Olavius cornuatus sp.n. (Oligochaeta, Tubificidae) from Georges Bank (NW Atlantic)

Olavius cornuatus sp.n. is described from Georges Bank off Massachusetts. It is distinguished from other species of Olavius Erséus, 1984 by its curved, horn-shaped atria ending in large, protrusible penes and up to six ectally hooked penial setae per bundle.     

Taxonomic Revision of the Marine Genera Bathydrilus Cook and Macroseta Erséus (Oligochaeta, Tubificidae), with Descriptions of Six New Species and Subspecies

Macroseta Erséus, 1975 is considered a junior synonym to Bathydrilus Cook, 1970 after examination of the type material of B. asymmetricus Cook, 1970. Bathydrilus is defined mainly by its organization of the male duct: vas deferens entering ectal half of atrium, one prostate gland attached to ectal part of atrium, near entrance of vas deferens, another prostate attached to apical, ental end of atrium. B. atlanticus sp.n., B. hadalis sp.n., B. meridianus sp.n., B. graciliatriatus sp.n. and B. longus sp.n. are described from specimens collected from various parts of the world; most of the species are deep-sea forms. Phallodrilus rohdei Jamieson, 1977 and Phallodrilus adriaticus Hrabe', 1971 are transferred to Bathydrilus , and a Caribbean/Bermudian form is described as B. adriaticus trisetosus subsp.n. The taxonomy and morphology of the different species, including B. rarisetis (Erséus, 1975) comb.n., are discussed. Bathydrilus is included in the sub-family Phallodrilinae. It appears closely related to Phallodrilus Pierantoni, 1902, from which it is distinguished principally by differences in the relative positions of the vasa deferentia, the atria and the prostate glands.     

Six new species of marine Tubificidae (Oligochaeta) from the continental shelf off Peru

Six species of marine tubifieids are described from the continental shelf off Peru. Two of them are members of the gutless genus Olavius Erséus, 1984 (subfamily Phallodrilinae). Olavius bullatus sp.n. possesses 2 pairs of large penial setae in voluminous copulatory sacs, tiny atria, and spermathecal pores in line with dorsal setae. Olavius crassitunicatus sp.n. is characterized by small atria with thin muscular layer, spermathecae with short ducts, opening dorso-laterally, and lack of penial setae. Four species arc members of the subfamily Limnodriloidinae. Limnodriloides busilicus sp.n. belongs to the appendiculatus-group (sensu Erséus). It is discriminated by somatic setae with subdental ligaments, and its voluminous elongate prostatic pads. Limnodriloides clavellatus sp.n. is distinguished from its congeners by a peculiar bulge in the cavity of each atrial ampulla, and spermatozeugmata imbedded in the walls of the spermathecae. Tectidrilus intermixtus sp.n. is similar to T. bori (Righi & Kanner, 1979); it is distinguished from the latter by having trisetal bundles in segment V or V-VI and by lacking copulatory glands. Marcusaedrilus peruanus sp.n. is characterized by nongranulated atrial ducts and bipartite spermathecae.     

Redescriptions of Phallodrilus parthenopaeus Pierantoni and P. obscurus Cook (Oligochaeta, Tubificidae)

The type species of Phallodrilus Pierantoni, 1902, P. parthenopaeus Pierantoni, 1902; is redescribed on the basis of new material from W Scotland. The species is characterized by its specific spermathecal and penial setae, its very slender atria, and its long spermathecal ducts. Sexually fully mature specimens of P. obscurus Cook, 1969 are described for the first time, on the basis of material from New Jersey (E USA). The new material shows that this species possesses 5 to 9 penial setae in each ventral bundle of segment XI, and that its atria are coated with a relatively thick musculature.     

