The World of Protozoa, Rotifera, Nematoda and Oligochaeta
Marionina
Marionina Michaelsen, 1889 (ref. ID; 1257, 3692, 5967)
Quote from ref. ID; 5001
Family Enchytraeidae (ref. ID; 1257, 1928, 7502, 7620)
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 undivided and massive. (ref. ID; 1923)
ref. ID; 5967
Remarks
Marionina is a heterogeneous genus and it is difficult to characterize the species by one or a few characteristic morphological conditions. Blood is usually colourless so that living specimens appear white or transparent, but it may be coloured. Seminal vesicles may or may not be present; however, if present they are compact. It is thought as well that the seminal vesicles may deteriorate at advanced sexual maturity. Nielsen and Christensen (1959) provide the best existing diagnosis of Marionina. The seven marine species of Marionina found in the 1976-1979 British Columbia collecitons have all been small, less than 7 or 8 mm, and were opaque white or transparent when alive. Marionina vancouverensis, M. neroutsensis, M. sjaelandica, M. appendiculata, and M. subterranea were observed to have "marionine" anterior blood systems like that discussed by Giere (1974). These species form, along with other small intertidal species such as M. spicula (Leuckart, 1847), M. istriae Giere, 1974, and M. achaeta Lasserre, 1964, a morphologically consistent group within Marionina. Nielsen and Christensen (1959) transferred numerous poorly described enchytraeid species to Marionina. The North American Pacific coast species placed there were Enchytraeus modestus, a synonym for E. kincaidi; E. citrinus, E. metlakatlensis, and E. saxicola, of dubious status but unlikely to belong in Marionina; Enchytraeus multiannulatus and E. multiannulatoides, both good Enchytraeus species; Enchytraeus pugetensis, a synonym of Lumbricillus annulatus; and Michaelsena paucispina, a species of Grania. These generic modifications were not well founded and contributed to the heterogeneity and confusion within Marionina. (ref. ID; 5967)
Distribution
Cosmopolitan; marine intertidal (few species subtidal), limnic, and terrestrial. (ref. ID; 5967)
The species as described (Eisen 1904) is indeterminate. No genus of enchytraeid is known which has dorsal pores in II to IV and the spermathecae described are more typical of species of Buchholzia or Fridericia than of Marionina. (ref. ID; 5967)
Eisen (1904) described this species from a single specimen of an unstated level of maturity, which he reported to be in a poor state of preservation. Many taxonomically important characters were not described in detail. The testes are "entire" (Eisen 1904, p.93), which does not fit the generic definition of Lumbricillus (Nielsen and Christensen, 1959) in which it was placed both by Cernosvitov and Nielsen and Christensen. The species is of an indeterminate taxonomic status. (ref. ID; 5967)
The British Columbia specimens differ slightly from those originally described (Nielsen and Christensen 1959). The third pair of pharyngeal glands is usually very small and setal numbers are slightly higher in the British Columbia material. Additional to the original description, it was oberved by scanning electron microscopy that an external epidermal groove extends ventrally between the male orifices. Marionina appendiculata can be recognized most easily by its slightly sigmoid setae distributed in bundles of up to six, usually four or five, setae. No seminal vesicle is present and the tests and sperm funnels are very small. The penial bulb is characteristic, consisting, in the British Columbia specimens, of one, small, compact structure attached to the body wall immediately posterior to the male pore. (ref. ID; 5967)
Habitat
Coarse sand, fine sediments and amongst emergent vegetation near high water level. British Columbia specimens were found in upper intertidal silty, or poorly sorted sediments with a large percentage of fine particles. (ref. ID; 5967)
Marionina charlottensis Coates, 1980 (ref. ID; 5953 original paper, 5967) reported author and year? (ref. ID; 5790)
Descriptions
Small intertidal worms, fixed length 4 to 6 mm, 25 to 32, 35 segments, usually 26 to 29. Small cutaneous glands arranged in four or five transverse rows per anterior segments. Setae straight with an ental hook, robust, two per bundle, laterals lacking in II and rarely in III, XII lacking laterals and ventrals. Brain truncate posteriorly. Transition from oesophagus to intestine gradual, oesophagus without diverticula. Three pairs of septal glands present at 4/5, 5/6, and 6/7, anterior two pairs united dorsally, second with ventral lobes; posterior pair free, with large ventral lobes. Lymphocytes few, granulate, nucleate, ovoid or spindle shaped, about one-third length of setae. Chloragocytes dense from VI, occasionally, a few cells in V; large, with many refractile droplets, sometimes gold coloured, one-third to one-half length of setae. Dorsal vessel originating in posterior of XII or XIII. Blood colourless in fixed specimens. Nephridia present from 7/8 but not in all postclitellar segments. Anteseptale as long as one-quarter of whole organ, including funnel and a few coils of the nephridial canal, efferent duct short, wide, terminal. Clitellum extending over half XII to half XIII, incomplete ventrally, not thick; gland cells large, rectangular, in irregular transverse rows. Testes small, compact at 10/11. Seminal vesicle present, extending to X. (Female and male structures following are laterally paired). Sperm funnel regular, length just less than twice width; sperm collar prominent width just less than funnel width. Vas deferens narrow, irregularly coiled in XII. Penial bulb small, compact; male orifice large. Spermatheca simple, with distinct ectal and ental ducts. Ectal pore at 4/5, ventral to lateral setal bundles, duct surrounded by glands of irregular size, no basal rosette. Ampullar walls thick. Ampulla broadly connected to oesophagus at half IV by distinct ental canal. (ref. ID; 5953)
