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Enchytraeus polatdemiri Arslan & Timm, sp. nov. (Figures 2, 3) Holotype. ESOGU-Epolatdemiri-01, adult specimen, unstained whole mount, anterior end (14 segments), mounted between two coverslips, posterior end processed for DNA sequencing (specimen ID EP2, see Table 2, Fig. 4), Turkey, Lake Van, N38��21.466' E043��04.345', profundal sediment at 50 m depth, station 3 (Table 1), 16.06.2012, N. Arslan. Paratypes. ESOGU-Epolatdemiri-02���04, 3 adult and subadult specimens, same data as holotype. ESOGU- Epolatdemiri-05���08, 4 adult and subadult specimens, processed as holotype, N38��36.502' E043��07.343', profundal sediment at 36 m depth, station 14 (Table 1), 16.6.2012, N. Arslan. ESOGU-Epolatdemiri-09, 9 specimens, 4 adult, 3 subadult, 2 juvenile, unstained whole mounts on one slide, Station 24, 14.10.2011, N. Arslan (Table 1). ESOGU- Epolatdemiri-10, 9 specimens, 5 adult, 2 subadult, 2 juvenile, unstained whole mounts on one slide, Station 2 (Table 1), 16.6.2012, N. Arslan. ESOGU-Epolatdemiri-11, 4 specimens, 3 adult, 1 subadult, slightly stained, partly fragmented, Station 32, 16.06.2012, N. Arslan (Table 1). VLS-S4258, 3 specimens, 1 adult, 2 subadult, stained whole mounts on one slide, Station 20 (Table 1), 19.8.2011, N. Arslan. VLS-S4 259, 8 specimens, 3 adult, 5 subadult, stained whole mounts on one slide, Station 21, 14.10.2011, N. Arslan (Table 1). Further material. ESOGU-Epolatdemiri-12, Station 18, ca. 8 specimens in 96% ethanol, partly fragmented, subadult and juvenile, 14.10.2011, N. Arslan. ESOGU-Epolatdemiri-13, Station 20, 10 specimens in 96% ethanol, subadult and juvenile, 14.10.2011, N. Arslan. ESOGU-Epolatdemiri-14, Station 21, 6 specimens in 96% ethanol, subadult and juvenile, 14.10.2011, N. Arslan. ESOGU-Epolatdemiri-15, Station 3, ca. 7 specimens in 96% ethanol, partly fragmented, subadult and juvenile, 16.06.2012, N. Arslan. ESOGU-Epolatdemiri-16, Station 14, ca. 3 specimens in 96% ethanol, partly fragmented, subadult and juvenile, 16.06.2012, N. Arslan. ESOGU-Epolatdemiri- 17, Station 24, multiple fragments in 96% ethanol, not counted, 16.06.2012, N. Arslan. VLS-S4263, 1 specimen, sagittal sections on 2 slides, Station 24, 14.10.2011, N. Arslan. VLS-S4264, 1 specimen, sagittal sections on 2 slides, Station 24, 14.10.2011, N. Arslan. VLS-S4260, 1 specimen, horizontal sections on 2 slides, Station 23, 14.10.2011, N. Arslan. VLS-S4261, 1 specimen, horizontal sections on 3 slides, Station 23, 14.10.2011, N. Arslan. Etymology. The species is named in memory of Şevket Polatdemir (1940-1997), Prof. Dr. Naime Arslan's father, an outstanding person. Description. Body length 7.5���11 mm, diameter c. 0.4 mm, in clitellar region c. 0.5 mm (0.32���0.38 mm at V, 0.41���0.58 mm at XII, 0.42���0.49 mm at XX, N=10). Segment number 40���55 (N=10). Chaetae 2 or 3 per bundle, rarely more (6 in 1 bundle of 1 specimen, Fig. 2A, segment VIL), occasionally 1 or 0; at XII absent ventrally, present laterally, 2 per bundle. Chaetal formula 2,3 ��� 2,3: 2,3 ��� 2,3. Lateral preclitellar bundles: 2 and 3 irregularly alternating, some specimens with mostly 2, others with mostly 3 chaetae, or intermediate states. Ventral preclitellar bundles: mostly 3 chaetae, 2 chaetae near clitellum (X, XI). Lateral and ventral postclitellar bundles: 2 in segments following clitellum, caudad increasingly 3. Chaetae straight with ental hook, ectally pointed: 80���110 ��m long, 6���7 ��m wide, smallest at II, XII and XIII (length 70���90 ��m, diameter 3.5���5 ��m). Caudal chaetae not or only slightly enlarged. Epidermal gland cells not seen. Brain c. 1.5x as long as wide, rounded posteriorly, sides slightly merging anteriad. Oesophageal appendages a pair of blind-ending tubes in III/IV, c. 140 ��m long and 40 ��m wide, with common root inserting dorsally in III behind pharyngeal pad, tubes curved, not meandering, not branched, not tapering towards blind end. Pharyngeal glands in IV���VI, of equal size, with dorsal and ventral lobes, connected dorsally in IV, connected or separate dorsally in V, VI. Nephridial anteseptale with funnel only, postseptale bulged, efferent duct short, no terminal vesicle; present at 6/7���9/10 (4 preclitellar pairs) and in postclitellar segments. Dorsal blood vessel from XIII, XIV, (XV), often conspicuous (Fig. 3B). Anterior bifurcation near prostomium. Coelomocytes flattened, 3x as long as wide, c. 20���30 ��m long, oval or narrowed at one end, texture not ascertained. Clitellum in XII���XIII, hyalocytes and granulocytes from level of septum 11/12 to level of chaetae XIII. Anteriorly and posteriorly a variously extending sleeve of hyaline border cells (Fig. 2A,E: cb). Clitellum welldeveloped dorsally and laterally, height c. 30 ��m, cell diameter c. 15 ��m, hyalocytes and granulocytes arranged in reticulate fashion, mostly interrupted mid-ventrally, continuous at posterior border; area without clitellum widest at level of male pores, including the exterior lips (Fig. 2E: ml); exclusively granulocytes at ventro-lateral borders and dorsally of the male exterior lips. Testes and sperm funnels in XI, ovaries, vasa deferentia, male pores and glands in XII. Seminal vesicle unpaired, in X���XI. Mature spermatozoa aligned on top of sperm funnel; length of sperm heads not measured. Sperm funnel c. 5x as long as wide, variable in shape due to soft tissue of glandular funnel body. Collar about as wide as funnel body. Vasa deferentia occasionally extending into posterior segments (XIII: 2 specimens; XIV: 1 specimen; XV: 1 specimen; N=38), but mostly confined to XII in a dense irregular coil that fills almost the entire segment; wider entally near sperm funnel (Fig. 3C: en) than ectally near male pore (Fig. 3C: ec), ciliated throughout, ectally passing ventrad between inner face of body wall and primary male glandular bulb en route to male pore. Male glands multiple: one large rounded primary bulb near male pore, and ca. 9 smaller secondary papillae medially, arranged roughly in a semicircle around male pore and primary bulb (Figs 2D, 3C,D). Male pores opening each into a bursa, a lateral invagination of the body wall that is covered by a lip-like fold (bursal fold) directed mediad (Fig. 2E: ml). Spermathecae with ectal duct, ampulla, no diverticula, separate openings into esophagus. Ectal pores at 4/5, slightly below lateral lines, not widened. Ectal duct slightly longer than wide (80���85 ��m by 55���80 ��m), covered completely with glands, canal widening entad, lined with cuticle; ampulla spherical, diameter c. 120���150 ��m, wider than ectal duct, with distinct, smooth walls, lumen completely filled with masses of spermatozoa; ental duct short, connecting laterally with esophagus. Remarks. Enchytraeus polatdemiri sp. nov. belongs to the Enchytraeus albidus group as circumscribed in Schmelz & Collado (2010). This group of species is distinguished within the genus by comparatively large body size (length> 10mm, up to 30 mm), high segment number (>40, usually>50), short and tube-like oesophageal appendages, and a large and well-developed male reproductive system that, apart from a male glandular ("penial") bulb, often includes a number of smaller accessory glands around the male pores. Furthermore, several species, among them the common and widespread Enchytraeus albidus Henle, 1837, have more than 3 chaetae per bundle, elongate longitudinal body folds ventrally in the clitellar region that enclose the male pores, and vasa deferentia that extend beyond the clitellar region into posterior segments. Species of this mainly holarctic group live in the marine upper littoral, in brackish water, and in organically enriched habitats such as compost heaps. Apart from E. albidus it currently includes E. capitatus von B��low, 1957, E. kincaidi Eisen, 1904, E. mediterraneus Michaelsen, 1926, E. multiannulatus Altman, 1936, and E. multiannulatoides Altman, 1936, together with a number of junior synonyms. Within this group, E. polatdemiri sp. nov. is distinguished by the following combination of characters: (1) Comparatively small body size, (2) each specimen with bundles of two and three chaetae in varying arrangement, (3) vasa deferentia usually not extending beyond clitellum, and if so, only 1���2 segments, (4) spermathecal ectal duct completely glandular, (5) spermathecal ampulla spherical, wider than ectal duct, without diverticula. Further possibly distinguishing characters refer presently to E. albidus only, because they are still unkown in the other species: (6) clitellum ventrally almost absent, with a peculiar distribution of granular cells (Fig. 2E), (7) absence of a genital field, and (8) bursal folds not elongate. Among the six above-mentioned species of the E. albidus -group, only E. albidus has spermathecae that, at least in some specimens, are similar to those in E. polatdemiri sp. nov.: ectal duct completely glandular, ampulla spherical, no diverticulum (comp. Nielsen & Christensen 1959, Fig. 97). In the remaining five species, spermathecal ectal glands are absent or reduced to a crown around the ectal pore, and/or one or two diverticula are present. E. albidus differs in several characters from E. polatdemiri sp. nov.: more than 3 chaetae per bundle (up to 6 ventrally), bursal folds elongate, genital field present, vasa deferentia extending beyond clitellum (Schmelz & Collado 2010). Furthermore, specimens are usually much larger and up to 3 cm long. However, the species is highly variable and the evaluation of taxonomic characters is not straightforward. To give examples: Spermathecae are reported that include forms that are common for the other species (Nielsen & Christensen 1959). For example, all specimens of E. albidus sequenced in this study had a spermatheca with a large, asymmetrical dorsal diverticulum and an incomplete glandular coverage of the ectal duct (comp. Nielsen & Christensen 1959: 96). It is also unknown whether the posterior extension of the vas deferens applies to all specimens without exception; in addition, some specimens of the new species share this character state. Furthermore, E. sabulosus Southern, 1906, tentatively included in E. albidus by Nielsen and Christensen (1959), has also a maximum of 3 chaetae per bundle and a body size (10���15 mm when alive) comparable to that of the new species. Finally, there is molecular evidence that E. albidus consists of more than one species (Ers��us & Gustafsson 2009). On this background of taxonomic uncertainty is was advantageous to have a second dataset, i.e. molecular evidence, to evaluate the taxonomic status of the Lake Van population. Molecular analysis. The individual gene trees of COI and H3, as well as the combined phylogenetic analysis of both tree, recovered Enchytraeus polatdemiri as a monophyletic group within the genus Enchytraeus, with the highest support (pp = 1) (Fig. 4). In the concatenated tree, E. polatdemiri forms a well-supported sister group to the clade comprising all E. albidus specimens (Fig. 4C), whereas in the individual gene trees, relationships between E. polatdemiri sp. nov., E. albidus, E. japonensis (only COI), and E. bulbosus are poorly resolved (Fig. 4A,B). The maximum uncorrected p-distance between the specimens of Enchytraeus polatdemiri sp. nov. was 1.0 % for the COI gene, while H3 sequences were identical in this group (Tables 3 and 4). Average p-distances for COI between Enchytraeus polatdemiri sp. nov. and the other congeneric species included in the analysis ranged from 14.5 % with E. albidus (excluding the specimen from Denmark) to 19.5 % with E. japonensis (Table 3). Average H3 distances were substantially lower, ranging from 5.4 % with E. albidus to 10.5 % with E. bulbosus (Table 4). A potential barcode gap was found for COI pairwise genetic distances between 5 % and 11 % (Fig. 5). ABGD���s initial partition delineated seven MOTUs, which are those indicated in Table 3. H3 pairwise distances did not show a clear barcode gap., Published as part of Arslan, Naime, Timm, Tarmo, Rojo, Ver��nica, Vizca��no, Ant��n & Schmelz, R��diger M., 2018, A new species of Enchytraeus (Enchytraeidae, Oligochaeta) from the profundal of Lake Van, the world's largest soda Lake (Turkey, East Anatolia), pp. 367-380 in Zootaxa 4382 (2) on pages 372-375, DOI: 10.11646/zootaxa.4382.2.8, http://zenodo.org/record/1182147, {"references":["Schmelz, R. M. & Collado, R. (2010) A guide to European terrestrial and freshwater species of Enchytraeidae (Oligochaeta). Soil Organisms, 82, 1 - 176. Available from: http: // www. senckenberg. de / root / index. php? page _ id = 13185 (Acesses 20 Feb. 2018)","Henle, F. G. J. (1837) Ueber Enchytraeus, eine neue Anneliden-Gattung. Mullers Archiv fur Anatomie, Physiologie und wissenschaftliche Medizin, Berlin, 1837, 74 - 90.","Nielsen, C. O. & Christensen, B. (1959) The Enchytraeidae. Critical revision and taxonomy of European species (Studies on Enchytraeidae VII). Natura Jutlandica, 8 - 9, 1 - 160.","Erseus, C. & Gustafsson, D. (2009) Cryptic speciation in clitellate model organisms. In: Shain, D. H. (Ed.), Annelids in Modern Biology. Wiley-Blackwell, Hoboken, NJ, pp. 31 - 46. https: // doi. org / 10.1002 / 9780470455203. ch 3"]}