A generic revision of the Phallodrilinae (Oligochaeta, Tubificidae)

The generic classification of the subfamily Phallodrilinae (with a total of 252 species) is revised according to phylogenetic principles. Morphological character patterns are assessed under the principle of maximum parsimony, but due to the poor resolution and instability in the resulting cladograms, the analysis is supplemented with intuitive weighting of synapomorphic similarities. Thirty-one genera are recognized. The former definition of the type genus Phallodrilus Picrantoni, 1902, is revised and the members of this genus are reclassified into 20 taxa, each hypothesized to be monophyletic: Phallodrilus s.str . (revised to contain the type species only), Somalidrilus gcn.n., Milliganius gen.n., Phallodriloides gen.n., Albanidrilus gen.n., Abyssidrilus gen.n., Paraktedrilus gen.n., Uniporodrilus Erséus, 1979, Inermidrilus gen.n., Gianius gen.n., Mexidrilus gen.n., Peosidrilus Baker & Erséus, 1979, Atlantidrilus Erséus, 1983, Thalassodrilus Brinkhurst, 1963, Pirodrilus gen.n., Pirodriloides gen.n., Marionidrilus gen.n., Pectinodrilus gen.n., Pacifidrilus gen.n. and Pseudospiridion gen.n. Nine new species are described from the Northwest Atlantic: Phallodriloides pinnulatus sp.n., Abyssidrilus opulentus sp.n., A. potens sp.n., A. altoides sp.n., Mexidrilus obtusus sp.n., Peosidrilus dalei sp.n., P. aduncus sp.n., P. riseri sp.n., and Atlantidrilus hamulus sp.n. The genus Discordiprostatus Baker, 1982, is synonymized with Nootkadrilus Baker, 1982.     

Inanidrilus bulbosus gen. et sp.n., a Marine Tubificid (Oligochaeta) from Florida, USA

Inanidrilus bulbosus gen. et sp.n. is described from subtidal sands near Miami, Florida. The species lacks alimentary canal, but it possesses highly modified penial setae and an unpaired dorsal spermatheca. It appears closely related to Phallodrilus Pierantoni, but since it is devoid of anterior prostate glands, which are attached to the ental ends of the atria in Phallodrilus , a new monotypic genus, Inanidrilus gen.n., is established.     

Marine Tubificidae (Oligochaeta) of Antarctica, with descriptions of three new species of Phallodrilinae

Five species of Tubificidae are recorded from Antarctic waters: Torodrilus lowryi Cook, 1970, Torodrilus sp. (subfamily Rhyacodrilinae), Marionidrilus antarcticus sp. n., M. weddellensis sp. n., and Thalassodrilus bicki sp. n. (subfamily Phallodrilinae); only T. lowryi was known from Antarctica before. The status of the two Southern Hemisphere genera Torodrilus Cook, 1970, and Marionidrilus Erséus, 1992, are discussed; although the taxonomic position of the two new species of Marionidrilus is somewhat uncertain. Thalassodrilus bicki , however, appears closely related to Northern Hemisphere taxa.     

Taxonomic Studies of Phallodrilinae (Oligochaeta, Tubificidae) from the Great Barrier Reef and the Comoro Islands with Descriptions of Ten New Species and One New Genus

A taxonomic and morphological account of a collection of Phallodrilinae from coralline sands at Heron Island in Australia's Great Barrier Reef is given. Ten new forms are described: Jamiesoniella athecata gen. et sp.n. Phallodrilus rectisetosus heronensis subsp.n. P. geniculatus sp.n. P. filithecatus sp.n. P. clavatus sp.n. (latter three all devoid of an alimentary canal), Bathydrilus superiovasatus sp.n. Coralliodrilus atriobifidus sp.n. C. oviatriatus sp.n. C. parvigenitalis sp.n. and the gutless C. avisceralis sp.n. Morphological notes are given for Phallodrilus albidus Jamieson, 1977 and Bathydrilus rohdei (Jamieson, 1977). The new meiobenthic genus Jamiesoniella is characterized by having simple atria with only one pair of prostate glands, which are attached to ectal parts of atria, and by lacking spermathecae and penial setae, The genus is probably related to Aktedrilus Knöllner and Bacescuella Hrabě. A gutless species, Phallodrilus comorensis sp.n. is described from the Comoro Islands in the Mozambique Channel (W Indian Ocean).     

