New Nearctic Eremidrilus species (Clitellata: Lumbriculidae). Part 2, western species with one spermathecal segment.

Three new species of the lumbriculid genus Eremidrilus Fend Rodriguez, 2003 are described from rivers in California, Nevada and Idaho, USA. The new species share the typical characters of the genus: a filiform proboscis, semiprosoporous male ducts and atria in X, spermathecae paired in the postatrial segment and male pores usually associated with distinct porophores. Species in Eremidrilus have either two spermathecal segments, XI and XII, or only one, XI. Compared with other Eremidrilus species having only one pair of spermathecae in XI, E. chalonensis n. sp. is mainly distinguished by the presence of a ring of glands at the spermathecal pore; E. pinedai n. sp. has a very small atrium and porophore; E. owyhee n. sp. has a thin, cylindrical atrium and a very long porophore. The spermathecal pores open in line with ventral chaetae in E. chalonensis, a position which separates it from previously described California species where the pore opens in a more lateral position. In contrast, the spermathecal pores in E. owyhee open at the lateral line in a transverse body fold. Most Eremidrilus species described so far show narrow endemicity, and their geographic distributions are restricted to small hydrographic areas. Only E. felini and E. owyhee have distribution ranges greater than 200 km. We provide a key to all known species of Eremidrilus.

On spermatophore-producing aquatic microdrile oligochaetes (Annelida: Clitellata).

The formation of encapsulated spermatophores is exceptional among aquatic oligochaetes, although it seems to have occurred independently in several unrelated taxa. Among the microdriles, some variations appear unique to single species. The recently described lumbriculid Uktena riparia Fend et al. forms spermatophores in the male duct and attaches them within a deep spermathecal bursa. The attachment of spermatophores to the body wall, in the clitellar region, or in the vicinity of male or spermathecal pores has been reported in Paranadrilus Gavrilov, Bothrioneurum Stolc and some Aktedrilus Knöllner species. Anatomical comparison of reproductive organs suggests convergent development of glandular organs in the male duct or spermathecae, used for the formation, transfer and/or attachment of spermatophores to the concopulant worms. The presence of similar organs in Smithsonidrilus Brinkhurst, where spermatophores have not been reported, is also discussed. Furthermore, a lectotype is designated for Paranadrilus descolei Gavrilov, 1955.

Multi-locus phylogenetic analysis of the genus Limnodrilus (Annelida: Clitellata: Naididae).

Limnodrilus species are annelid worms distributed worldwide in various freshwater sediments. The systematics of Limnodrilus has chiefly been based on morphology, but the genus has not been subject to any closer phylogenetic studies over the past two decades. To reconstruct the evolutionary history of Limnodrilus, and to assess the monophyly of this genus and its systematic position within the subfamily Tubificinae (Annelida: Clitellata: Naididae), 45 Limnodrilus specimens, representing 19 species, and 35 other naidid species (representing 24 genera) were sampled. The data consisted of sequences of three mitochondrial genes (COI, 12S and 16S rDNA) and four nuclear markers (18S and 28S rDNA, Histone 3, and ITS). The phylogeny was estimated, using Maximum Likelihood and Bayesian analyses of concatenated data of seven DNA loci, as well as a multi-locus coalescent-based approach. All analyses strongly suggest that Limnodrilus is monophyletic, but only if the morphospecies L. rubripenis is removed from it. Limnodrilus rubripenis and (at least) Baltidrilus, Lophochaeta and some species attributed to Varichaetadrilus comprise the sister group to the clade Limnodrilus sensu stricto, and the latter is further divided into three well-supported groups. One of them contains morphospecies characterized by short cuticular penis sheaths and enlarged chaetae in anterior segments (L. udekemianus, L. silvani and L. grandisetosus). The second is a small group of species with moderately long penis sheaths, i.e., L. sulphurensis and L. profundicola. The third, and largest group, includes not only the multitude of cryptic species in the L. hoffmeisteri complex, but also other, morphologically distinct, species nested within this complex. All studied species in this large group have long penis sheaths, which are exceptionally long in L. claparedianus, L. maumeensis, and a form morphologically intermediate between L. claparedianus and L. cervix. The identification and classification of these groups provide a framework for directed sampling in further phylogenetic studies, and for revisionary work on the L. hoffmeisteri complex and other unresolved Limnodrilus species.