Remarks
Of other known Marionina species two, M. southerni (Cernosvitov, 1937) and M. argentea (Michaelsen, 1889), are consistently without setae in lateral bundles of II and have only two (rarely three) straight setae in other bundles with the exception of some clitellar segments. Of these two, M. southerni has a distinctive sperm distribution in the spermathecal ampulla and in this and other characteristics it is morphologically similar to a few other, small, intertidal Marionina species. Marionina charlottensis is morphologically similar to M. argentea, in particular, and, in general, to the group of species discussed in this paper with reference to M. vancouverensis. Marionina argentea has a distinct ring of small glands at the ectal orifice of the spermatheca and is not known to have a well-developed seminal vesicle. Nielsen and Christensen (1959) also described M. argentea as having numerous lymphocytes which result in its characteristic opaque while appearance. The fixed specimens of M. charlottensis have notably few lymphocytes. (ref. ID; 5953)
Etymology
From Queen Charlotte Sound, access to the Pacific Ocean west of Ocean Falls. (ref. ID; 5953)
Habitat
Northern inlets, intertidal habitats, less than maximum coastal salinities. Frequently associated with attached upper intertidal zone algae, i.e. Fucus, Bangia, and Entermorpha. (ref. ID; 5953)
Type locality
Ocean Falls on Cousins Inlet, British Columbia, 52 degrees 21.55'N, 127 degrees 43.15'W, 3.4 m above chart datum, in sand-slit with Bangia and Enteromorpha. Collected February 26, 1977, by K.A. Coates. (ref. ID; 5953)
Examined materials
Holotype: NMCIC1979-1601, a fully mature specimen, stained with Grenacher's alcoholic borax carmine, whole mounted in Canada balsam. (ref. ID; 5953)
Paratypes: NMCIC1979-1602a to 1979-1602d, four whole-mounted specimens from type locality and USNM 58914, four whole-mounted specimens from type locality. (ref. ID; 5953)
Other material: Alice Arm and Hastings Arm off Portland Canal (Dobrocky Seatech collection) and additional material from Ocean Falls, British Columbia (author's collection). (ref. ID; 5953)
Other than their smaller size, the British Columbia specimens are the same as Scandinavian material described by Nielsen and Christensen (1959, 1961). A careful study by Kossmagk-Stephan (1982) of European populations of M. subterranea and M. glandulifera has clarified the structural differences between the species. The differences between the spermathecae of M. subterranea and M. glandulifera can be seen in Figs.9 and 10. Marionina subterranea has no external gland cells along the spermathecal ectal duct whereas M. glandulifera has large gland cells along at least the ectal one-half of the duct. Kossmagk-Stephan (1983) described adhesive glands present from VII, replacing the dorsal setal bundles. In the British Columbia specimens epidermal adhesive glands were located just lateral to the ventral setal bundles. The seminal vesicle was unpaired and dorsal. The pattern of the anterio-dorsal blood vessels was marionine and the vas deferens exited lateral to the glandular penial bulb, as in M. nevisensis (q.v.). (ref. ID; 7204)
Habitat
High to low intertidal in coarse sediments; subtidal in sandy bottom of river estuary. (ref. ID; 7204)
Material examined
Collected at Elk Falls, British Columbia, 50 degrees 04.29'- 50 degrees 04.34'N, 125 degrees 16.40'W and in the Fraser River estuary at Steveston, British Columbia, by personnel of Dobrocky Seatech Ltd. (NMCIC1982-0778 and 1982-0779; USNM073895). (ref. ID; 7204)
Marionina klaskisharum Coates, 1983 (ref. ID; 7204 original paper) reported author and year? (ref. ID; 5790)
Descriptions
Thin, transparent specimens, cuticle not rigid. Mature specimens, with one or two eggs, with 34 to 40 segments; fixed, slightly compressed specimens 5.8 to 8.8 mm long, approximately 0.18 to 0.24 mm wide at XII, narrowing to 0.16 to 0.20 mm at V. Setate in (II), III and IV(V) as unisetal ventral "bundles", other segments lacking setae, epidermal pads, and setal sac. Setae 27 to 28 µm long, thin; broadening of slight ental hook, protruding only slightly above cuticle. Epidermal gland cell small, in regular transverse rows. Spermathecal pores midlateral at 4/5. Clitellum over XII and XIII, cells not very tall, with slightly irregular shape and distribution. Anterior part of brain dorsoventrally compressed and narrow, posterior part thickened, approximately twice as wide as anterior part, incised. Dorsal blood vessel originating near 15/16 or 16/17; dorsal vessel exending foward under brain then branching into a pair of circumpharyngeal vessels in the prostomium, branches extending posterioventrad uniting as ventral vessel. Pharyngeal glands paired at 4/5, 5/6, and in VI and VII, joining dorsally at 4/5 and 5/6, all pairs with ventral lobes, ventral lobes extending through VI into VII massive. Transition of oesophagus to intestine gradual, no oesophageal or intestinal diverticula. Coelomocytes few, flattened oval, nucleate. Chloragocytes dense from VII or VIII, containing many golden-brown refractile granules. Anteseptal part of nephridium large, including funnel plus obvious canal, funnel comprising one-third of anteseptal part. Whole anteseptale about one-fifth total length of nephridium. Postseptale constricted at septum, without a distinct transition to efferent duct. Nephridia only in some of postclitellar segments. Seminal vesicle well-developed, single, dorsal, extending anteriad from 10/11, usually as far as 9/10. Sperm funnel length 2 to 4 times width, glandular part very granular. Collar narrower than funnel, pronounced; glandular cells differently developed around duct. Vas deferens narrowing from glandular part to male pore, moderately long; irregularly coiled in XII. Penial bulb with a large glandular expansion medio-dorsal to internal male bursa, vas deferens runs along anteriolateral part of bulb, exiting separately, via small pore, ino bursa. Spermatheca in V, attached to oesophagus in anterior one-half of V. Ampulla more or less oval with some small, irregular lobes along ectolateral edges; ental duct not distinct from unlobed ental part of ampulla. Sperm heads embedding in walls of small diverticula, tails extending into cavity of ampulla. Ectal duct length just less than that of ampulla plus ental duct; ectal duct surrounded by small, irregular gland cells; a few glands near ectal pore longer than the rest. (ref. ID; 7204)