The marine Tubificidae (Oligochaeta) of the barrier reef ecosystems at Carrie Bow Cay, Belize, and other parts of the Caribbean Sea, with descriptions of twenty-seven new species and revision of Heterodrilus, Thalassodrilides and Smithsonidrilus

Fifty-nine species of Tubifieidae are recorded from Belize on the Caribbean side of Central America. Twenty-five are new to science: Ainudrilus geminus sp.n., Heronidrilus gravidus sp.n., Heterodrilus flexuosus sp.n., H. ranus sp.n., H. modestus sp.n., Coralliodrilus rugosus sp.n., C. randyi sp.n., Phallodrilus compactus sp. n., P. singularis sp.n., P. vicinus sp.n., P. nasutus sp.n., P. bipartitus sp.n., Inanidrilus reginae sp.n., Olavius finitimus sp.n., O. vacuus sp.n., O. (Olavius) pravus sp.n., Bathydrilus vetustus sp.n., B. egenus sp.n., Thalassodrilides bruneti sp.n., Limnodriloides anxius sp.n., L. major sp.n., L. sacculus sp.n., L. adversus sp.n., Smithsonidrilus appositus sp.n., S. involutus sp.n. Phylogenetic analyses indicate that Heterodrilus Pierantoni, 1902, Thalassodrilides Brinkhurst & Baker, 1979, and Marcusaedrilus Righi & Kanner, 1979, are paraphyletic taxa as previously defined. They are therefore revised to include also apomorphic species earlier attributed to separate genera; in so doing, Marcusaedrilus becomes a junior synonym of Smithsonidrilus Brinkhurst, 1966. Smithsonidrilus pauper sp.n. (Peru), and S. multiglandularis sp.n. (Florida and Puerto Rico) are also described, and Limnodriloides claviger Erséus, 1982 is regarded as a synonym of L. pierantonii (Hrabě, 1971).     

New records of Adelodrilus (Oligochaeta,Tubificidae), with descriptions of two new species from the North-west Atlantic

Adelodrilus fimbriatus sp.n. is described from bathyal depths off the continental shelf of Eastern United States, A. cristatus sp.n. from sublittoral bottoms on this shelf (off Maryland and New Jersey). The two species are principally distinguished from their congeners by the morphology of their penial setae. New geographical records and morphological notes are given for A. pusillus Erséus (including the first find of Adelodrilus in the Mediterranean Sea), A. cooki Erséus, A. voraginus (Cook), and A. acochlearis Erséus & Loden. A pictorial key to all eleven species of Adelodrilus is provided.     

The family Capilloventridae (Annelida, Clitellata) in Australia, with descriptions of two new species of Capilloventer

Two new freshwater species of the small family Capilloventridae are recorded from rivers in south-eastern Australia. Capilloventer acheronensis sp. n. has all hair setae anteriorly and dissimilar crotchet setae without hairs posteriorly while Capilloventer longicapitus sp. n. has an elongate prostomium, very long hair setae anteriorly and all bifid setae posteriorly. Immature specimens from south-west Western Australia appear to represent a further species and are tentatively included within the family. Additional specimens of Capilloventer australis Erséus, 1993 have allowed some features (setae, segmental glands, genitalia and ventral buccal organ) to be described in more detail. The new C. australis specimens are from the upper reaches of Victorian rivers, showing that it is a freshwater species, rather than a marine incursion as the estuarine type locality suggested. There are now more freshwater species than marine species, although the origin and phylogenetic relationships of the family remain unclear.     

