intertidal; salt marsh; along shoreline sewer effluent; sand, gravel, shingle; decaying drift algae. Estuarine; lower to upper midintertidal; in gravel, sand with living Fucus and (or) emergent vegetation. (ref. ID; 5967)

    Lumbricillus verrucosus (Claparede) (ref. ID; 1928)

    Descriptions

    This species is a common littoral form, and recorded it from several fresh-water localities. There are copulatory glands in the 14th and 15th segments. (ref. ID; 1928)