Remarks
The most distinctive, superficial characteristic of Marionina klaskisharum is the distribution of its setae in unisetal bundles in only segments II to V. Marionina preclitellochaeta Nielsen and Christensen, 1963, M. subachaeta Shurova, 1979, M. oligosetosa Kossmagk-Stephan, 1983, and M. subtilis (Ude, 1986) are also characterized by restriction of the ventral setae of a few anterior segments and by the absence of dorsal bundles. The species can be distinguished on the basis of the characteristics shown in Fig.5. Shurova (1979) dos not specify that there are two setae in each of the few setal bundles present in M. subachaeta but she does not mention a difference in setal numbers in her brief comparison of M. subachaeta to M. preclitellochaeta. Marionina weilli Lasserre, 1964 and M. welchi Lasserre, 1971 hav both bisetate and unisetae bundles; however, their overall setal distribution are unique. The typical marionine form of the nepharidia has the preseptal part including the nephrostome (funnel) plus a part of the nephridial canal. In M. klaskisharum and M. oligosetosa the unpaired seminal vesicles are dorsal. The presence of a seminal vesicle in M. preclitellochaeta was doubtful according to Nielsen and Christesen (1963) but Lasserre (1971) confirmed its presence. In M. subtilis sperm morulate float freely in the coelom. (ref. ID; 7204)
Etymology
Derived from the name of the type locality, Klaskish River, B.C. (ref. ID; 7204)
Habitat
Near high tide driftline or at emergent vegetation, in coarse clean to silty sand sediments usually with wood debris; freshwater present. (ref. ID; 7204)
Type materials
Holotype: NMCIC (the Invertebrate Collection of the Canadian National Museum) 1982-0774, from the Klaskish River estuary, Vancouver Island, 50 degrees 15.94'N, 127 degrees 43.8'W, in sand-silt with wood debris and emergent vegetation, collected on July 26, 1976 by the author. (ref. ID; 7204)
Patatypes: NMCIC1982-0775, one specimen from the type location; USNM (the United Stated National Museum) 073893, from the type location; and USNM 073894, from Watson's Bay, Tribune Channel, 50 degrees 50.9'N, 126 degrees 17.9'W, small stream, coarse granite sand with wood chips and other debris, collected March 22, 1980 by the author and H.R. Baker. (ref. ID; 7204)
Other material: Collected from sites between 50 degrees 15'N and 53 degrees 04'N, from Vancouver Island, Queen Charlotte Islands, and the British Columbia mainland. (ref. ID; 7204)
Marionina neroutsensis Coates, 1980 (ref. ID; 5953 original paper, 5967) reported author and year? (ref. ID; 5790)
Descriptions
Small, intertidal worms, 2 to 4 mm, 26 to 33 segments, usually 27 to 29. Cutaneous glands arrayed in 10 or 11 transverse rows per segment. Setae straight with ental hook, two per bundle throughout, rarely three, ventrals and laterals lacking in XII. Posterior setae just slightly longer than anterior. Brain concave posteriorly, lateral margins converge anteriorly, length approximately 1.6 times width. Transition from oesophagus to intestine gradual oesophagus without diverticula. Three pairs septal glands present, anterior two united at 4/5 and 5/6 without ventral lobes, third pair free, with elongate ventral lobes in VI. Lymphocytes coarsely granulate, nucleate, ovoid to disc-shaped, about one-sixth length of setae. Chloragocytes small, pale golden brown or colourless, dense from VI. Dorsal vessel originating at 12/13 or in XIII. Blood colourless both in living and fixed specimens. Anteseptale large, including funnel plus a few loops of nephridial canal, comprising one-third total of length of nephridium excluding efferent duct. Efferent duct posteroventral not precisely demarcated from postseptale. Clitellum extending over XII to half XIII, gland cells rectangular, in almost regular rows, incomplete ventrally. Seminal vesicle present, extending forward to X. The following male and female structures are laterally paired. Sperm funnel tapering toward vas deferens (somewhat pear shaped), length approximately one and a half times maximum width, length approximately two-thirds body width at XI. Sperm collar width equal funnel but sperm attaching only in an area at middle of collar. Vas deferens coiled in XII. Penial bulb small, depressed. Spermatheca simple. Ampulla round, walls not very thick, connected to oesophagus in posterior one-half of V very short ental duct. Ectal duct distinct, surrounded along entire length by small, fused (?) glands, no distinct rosette. Ectal orifice posterior to septum 4/5 and just ventral to the line of the lateral setal bundles. Only one mature egg present at a time. (ref. ID; 5953)
Remarks