Limnodrilus sulphurensis n. sp., from a sulfur cave in Colorado, USA, with notes on the morphologically similar L. profundicola (Clitellata, Naididae, Tubificinae).

A new species of the tubificine genus Limnodrilus is described and COI barcoded from Sulphur Cave and associated springs in Colorado, USA. The habitats are characterized by high sulfide concentrations. The new species, L. sulphurensis, is distinguished from all congeners by the elongate, nearly parallel teeth of chaetae in its anterior segments. It has a penis sheath resembling that of L. profundicola; consequently, museum specimens and new collections are examined here to resolve some of the taxonomic confusion surrounding that widespread, but uncommon species.

Uktena riparia n. gen., n. sp. (Annelida, Clitellata, Lumbriculidae), a new spermatophore-producing oligochaete.

Uktena riparia n. gen., n. sp. has been collected in hyporheic habitats at several sites in North Carolina, southeastern USA. The genus is defined by unusual characters related to reproductive structures, including the formation of encapsulated spermatophores for sperm transfer and large bundles of genital chaetae, both previously unknown in the Lumbriculidae. The simultaneous occurrence of both spermatophores and spermathecae is rare in the microdrile oligochaetes. Uktena spermatophores appear more similar to those reported in leeches than to those in other microdrile oligochaete families. Possible synapomorphies associating Uktena with the genera Kincaidiana and Guestphalinus include a filiform, ringed proboscis, a forward shift of reproductive organs relative to the usual position in the family, and spermathecae in the atrial segment. The new species adds to the already diverse, endemic lumbriculid fauna of the North Carolina Sandhills ecoregion.

Sylphellapuccoon gen. n., sp. n. and two additional new species of aquatic oligochaetes (Lumbriculidae, Clitellata) from poorly-known lotic habitats in North Carolina (USA).

Three new species of Lumbriculidae were collected from floodplain seeps and small streams in southeastern North America. Some of these habitats are naturally acidic. Sylphellapuccoon gen. n., sp. n. has prosoporous male ducts in X-XI, and spermathecae in XII-XIII. Muscular, spherical atrial ampullae and acuminate penial sheaths distinguish this monotypic new genus from other lumbriculid genera having similar arrangements of reproductive organs. Cookidriluspocosinus sp. n. resembles its two subterranean, Palearctic congeners in the arrangement of reproductive organs, but is easily distinguished by the position of the spermathecal pores in front of the chaetae in X-XIII. Stylodriluscoreyi sp. n. differs from congeners having simple-pointed chaetae and elongate atria primarily by the structure of the male duct and the large clusters of prostate cells. Streams and wetlands of Southeastern USA have a remarkably high diversity of endemic lumbriculids, and these poorly-known invertebrates should be considered in conservation efforts.

Rhynchelmis subgenus Sutroa Eisen new rank, with two new species from western North America (Annelida, Clitellata, Lumbriculidae).

The lumbriculid Rhynchelmis subgenus Sutroa Eisen, 1888 new rank is defined for a group of Nearctic species having multiple diverticula originating at the spermathecal ducts and eversible penial bulbs. Characters are confirmed in specimens of the type species, Rhynchelmis (Sutroa) rostrata (Eisen, 1888), collected from the type locality. Rhynchelmis (Sutroa) klamathensis Fend n. sp. is described from open water benthic habitats in Upper Klamath Lake, Oregon, USA. It resembles other R. (Sutroa) species in the paired spermathecal diverticula, the spermathecal and penial bulbs, the histological structure of the atria, the nonfunctional anterior male funnels, and the arrangement of blood vessels. Rhynchelmis klamathensis differs from all Nearctic Rhynchelmis in lacking a filiform proboscis. The combination of large body size, the elongate spermathecal ducts with paired and usually unbranched diverticula, the highly contorted atria, and the complex male pores with conical penes also distinguish typical R. klamathensis from other Rhynchelmis species. Smaller specimens with otherwise similar morphology, from the Sacramento River Delta, California, are also assigned to this species. Rhynchelmis (Sutroa) diespluviae Fend n. sp. is described from several stream sites, mostly in northern Idaho, USA. Rhynchelmis diespluviae differs from closely related species in morphology of the conical penes, and in the structure and anterolateral position of the paired spermathecae.