Hemienchytraeus sp. 1 Material examined. IG 327183 - 21, juvenile specimen, Honduras, Cusuco National Park [15.506299 -88.23704], 1457 m asl, M. Jocque 07/07/ 2013. Description. Length 1.5 mm, diameter 0.12 mm at V, 0.15 mm at XIII; 21 segments. Two chaetae per bundle, distally straight; chaetae slightly enlarged in caudal segments, ventrally c. 45: 4 ��m in caudal segments, max. c. 35���40: 3.5 ��m in anterior segments. Ventral chaetae thicker than lateral ones, length of laterals not measurable. Epidermal glands inconspicuous, one row of pale enlarged cells at level of chaetae. Oesophageal appendages bifurcating twice, with two elongate secondary branches on each side. Pharyngeal glands 3 pairs in IV���VI, all connected dorsally, a pair of small secondary ventral lobes in V. Dorsal blood vessel from IX, midgut pars tumida in XV���XVI. Five pairs of preclitellar nephridia, from 5 / 6 to 9 / 10, anteseptale as large as postseptale, c. 25 ��m long. Coeolomocytes very scarce, small, length c. 10 ��m. Remarks. This juvenile specimen cannot be identified to species, but by virtue of the oesophageal appendages, the pharyngeal glands and the preclitellar nephridia it belongs to a species different from H. phytotelmatus sp. nov., Published as part of Schmelz, R��diger M., Jocque, Merlijn & Collado, Rut, 2015, Microdrile Oligochaeta in bromeliad pools of a Honduran cloud forest, pp. 508-526 in Zootaxa 3947 (4) on page 516, DOI: 10.11646/zootaxa.3947.4.3, http://zenodo.org/record/242615

Genus Hemienchytraeus Černosvitov, 1934 Type species: Enchytraeus stephensoni Cognetti, 1927 (nom. nov. pro Enchytraeus cavicola Stephenson, 1924 non E. cavicola Joseph, 1880), Published as part of Schmelz, R��diger M., Jocque, Merlijn & Collado, Rut, 2015, Microdrile Oligochaeta in bromeliad pools of a Honduran cloud forest, pp. 508-526 in Zootaxa 3947 (4) on page 513, DOI: 10.11646/zootaxa.3947.4.3, http://zenodo.org/record/242615, {"references":["Cernosvitov, L. (1934) Zur Kenntnis der Enchytraeiden. II. Zoologischer Anzeiger, 105, 295 - 305.","Cognetti de Martiis, L. (1927) Lumbricidi dei Carpazi. Bollettino dei Musei di Zoologia e Anatomia comparata della Reale Universita di Genova, 7, 1 - 8.","Stephenson, J. (1924) Oligochaeta of the Siju Cave, Garo Hills, Assam. Records of the Indian Museum of Calcutta, 26, 127 - 137."]}

Cernosvitoviella atrata (Bretscher, 1903) Marionina atrata Bretscher, 1903: 17. Enchytraeoides atratus (Bretscher). Černosvitov 1928: 14 ���16, text fig. 3, pl. 1 figs 8, 9. Cernosvitoviella atrata (Bretscher). Nielsen & Christensen 1959: 38 ���39, figs 15���18. Chalupsk�� 1992: 140 ���141, fig. 7. Rota & Healy 1999: 31. Timm 2009: 148 ���149. Schmelz & Collado 2010: 57, 62. Material examined. IG 327183 - 22, one adult specimen, unstained whole mount. Honduras, Cusuco National Park [15.505813 -88.21473], 2061 m asl, M. Jocque 29 /06/ 2013. Description. Length 1.8 mm, diameter at V 0.12 mm, 0.17 at XII, 0.1 mm at XX, 23 segments. Chaetae sigmoid with nodulus, formula 4,5-4,3: 5,6-4,3. Length c. 25 ��m, diameter c. 1.5 ��m in caudal segments, chaetae appear smaller in anterior segments, length not measurable. Lateral chaetae slightly shifted dorsally, bundles completely in dorsal body half; distance of dorsal bundles to lateral line about half of that of ventral bundles (c. 17 vs. 33 ��m). Epidermal gland cells not seen, circumferal epidermis somewhat rugose at chaetal level. Body wall 4 ��m thick dorsally, up to 6 ��m ventrally. Prostomium 60 ��m high at base, 45 ��m long, hollow, very few transverse muscles inside, no papillae or protrusions, inner epithelial surface slightly wavy. Prostomium separated from peristomium by a medial dorsal-ventral ligament (musculature?). Dorsal and ventral chaetal bundles of one side in a segment interconnected with two transverse intracoelomic ligaments, possibly retractor muscles. Brain deeply incised posteriorly, 95 ��m long, width not measured. No oesophageal appendages, no intestinal diverticula. Four pairs pharyngeal glands, two primary pairs in IV, V, broadly attached to dorso-anterior face of respective septa, two ventral secondary pairs in V, VI, larger than primary glands, immediately behind but not attached to septa. Chloragocytes from V, dense coverage from VIII, cell diameter c. 10 ��m. Dorsal blood vessel from XIII, anterior bifurcation prostomial. Midgut pars tumida not distinguished. Ventral nerve cord medullar, with promiment segmental lateral swellings in IV���VI, swellings continuing in VII f., but gradually fading out. Preclitellar nephridia unpaired at 6 / 7 and 8 / 9 to the left, at 8 / 9 to the right. Anteseptale an elongate funnel, postseptale covering entire segment length, lobed, loops of canals visible. Coelomocytes few in number, different anteriorly and posteriorly; in anterior segments small and inconspicuous, oval to rhomboid, with fine irregular vesicles, length c. 12 ��m; in hindmost 4 segments an aggregation of larger cells, length 16���19 ��m, with conspicuous large vesicles of equal size at periphery; both types nucleated. Clitellum absent ventrally and dorsally, in XII���XIII, 2 complete segment lengths, beginning 1 epidermal cell row behind chaetae of XI, ending 1 cell row behind chaetae of XIII, consisting of c. 31 transverse rows. Seminal vesicle absent, few cysts free dorsally in XI. Sperm funnel funnel-shaped, 2 x as long as wide, c. 30 ��m long, tapering from 15 ��m entally to 8 ��m near septum; vas deferens thin-walled, widened to 6���8 ��m diameter behind septum 11 / 12, narrowing ectally to 3.5 ��m diameter near ectal pore. Male pores on body surface, with irregular glandular mass (diameter c. 16 ��m) surrounding most distal part of vas deferens. One mature egg. Spermathecae not attached to oesophagus, a simple tube c. 4 x as long as wide, with ectal duct 8���10 ��m wide and elongate ampulla 10���12 ��m wide; no glands or other modifications near ectal pore. Remarks. Cernosvitoviella is a Holarctic aquatic genus, common in temperate and boreal sediments of limnic and river habitats, and in moist soil. Its presence in phytotelmata of a tropical cloud forest is surprising from a biogeographical point of view but not in contradiction with the known habitat requirements. There are few records of the genus outside the Holarctic. Botea (1983) identified a single specimen from groundwater habitats in Cuba as " Cernosvitoviella sp." but the illustrations (Botea 1983, fig. 18 A,B) show a specimen with only two chaetae per bundle, which suggests that the specimen was misidentified. Reference material is no longer available (V. Pop. pers. com.). Cernosvitoviella has further been recorded from rain forest soil in Papua New Guinea (Standen 1988). Recently it was also found in peat soil on mountain peaks of the Mata Atl��ntica in Paran��, Brazil (R. Schmelz, C. Niva, unpublished). We consider our find as the first record of the genus from Central America. The single specimen is well-preserved and allows investigation of all relevant characters except presence/ absence of dark coelomocyte granules, seen only in living specimens. Our identification of specimens as C. atrata is mainly based on the shape of the spermatheca and the male efferent apparatus, the latter matching closely the redescription in Nielsen & Christensen (1959, Fig. 18, reproduced in Schmelz & Collado 2010). Further characters also fit the species diagnosis: coelomocytes differing in shape anteriorly and posteriorly, clitellum open ventrally and dorsally (Schmelz pers. obs.), 2 + 2 pattern of pharyngeal glands. C. atrata is highly variable and probably a species complex (Timm 2009, Schmelz & Collado 2010). Several forms which may be separate species have been distinguished, based on body size, coelomocyte variations and size of spermathecae (Nielsen & Christensen 1959, Chalupsk�� 1992), but none of the forms are well-described; furthermore the original description (Bretscher 1903) is poor and there are no types. Therefore it remains undecided whether this Honduras specimen belongs to one of the microspecies within C. atrata as known from the northern hemisphere, which would suggest a cosmopolitan distribution, or whether it is a species of its own. Nevertheless, referring to current taxonomic knowledge, this specimen belongs to C. atrata., Published as part of Schmelz, R��diger M., Jocque, Merlijn & Collado, Rut, 2015, Microdrile Oligochaeta in bromeliad pools of a Honduran cloud forest, pp. 508-526 in Zootaxa 3947 (4) on pages 516-517, DOI: 10.11646/zootaxa.3947.4.3, http://zenodo.org/record/242615, {"references":["Bretscher, K. (1903) Beobachtungen uber die Oligochaeten der Schweiz, VII. Folge. Revue Suisse de Zoologie, 11, 1 - 21.","Cernosvitov, L. (1928) Die Oligochaetenfauna der Karpathen. Zoologische Jahrbucher. Abteilung fur Systematik, Geographie und Biologie der Tiere. Jena, 55, 1 - 28.","Nielsen, C. O. & Christensen, B. (1959) The Enchytraeidae. Critical revision and taxonomy of European species (Studies on Enchytraeidae VII). Natura Jutlandica, 8 - 9, 1 - 160.","Chalupsky, J. (1992) Terrestrial Enchytraeidae (Oligochaeta) and Parergodrilidae (Polychaeta) from Sweden, with description of a new enchytraeid species. Zoologica Scripta, 21, 133 - 150. http: // dx. doi. org / 10.1111 / j. 1463 - 6409.1992. tb 00316. x","Rota, E. & Healy, B. (1999) A taxonomic study of some Swedish Enchytraeidae (Oligochaeta), with descriptions of four new species and notes on the genus Fridericia. Journal of Natural History, 33, 29 - 64. http: // dx. doi. org / 10.1080 / 002229399300461","Timm, T. (2009) A guide to the freshwater Oligochaeta and Polychaeta of Northern and Central Europe. Lauterbornia, 66, 1 - 235.","Schmelz, R. M. & Collado, R. (2010) A guide to European terrestrial and freshwater species of Enchytraeidae (Oligochaeta). Soil Organisms, 82, 1 - 176.","Botea, F. (1983) Oligochetes souterraines de Cuba. Resultats des expeditions biospeologiques cubano-roumaines a Cuba, 4, 19 - 38.","Standen, V. (1988) Oligochaetes in fire climax grassland and conifer plantations in Papua New Guinea. Journal of Tropical Ecology, 5, 39 - 48. http: // dx. doi. org / 10.1017 / S 0266467400002480"]}