As for the two Marionina species already described, M. neroutsensis is morphologically similar to a group of species including M. istriae Giere and M. argentea (Michaelsen). Marionina argentea, which is usually restricted to supralittoral or terrestrial habitats, is characterized, as previously indicated, by the absence of lateral setae in II. Marionina istriae is considerably larger than the other two species and as well is distingished by its spermathecal morphology. Another species, M. cana Marcus described by Marcus (1965) from a brackish beach habitat, with "soil-like" sediments is also morphologically similar to M. neroutsensis. However, the origin of the dorsal blood vessel of M. cana is preclitellar (IX) and this species is described as having oesophageal diverticula like those of M. simillima Nielsen and Christensen. This characteristisc structure is not illustrated by Marcus (1965, Fig.24). Marionina neroutsensis was reported (Coates and Ellis 1980) as Marionina n. sp. No.2 in an Enchytraeid species association near a pulp mill discharge. (ref. ID; 5953)
Etymology
From Neroutsos Inlet, British Columbia. (ref. ID; 5953)
Habitat
Upper intertidal, finer sediments, saturted with water. (ref. ID; 5953)
Type locality
Neroutsos Inlet, near Port Alice, British Columbia, 50 degrees 31.12'N, 127 degrees 37.10'N at high water mark. Collected May 2, 1978, by K.A. Coates. (ref. ID; 5953)
Examined materials
Holotype: NMCIC1979-1596, a fully mature specimen, stained with Grenacher's alcoholic borax carmine and mounted whole in Canada balsam. (ref. ID; 5953)
Paratypes: NMCIC1979-1597 one whole specimen in alcohol, NMCIC1979-1598a to 1979-1598d, four whole-mounted specimens, as holotype; all from type locality. (ref. ID; 5953)
Other material: Author's collection; 48 whole specimens from type locality and 100+ whole specimens collected at other locations along the shores of Neroutsos Inlet. (ref. ID; 5953)
The original specimen of Marionina nevisensis was from Fort Charles, south of Charlestown on Nevis Island in the Caribbean. It was collected in June 1949 and no subsequent records are known. In spite of the large geographic displacement of the original and the British Columbia specimens no substantial structural differences could be found. Unfortunately, I was unable to borrow the origial material but the specimens collected in British Columbia allow the addition of a few details and extensions to the ranges in the original description (Righi & Kanner 1979).
The British Columbia specimens were nematodelike probably because of a thikened (3-4 µm) cuticle. Mature fixed specimens, 30-39 segments, were 3.0-7.7 mm long; the Caribbean specimens had 40 segments and was 4.5 mm long. In the British Columbia specimens the patten of the anteriodorsal blood vessel was marionine, bifurcating in III or IV, then extending anteriad as a loop on either side of the pharynx and brain. Glandular proliferations were present on the ventral nerve cord at the ganglia of XIII, XIV, and frequently XV. The single, dorsal seminal vesicle extended as far anteriad as VII. The glandular sperm funnel ranged from two to four times as long as wide and the glandular cells were differently developed resulting in an eccentric duct location. The vas deferens was closely applied to the lateral surface of the compact penial bulb but opened separately into the small male bursa. The glands of the bulb opened directly into the male bursa. The ectal duct of the spermatheca was covered along its entire length by gland cells but those originating at the orifice were longer and more prominent than the rest. Three species of Marionina without setae are known to live in upper intertidal and supralittoral beach habitats. Marionina achaeta Lasserre, 1964 was recorded (Lasserre 1964, 1966) from the Atlantic coast of France and subsequently recorded from the United States (North Carolina and Massachusetts), North Africa, numerous locations in Europe (Lasserre 1971), Bermuda (Lasserre and Erseus 1976) and Ireland (Healy 1977, 1979). Marionina arenaria Healy, 1979 is reported from sand dunes and storm beach habitats of Ireland (Healy 1979). The spermathecae of M. arenaria are not attached to the oesophagus.
Differences exist between M. nevisensis and M. achaeta in gland cell distributions around their spermathecal ectal ducts, in origins of the dorsal blood vessels and in the form of the seminal vesicles. The degree of structural difference between M. arenaria and M. achaeta is similar to that found between M. nevisensis and M. achaeta (Righi and Kanner 1979). The sympatric occurrence and subsequent specific classification (Healy 1979) of the former species requires a consistent, specific, taxonomic status for M. nevisensis. (ref. ID; 7204)
Habitat
Sandstone and coral sand, intertidal in the Caribbean; upper to lower intertidal usually in coarse, mixed gravel-sand-silt or shell sediments, usually with Fucus Bangia, Enteromorpha or wood debris in British Columbia. (ref. ID; 7204)
Material examined
Collected from numerous sites between 49 degrees 25'N and 53 degrees 03'N, on Vancouver Island, Queen Charlotte Islands, and the British Columbia mainland. (ref. ID; 7204)
Marionina pituca Righi, 1974 (ref. ID; 7620 original paper) reported author and year? (ref. ID; 5790)
Descriptions
The species is one of the smallest of the genus. The number of segments is from 23 to 25. There is no pigment. Under reflected light the granules of the coelomocytes and mainly those of the chloragogen cells present a milk white colour. Under transmitted light the granules are highly refractile with dark edges, and on the whole they appear ash coloured. The prostomium is triangular in outline, its length is somewhat less than half its basal breadth. The head pore lies on the apex of the prostomium. Dorsal pores are wanting. The clitellum occupies segments 12-1/2 13. It is saddle-shaped, opened ventrally, and the glandular cells are disposed in transverse rows. There are two setae per bundle. Dorsal and ventral bundles occur in segments 2-6, from 7 backwards there are only the ventral ones. The setae are straight, without nodulus, the inner end is curved and the outer one is single pointed. The length of the preclitellar ventral setae varies from 20-23 µm, and that of the postclitellar ones from 28-31 µm. The cerebral ganglion is about three times