A new species of Helobdella (Hirudinida: Glossiphoniidae) from Oregon, USA.

Helobdella bowermani n. sp. is described from specimens collected in fine sediment of open water benthos of Upper Klamath Lake, Klamath County, Oregon. The new species has pale yellow/buff coloration with scattered chromatophore blotches throughout the dorsal surface, lateral extensions or papillae only on the a2 annulus, dorsal medial row of papillae with small papilla on al and larger papillae on a2 and a3, and a small oval scute (rarely triangular). Helobdella bowermani n. sp. is morphologically similar to Helobdella atli and Helobdella simplex. Molecular comparison of CO-I sequence data from H. bowermani n. sp. revealed differences of 10.6/--10.8% with Helobdella californica, differences of 12.2%-13.7% with H. atli, and differences of 12.7%-13.2% with H. simplex.

New species of Rhyacodrilus (Annelida: Clitellata: Rhyacodrilinae) of North America, with re-description of R. sodalis (Eisen, 1879).

Six new Nearctic species of the aquatic oligochaete genus Rhyacodrilus (Annelida, Clitellata, Rhyacodrilinae), are described, five (R. saelonae sp. n., R. quileuticus sp. n., R. clio sp. n., R. alcyoneus sp. n. and R. longichaeta sp. n.) from western and one (R. propiporus sp. n.) from eastern North America. The taxonomy of the most common Rhyacodrilus species reported in the Nearctic region has been based largely on chaetal characters, which has generated certain confusion. The new species give a new perspective on the genus Rhyacodrilus in North America, suggesting a much higher diversity than previously expected. The description of R. longichaeta sp. n. questions the taxonomic status of R. montana (Brinkhurst), which is here regarded as species inquirenda. The taxonomic status of R. sodalis (Eisen) is discussed based on characters of the reproductive system, the existing Lake Tahoe neotype series is invalidated, and a neotype is described from Eisen's type locality. Based on the discussion of the characters of the genus Rhyacodrilus, the genus Stochidrilus Martinez-Ansemil et al. is proposed as a junior synonym of that genus. The presence of the widely reported species R. coccineus has not been confirmed in the study collections, although the species requires a sound revision. A key to the species bearing hair chaetae is provided, based mainly on features of the reproductive system.

Yamaguchia toyensis n. sp., n. gen. (Annelida, Clitellata, Lumbriculidae) from profundal Lake habitat in Japan.

Yamaguchia toyensis n. sp., n. gen. is described from an oligotrophic caldera lake, Lake Toya, Hokkaido, Japan. Although the taxonomic affinities are unknown, the genus differs from all other Lumbriculidae in having the combination of testes and atria in X, a single, prosoporous male funnel per atrium, and spermathecae in XI. Unlike other Japanese lakes that have thus far been surveyed, Lake Toya supports abundant populations of lumbriculids in the profundal benthos.

Importance of sediment-water interactions in Coeur d'Alene Lake, Idaho, USA: management implications.

A field study at Coeur d'Alene Lake, Idaho, USA, was conducted between October 1998 and August 2001 to examine the potential importance of sediment-water interactions on contaminant transport and to provide the first direct measurements of the benthic flux of dissolved solutes of environmental concern in this lake. Because of potential ecological effects, dissolved zinc and orthophosphate were the solutes of primary interest. Results from deployments of an in situ flux chamber indicated that benthic fluxes of dissolved Zn and orthophosphate were comparable in magnitude to riverine inputs. Tracer analyses and benthic-community metrics provided evidence that solute benthic flux were diffusion-controlled at the flux-chamber deployment sites. That is, effects of biomixing (or bioturbation) and ground-water interactions did not strongly influence benthic flux. Remediation efforts in the river might not produce desired water-quality effects in the lake because imposed shifts in concentration gradients near the sediment-water interface would generate a benthic feedback response. Therefore, development of water-quality models to justify remediation strategies requires consideration of contaminant flux between the water column and underlying sediment in basins that have been affected by long-term (decadal) anthropogenic activities.