Pristina sp. 1 Material examined. One juvenile specimen, not contracted, unstained whole mount, IG 327183 - 34 [15.506299 -88.23704], 1457 m asl, M. Jocque 07/07/ 2013. Description. Length 1.5 mm, diameter c. 0.1 mm, 20 segments. Dorsal and ventral chaetae from II, dorsally 1 hair and 1 needle per bundle, ventrally 4 bifids per bundle in II���XII, from XIII 3 per bundle. Hairs smooth, max. length 160 ��m (many with distal part broken off); needles simple-pointed, 25���30 ��m long. Ventrals with teeth equally long anteriorly (c. 1 ��m); posteriorly lower tooth slightly longer than upper tooth; length of ventrals 26 ��m in II, 28 ��m in VII, posteriorly ca. 30 ��m; anteriorly 4 or 5 per bundle, posteriorly mostly 3, in last 2 segments 2. Prostomium 40 ��m long, 60 ��m high at base, without inner protuberances, i.e. inner epithelial surface smooth. Pharyngeal glands at 4 /5, 5/ 6, first nephridium in VIII. Gut widening gradually in VIII. Coelomocytes not seen. Body wall incrusted with numerous conspicuous bodies, probably monocystid parasites, with smooth, wavy, surface, as if with multiple cuticle, oval, slightly longer than wide (length 12���14 ��m, diameter 8���10 ��m), with irregular non-vesicular rugged texture. Remarks. This specimen is similar to P. terrena but differs in fewer and smaller ventral chaetae and in smooth hairs. Acanthodrilidae, Benhamiinae Dichogaster (Diplothecodrilus) annae (Horst, 1893) (Ref.: Csuzdi & Pavlicek 2009) Dichogaster (Diplothecodrilus) bolaui (Michaelsen, 1891) (Ref.: James & Brown 2006) Dichogaster (Diplothecodrilus) tristani Cognetti, 1907 (Ref.: Michaelsen 1912; Picado 1913, as D. pitahayana Michaelsen) Eutrigaster (Eutrigaster) athena (James, 1996) Eutrigaster (Eutrigaster) bromeliocola (James, 2004) Eutrigaster (Graffiona) arborea (James, 1996) Eutrigaster (Graffiona) azul Sherlock & Csuzdi, 2011 Eutrigaster (Graffiona) basseterrensis (James, 1996) Eutrigaster (Graffiona) caesiticifusca (James, 1996) Eutrigaster (Graffiona) callaina (James, 1996) Eutrigaster (Graffiona) georgei Sherlock & Csuzdi, 2013 Eutrigaster (Graffiona) girija (James, 1996) Eutrigaster (Graffiona) johnsoni (James, 2004) Eutrigaster (Graffiona) picadoi (Michaelsen, 1912) (Ref.: Picado 1913) Eutrigaster (Graffiona) sporadonephra (Cognetti, 1905) (Ref.: Picado 1913, Fragoso & Rojas-Fern��ndez 1996) Acanthodrilidae, Acanthodrilinae Kaxdrilus sherbutti Sherlock & Csuzdi, 2013 Megascolecidae Amynthas rodericensis (Grube, 1879) (Ref.: Csuzdi & Pavlicek 2009) Glossoscolecidae Andiodrilus biolleyi Cognetti, 1904 (Ref.: Picado 1913) Andiodrilus cabalensis Righi, 1995 (Ref.: Feijoo & Celis 2012) Andiodrilus orosiensis Michaelsen, 1912 (Ref.: Picado 1913) Andiorrhinus planaria Michaelsen, 1934 (Ref.: Adis & Righi 1989; James & Brown 2006) Andiorrhinus tarumanis Righi et al., 1976 (Ref.: Adis & Righi 1989; James & Brown 2006) Fimoscolex ohausi Michaelsen, 1900 (Ref.: James & Brown 2006) Ocnerodrilidae Kerriona limae Righi, 1980 (Ref.: James & Brown 2006) Kerriona luederwaldti Michaelsen, 1926 (Ref.: James & Brown 2006) Enchytraeidae Bryodrilus hondurensis sp. nov. Hemienchytraeus phytotelmatus sp. nov. Naididae Aulophorus superterrenus Michaelsen, 1912 (Ref.: Picado 1913, Marcus 1943, Righi 1984, Christoffersen 2007) Pristina acuminata Liang, 1958 (Ref.: Harman et al. 1988) Pristina jenkinae (Stephenson, 1914) (Ref.: Marcus 1943, as Naidium roseum Piguet) Pristina menoni (Aiyer, 1930) (Ref.: C��r��ghino et al. 2011) Pristina notopora Černosvitov, 1937) (Ref.: C��r��ghino et al. 2011) Pristina osborni (Walton, 1906) (Ref.: C��r��ghino et al. 2011) Pristina terrena Collado & Schmelz, 2000, Published as part of Schmelz, R��diger M., Jocque, Merlijn & Collado, Rut, 2015, Microdrile Oligochaeta in bromeliad pools of a Honduran cloud forest, pp. 508-526 in Zootaxa 3947 (4) on pages 520-521, DOI: 10.11646/zootaxa.3947.4.3, http://zenodo.org/record/242615, {"references":["Csuzdi, C. & Pavlicek, T. (2009) New records of earthworms from Guadeloupe with description of a new species (Oligochaeta: Glossoscolecidae, Acanthodrilidae, Megascolecidae and Eudrilidae). Opuscula Zoologica Budapest, 40, 9 - 15.","James, S. W. & Brown, G. G. (2006) Earthworm ecology and diversity in Brazil. In: Moreira, F. M. S., Siqueira, J. O. & Brussaard, L. (Eds.), Soil biodiversity in Amazonian and other Brazilian ecosystems. CABI Publishing, Cambridge, MA, pp. 56 - 116. http: // dx. doi. org / 10.1079 / 9781845930325.0056","Michaelsen, W. (1912) Uber einige zentralamerikanische Oligochaten. Archiv fur Naturgeschichte, 78 A, 112 - 129.","Picado, C. (1913) Les bromeliacees epiphytes considerees comme milieu biologique. Bulletin Scientifique de la France et de la Belgique, 47, 215 - 360.","James, S. W. (1996) Nine new species of Dichogaster (Oligochaeta, Megascolecidae) from Guadeloupe (French West Indies). Zoologica Scripta, 25, 21 - 34. http: // dx. doi. org / 10.1111 / j. 1463 - 6409.1996. tb 00149. x","James, S. W. (2004) Earthworms (Clitellata, Acanthodrilidae) of the mountains of Eastern Jamaica. Organisms, Diversity & Evolution, 4, 277 - 294. http: // dx. doi. org / 10.1016 / j. ode. 2004.04.003","Sherlock, E., Lee, S., McPhee, S., Steer, M., Maes, J. M. & Csuzdi, C. (2011) The first earthworm collections from Nicaragua with description of two new species (Oligochaeta). Zootaxa, 2732, 49 - 58.","Sherlock, E. & Csuzdi, C. (2013) Two new earthworm species from Belize (Oligochaeta: Acanthodrilidae). Journal of Natural History, 47, 1911 - 1919. http: // dx. doi. org / 10.1080 / 00222933.2013.770931","Fragoso, C. & Rojas-Fernandez, P. (1996) Earthworms inhabiting bromeliads in Mexican tropical rainforests: ecological and historical determinants. Journal of Tropical Ecology, 12, 729 - 734. http: // dx. doi. org / 10.1017 / S 0266467400009925","Righi, G. (1995) Colombian earthworms. In: Van der Hammen, T. & Dos Santos, A. G. (Eds.), Studies on Tropical Andean Ecosystems / Estudios de Ecosistemas Tropandinos. Vol. 4. Cramer, Berlin, pp. 485 - 607.","Feijoo, M. A. & Celis, L. V. (2012) New species of earthworms (Oligochaeta: Glossoscolecidae) in the Amazon region of Colombia. Zootaxa, 103 - 119.","Adis, J. & Righi, G. (1989) Mass migration and life cycle adaptation - a survival strategy of terrestrial earthworms in Central Amazonian inundation forests. Amazoniana, 11, 23 - 30.","Righi, G., Ayres, I. & Bittencourt, E. (1976) Glossoscolecidae (Oligochaeta) do Instituto Nacional de Pesquisas da Amazonia. Acta Amazonica, 6, 335 - 367.","Marcus, E. (1943) Sobre Naididae do Brasil. Boletins da Faculdade de Filosofia, Ciencias e Letras da Universidade de Sao Paulo, 7, 3 - 247.","Righi, G. (1984) Oligochaeta. In: Schaden, R. (Ed.), Manual de Identificacao de Invertebrados Limnicos do Brasil, 17, pp. 1 - 48. [CNPq, Conselho Nacional de Desenvolvimiento Cientifico e Tecnologico, Brasilia]","Christoffersen, M. L. (2007) A catalogue of aquatic microdrile oligochaetes (Annelida: Clitellata) from South America. Acta Hydrobiologica Sinica, 31 (Supplement), 59 - 86.","Harman, W. J., Brinkhurst, R. O. & Marchese, M. (1988) A contribution to the taxonomy of the aquatic Oligochaeta (Naididae) of South America. Canadian Journal of Zoology, 66, 2233 - 2242. http: // dx. doi. org / 10.1139 / z 88 - 331","Stephenson, J. (1914) On a collection of Oligochaeta mainly from Northern India. Records of the Indian Museum of Calcutta, 10, 321 - 366.","Cereghino, R., Leroy, C., Carrias, J. - F., Pelozuelo, L., Segura, C., Bosc, C., Dejean, A., Corbara, B. (2011) Ant-plant mutualisms promote functional diversity in phytotelm communities. Functional Ecology, 25, 954 - 963. http: // dx. doi. org / 10.1111 / j. 1365 - 2435.2011.01863. x","Walton, L. B. (1906) Naididae of Cedar Point, Ohio. The American Naturalist, 40, 683 - 706. http: // dx. doi. org / 10.1086 / 278670","Collado, R. & Schmelz, R. M. (2000) Pristina silvicola and Pristina terrena spp. nov., two new soil-dwelling species of Naididae (Oligochaeta, Annelida) from the tropical rain forest near Manaus, Brazil, with comments on the genus Pristinella. The Journal of Zoology, London, 251, 509 - 516. http: // dx. doi. org / 10.1111 / j. 1469 - 7998.2000. tb 00806. x"]}

Genus Cernosvitoviella Nielsen & Christensen, 1959 Type species: Marionina atrata Bretscher, 1903, Published as part of Schmelz, R��diger M., Jocque, Merlijn & Collado, Rut, 2015, Microdrile Oligochaeta in bromeliad pools of a Honduran cloud forest, pp. 508-526 in Zootaxa 3947 (4) on page 516, DOI: 10.11646/zootaxa.3947.4.3, http://zenodo.org/record/242615, {"references":["Nielsen, C. O. & Christensen, B. (1959) The Enchytraeidae. Critical revision and taxonomy of European species (Studies on Enchytraeidae VII). Natura Jutlandica, 8 - 9, 1 - 160.","Bretscher, K. (1903) Beobachtungen uber die Oligochaeten der Schweiz, VII. Folge. Revue Suisse de Zoologie, 11, 1 - 21."]}

Genus Bryodrilus Ude, 1892 Type species: Bryodrilus ehlersi Ude, 1892 (by monotypy), Published as part of Schmelz, R��diger M., Jocque, Merlijn & Collado, Rut, 2015, Microdrile Oligochaeta in bromeliad pools of a Honduran cloud forest, pp. 508-526 in Zootaxa 3947 (4) on page 510, DOI: 10.11646/zootaxa.3947.4.3, http://zenodo.org/record/242615, {"references":["Ude, H. (1892) Ein neues Enchytraeiden-Genus. Zoologischer Anzeiger, 401, 344 - 345."]}

Enchylea heteroducta Nielsen & Christensen, 1963 (Fig. 7 A���C) Enchylea heteroducta Nielsen & Christensen, 1963: 11 ���12, Figs 14���17. Schmelz & Collado 2012: 54 f. Holotype and paratypes. Not designated in original description, not located, probably lost. Investigated material. Three spms, 1 adult, 2 early-stage subadults. Portugal, Coimbra, in soil from the experimental field area of the Coimbra Agricultural School (ESAC), crop site (Table 2); IV 2010. One of the subadults cut in halves: posterior half used for DNA extraction and sequence generation; anterior half as voucher in 100 % ethanol. Rest fixed in Bouin's fluid, stained with paracarmine, whole-mounted in Canada balsam. Description. Dimensions are from the single adult whole-mounted specimen, unless stated otherwise. Specific traits found only in one of the whole-mounted specimens are marked with "AD" (adult specimen) or "SA" (subadult specimen). Differences of the original description are inserted, marked by "NC". Length c. 5 mm (viv), 2.5���4 mm (fix); diameter (fix) 0.22 mm at V, XX, 0.26 mm at XII. Segments 25 (N= 2), NC: (27)��� 31 ���(35). Chaetae straight with weak ental hook, 2 and 3 per bundle, ventrally only 3. Formula 2,3 ��� 2,3: 3 ��� 3. In lateral bundles 2 chaetae mostly near clitellum. Chaetae absent at XII. Lateral chaetae in truly lateral position, not shifted dorsally. Chaetae mostly 40���50 ��m long and 3���4 ��m wide, largest in caudal segments (> 50 ��m), smallest in II (c. 35: 3 ��m); size differences between ventral and lateral chaetae not ascertained. Loose packages of detached chaetae present in the coelom in the adult specimen. Head pore mid-dorsally at 0/I, a longitudinal slit, length c. 15 ��m. Prostomium rounded, broadly conical, about as long as wide or high, shorter in fixed specimens. Epithelium without frontal recess, increasing in height towards the front, lined basally with musculature (not shown in Fig. 7 A). No sensory papillae apparent (fix), no prostomial papillae seen. Fibers of subepithelial muscular lining extending across lumen of prostomium in various longitudinal, transverse and oblique directions (not shown in Fig. 7 A). Peristomium c. 3 / 5 the length of following segments (II, III etc.). Epidermal gland cells not observed. No segmental dorsal pores. Body wall thin, c. 10 ��m, up to 20 ��m in anterior segments. Longitudinal muscle layer about as thick as layers of ring muscles, epidermis, and cuticle together. Cuticle very thin, ring muscles in irregular arrangement (optical section). Septa very thin, slightly thickened at 4 / 5���6 / 7. Brain 85: 60 ��m (fix), rounded or truncate posteriorly, sides converging anteriad, anteriorly concave; prostomial nerves with a pair of ganglia (SA), or ganglia not seen (AD). Ventral nerve cord medullar, slightly constricted and with fewer perikarya near the septa. Inner pair of postpharyngeal ganglia situated on top of oesophageal appendages; outer pair on afferent fascicles, enlarged and apparently with pharyngeal gland tissue in adult specimen (Fig. 7 A, arrow). Pharyngeal pad comparatively large, 130 ��m long and 90 ��m wide in adult specimen (fix), heavily invested with musculature dorsally, fibers extend backwards into V. Oesophageal appendages in III���IV, a pair of short, unbranched, free-floating, sac-like tubes with wide lumen, attached separately and adjacently to oesophagus dorsally behind pharyngeal pad and below post-pharyngeal ganglia, extending obliquely backwards and downwards, blind-ending ventrally below pharyngeal gland lobes of IV (AD); maximum diameter 40 ��m, estimated length c. 160 ��m; epithelium 3���4 ��m thick, not folded, smooth towards coelom, slightly rugose towards inner lumen. Pharyngeal glands in IV���VI, decreasing in size from IV to VI (AD) or of equal size in IV���VI (SA); dorsal lobes separate or with narrow dorsal connection, ventral lobes present from IV���VI, no secondary glands. Intestinal diverticula in VII, dorsally and laterally of oesophagus, with common unpaired opening into oesophagus dorsally, lateral wings widely connected dorsally of oesophagus, laterally flattened, extending forwards and backwards to the same degree; diverticular epithelium smooth, not folded, lined with blood sinus and chloragocytes, with histological properties different from adjacent oesophageal and intestinal epithelium: somewhat yellowish in whole mounts, not staining