longer than large. The posterior margin has a median incision separating rounded lobes. The lateral margins converge slightly forward, and the anterior border is convex. On each side of the anterior margin arises an egg-shaped structure from wich depart the circumpharyngeal connectives. In resting animals, and during slow advancing, these structure stay side by side, covering the anterior third of the ganglion, so that it is perfectly seen only while it struggles. In segments 2-4 the ventral nerve-cord appears as a large rectangular plate, individulalized ganglion occur from segment 5 backwards. Associated with the pharygeal mass there is a pair of post-pharyngeal bulbs. Pharyngeal and esophageal peptonephrida are wanting. The transition between esophagus and intestine is gradual, without appendages. The intestine is slightly sinuous in 8. Chloragogen cells are scatter from 8 backwards. In living animals slightly compressd by the cover glass, the movements of the intestine dislodge the granules of the chloragogen cells to the sides, where they force the cellular membranes to take a hemispherical shape. The limits of the chloragogen cells are not recognizable on the surface, it appears that they are arranged transversely on the intestine. Septal glands are attached to the anterior face of the septa 4/5-6/7. The dorsal lobes of the symmetrical glands of 4 and 5 are confluent over the dorsal vessel. In 5 and 6 there is a ventral finger-shaped lobe on each side of the esophagus. The blood is colourless. The dorsal vessel begins in segment 12. One pair of commissural dorso-ventral vessel is immediately anterior to the septal glands of 4. Two other pairs of commissural vessels have their common origin in 3. In front of these commissures the diameter of the dorsal vessel diminishes to half the posterior one. The coelomocytes are discoid and flattened with the plasma full of refractile granules. Their length varies from 10 to 13 µm. Pairs of nephridia are attach to the septa 6/7, 7/8, 14/15, 16/17, and 20/21. There are alike throughout the body. The small preseptal portion contains the funnel and a few loops of the nephridial canal. The cylindrical and voluminous postseptal portion is five times longer than the preseptal one. The short excretory duct arises posteriorly, its canal opens into a terminal vesicle. The nephridiophores lies in line with setae ab. The rounded or slightly egg-shaped testes are linked to septum 10/11. The sperm funnels are conical, their wide collar is separated from the glandular portion by a strong constriction. In each funnel the length is a little greater than the breadth. The vasa deferentia are short, they bend irregularly into segment 12. In irritated worms each duct winds like a spring. The vasa deferetia open directly on the surface, without a penal bulb. Glandular club cells open around the male apertures. Spermathecae are wanting. In segments 12-13 of several specimens there is one egg full of yolk and some others in differentiation. (ref. ID; 7620)
Remarks
Due the combination of straight setae without nodulus and few per bundle, absence of peptonephrida and esophageal chambers, dorsal vessel starting from the clitellar segments, short antseptal portion of the nephridia with the funnel and few coils of the canal, and compact testes, the present species is classified as Marionina. From the 48 known species, 12 are recorded from South America and regions zoogeographically related (Michaelsen 1888, 1905, 1907; Ude 1896; Benham 1905, 1922; Stephenson 1932; Marcus 1965), and all come from marine environments. The genus is now for the first time indicated from the South American soil. Marionina pituca is distinguished from the congeneric species by the position of the head pore, shape of the clitellum, absence of spermathecae and disposition of the setae. (ref. ID; 7620)
Type locality
Samples of moist soil (Serra do Cipo (about 19 degrees 30'S - 43 degrees 45'W), state of Minas Gerais, Brazil) under dead leaves in a gallery forest in September 1972. (ref. ID; 7620)
Deposited materials
The Department of Zoology, University of Sao Paulo (ZU-270). (ref. ID; 7620)
Measurements
Mature animals: Its length varies from 2 to 3 mm, the diameter of the clitellum from 132 to 158 µm, and in the middle body region from 80 to 100 µm (n=20). (ref. ID; 7620)
Fixed length 3 mm; segments 26. Body reddish when living. Chaetae sigmoid, 2-4 per bundle, mostly 3. Epidermal gland cells 2-4 rows per segment, reddish when living. Clitellum over 1/2XI-1/2XIII, with gland cells irregularly arranged. Brain deeply incised posteriorly. Only first pair of septal glands united dorsally. Dorsal vessel originating in XIII, with reddish blood when living. Efferent ducts of nephridia originating from posterior of postseptal parts. Coelomocytes spindle-shaped and abundant. Sperm funnels cylindrical, twice as long as width, with collar narrower than funnel. Vasa deferentia confined to XII. Penial bulbs large. No seminal vesicles. The connection between spermatheca and oesophagus rather thin. Spermathecal ampullae oval, ca. 26 µm in length, ca. 18 µm in maximum width. Ectal ducts ca. 90 µm long and 10 µm wide, with a gland at ectal pore. (ref. ID; 6650)
Remarks
In specimens described by Neilsen & Christensen (1959), the spermathecal ampullae are larger than those of our specimens and the clitellar gland cells are arranged in transverse rows. (ref. ID; 6650)
Material examined
Five whole mounts and two live specimens from A1, A3, A4, A5, A6. (ref. ID; 6650)