Introduction to Physidae (Gastropoda: Hygrophila); biogeography, classification, morphology

Physidae, a world-wide family of freshwater snails with about 80 species, are reclassified by progressive characters of the penial complex (the terminal male reproductive system): form and composition of penial sheath and preputium, proportions and structure of penis, presence or absence of penial stylet, site of pore of penial canal, and number and insertions of penial retractor muscles. Observation of these characters, many not recognized previously, has been possible only by the technique used in anesthetizing, fixing, and preserving. These progressive characters are the principal basis of 23 genera, four grades and four clades within the family. The two established subfamilies are divided into seven new tribes including 11 new genera, with diagnoses and lists of species referred to each. Proposed as new are: in Aplexinae, Austrinautini, with Austrinauta g.n. and Caribnautu harryi g.n., nom.nov.; Aplexini; Amecanautini with Amecanauta jaliscoensis g.n., sp.n., Mexinauta g.n., and Mayabina g.n., with M. petenensis, polita, sanctijohannis, tempisquensis spp.nn., Tropinauta sinusdulcensis g.n., sp.n.; and Stenophysini, with Stenophysa spathidophallus sp.n.; in Physinae, Haitiini, with Haitia moreleti sp.n.; Physini, with Laurentiphysa chippevarum g.n., sp.n., Physa mirollii nom.nou.; and Physellini, with Chiapaphysa g.n., and C. grijalvae, C. pacifica spp.nn., Utahphysa g.n., Archiphysa g.n., with A. ashmuni, A. sonomae spp.nn., Physella hemphilli sp.n., and Ultraphysella sinaloae g.n., sp.n. The simplest reproductive system is found in Austrinauta of the Aplexinae; its penial complex approaches that in the related family Lymnaeidae. Within Physinae a close approximation is found in Haitia. By these two genera the two subfamilies are drawn close together. Four grades of progressive complexity are recognized: (I) penial sheath entirely muscular; (II) penial sheath with both glandular and muscular tissue; (III) penis with penial stylet or other specialization of the tip of the penis; and (IV) pore of penial canal lateral rather than terminal as in the lower grades. In both subfamilies there are clades with glandular tissue in the penial sheath, a penial sheath subdivided into two parts, and tip of penis specialized in various ways. These clades are formalized as new tribes. Of 23 genera of Physidae, 17 occur in Pacific drainages of North and Central America, eight of these restricted to the region. Concentration of primitive genera along the Pacific coast from Mexico to Costa Rica conforms to previous observations that primitive pulmonate families are concentrated within, or along the continental margins of, the Pacific Ocean. An ancestral origin of Physidae along an ancient eastern Pacific coast is probable. From this region the several lineages have spread north, south and east in the Americas, and through Siberia to Europe. Although Physinae have fewer genera than Aplexinae (11 v. 12), they have more species (47 v. 34). Greater land area in the temperate zone has provided more opportunity for speciation of Physinae, in contrast to the generally tropical and warm-temperate range of Aplexinae. Furthermore, 10 species of Physinae are localized in individual lakes, whereas Aplexinae are not lake-dwellers. Both well-developed egg strings and capsular strings are found in the spawn of Sibirenauta elongatus. These structures have been known in Lymnaeidae, but not hitherto in Physidae; they are a link with some marine groups, such as Siphonariidae. Spiral color bands and white streaks in the shell of Mexinauta recall those in Lancidae (Lymnaeacea), whereas the radula of Physidae is like that of Chilinidae. Physidae thus show affinities to various basal stocks of aquatic pulmonates; no clear-cut sister-group can be recognized. Most species have a restricted range; out of 55 with sufficiently detailed information for analysis, 25 are limited to a single 1 degrees x 1 degrees quadrangle. Only a few species are widespread, on one or even two continents. Accordingly, more species of Physidae are threatened by habitat destruction than in other families of Hygrophila with generally wider distributions. Other features are a key to genera; catalog of more than 430 names applied to living Physidae, with original reference, type locality, and location of type specimens; summary of museums with types; and glossary.     