with paracarmine. Intestine widening strongly from c. 50 ��m diameter at 6 / 7 to c. 150 ��m in VIII; widening abrupt at 7 / 8 in contracted specimens. Pars tumida of midgut not distinguished. Intestine filled with amorphous humous substance mixed with sand grains up to 40 ��m in diameter. Chloragocytes with dense layer from VI, covering also intestinal diverticula; cell diameter c. 10 ��m (fix); in postclitellar segments cells often higher than wide (fix). Blood colourless, dorsal blood vessel from VII, large and pulsating, bifurcating in I below brain into circumesophageal connectives, these uniting into ventral vessel in IV. Two pairs of commissural vessels seen in III posteriorly, branching off dorsal vessel close to each other. Preclitellar nephridia 6 pairs, from 4 / 5 to 9 / 10. Postclitellar nephridia from 13 / 14, absent at several positions around XVII���XX; length c. 80 ��m; anteseptale with funnel only, postseptale laterally flattened, c. 1.5 x as long as high, with well-developed interstitial tissue, efferent duct about half the length of postseptale, rising terminally, nephroporus with small terminal vesicle, conspicuous in vivo. Coelomocytes one type, mucocytes, flat, broadly oval or broadly pear-shaped, c. 20 ��m long (viv), 16���22 ��m long and 10���12 ��m wide (fix), filled with small, slightly refractile vesicles (viv), nucleus and vesicles distinct (fix). Cells numerous, but not in dense aggregations. Clitellum in XII and XIII (NC: XII��� 1 / 2 XIII), extending beyond chatae of XIII by 2���3 transverse cell rows; girdle-shaped, cells in c. 32 dense rows, hyalocytes evenly distributed on all sides, present also mid-ventrally; celldiameter c. 8���10 ��m (fix), cells higher than wide laterally (c. 15 ��m, fix). Testes and sperm funnels in XI, male pores and ovaries in XII. Testes and developing sperm enclosed by a common membrane, i.e. sperm developing in testis sacs and not freely in the coelom. Two testis sacs present, one on each side, not lobed, confined to lateral sides of XI, occupying up to one segment length. Spermatozoa c. 50 ��m long, heads c. 12 ��m (fix); lengths in vivo not measured. Sperm funnel small, 1.5��� 2 x as long as wide (e.g. 60: 36 ��m, fix), tapering distad, collar as wide as funnel body. Vas deferens short (estimated length c. 400 ��m), not winded into coils, proximal region with wide ciliated lumen, tube diameter here c. 20 ��m, tapering distad more or less continuously down to 7���8 ��m near male pore. Tube walls c. 2 ��m thick throughout, cilia in lumen conspicuous. Male copulatory organ with bursa, gland, and copulatory muscles. Gland rounded and compact, c. 50 ��m long, 40 ��m wide, 30 ��m high (fix), bursal slit longitudinal, c. 25 ��m long, transverse copulatory muscles only in immediate vicinity of male gland. Subneural glands absent. Spermathecae as simple, mid-dorsally united tubes. Each tube with a double swelling, the ectal one possibly representing ectal duct, but cuticular lining not distinguished here and sperm present in both swellings. Ental swelling wider (c. 25 ��m) than ectal swelling (c. 20 ��m). No diverticula, no ectal glands. (NC: Ectal duct long and narrow, with distinct canal, sometimes with few separate gland cells near ectal pore). Ectal porus minute, at halflength between lateral chaetae of IV and V, slightly shifted ventrally. Two mature eggs in the adult specimen, each occupying about one segment length (NC: 1��� 8 eggs at a time). No intestinal or coelomic parasites observed. Remarks. Enchylea heteroducta had been described from laboratory cultures of unknown origin (Nielsen & Christensen 1963). This is the first record of the species from a natural habitat; the note in Schmelz and Collado (2012) refers to the same record. Specimens underlying the original description had more segments (27���35), up to 8 mature oocytes at a time, and a slightly different spermatheca: There was an ectal duct at the same place where we found a sperm-containing swelling. For the time being we assume that these differences are intraspecifi c. Several structures differ in appearance between the adult and the subadult specimen; more material is necessary to decide which character state is the 'valid' one. The reduced size of the hindmost pair of pharyngeal glands in the adult specimen may be a case of degeneration, although it agrees with the original description. Peculiar in this species are the combined presence, position and shape of oesophageal appendages and intestinal diverticula, and the short and stout vasa deferentia. Gut appendages are among the most important characters to distinguish genera among enchytraeids, therefore the new species was accomodated in a genus of its own. The oesophageal appendages are very similar to those in the Enchytraeus albidus group sensu Schmelz and Collado (2010), the only difference in the latter being their smaller size relative to the rest of the body and a thicker epithelium relative to the lumen diameter. Enchytraeus differs from Enchylea in the absence of intestinal diverticula, among other characters, and species of the E. albidus group are much larger than Enchylea, with body lengths around 3 cm. The intestinal diverticula in E. heteroducta are almost unique in the family: paired in VII laterally, broadly interconnected dorsally and with one common opening into the intestine. Perhaps they are better described in the singular form, as one dorsal diverticulum, bifurcating twice, first in lateral, then in antero-posterior direction. Such a structure is also described in Guaranidrilus sawayai Righi, 1973, the only species in this predominantly neotropical genus where the diverticula are not separate. Guaranidrilus differs from Enchylea in the oesophageal appendages (rounded and compact bodies in VI, when present), in not more than 2 chaetae per bundle and in free spermathecae, among other characters. Crop site Meadow site * Enchytraeus bulbosus Nielsen & Christensen, 1963 x x * Enchytraeus dichaetus Schmelz & Collado, 2010 x x = Enchytraeus christenseni bisetosus Rota & Healy, 1994 1 Form without refractile granules in coelomocytes (Schmelz & Collado 2010), possibly comprising several species. 2 Form with straight canal of spermathecal ectal duct near ampulla (Schmelz 2003: 291 f.). 3 Possibly new species but without reference material of sufficient quality. 4 Form with refractile granules in coelomocytes (Schmelz & Collado 2010), possibly comprising several species. 5 Form with spiral canal of spermathecal ectal duct near ampulla (Schmelz 2003: 291 f.). 6 The recent split of M. argentea into four species (Rota 2012) could not be considered here. Nielsen and Christensen (1963) suggested an affinity of Enchylea to Enchytraeus, because of the shape and position of the oesophageal appendages. The presence of testis sacs, one of the traits not mentioned originally but present in our material, supports their assumption. Testis sacs are known so far only in Enchytraeus and in Lumbricillus, the latter without oesophageal appendages. These two genera further agree with Enchlyea in the short nephridial anteseptale, which consists of not more than the funnel. However, Buchholzia is also a likely candidate. Species of this genus have oesophageal appendages and intestinal diverticula, and these are sufficiently similar to the respective structures in Enchylea to claim a homology. Noteworthy is the dorsal insertion of the intestinal diverticula, because lateral insertion is more common in enchytraeids (genera Henlea, Enchytronia, Guaranidrilus). However, diverticula in Buchholzia are compact bodies in VIII without lateral wings and with narrow canals. Oesophageal appendages differ in details of shape (more sac-like) and insertion (laterally and more posteriorly in IV), but other details (simple tubes with wide lumen inserting in IV) are similar. Other structures differ from Enchylea: The nephridial anteseptale is with small parts of the nephridial body, testis sacs are absent, and there are two types of coelomocytes. Further peculiarities in Enchylea, not shared by species of Enchytraeus or Buchholzia, are the short and stout vasa deferentia (present in Cernosvitoviella and some tropical Achaeta species), and the anterior location of the first nephridium (4 / 5, present also in some species of Henlea and Bryodrilus). Enchytraeus and Buchholzia occupy different branches in the molecular phylogenetic trees generated by Ers��us et al (2010). DNA has been extracted from one of the three specimens of Enchylea that we found, and future sequencing may help to clarify the phylogenetic relationships of Enchylea. Enchylea heteroducta was found in low numbers (3 specimens among thousands of enchytraeids), which means that the preferred habitat type of the species may differ from the types represented by the sites sampled in this study. The high number of oocytes mentioned in the original description (up to 8 per individual) and its first appearance in laboratory cultures suggests a lifestyle similar to that of other easily culturable enchytraeid species, all of them belonging to Enchytraeus, r-strategists adapted to exploiting high amounts of all kinds of organic material. We therefore believe that higher densities of this interesting species will be found in mediterranean compost heaps., Published as part of Schmelz, R��diger M. & Collado, Rut, 2013, Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963, pp. 307-328 in Zootaxa 3647 (2) on pages 322-326, DOI: 10.11646/zootaxa.3647.2.4, http://zenodo.org/record/218989

Fridericia roembkei sp. nov. (Figs 2 A���E, 5 B, Table 3) Holotype. MNHML MB 29 -0 0 0 302, adult spcm, stained whole mount. Portugal, Coimbra, in soil from the experimental field area of the Coimbra Higher School of Agriculture (ESAC), meadow site (Table 2); II 2012. Paratypes. 60 spms. MNHML MB 29 -000303-311, 9 adult spms, stained whole mounts. ZMH OL 14520, stained whole mounts: 10 spms, 9 adults, 1 subadults. ZMH OL 14521, fixed in Bouin's fluid, preserved in 70 % ethanol: 8 spms. ZMH OL 14522, fixed in 70 % ethanol, preserved in 100 % ethanol: 10 spms. Other material. 134 spms investigated in vivo, preserved in collective sample vials, in the authors' collection. Etymology. Named in honour of J��rg R��mbke, enchytraeidologist, soil ecologist and ecotoxicologist at ECT Oekotoxikologie GmbH Fl��rsheim, research director of the TME experiments, in grateful recognition of his varied initiatives to support taxonomic work with enchytraeids. Diagnosis. Less than 40 segments, max. 4 chaetae per bundle, clitellum girdle-shaped, cells absent between bursal slits, coelomo-mucocytes without refractile vesicles, nephridia present at 10 / 11, chylus cells post-clitellar, seminal vesicle absent, sperm funnel small, spermathecae joint entally, without ectal gland, ectal duct longer than body diameter, two stalked diverticula with ciliated subchamber and ciliar movement. Description. Small Fridericia species. Length 8���10 mm when relaxed, 4���5 mm when contracted (viv), 6���7.5 mm (fix), diameter 0.2���0.25 mm (viv), 0.2���0.26 at V, 0.28���0.36 mm at XII���XIII, 0.22���0.29 mm in postclitellar segments (fix). Segment number (29)��� 34���38 (N = 40), mostly (75 %) 36���38. Chaetae max. 4 per bundle, formula 2,3, 4 ��� 4,3, 2: (2,3,) 4 ��� 4,3, 2. Posterior 12���20 segments with only 2 chaetae per bundle; often all lateral postclitellar bundles with 2 chaetae. Chaetae in caudal segments largest, 65���70: 5.5 ��m, size of largest anterior chaetae c. 52���55: 5.5 ��m. Inner chaetae in bundles of 4 always smaller than outer, c. 2 / 3 the size of outer chaetae. Epidermal gland cells (Fig. 2 C,D) in 1���3 rows per segment, mid-row at chaetal level, cells pale or yellowish, in regular, alternating pattern, pale cells with clear outline, quasi-rectangular, yellow cells with indistinct outline, smaller; gland cells indistinct or seemingly absent in some specimens. Body wall comparatively thin, 10���15 ��m thick, longitudinal muscle layer not or only slightly thicker than layer of ring muscles plus epidermis; cuticle very thin (c. 125 ��m (fix), length ratio anteseptale: postseptale 2: 5, no constriction at septum, medial rise of efferent duct; postclitellar nephridia from 13 / 14. Coelomo-mucocytes pale with blurred vesicles, lenticytes minute, very numerous, optically dominating. Clitellum