This littoral species was originally described by Claparede from specimens found on the Island of Sky in the Hebrides. The description given, though not complete, is sufficient to characterize the species. It has not been recorded since, till I found it on Lambay (tom, cit.). It seems desirable to complete the description. The Irish specimens were 10 mm long, the Scotch 18-25 mm. Claparede gives 8-10 mm. The colour is reddish-yellow. Red glands were sometimes present on the epidermis. The clitellum is composed of close-set glands, and occupies the 12th, and adjacent parts of the 11th and 13th segments. There are 4-5 setae in a bundle. The brain is somewhat concave before and behind, longer than broad, and much broader behind than in front. The ventral ganglia of the anterior segments are kidney-shaped, and very large, as is often the case in this genus. Small oval copulatory glands occur on the 13th, 14th, 15th, and 17th segments, or in some of them. The male organs and spermathecae agree closely with Claparede's figures. There are five pairs of large septal glands in the 4th-7th segments, those on the 6th and 7th being the largest. In the 5th segment there is a dorsal and a ventral pair. The dorsal vessel rises in the 13th segment. The peritoneal cells of the gut are filled with dark contents. The penial bulbs are large and cylindrical. (ref. ID; 1928)
A few morphological differences, probably attributable both to geographic separation and to genetic, infrasubspecific variability, exist between the British Columbia specimens and the originally described European specimens (Nielsen & Christensen 1959). In the British Columbia populations, M. sjaelandica has more segments than in European populations; there is, consistently, a small ventral septal gland lobe in V of British Columbia specimens; and mature specimens of M. sjaelandica found in British Columbia have a well-developed ventral gland on the nerve cord in XIV, a condition sometimes reported for M. southerni (Southern, 1909). Because of the essential identity of the reproductive organ morphologies of British Columbia and European specimens these are considered to be the same species. Marionina sjaelandica is the only intertidal Marionina species reported from British Columbia that has numerous (M. vancouverensis rarely has one or two) discrete, round sperm bundles embedded in the walls of the spermathecal ampulla; this combined with its large seminal vesicles and bisetate condition make it easy to distinguish. Marionina sjaelandica was often the most abundant species in polluted, estuarine localities. (ref. ID; 5967)
Habitat
Beaches, upper intertidal in coarse sediments with abundant decaying algae; estuaries, in finer sediments. (ref. ID; 5967)
Marionina southerni was variously described prior to its revision by Nielsen and Christensen (1959). The consistent absence of lateral setal bundles in II (and occasionally III) was confirmed by Nielsen and Christensen (1959). Setae in the anterior bundles of the specimens from British Columbia were usually shorter than those in posterior bundles, 28 to 50 µm and 49 to 55 µm, respectively. Knollner (1935) also noted (for Fridericia pseudoargentea) that the posterior setae were, in general, longer than the anterior ones. Nielsen and Christensen described M. southerni as having large well-developed seminal vesicles. The size of the seminal vesicle was quite variable in specimens found in British Columbia; however, free sperm morulae were not seen. The specimens of Nielsen and Christensen were larger than those found in British Columbia, the populations having 28-36 and 24-32 segments, respectively. The individuals of the population described by Knollner (1935) had 18-34 segments. Stephenson (1911) noted (for Enchytraeus nodosus) an anterior, lumbricilline bifurcation of the dorsal blood vessel, also a characteristic of the British Columbia specimens and the single specimen from the Swale River estuary, England. The description by Stephenson of the glandular sperm funnel with its lumen nearer the medial side coincided with observation on the present material. Viewed end on the funnel was kidney-shaped, the gland cells being much shorter along one side. It was noted from the British Columbia material that the vasa deferentia penetrated the compact penial bulbs just ental to the small male bursae and that the gland cells emptied into the terminal part of the vas deferens. (ref. ID; 7204)
Habitat
Mid to upper intertidal in coarse mixed sediments, with wood debris and other plant material; sometimes where freshwater runs over the sediments. (ref. ID; 7204)
Material examined
Specimens were collected in British Columbia from numerous sites between 49 degrees 41'N and 55 degrees 28'N, on the mainland, Vancouver Island, and the Queen Charlotte Islands. (ref. ID; 7204)
It is almost invariably to be found in the soil of moors and hills above 500 feet. Specimens are very rarely found in the mature stage. The length varies from 5 to 20 mm. Setae never more than three. The egg-sac is very large. In immature forms, the intestine is usually covered with large cells full of oil-drops. The blood is usually only very faintly coloured, and in some cases is quite colourless. (ref. ID; 1928)
Marionina subterranea (Knollner, 1934) (ref. ID; 6655) or 1935 (ref. ID; 5967, 7204) reported author and year? (ref. ID; 5790)
Remarks
The specimens found in British Columbia differ slightly from those described by Nielsen and Christensen (1959) in having more developing eggs at one time. Nielsen and Christensen (1959) made no comment regrading ventral nerve cord glands in this species. Small glands were found on the ventral nerve cord in XIII and XIV of the British Columbia specimens. Marionina subterranea can be quickly recognized by the absence of lateral setae and presence of ventral setal bundles with only two setae. Detailed observations of the spermathecal morphology confirm the identification. K. Kossmagk (personal communication) is preparing a significant revision of Marionina subterranea in which is synonymy with Michaelsena glandulifera will be reevaluated. (ref. ID; 5967)