Tubificidae (oligochaeta) from the Ross Sea (Antarctica), with descriptions of one new genus and two new species

Three species,Torodrilus gelidus sp. nov. (subfamily Rhyacodrilinae),Rossidrilus terraenovae gen. et sp. nov. (Limnodriloidinae), and a second unnamed species of Limnodriloidinae, are reported from marine sediments in Terra Nova Bay, Ross Sea. Torodrilus gelidus is distinguished fromT. lowryi Cook, 1970 by its setal pattern (with few exceptions, both anterior and posterior setae are single-pointed in sexually mature specimens ofT. gelidus) and the morphology of its male protuberances (the latter folded over a midventral bursa in segment XI).Rossidrilus terraenovae is characterized by large diverticula attached to the oesophagus in the posterior part of segment IX, unpaired male and spermathecal pores, heavily muscular and entally ciliated atrial ampullae, elongate prostatic pads, and a deep, unpaired and muscular, copulatory sac. It is the first species of its subfamily to be described from Antarctic waters.     

Taxonomic Revision of the Marine Genus Heterodrilus Pierantoni (Oligochaeta,Tubificidae)1

Marine tubificids possessing trifid anterior setae are morphologically and taxonomically reviewed, on the basis of material from the Atlantic and Pacific Oceans. Heterodrilus Pierantoni, 1902 is revised to include thirteen species, H. arenicolus Pierantoni, 1902, H. minisetosus sp.n.H. ascensionensis sp.n. H. queenslandicus (Jamieson, 1977), H. lacertosus sp.n. H. scitus sp.n. H. keenani sp.n. H. claviatriatus sp.n. H. subtilis (Pierantoni, 1917), H. jamiesoni sp.n. H. occidentalis sp.n. H. pentcheffi sp.n. and H. bulbiporus sp.n. The genus is briefly defined: marine tubificids with trifid setae in preclitellar segments (with H. subtilis as the only exception); paired spermathecae located in segment X; vasa deferentia entering apical, ental ends of slender, ciliated atria, which bear broadly attached masses of prostate glands; paired male pores, and generally with penial setae arranged in bisetal bundles. Heterodriloides gen.n. is established for H. quadrithecatus sp.n. distinguished from Heterodrilus by two main features: its spermathecae are located in XII, with a supplementary pair generally located in XI; and its vasa enter the ectal part of the atria. Giereidrilus gen.n. a third genus with trifid setae, is established to include Phallodrilus ersei Giere, 1979 and G. inermis sp.n. Both species have unpaired spermathecal and male pores, and their atria are not ciliated. Heterodrilus, Heterodriloides and Giereidrilus are placed in the subfamily Rhyacodrilinae Hrabě, 1963.     

Taxonomic Studies on the Marine Genera Aktedrilus Knöllner and Bacescuella Hrabe' (Oligochaeta, Tubificidae), with Descriptions of Seven New Species

The taxonomy and morphology of Aktedrilus Knöllner, 1935 and Bacescuella Hrabe' 1973 (subfamily Phallodrilinae), both common genera of the marine, littoral meiofauna, were studied on the basis of material from various geographical areas. A. monospermathecus Knöllner, 1935 is redescribed from France and Scotland. A. magnus sp.n., A. hrevis sp.n., A. curvipenis sp.n. and A. floridensis sp.n. are described from Italy, Brazil, France and Florida, respectively. Two very closely related forms are described from the Pacific: A. locyi sp.n. (California) and A. parviprostatus sp.n. (Great Barrier Reef). B. mediterranea sp.n. is described from Italy, and new European records are given forB. arclica Erséus, 1978 andfi. parvithecata Erséus, 1978. Both genera are characterized by having two pairs of prostate glands, well developed penes, and unpaired, mid-dorsal spermatheca (if present). The species of Bacescuella transfer their sperm by means of external spermatophores, structures that are not developed in Aktedrilus. The eight species of Aktedrilus are largely distinguished by means of the morphology of the spermatheca, penes and prostates. The four species of Bacescuella differ principally from each other in the length of the vasa deferentia, and in the morphology of the prostates and copulatory organs. Most Bacescuella species lack spermatheca.     