girdle-shaped, absent between bursal slits, prominent, distinctly higher (c. 5��� 7 x) than non-clitellar epidermis, cells in dense to indefinite rows, hyalocytes and granulocytes alternating, laterally of bursal slits only granulocytes; average diameter of hyalocytes 15 ��m, of granulocytes 10 ��m, cell height 15���20 ���(25) ��m (fix). Seminal vesicle absent, developing sperm anteriorly in XI. Spermatozoa not numerous on sperm funnel, sperm heads not measurable in vivo, at least 60 ��m (fix), probably longer. Sperm funnel small, c. 2 x as long as wide (e.g. 100: 50 ��m, or 120: 60 ��m, fix), tapering distad, collar about half as wide as funnel body. Vas deferens in wide loops parallel to ventro-lateral body wall, therefore difficult to see, 8 ��m wide. Male copulatory organ: bursa longitudinal, male gland c. 80 ��m long, 50 ��m wide, 40 ��m high (fix), kidney-shaped, with thick-walled bursa in concavity. Subneural glands and other accessory glands absent. Spermatheca: no ectal gland; ectal duct c. 250 ��m long, longer than body diameter, c. 4 x as long as ampulla, coiled in contracted specimens, diameter 12 ��m, with distal swelling up to 20 ��m; proximal endpiece (ental bulb) not widened; ampulla with two stalked diverticula oriented ectad, diverticula with ciliated subchamber and rotating spermatozoa in peripheral chamber; ampullae joint entally and with one common attachment to oesophagus dorsally or dorso-laterally. Aspect of joint ampullae plus diverticula varied: common lumen inflated, or collapsed, inner surface smooth or wavy, organs contorted with left ampulla to the right and vice versa, or ampullae compressed in dorso-ventral position with diverticula approached and in staggered position (Fig. 2 A,B). One mature oocyte at a time, occupying up to 3 segment lengths. Remarks. Fridericia roembkei sp. nov. belongs to a group of Fridericia species characterized by two spermathecal diverticula with a ciliated subchamber: F. perrieri (Vejdovsk��, 1878), F. o m e r i Stephenson, 1932, F. rendsinata D��zsa-Farkas, 1972, F. u l r i k a e Rota & Healy, 1999, F. h e a l y a e Schmelz, 2003, F. dozsae Schmelz, 2003 (Schmelz 2003: 296, 341 f.), and partly also F. galba (Hoffmeister, 1843). It differs, together with F. m a rg i na ta described below, from all species of this group in the proximal fusion of the spermathecal ampullae. Using the tabular comparison of Fridericia species with two spermathecal diverticula in D��zsa-Farkas (2009), F. roembkei sp. nov. belongs to a group of 10 species with proximally fused spermathecae. It differs from all species of this group in the absence of spermathecal ectal glands, in an exceptionally long spermathecal ectal duct, and in the ciliation of the diverticula. Differences to F. marginata sp. nov., a remarkably similar species, are dealt with in the remarks section of that species, see also Figure 5 and Table 2. The aspect of the spermathecae in F. roembkei varies considerably. The ampullae are often compressed, twisted, and in upright position, as shown in Fig. 2 A, or they are inflated into a spherical chamber���much more than shown in Fig. 2 B���mimicking the aspect in F. gamotheca (Fig. 2 F). Both aspects can be found at different times in the same living specimen. These unusual variations may be related to the thin and apparently soft ampullar walls. In the key to species of Fridericia in Schmelz (2003), F. roembkei would key out together with F. g a m o t h e c a Issel, 1905. F. gamotheca as originally described (Issel 1905) and redescribed (Rota 1995) from several Italian localities is distinguishable from F. roembkei by the more oval spermathecal diverticula, a shorter spermathecal ectal duct (1.5 x times as long as the ampulla), a larger sperm funnel, and higher segment number (40���44). Furthermore, diverticula are not ciliated (Schmelz pers. obs.) and the inflated aspect of the joint ampullae is a constant feature, not a transitory state as in F. roembkei. Some further possible differences are not straightforward because of variations noted in Rota (1995) regarding shape of diverticula, seminal vesicles and spermathecal ectal glands. Another variant of F. gamotheca from Morocco of uncertain taxonomic status (Schmelz 2003: 193 f.) has spherical diverticula, only 28���33 segments (D��zsa-Farkas 1989), and a ventral clitellum as described for F. roembkei: girdle-shaped, absent between bursal slits (Schmelz 2003). It differs from F. roembkei in a short and thick ectal duct and in the presence of spermathecal ectal glands. The morphological variability documented in the redescriptions of D��zsa-Farkas (1989) and Rota (1995) have led Collado et al. (2012) to assume that F. g a m o t h e c a is a species complex. Remarkable in F. roembkei is further the presence of both pale and yellow epidermal gland cells (Fig. 2 C,D), distinct only in living specimens, and clearly representing two different cell types. The pale cells are quasirectangular, filled with pale vesicles and mostly arranged in one row at chaetal level, while the yellow cells are smaller spots of irregular outline, not arranged in one row but nonetheless distributed with some regularity., Published as part of Schmelz, R��diger M. & Collado, Rut, 2013, Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963, pp. 307-328 in Zootaxa 3647 (2) on pages 312-314, DOI: 10.11646/zootaxa.3647.2.4, http://zenodo.org/record/218989

Fridericia marginata sp. nov. (Figs 3 A���D, 5 D, Table 3) Holotype. MNHML MB 29 -000312, adult spcm, stained whole mount. Portugal, Coimbra, in soil from the experimental field area of the Coimbra Higher School of Agriculture (ESAC), meadow site (Table 2); II 2012. Paratypes. 15 spms. MNHML MB 29 -000313 ��� 315, stained whole mounts, 2 adults, 1 subadult. ZMH OL 14523: 4 juvenile spms, stained whole mounts. ZMH OL 14524, fixed in Bouin's fluid, preserved in 70 % ethanol: 8 spms. ZMH OL 14525, fixed in 70 % ethanol, preserved in 100 % ethanol: 5 spms. Other material. 37 spms investigated in vivo, preserved in collective sample vials, in the authors' collection. Etymology. Named for the ventro-lateral margins of the clitellum, consisting only of granulocytes. Diagnosis. Less than 40 segments, max. 4 chaetae per bundle, clitellum saddle-shaped with ventral margins consisting of granulocytes only, coelomo-mucocytes with refractile vesicles, nephridia present at 10 / 11, chylus cells post-clitellar, no seminal vesicle, sperm heads 90 ��m, sperm funnel small, spermathecae joint entally, without ectal gland, two stalked diverticula with ciliated subchamber, ampulla and diverticula elongate. Description. Body colour opaque grey-white, often with a yellowish tinge (viv), slow body movements. Length c. 7���8 mm (viv), diameter c. 0.3 mm (viv). Segments 35���39. Chaetae max. 4 per bundle, formula 3,4 ��� 4,3, 2: (3), 4 ��� 4,3, 2. Ventrally 2 per bundle from XXV to posterior end. Inner chaetae shorter than outer. Chaetae in caudal segments largest, 65���70: 5.5 ��m, size of largest anterior chaetae c. 55: 4.5 ��m. Epidermal gland cells scarce, pale, one row at chaetal level, cells quadrangular or without clear outline. Body wall c. 20 ��m thick with longitudinal muscle layer thickest (c. 15 ��m), cuticle thin (c. 1 ��m, matrix transparent; cells very numerous, filling all spaces, making body opaque with grey-yellow tinge (viv); lenticytes minute, length c. 6���8 ��m, sparse. Clitellum saddle-shaped, prominent, cells in c. 35 dense rows with granulocytes and hyalocytes alternating; mid-ventral aclitellar field as wide as distance between bursal slits; hyalocytes absent at ventro-lateral borders and in a semi-circular field laterally of and around bursal slits; here only granulocytes in indefinite rows or in reticulate arrangement. (Single hyalocytes near ventral border between male and female pores observed in one specimen). Seminal vesicle absent; occasionally dense masses of developing sperm in anterior half of XI, and septum 10 / 11 bulged forward dorsally, unpaired. Spermatozoa> 200 ��m long, heads c. 90 ��m, not numerous on top of sperm funnel (Fig. 3 C). Sperm funnel comparatively small, hardly visible among coelomocytes, developing sperm, and oogonia (viv), about half as long as body diameter, tapering proximally, c. 2.5 x as long as wide (e.g. 140: 60 ��m), collar not wider than funnel body, not folded outwards. Vas deferens in dense irregular coils ventro-laterally, 8 ��m wide. Male copulatory organ elevating body suface even when completely retracted (viv), transverse copulatory muscles strongly developed. Male glands broadly oval in ventral view, flat in side view, 1.5 x as long as wide (e.g. 120 ��m long, 80 ��m wide, 50 ��m high) bursal slits longitudinal, bursa thick-walled, with small and flat modiolus (comp. Schmelz 2003: 49) around ectal opening of vas deferens (the primary male pore); bursa plus male gland almost spherical. Subneural glands and other accessory glands absent. Spermathecae: no ectal gland; ectal duct about as long as body diameter, slightly widened in distal part, tapering again near ectal pore, ental bulb not projecting into ampullar lumen, only slightly wider than ectal duct, ectal duct canal straight throughout. Ampullae 1.5��� 2 x as long as wide, tapering entad, ental tips fused, creating a common V-shaped lumen. Ampullar walls 4���6 ��m thick, uniform, no histological differences observed, with smooth outer and inner surface, inner surface occasionally wavy, as a transitory state during contraction. Diverticula inserting on opposite sides of ental bulb at base of ampulla and projecting obliquely ectad, distance (= maximum width of spermatheca) c. 80 ��m, c. 1 / 3 body diameter. Each diverticulum 2 x as long as wide, diameter c. 20 ��m, lumen subdivided into peripheral spermcontaining chamber and ciliated sub-chamber; both chambers connected by a porus; peripheral chamber wider than long, with dense sperm roll circulating; sub-chamber longer than wide, with ciliar movement, widely connected with ampullar lumen. No ciliar movement in ampulla. One mature oocyte at a time, extending over c. 2 segment lengths. Remarks. The new species owes its name to the saddle-shaped clitellum with ventro-lateral margins consisting only of granulocytes, a character new in Fridericia, but common in some other genera, e.g. Guaranidrilus, Hemienchytraeus, Achaeta, Marionina. In the other Fridericia species with saddle-shaped clitellum the ventro-lateral borders do to our knowledge not differ from the rest and are composed of granulocytes and hyalocytes. However, not all species of the genus have been examined for that character. A further peculiarity is the slenderness of ampulla and diverticula, both about twice as long as wide. Using the tabular comparison of Fridericia species with two spermathecal diverticula in D��zsa-Farkas (2009), F. marginata sp. nov. belongs to a group of 10 species characterized by proximally fused spermathecae. None of these species has the character combination of max. 4 chaetae per bundle, saddle-shaped clitellum, and stalked spermathecal diverticula. Using the key to Friderica species in Schmelz (2003), F. marginata would key out together with F. gamotheca Issel, 1905. This species has also coelomocytes with few refractile vesicles, as F. marginata (Rota 1995, Schmelz pers. obs.). It differs from F. marginata in larger body size and higher segment number (37���50 according to Rota 1995), in a girdle-shaped clitellum according to Schmelz and Collado (2010), and in a complete fusion of the two spermathecal ampullae. A variant of F. gamotheca described from Morocco (D��zsa-Farkas 1989) and of uncertain taxonomic status (Schmelz 2003) is more in the size range of F. m a rg i na ta (length 5���7 mm, 28���33 segments), but differs in a girdle-shaped clitellum, coelomocytes without refractile vesicles and in the presence of a spermathecal ectal gland, among other characters. F. gamotheca is a problematic species, see F. roembkei sp. nov., remarks. F. m a rg i n a t a and F. roembkei spp. nov. are quite similar at first sight, but they are easily distinguished by the coelomocytes���with refractile vesicles in F. m a rg i n a t a and without them in F. roembkei ���and by the ventral pattern of the clitellum, see Figure 5. Further differences are more difficult to observe and may not be evident unless specimens of both taxa are available for comparison (in the following, character states of F. marginata first): spermathecal diverticula and ampulla elongate/not elongate; spermathecal ectal duct not longer/longer than body diameter; yellow epidermal gland cells absent/often present; pharyngeal glands in VI united/separate dorsally (Table 2)., Published as part of Schmelz, R��diger M. & Collado, Rut, 2013, Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963, pp. 307-328 in Zootaxa 3647 (2) on pages 314-316, DOI: 10.11646/zootaxa.3647.2.4, http://zenodo.org/record/218989