Details regarding setal distribution, nephridial shape, dorsal blood vessel origin and the structure and shape of the spermathecae of the specimens examined from British Columbia and from New Jersey were the same as those of Marionina subterranea (Knollner) s.s. The setae of the specimens examined were of a more or less equal size throughout. Knollner (1935) and Kossmagk-Stephan (1983) found that the posterior setae could be one and one-quarter to one and one-third times as long as the anterior setae in M. subterranea from Kiel Bay and the Isle of Sylt. The connection of the posterior pair of pharyngeal glands is apparently a variable characteristic ranging from narrow (Kossmagk-Stephan 1983) to absent (personal observation and Knollner 1935). A single, dorsal seminal vesicle was observed in the material from British Columbia and New Jersey; Knollner (1935) observed no seminal vesicle. Previously undescribed details of the dorsal anterior blood vessels and male pore apparatus may prove to be important systematic charateristics. The anteriodorsal blood vessel pattern observed in the material examined from British Columbia and New Jersey is the type described as marionine (Giere 1974; Coates and Ellis 1981). The vasa deferentia exit lateral to the associated, compact, glandular bulbs into small male bursae as in M. nevisensis. The gland cells may empty either into the terminal part of the associated vas deferens or directly into the bursa. (ref. ID; 7204)
Habitat
Littoral and sublittoral in sand and other coarse sediments. (ref. ID; 5967)
On the lower part of the beach at groundwater level; upper to lower intertidal in organic sand-silts or muds; subtidal to 37 m in medium fine sand. (ref. ID; 7204)
Material examined
Specimens were collected intertidally in British Columbia from 49 degrees 52'N to 54 degrees 19'N on Vancouver Island and the mainlnad coast. A single specimen (USNM 073900) received from R. Diaz was from the Atlantic, off the coast of New Jersey, 39 degrees 14.5'N, 74 degrees 06.1'W at a depth of 33-37 m. (ref. ID; 7204)
Long, thin intertidal worms, fixed length approximately 6 to 7.5 mm, 35 to 44, usually 39 to 42, segments. Setae slightly sigmoid with an ental hook, present dorsally and ventrally from II. Distribution: preclitellar lateral (2), preclitellar ventral (2-3), postclitellar lateral (2), and postclitellar ventral (2). Only ventrals missing in XII. Brain slightly concave posteriorly, lateral margins converge anteriad, length approxismately one and a half times width. Transition from oesophagus to intestine gradual, oesophagus without diverticula. Peptonephridia absent; however, there is large mass of solid tissue extending posterodorsad from the pharyngeal pad in IV. Three pairs of septal glands, present, 4/5 to 6/7, united dorsally at 4/5, 5/6, and 6/7, all pairs with distinct, almost separate, ventral lobes. Lymphocytes granulate, nucleate, spindle shaped, greater than one-half length of setae. Chloragocytes dense from VII, cells not granulate height of posterior cells just greater than length of setae. Dorsal blood vessel originating near half XIII. Blood not coloured in fixed specimens. Nephridia present from 6/7, absent in clitellar and many postclitellar segments. Anteseptale including nephrostome only, postseptale with distinctly demarcated, terminal ectal duct, duct almost as long as postseptale. Clitellum extending over posterior of XI to XII, gland cells arrayed in irregular transverse rows rows. Seminal vesicle well developed, extending posteriad from X to XIV. Following male and female structures are laterally paired. Sperm funnel length just greater than twice width. Sperm collar equalling funnel width. Vas deferens narrow, irregularly coiled in XII. Penial bulb very small and depressed. Spermatheca not attached to oesophagus but confined to V, ectal orifice at IV/V. Ectal duct uniformly narrow, muscular (?), aglandular, with four large, hyaline glands at ectal end. Ampulla of spermatheca consisting of two, sequential, sphaerical or heart-shaped chambers, the distal with thick walls and no distinct cavity and the proximal with a thin wall and spermatozoa arranged in an approximately regular central ring. Three or four eggs present in body at one time, egg sac extending posteriad to XVI. (ref. ID; 5953)
Remarks
Marionina trevori, unlike the other new species, possesses some characteristics that are uncommon or atypical for Marionina species. A few species (M. aberrans Finogenova, 1973, M. cambrensis O'Conner, M. tubifera Nielsen and Christensen) have the nephridial anteseptale consisting only of the nephrostome. A very few species (M. libra Nielsen and Christensen and M. mesopsamma Lasserre) have free spermathecae. A "post-pharyngeal bulb" (Righi 1974) occurs in M. pituca Righi but no details of its function or structure are available. Marionina trevori, however, lacks oesophageal and intestinal diverticula, peptonepharidia, and noduli on the setae, and has a gradual oesophageal-intestinal transition, a terminal nephridial efferent duct, and setal shape and distribution which, in toto, preclude it from all enchytraeid genera other than Marionina. A well-developed seminal vesicle extending posteriad from X is without previous record in Marionina. Within Marionina, M. trevori is distinctly characterized by its spermathecal morphology. (ref. ID; 5953)
This unusual Marionina species can be readily distinguished from the other intertidal Marionina species found by its large, pre- and post-clitellar seminal vesicle and unusual spermathecal morphology, a bipartite ampulla, and a "muscular", naked ectal duct. The anterior blood vascular system of M. trevori is of the type described by Giere (1974) as "lumbricilline". It was previously incorrectly described as of the typical "marionine" type (Coates 1980, p.1316). (ref. ID; 5967)
Etymology
For Trevor A. Smyth who collected the type material. (ref. ID; 5953)
Habitat