New species of the genus Trichodrilus (Oligochaeta, Lumbriculidae)

Five new species of the genus Trichodrilus are described: Trichodrilus longipenis. sp. n. T: rouchi sp. n. T. capilliformis sp. n. and T. angelieri sp. n. from hyporhcic waters of the Iberian Peninsula and France. and T. bonheurensis sp. n. from superticial waters of France. The new taxa contribute with some new features to an amended diagnosis of the genus: presence of modified penial setae, four pairs of spermathecae, hair-like dorsal setae and glandular pads in the epidermis.     

Marine Oligochaeta from the Koster Area, West Coast of Sweden, with Descriptions of Two New Enchytraeid Species

A total of 28 species of marine Oligochaeta (belonging to the families Naididae, Tubificidae and Enchytraeidae) are reported from eulittoral and sublittoral sediment samples taken in the Koster area on the Swedish West Coast. Lumbricillus algensis sp.n. and Grania ovitheca sp.n. are described and their morphological and taxonomical relations to other enchytraeids are discussed. L. algensis is characterized mainly by its spermathecae, which project into segment VI, and by its very large penial bulbs. G. ovitheca possesses no dorsal setae, the ventral setae being present posterior to the clitellum only, and its spermathecae consist of a narrow duct and a large egg-shaped to oval ampulla, filling most of the coelom of segment V. Four sublittoral species, Limnodriloides barnardi (Tubificidae), Lumbricillus semifuscus, Marionina sublitoralis and Grania roscoffensis (all Enchytraeidae) are reported as new to the Swedish fauna.     

Taxonomy and Phylogeny of the Gutless Phallodrilinae (Oligochaeta,Tubificidae), with Descriptions of One New Genus and Twenty-Two New Species*

The gutless members of the subfamily Phallodrilinae are taxonomically revised and generically split off from their gut-bearing relatives. Inanidrilus Erséus, 1979, a much modified definition of which is presented, comprises the species leukodermatus (Giere, 1979) comb.n., speroi sp.n., fijiensis sp.n., bonomii sp.n., carterensis sp.n., gustavsoni sp.n., wasseri sp.n., scalprum sp.n., belizensis sp.n., aduncosetis sp.n., vacivus sp.n., triangulatus sp.n., ernesti sp.n., manae sp.n., falcifer Erséus & Baker, 1982, renaudae sp.n., extrmus (Erséus 1979) comb.n., mexicanus Erséus & Haker, 1982, and bulbosus Erséus, 1979. Olavius sp.n. is established to accommodate the species geniculatus (Erséus, 1981) comb.n., imperfectus sp.n., filithecatus (Erséus. 1981) comb.n. albidus (Jamieson, 1977) comb.n., propinquus sp.n., comorensis (Erséus. 1981) comb.n., cornuatus Davis, 1984, pellucidus sp.n., tenuissimus (Erséus, 1979) macer sp.n., caudatus (Erséus. 1979) comb.n., planus (Erséus, 1979) comb.n., clavatus (Erséus. 1981) comb.n., tantulus sp.n., longissimus (Giere, 1979) comb.n., alius sp.n., avisceralis (Erséus, 1981) comb.n., loisae sp.n., and hanssoni sp.n., Two subgenera are erected for tantulus—longissimus–alius (nominate subgenus Olavius) and avisceralis-loisae-hanssoni (Coralliodriloides subgen.n.), respectively. A tentative phylogeny of the gutless group, which appears monophyletic, is inferred on the basis of a partly computerized Wagner-tree analysis of all 38 species. According to this analysis. According to this analysis, Inanidrilus can be defined by one synapomorphy, more or less sickle-shaped penial setae, whereas Olavius can be discriminated by two such characters, (1) atrium comma-shaped/horizontal rather than erect and (2) copulatory sacs present.