Fridericia sousai sp. nov. (Figs 1 A���C, 5 A, Table 3) Holotype. MNHML MB 29 -000297, adult spcm, stained whole mount. Portugal, Coimbra, in soil from the experimental field area of the Coimbra Higher School of Agriculture (ESAC), meadow site (Table 2); II 2012. Paratypes. Eight spms. MNHML MB 29 -000298 ��� 301, stained whole mounts: 3 adults, 1 juvenile. ZMH OL 14518, fixed in Bouin's fluid, preserved in 70 % ethanol: 1 adult, 1 juvenile. ZMH OL 14519, fixed in 70 % ethanol, preserved in 100 % ethanol: 2 spms. Other material. 7 spms, investigated in vivo, preserved in collective sample vials, in the authors' collection. Etymology. Named in honour of Paulo Sousa, soil zoologist, ecologist and ecotoxicologist at the University of Coimbra, research director of the TME experiments. Diagnosis. Length Description. Small Fridericia worms. Length 6���8 mm (viv), 4.5���6 mm (fix); diameter 0.15 mm (viv), 0.16 mm (fix), 0.2 mm at clitellum. Segment number 30���35. Chaetae max. 4 per bundle. Formula 2,3, 4 ��� 4,3, 2: 2,3, 4 ��� 4,3, 2. In preclitellar bundles mostly 4 chaetae, in postclitellar bundles 4 and 3 chaetae, more 3 than 4 in many specimens; two chaetae only in II and in the hindmost 1 or 2 segments. Anteriorly inner chaetae almost as large as outer, posteriorly all chaetae in a bundle of same size. Posterior chaetae slightly larger than anterior (52���55: 3���3.5 ��m and c. 30���40: 2���3 ��m, respectively). Chaetae increasing in size from II to VII (from c. 30 ��m to c. 45 ��m); chaetae in segments following clitellum as large as preclitellar; caudal chaetae largest (up to 60: 4 ��m). Epidermal gland cells one row per segment, at chaetal level, few cells, one each dorsally of lateral chaetae. Body wall c. 15 ��m thick, longitudinal muscle layer comparatively thin, 1���1.5 x as thick as layer of ring muscles plus epidermis, cuticle very thin, barely visible at 400 x magnification, estimated thickness c. 17 / 18, shape as preclitellar, terminal rise of efferent duct. Coelomomucocytes with small refractile vesicles at periphery, cell length 15���20 ��m, matrix almost hyaline (viv); lenticytes numerous but inconspicuous (viv), length 5���6 ��m, i.e. about 1 / 3 of mucocyte length. Clitellum girdle-shaped, cells in c. 25 dense rows, flat (i.e. cells wider than high), conspicuous in vivo, inconspicuous in whole mounts, also present between bursal slits; hyalocytes and granulocytes of same size, diameter 8���10 ��m (fix). Seminal vesicle absent, developing sperm in anterior half of XI. Spermatozoa c. 80 ��m long, heads 20���25 ��m long. Sperm funnel small, length c. 40 ��m long, 30 ��m wide, 30 ��m high (fix). Subneural glands and other accessory glands absent. Spermatheca: small ectal gland, ectal duct shorter than body diameter, c. 1.5 x as long as ampulla, c. 90 ��m long, diameter c. 12 ��m, proximal endpiece slightly projecting into ampulla, not widened; two broadly sessile diverticula or diverticula-like protrusions, diameter c. 18 ��m, sperm-containing; ampulla tapering proximad, diverticula and ampulla apparently solid (viv) but lumina visible in whole mounts. One mature oocyte at a time, extending over 1.5���2 segment lengths. Remarks. Using the tabular comparison of Fridericia species with two spermathecal diverticula in D��zsa- Farkas (2009), F. sousai sp. nov. belongs to a group of 11 species, characterized by separate spermathecae and globular, hemispherical, or short-stalked diverticula. In this group, only F. isseli Rota, 1994 shares the following combination of characters with F. sousai: length F. isseli differs from the new species in the following traits, among others: (1) chylus cell canals branched, not widened (vs. unbranched, sac-like), (2) clitellum saddle-shaped (vs. girdle-shaped), (3) male gland tripartite with two small accessory glands (vs. accessory glands absent). In the key to Fridericia species in Schmelz (2003), F. sousai would key out together with F. isseli and with F. w a l d e n s t ro e m i Rota & Healy, 1999. The latter differs from F. s o u s a i in having 40���54 segments, five preclitellar pairs of nephridia, a small subneural gland in XIV, and proximally fused spermathecae with long-stalked diverticula, among other characters. Figure 5 and Table 2 give a comparison of F. sousai with the other three new species of Fridericia from the same site., Published as part of Schmelz, R��diger M. & Collado, Rut, 2013, Enchytraeidae (Oligochaeta, Annelida) from a field site in Portugal, with the description of five new species and a redescription of Enchylea heteroducta Nielsen & Christensen, 1963, pp. 307-328 in Zootaxa 3647 (2) on pages 310-311, DOI: 10.11646/zootaxa.3647.2.4, http://zenodo.org/record/218989

Guaranidrilus hoeferi sp. nov. (Figs 4, 8 D) Holotype. UFPR OL- 13, adult/subadult specimen, stained whole mount, Guareque��aba, Itaqui, 25 �� 14 '45,3''S, 48 �� 30 '19,4''W, 27 m a.s.l., medium old secondary forest on Cambisol [site 38], Sep 2007, leg. P. Heine, R.M. Schmelz. Paratypes. MZUSP, 6 specimens, Antonina, Cachoeira, stained whole mounts: MZUSP 1230, 2 specimens, 25 �� 14 ' 38 '', 48 �� 40 ' 10 '', 140 m a.s.l., old-growth forest on Cambisol [site 13], May 2003, leg. J. R��mbke, R. M. Schmelz. MZUSP 1231, 4 specimens, 25 �� 19 '00"S, 48 �� 40 ' 14 "W, and 25 �� 19 ' 41 "S, 48 �� 40 ' 36 "W, 30 and 120 m a.s.l., abandoned pasture and medium old secondary forest on Cambisol [sites 6, 12], Oct 2004. leg. B. F��rster, R.M. Schmelz. UFPR, 44 specimens, Guareque��aba, Itaqui, stained whole mounts (35) and ethanol-preserved (9): UFPR OL-14, 5 specimens, same data as holotype. UFPR OL-15, 13 specimens, 25 �� 15 ' 43.8 ''S, 48 �� 29 ' 14.8 ''W, ca. 20 m a.s.l., young forest on Cambisol [site 36], Sep 2007, leg. P.Heine, R. M. Schmelz. UFPR OL-16, 17 specimens, 25 �� 15 ' 32.7 ''S, 48 �� 30 ' 31.9 ''W and 25 �� 16 ' 23.7 ''S, 48 �� 29 ' 13.7 ''W, 20 and 31 m a.s.l., respectively, old-growth forest on Cambisol [sites 41, 42], Sep 2007, leg. P.Heine, R. M. Schmelz. UFPR OL-17, 9 specimens, ethanol-preserved. Additional material. Fifteen specimens from sites in Itaqui, examined in vivo, not preserved. Etymology. Named in honour of Hubert H��fer (State Museum of Natural History, Karlsruhe), initiator and German coordinator of the SOLOBIOMA project, in grateful recognition of the tremendous amout of work to launch and to maintain this project, which allowed the discovery and description of so many enchytraeid species. Description. Slow body movements. Body dimensions. Mature specimens about 12���15 mm long in vivo, live diameter ca. 0.2 mm, submatures often distinctly smaller (Segment number 39���48. Chaetae distally straight, with weak proximal hook, not thickened proximally. Preclitellar chaetae small and stout, ca. 22���28 ��m long and and 2.5���3 ��m thick. Postclitellar chaetae behind clitellum small and inconspicuous, ca. 20 ��m long and 2 ��m wide; chaetae increasing gradually in size towards posterior end. In caudal segments chaetae stout again, 40���50 ��m long and 6 ��m thick. Epidermal gland cells inconspicuous in vivo. In whole mounts all epidermis apparently glandular, cells transversely elongate, hyaline, in about 11 rows per segment, almost continuous except mid-ventrally. Clitellum (Fig. 4 H) saddle-shaped, not developed ventrally; on dorsal half granulocytes and hyalocytes, granulocytes isolated, hyalocytes large, seen as large free interspaces in living specimens; ventro-laterally only granulocytes. On dorsal half cells in indefinite rows or reticulate; latero-ventrally cells in dense rows. In adults accessory glands present, a glandular thickening of the epidermis mid-ventrally in XIII or in XI and XIII, immediately in front of and behind clitellum (Fig. 4 A,K); glandular field between the ventral chaetae and the respective posterior septum. Head pore on prostomium mid-dorsally. Prostomium without inner papillae. Pygidium with strongly developed rectal musculature, filling body cavity. Body wall thin (ca. 10���20 ��m), ring muscles in one regular layer. Cuticle mostly thick compared to rest of body wall (up to 9 ��m), thicker than epidermis + ring muscle layer. Anterior septa not thickened. Brain posteriorly concave; a few perikarya on prostomial nerves, no ganglionic swelling. Pharyngeal glands (Fig. 4 A,B) in IV���VI, unpaired and fused dorsally; dorsal lobes of V and VI with ventral anterior projections (primary ventral lobes), large in VI, small in V; secondary ventral lobes present in V and VI, spherical. Oesophageal appendages (Fig. 4 A,B) paired dorso-laterally in anterior half of VI, elongate, with irregular outline. Intestinal diverticula (Fig. 4 A,B) in VII, as pouches with wide lumen, directed anteriad, i.e. connection with intestine posteriorly; walls with radial striation, possibly folds of inner wall; chloragocytes present on outer surface. Dorsal blood vessel (Fig. 4 A,B) from 1 / 2 VIII, pulsating. Preclitellar nephridia (Fig. 4 A,B,I) two pairs, at 7 / 8 and 8 / 9. Anteseptale large, with coils of canal, outline spherical in side view, postseptale elongate and narrower than anteseptale, with dorsal vesicle, gradually merging into short and stout efferent duct, no terminal vesicle. Postclitellar nephridia (Fig. 4 J) of similar shape but slightly longer, anteseptale longer than wide. Nephridia missing at several positions. Coelomocytes (Fig. 4 G) numerous, darkened in aggregations. Cells flattened, outline broad-oval, length ca. 20���40 ��m. A few dark and refractile granules of irregular outline scattered in a finely and regularly vesicular, pale-brown matrix. Seminal vesicle (Fig. 4 A) large, extending over up to 5 segments, caused by forward bulge of septum 10 / 11 and backward bulge of septum 11 / 12. Spermatozoa very numerous, heads 33 ��m long (measured in vivo). Sperm funnel (Fig. 4 A) longer than body diameter, ca. 2 x as long as wide (or longer), tapering distally; collar conspicuous, with free extensions beyond funnel body, often U-shaped, conspicuous by attached sperm. Each extension may have stump-like sub-lobes. Vas deferens (Fig. 4 A) very long and irregularly coiled, of equal diameter throughout (8 ��m in fixed material. Male copulatory organ (Fig. 4 A) with male pores on body surface, each withdrawn in a longitudinal eversible furrow; both furrows creating a mid-ventral "penial plate" when withdrawn. In all fixed specimens furrows everted into elongate-conical projections; male pores widely separate, distance wider than distance between ventral chaetal bundles of a segment. No male glands around male pore, but epidermis thickened here and also in a field between the pores, the "penial plate". Copulatory muscles (Fig. 1 A) conspicuous in preserved specimens as a parasagittal series of fine, dorso-ventral strands, arranged in paralles (side view): ventral insertion of muscles at inner wall of papilla, dorsal insertion at inner body wall surface latero-dorsally, slightly above longitudinal row of lateral chaetae. Spermatheca (Fig. 4 A,C,D,E,F) very large, extending into IX. Ectal duct short, distal part of ampulla conspicuous, thin-walled, with sperm and surrounding free lumen; ental reservoir extending over 3 segments, into IX, here elongately club-shaped, thick-walled; walls with large, non-staining inclusions of irregular outline (Fig. 4 F, "se"); lumen completely filled with spermatozoa, forming a long, compact and iridescent strand. One egg at a time seen, but no specimen with fully developed egg found. Habitat. G. hoeferi was found at all stages of forest regeneration. It was absent in pastures and only one specimen was found at the agroforestry sites. Remarks. Among all species present at the study sites, G. hoeferi was instantly distinguished by (1) the peculiar coelomocytes. Further traits that confirmed the identification