Upper intertidal, i n coares sand mixed with an under decaying algal debris, near a sewage outfall (beach nearby posted as polluted by domestic effluent). (ref. ID; 5953)
Type locality
Qualicum Beach, Vancouver Island, British Columbia, 49 degrees 21.38'N, 124 degrees 26.50'W, at 4.8 m above chart datum (approximate L.W.) in coarse sand mixed with decaying algal debris. Collected June 10, 1976, by Mr. Trevor A. Smyth. (ref. ID; 5953)
Examined materials
Holotype: NMCIC1979-1599, a fully mature specimen, stained with Grenacher's alcoholic borax carmine and mounted whole in Canada balsam. (ref. ID; 5953)
Paratypes: NMCIC1979-1600, seven whole-mounted specimens (one slide) and USNM58913, four whole-mounted specimens, as holotype; all from type locality. (ref. ID; 5953)
Other material: Author's collection, 15 whole specimens in alcohol, from type locality. (ref. ID; 5953)
In consideration of the recognized heterogeneity of Marionina a detailed description of the specimens' characteristic is given. Small, intensely white, intertidal oligochaetes, living specimens as long as 6 mm, fixed, only 2 or 3 mm, 28, 31-38 segments. Cutaneous glands in three to four distinct transverse rows per segment. Setae straight or slightly curved with ental hook, present dorsally and ventrally from II. Distribution: preclitellar lateral (1-4), preclitellar ventral (2-4); postclitellar lateral ((1) 2-3 (4)), and postclitellar ventral (2-4). Inner seta(e) of bundle frequently shorter than outer. Setae absent from XII, ventrally and laterally. Brain incised or concave posteriorly, suboesophageal ganglion very large. Transition from oesophagus to intestine gradual, oesophagus without diverticula. Three pairs of septal glands, the two anterior pairs united dorsally at 4/5 and 5/6 sometimes with small ventral lobes, medium-sized ventral lobes in VI. Lymphocytes ovoid, approximately equal to half of setal length, nucleate, grandulate, very abundant; live specimens as a consequence are opaque white. Chloragocytes present from IV or V, dense from VI. Dorsal vessel originating in XII to XIV, blood colourless. Nephridia present from 6/7. Anteseptale including funnel plus a few coils of the nephridial canal, postseptale pear shaped, wide efferent duct arising posteroventrally. Clitellum extending from XII to half XIII, gland cells large, rectangular, in transverse rows, incomplete ventrally. Seminal vesicles absent, small testes attached at 10/11. Following structures of male and female systems are laterally paired. Sperm funnel length approximately twice width. Outline irregular. Sperm collar approximately one-third to one-half width of funnel. Vas deferens short, wide, length one and a half to twice length of funnel. Penial bulb relatively small, compact. Often a small, eternal ridge is visible ventro-lateral to the male pore. Spermatheca simple. Ectal duct surrouded by small gland cells diminishing in size distally, canal distinct. Ampulla sac shaped or more regularly rounded, thick walled. Attached to oesophagus by short ental canal. Sperm present in the ampulla in an irregular mass. Spermathecal pore in intersegmental groove 4/5. As many as two mature eggs present at one time. (ref. ID; 5953)
Remarks
Marionina vancouverensis is morphologically related to M. spicula (Leuckart, 1847), M. brevis Finogenova, 1972, M. istriae Giere, 1974, and M. sublitoralis Erseus, 1976. Marionina spicula, a widely distributed species in upper to lower intertidal habitats of sandy beaches, is recognized by the large gland at the orifice of the spermathecal duct. The absence of this large gland is not a variation recorded in populations of M. spicula (Nielsen and Christensen 1959; Lasserre 1971; Erseus 1976). Finogenova (1972) described M. brevis from the Dnieper-Bug Estuary, U.S.S.R. The spermathecal ampulla is very large and its ectal orifice is near the middle of V; the vas deferens is relatively short, as in M. vancouverensis and M. spicula, but is narrow not wide (Finogenova 1972, Fig.37), Marionina istriae was recorded (Giere 1974) from the marine supralittoral of an island in the Adriatic Sea, just off the coast of Jugoslavia. It is a larger species than M. vancouverensis and differs considerably with regards to setal distributions and sperm funnel morphology. Marionina sublitoralis described (Erseus 1976) from subtidal shell-sand off the western coast of Norway is a very small species (2 mm) with a distinct setal distribution and shape. Marionina vancouverensis was reported (Coates and Ellis 1980) as Marionina n. sp. No. 1 in an Enchytraeid species association near a pulp mill discharge. (ref. ID; 5953)
Marionina vancouverensis Coates is similar to M. spicula (Leuckart) but can distinguished from it and the small, intertidal Marionina species found in British Columbia by characteristics shown in Fig. 4 and discussed by Coates (1980). (ref. ID; 5967)
Etymology
From Vancouver Island, British Columbia. (ref. ID; 5953)
Habitat
Intertidal estuarine, from lower to upper tide levels; less abundant in the upper intertidal. In sand under stones and gravel. (ref. ID; 5953)
Type locality
Tsitika River Estuary, British Columbia, 50 degrees 28.92'N, 126 degrees 34.98'W, 3.0 m above chart datum (approximate low water level (L.W.)), in sand-stone sediments, Fucus zone. Collected July 18, 1976, by K.A. Coates. (ref. ID; 5953)
Examined materials
Holotype: NMCIC1979-1594, a fully mature specimen, stained with Grenacher's alcoholic borax carmine and mounted whole in Canada balsam. (ref. ID; 5953)
Paratypes: NMCIC1979-1595a, 1979-1595b, two whole-mounted specimens from the type locality and USNM58911 two whole-mounted specimens from the type locality. (ref. ID; 5953)
Other material: Author's collection and USNM. Numerous specimens from Cayeghle Estuary and other sites on Neroutsos Inlet; Ocean Falls; Tsitika River Estuary. (ref. ID; 5953)