were: (2) only 2 pairs of anterior nephridia, at 8 / 9 and 9 / 10, (3) dorsal blood vessel from VIII, (4) intestinal diverticula confined to VII, mostly directed forwards. With this character combination, specimens of all age stages, including very small juvenile ones, could be identified. The striking gigantism of sperm-related organs is shared by several other Guaranidrilus species that possess oesophageal appendages (G. atlanticus Christoffersen, 1977, G. cernosvitovi Healy, 1979, G. joanae Christoffersen, 1977, G. m b o i Righi, 1975), and hence is little help in the identification of the species. G. hoeferi is most similar to G. joanae Christoffersen, 1977, another species of the Brazilian Atlantic rain forest, found ca. 250 km up north-east near the city of S��o Paulo. Both may be indeed be sister species; at least they belong to a common sub-group or clade within the genus. The excellent original description of G. joanae allows a detailed comparison. Both species agree in body size, behaviour, all details of pharyngeal glands, oesophageal appendages, intestinal diverticula, and in size and organization of spermathecae and the entire male reproductive system. They differ in four traits: (1) In G. joanae, chaetae are enlarged not only in caudal segments but also in the most anterior segments, and here (2) the ental bend is thickened in a claviform manner. (3) Preclitellar nephridia are present not only at 7 / 8 ��� 8 / 9 but also at 6 / 7, resulting in three preclitellar pairs instead of the two in G. hoeferi. (4) Coelomocytes of G. joanae appear to lack the conspicuous double texture present in G. hoeferi (few coarse granules embedded in a densely and finely vesicular pale-brown matrix), they are described as being "rather inconspicuously granulated" (Christoffersen 1977: 200). Furthermore segment number is slightly higher (49���55) and clitellar gland cells are irregularly arranged in G. joanae, but our specimens were not fully mature. Our specimens were relaxed and Christoffersen's specimens apparently contracted at the time of fixation, which may account for differences in body length (10���11 mm in G. joanae) and shape of preclitellar nephridia (efferent duct bent forwards in G. joanae). Epidermal gland cells are said to be slightly developed in G. joanae (Christoffersen 1977: 198), but the animals "... were always found enveloped in great amounts of mucus..." (ibid.), which suggests the same situation as in G. h o e f e r i: numerous glands cells, inconspicuous in vivo. In fact even in fixed material the glands are invisible unless interference contrast optics are used. The ectal pore of the spermathecae of G. joanae appears to be located ventrally in a figure (Christoffersen 1977, Fig. 24), but the description gives the usual and probably correct lateral location., Published as part of Schmelz, R��diger M., Collado, Rut & R��mbke, J��rg, 2011, Mata Atl��ntica enchytraeids (Paran��, Brazil): A new genus, Xetadrilus gen. nov., with three new species, and four new species of Guaranidrilus ��ernosvitov (Enchytraeidae, Oligochaeta), pp. 1-29 in Zootaxa 2838 on pages 10-13, DOI: 10.5281/zenodo.203260, {"references":["Christoffersen, M. L. (1977) New species of Guaranidrilus Cernosv. (Enchytraeidae, Oligochaeta) from Serra do Mar, Sao Paulo, Brazil. Studies on Neotropical Fauna and Environment, 12, 187 - 206.","Healy, B. (1979) Review of the genus Guaranidrilus (Oligochaeta, Enchytraeidae) with the description of two new species. Bulletin of the British Museum of Natural History (Zoology). London, 37, 7 - 15.","Righi, G. (1975) Algumas Enchytraeidae (Oligochaeta) Brasileiras. Ciencia e Cultura. Sao Paulo (Journal of the Brazilian Association for the Advancement of Science), 27, 143 - 150."]}

Xetadrilus gen. nov. Genus "X" ��� R��mbke et al. 2007: 144 f. Type species: Xetadrilus maacki sp. nov. Included species: Xetadrilus maacki sp. nov. Xetadrilus aphanus sp. nov. Xetadrilus fabryi sp. nov. Xetadrilus pitucus (Righi, 1974 a) (ex Marionina) Xetadrilus righianus (Xie & Rota, 2001) (ex Marionina) Xetadrilus ugandensis (Bell, 1954) (ex Stercutus) Genus diagnosis. Small worms, adults in vivo 2���5 mm long and 0.1��� 0.2 mm wide. Two chaetae per bundle, absent laterally from VIII on (absent from VII in X. pitucus), chaetae with proximal bend, distally mostly straight. Lateral chaetae not shifted dorsally. Epidermal gland cells pale, elliptical, not transversely elongate. Head pore on prostomium dorsally. Prostomial epithelium with frontal recess; prostomial papillae present. Brain incised posteriorly, longer than wide, with ganglia on prostomial nerves. Ventral nerve cord with suboesophageal ganglion in II��� IV and segmental ganglia from V on. Pharyngeal glands variable, secondary ventral lobes present or absent. Oesophageal appendages absent. Intestinal diverticula absent, present at VII in X. fabryi. Gut often widened abruptly at 6 / 7, with thickened epithelium in VII. Dorsal blood vessel origin in clitellar or postclitellar region, anterior bifurcation in segment I. Nephridial anteseptale with parts of nephridial body, no constriction at septum, efferent duct short. Coelomocytes one type, mucocytes. Male reproductive system simple, vas deferens not widened, male pores on body surface. Spermathecae not attached to oesophagus, with ectal duct and ampulla; ectal pore in lateral line and at half-length between lateral chaetae of IV and V; no diverticula, no ectal glands. Etymology. Named in honour of the Xet�� people, the last ethnia in the Brazilian state of Paran�� to come into contact with 'modern' civilization in the last century. Within a few years the Xet�� were almost eradicated by forest destruction for coffee plantations. Survivors and their descendants merged into the Brazilian population. Today five persons still speak Xet��, a language that belongs to the Tupi-Guaran�� language family. Common traits: (1) Not more than 2 chaetae per bundle (2) Head pore on prostomium (3) Ventral nerve cord ganglionic from V, ganglia II-IV fused into suboesophageal gan- glion (4) Nephridial anteseptale including parts of nephridial body (5) Spermathecae (when present) free, not attached to oesophagus (6) Spermathecae (when present) without diverticula (7) Brain incised posteriorly (truncate in T. marcusae) (8) Male copulatory organ without bursa, male pore on body surface ** * absent from XIII in T. lacteus Righi, 1974 b. ** bursa in T. marcusae. Remarks. Xetadrilus is closely related to Guaranidrilus and Tupidrilus Righi, 1974 b. Similarities and differences are listed in Table 1, see also below, discussion, and Figure 8. Within this triad of genera, Xetadrilus stands out by (1) lateral chaetae absent from VIII, (2) inner prostomial papillae, (3) conspicuous prostomial ganglia, (4) a frontal recess in the prostomial wall. Trait (1) is the presumed apomorphy of the genus sensu stricto. Traits (2,3,4) are probably plesiomorphies because they are also present in Achaeta (comp. Schmelz et al. 2008). Trait (4) occurs also in some species of the marine Grania (Rota et al. 2003). The peculiar similarity of one species of Enchytronia will be dealt with below, see discussion. All species of Xetadrilus lack oesophageal appendages, and intestinal diverticula are absent in all but one species. Tupidrilus differs from Xetadrilus in the presence of oesophageal appendages and in the presence of lateral chaetae from VIII on (they are absent from XIII in T. lacteus); traits (2),(3),(4) are unkown. Tupidrilus differs from Guaranidrilus only in the absence of intestinal diverticula, and its status was considered doubtful after a phylogenetic analysis (Coates 1989), but the taxonomy of Tupidrilus is beyond the scope of this paper. The genus includes four species, all from Brazil. Due to its similarity with Guaranidrilus and Achaeta, Xetadrilus may be accomodated in the Achaetinae Černosvitov, 1937. Achaetinae is recognized by a combination of characters (Table 1), there are currently no diagnostic traits, see below, discussion. Tupidrilus and Hemienchytraeus are also members of the subfamily. Two species of Marionina and one species of Stercutus are included in the new genus; criteria for their inclusions were: (1) compliance with all traits common to Guaranidrilus, Tupidrilus, and Xetadrilus (Table 1), (2) absence of oesophageal appendages, (3) absence of lateral chaetae from VIII on (M. pituca from VII), (4) prostomium with ganglia, inner papillae, and frontal epithelial recess. Traits related to the prostomium are unknown in X. righianus and X. ugandensis; types of the latter are lost, so fresh material will be necessary to confirm the new combination. Types of X. righianus were not investigated during the time of this study. In X. pitucus, large prostomial ganglia are originally described (Righi, 1974 a), although not recognized as such; and inner prostomial papillae and a frontal epithelial recess are present in the types (Schmelz & Collado, unpublished). The three species included in Xetadrilus do not comply in all details with the generic diagnosis (see Table 2), but they disagree considerably with the type species of their original genus (Stercutus niveus Michaelsen, 1888 and Marionina georgiana Michaelsen, 1888, respectively). S. ugandensis has been a species incertae sedis since Nielsen & Christensen (1959), and Marionina is a heterogeneous assemblage of species in need of revision (Rota et al. 2008, Schmelz & Collado 2008). The transfer of M. pituca and M. righiana to Xetadrilus is in agreement with the suggestion in Rota et al. (2008) to accomodate some species of Marionina in Achaetinae. Table 2 gives an overview of key traits of the included nominal species. Redescriptions of type material will be presented elsewhere. Three further South American species of Marionina, M. patua Righi, Ayres & Bittencourt, 1978, M. ecuadoriensis Righi, 1981 a, and M. tica Righi, 1981 b, are very close to Xetadrilus based on the following traits: free spermathecae, head pore on prostomium, two chaetae per bundle, no oesophageal appendages, no intestinal diverticula. They differ from Xetadrilus in the presence of lateral chaetae from VIII on. These species probably belong to Achaetinae, too, but their generic placement within the subfamily is not straightforward. Type-based redescriptions of these species, together with suggestions as to their generic position, will be presented elsewhere. With the African X. ugandensis and the Chinese X. righianus, Xetadrilus is a cosmopolitan genus. Its circumscription may change in the future when more species have been discovered and more species of Guaranidrilus, Tupidrilus, and Marionina have been reinvestigated., Published as part of Schmelz, R��diger M., Collado, Rut & R��mbke, J��rg, 2011, Mata Atl��ntica enchytraeids (Paran��, Brazil): A new genus, Xetadrilus gen. nov., with three new species, and four new species of Guaranidrilus ��ernosvitov (Enchytraeidae, Oligochaeta), pp. 1-29 in Zootaxa 2838 on pages 13-15, DOI: 10.5281/zenodo.203260, {"references":["Righi, G. (1974 a) Marionina pituca sp. n. (Oligochaeta, Enchytraeidae) from Brazilian Soil and its intestinal parasite Buetschliella marioninae sp. n. (Ciliata, Astomata). Zoologischer Anzeiger, 5 / 6, 414 - 419.","Xie, Z. & Rota, E. (2001) Four new terrestrial species of Marionina (Clitellata, Enchytraeidae) from China and re-examination of M. hoffbaueri Moller. Journal of Natural History, 35, 1417 - 1431.","Bell, A. W. (1954) Some enchytraeid worms (Oligochaeta) from East Africa. Transactions of the American Microscopical Society, 73, 279 - 311.","Righi, G. (1974 b) Notas sobre os Oligochaeta, Enchytraeidae do Brasil. Papeis Avulsos de Zoologia. Museu de Zoologia da Universidade de Sao Paulo, 28, 127 - 145.","Rota, E. & Erseus, C. (2003) New records of Grania (Clitellata, Enchytraeidae) in the Northeast Atlantic (from Tromso to the Canary Islands), with descriptions of seven new species. Sarsia, 88, 210 - 243.","Coates, K. A. (1989) Phylogeny and origins of Enchytraeidae. Hydrobiologia, 180, 17 - 33.","Michaelsen, W. (1888) Beitrage zur Kenntnis der deutschen Enchytraiden-fauna. Archiv fur Mikroskopische Anatomie, 31, 483 - 498.","Nielsen, C. O. & Christensen, B. (1959) The Enchytraeidae. Critical revision and taxonomy of European species (Studies on Enchytraeidae VII). Natura Jutlandica, 8 - 9, 1 - 160.","Rota, E., Matamoros, L. & Erseus, C. (2008) In search of Marionina (Clitellata, Enchytraeidae): A taxonomic history of the genus and re-description of the type species Pachydrilus georgianus Michaelsen, 1888. Italian Journal of Zoology, 75, 1 - 20 (online access).","Righi, G., Ayres, I. & Bittencourt, E. C. R. (1978) Oligochaeta (Annelida) do Instituto Nacional de Pesquisas da Amazonia. Acta Amazonica, 8, 1 - 49.","Righi, G. (1981 a) Alguns Oligochaeta cavernicolas do Equador. Papeis Avulsos de Zoologia. Museu de Zoologia da Universidade de Sao Paulo, 34, 235 - 249.","Righi, G. (1981 b) Notas sobre Enchytraeidae (Oligochaeta) Brasileiras. Revista Brasileira de Biologia. Rio de Janeiro, 41, 427 - 430."]}

Guaranidrilus cingulatus sp. nov. (Figs 3, 8 C) Holotype. MZUSP 1225, adult specimen, stained whole mount, Antonina, Cachoeira, medium old secondary forest on Cambisol [site 17], Mar 2004, leg. B. F��rster, R. M. Schmelz. Paratypes. MZSUP, 14 specimens, stained whole mounts: MZUSP 1226, 6 specimens, Antonina, Cachoeira, 25 �� 19 ' 59 "S ��� 48 �� 42 '02"W, young secondary forest on Gleysol [site 25], May 2003, leg. J. R��mbke, R. M. Schmelz. MZUSP 1227, 7 specimens, same data as holotype. MZUSP 1228, 3 specimens, Antonina, Cachoeira, 25 �� 19 '04"��� 41 ''S, 48 �� 40 ' 36 "��� 41 ' 51 "W, 20���120 m a.s.l., young and medium old secondary forests on Gleysol [sites 12, 24, 30], Oct 2004, leg. B. F��rster, R. M. Schmelz. MZUSP 1229, 1 specimen, Curitiba outskirts, 25 �� 19 ' 13 ''S, 49 ��06' 22 ''W, 957 m a.s.l., Araucaria forest on Podzol, property of R. Marques, Oct 2004, leg. R. Marques, R. M. Schmelz. UFPR, 6 specimens, 3 stained whole mounts, 3 ethanol-preserved, Guareque��aba, Itaqui: UFPR OL-10, 2 specimens, 25 �� 14 ' 45.3 ''S, 48 �� 30 ' 19.4 ''W, 27 m a.s.l., medium old secondary forest on Cambisol [site 38], Sep 2007, leg. P. Heine, R. M. Schmelz. UFPR OL-11, 1 specimen, 25 �� 18 ' 31.8 ''S, 48 �� 27 '54,4''W, 93 m a.s.l., old-growth forest on Cambisol [site 40], Sep 2007. leg. P. Heine, R. M. Schmelz. UFPR OL- 12: 3 specimens, ethanol-preserved, 25 �� 14 '51,1''S, 48 �� 29 ' 31.4 ''W, 8 m a.s.l., medium old secondary forest on Cambisol [site 39], Jan 2008, leg. J. R��mbke, R. M. Schmelz. Additional material. Eight specimens from Cachoeira and Itaqui sites, examined in vivo, not preserved. Etymology. 'cingulatus' (lat.) = girded; named in reference to the girdle-shaped clitellum, so far unique in the genus. Description. Body dimensions. Small species, living adults ca. 3.5���5 mm long and 0.1���0.15 mm wide, whole mounts ca. 2.5���4 mm long and 0.15���0.2 mm wide. Segment number 26���32. Chaetae (Fig. 3 B) two per bundle throughout, absent at XII, straight distally. In caudal segments chaetae 1.5��� 2 x as large as largest anterior chaetae, equally large ventrally and laterally (ca. 60 ��m long and 4���5 ��m wide); in anterior segments laterals smaller than ventrals (30 ��m vs. 40 ��m). Epidermal gland cells invisible in vivo; in well-preserved whole mounts two rows distinguishable of inconspicuous, transversely elongate cells in dorsal half of anterior segments. Clitellum (Fig. 3 E) well-developed, perfectly girdle-shaped, cells in ca. 34 transverse rows ventrally, dorsally rows indefinite when fully developed, here hyalocytes and granulocytes alternating, granulocytes smaller than hyalocytes and more numerous; ventrally only granulocytes; granulocytes almost square-shaped, giving a 'pavement'-like aspect. Head pore on prostomium mid-dorsally, prostomial inner papillae inconspicuous or absent. Body wall thick in vivo, 5���10 ��m thick in whole mounts, longitudinal muscle layer well-developed, especially in adults. Cuticle distinguishable, ca. 1.5 ��m thick. Septa slightly thickened in preclitellar segments, from 4 / 5 to 5 / 6 or 8 / 9. Brain concave posteriorly, twice as long as wide (length ca. 70 ��m, fix), prostomial ganglia very small or absent. Ventral nerve cord with suboesophageal ganglion in II���IV and segmental ganglia from V on. Oesophageal appendages (Fig. 3 A,C) in anterior of VI, paired, spherical in dorsal view, sessile, side by side dorsally on gut wall, not larger than secondary ventral pharyngeal gland lobes. Pharyngeal glands (Fig. 3 A,C) dorsally unpaired in IV, V, VI; primary and secondary ventral lobes in V and VI. Primary ventral lobes of VI largest. Intestinal diverticula (Fig. 3 A,C) paired, lateral to intestine, confined to VII, occupying entire segment length, rarely less (Fig. 3 A), separate from each other; each with large anterior and small stump-like posterior projection, the latter situated dorso-laterally, extending slightly into VIII in few specimens (Fig. 3 C); connections of diverticula with intestine in posterior of VII; outer and inner surface often irregularly lobed or wavy; walls ca. 7���10 ��m thick, no radial striation. Dorsal blood vessel from X ��� 1 / 2 XII. Preclitellar nephrida (Fig. 3 F) 3 pairs, at 6 / 7 ��� 8 / 9, occasionally absent at either 6 / 7 or 8 / 9. Anteseptale slightly longer than wide, with ca. 1 / 3 of entire nephridial body, postseptale slightly bulging dorsad, with dorsal vesicle, gradually merging into short efferent duct, terminal vesicle absent; ante- and postseptale separated by a slight constriction at level of septa. Postclitellar nephridia (Fig. 3 G) present in less than half of all positions; shape as in preclitellar nephridia, but more elongate. Coelomocytes slightly longer than wide, 17���28 ��m long, with very fine wavy or groovy texture, no granula distinguished; cell margins smooth, refractile in oblique position; cells abundant but not obscuring other inner structures. Seminal vesicle absent, free cysts in XI sparse. Spermatozoa ca. 47 ��m long, heads 18 ��m long. Sperm funnel (Fig. 3 E,H) small, length less than 1 / 4 body diameter, 1.2���1.5 x as long as wide, barrel-shaped, flattened. Vas deferens short with few loops, diameter 8 ��m proximally, 5 ��m distally. Male copulatory organ (Fig. 3 H,I) minute, as if absent, male glands small, not thicker than clitellum. Male pores on body surface, surrounded by male glands, no bursa. Transverse copulatory body muscles sparse, not seen in vivo. Accessory glands absent. Spermatheca (Fig. 3 C,D) simple, extending into V or VI, about as long as body diameter or slightly longer. Ectal duct not longer than 15 ��m, with cuticle-lined canal only in distal half; distal part of ampulla ca. 15 ��m wide, often without sperm; sperm present more proximally. Habitat. G. cingulatus was found in young to medium secondary forests and old-growth forest. It was absent in pastures and at agroforestry sites, and almost absent at recently abandoned pastures (1 specimen identified). Remarks. G. cingulatus is conspicuous (1) by the girdle-shaped clitellum, well-developed on all sides and quite prominent in living and preserved specimens. All other reproductive structures are small and inconspicuous. Further species-specific traits, diagnostic in combination: (2) epidermal gland cells appear absent; (3) cuticle conspicuous, longitudinal muscle layer well-developed, body wall often thick; (4) terminal (= caudal) chaetae up to twice as large as anterior chaetae; (5) oesophageal appendages in anterior of VI; (6) intestinal diverticula in VII, with posterior extension and often irregularly lobed; (7) three pairs of preclitellar nephridia, from 6 / 7 to 8 / 9. Traits 2���7 suffice to identify the species; they were used in the routine identification of animals, juveniles included. Most similar to G. cingulatus is the North-American G. oregonensis Coates & Diaz, 1988, described from fixed material. The species is similar in body length (3���4.5 mm), segment number (26���34), chaetae (enlarged in caudal segments), epidermal gland cells (few rows), origin of dorsal blood vessel (X���XII), position of preclitellar nephridia (3 pairs, 6 / 7���8 / 9), and sperm-related structures (small). G. oregonensis may also have a girdle-shaped clitellum, but the trait is described ambiguously and requires reinvestigation: "Clitellum... in transverse rows ventrolaterally; reduced but complete ventrally between male pores" (ibd.: 775). The species differs in the absence of oesophageal appendages (present in G. cingulatus) and in the extension over 2 segments of the intestinal diverticula (1 segment in G. cingulatus). Specimens had been sampled from freshwater marsh soils in the estuary of the Columbia river (Oregon, U.S.A.), see Coates & Diaz (1988) for details. Further similar species are G. rarus Černosvitov, 1937 a emend. Coates & Diaz (1988), and G. s a w a y a i Righi, 1973. Both species, which may be synonymous (Coates & Diaz 1988), differ from G. cingulatus in having a saddleshaped clitellum. Other differences are inconclusive and require a reinvestigation of the types., Published as part of Schmelz, R��diger M., Collado, Rut & R��mbke, J��rg, 2011, Mata Atl��ntica enchytraeids (Paran��, Brazil): A new genus, Xetadrilus gen. nov., with three new species, and four new species of Guaranidrilus ��ernosvitov (Enchytraeidae, Oligochaeta), pp. 1-29 in Zootaxa 2838 on pages 8-10, DOI: 10.5281/zenodo.203260, {"references":["Coates, K. A. & Diaz, R. J. (1988) Description of Guaranidrilus oregonensis (Oligochaeta: Enchytraeidae) from North America, with additional comments on the genus. Proceedings of the Biological Society of Washington, 101, 773 - 783.","Cernosvitov, L. (1937 a) Notes sur les Oligochaeta (Naididees et Enchytraeidees) de l'Argentine. Anales del Museo Argentino de Ciencias Naturales, Buenos Aires, 39, 135 - 157.","Righi, G. (1973) Sobre tres especies Brasileiras de Enchytraeidae (Oligochaeta). Boletim de Zoologia e Biologia Marina, Sao Paulo, 30, 469 - 482."]}

Xetadrilus maacki sp. nov. (Figs 5, 8 E, Table 3) Holotype. MZUSP 1206, adult specimen, stained and whole-mounted in Canada balsam, Paranagu��, Palmito, 25 �� 35 ' 41.4 ''S 48 �� 31 ' 13.7 ''W, 3���11 m a.s.l., young secondary forest on Podzol [site 43], Mar 2004, leg. B. F��rster, R. M. Schmelz. Paratypes. MZUSP, 18 specimens, stained whole mounts: MZUSP 1207, 6 specimens, same data as holotype. MZUSP 1208, 12 specimens, Antonina, Cachoeira, 25 �� 14 ��� 38 ������ 19 ' 21 "S, 48 �� 40 ��� 10 ������ 42 '08"W, 10���140 m a.s.l., medium old and old-growth forest on Cambisol and Gleysol [sites 10, 13, 28], May 2003, leg. J. R��mbke & R. M. Schmelz. UFPR, 8 specimens, stained whole mounts: UFPR OL-18, 6 specimens, Guareque��aba, Itaqui, 25 �� 14 ' 45.3 ''��� 15 '32,7'S, 48 �� 29 ' 31.4 ''��� 30 ' 31.9 ''W, 8 and 27 m a.s.l., medium old and old-growth forest on Cambisol [sites 38, 39, 41], Sep 2007, leg. P. Heine, R. M. Schmelz. UFPR OL-19, 2 specimens, Antonina, Rio Pequeno, 25 �� 14 ' 39.3 ''S, 48 �� 44 ' 48.2 ''W, 40 m a.s.l., agroforestry system on Gleysol, banana monoculture [site 46], Jan. 2008, leg. J. R��mbke, R. M. Schmelz. Additional material. Three specimens from type locality, examined in vivo, not preserved. Etymology. Named in honour of Reinhard Maack (1892���1969), German-Brazilian geographer and pioneer of environmental protection in Southern Brazil; seminal contributions to the physical geography of the state of Paran�� (e.g., Maack 1968). Maack discovered the Xet�� people in 1962. Vita in Fabry (2006). Description. Individuals somewhat chequered, transparent and milky/opaque body regions; coelom more or less free, inner structures well-distinguished. Body dimensions. Living adult specimens ca. 4 mm long and 0.1��� 0.12 mm wide, widest in clitellar region; fixed specimens 3���3.5 mm long and 0.12���0.18 mm wide. Segment number 21���28 in adults and subadults, most often 25���27 (N = 16), holotype 27. Subadults with Chaetae (Fig. 5 E,H) two per bundle, formula 2,0���0: 2 ��� 2. Lateral chaetae present in II���VII, absent from VIII on. Ventral chaetae from II on, absent in XII. Anterior chaetae faintly sigmoid, with slight proximal bend and almost straight distal tip, ventrally ca. 32���38 ��m long and 3 ��m wide, laterally smaller, dimensions not measured. Behind clitellum chaetae increasing in size towards posterior end, twice as large as anterior chaetae in ca. 8 hindmost segments, here 64���70 ��m long and 6���6.5 ��m wide. Chaetae slightly shifted posteriad with respect to septa, ratio of distance to anterior and posterior septum, respectively, about 4: 3. Epidermal gland cells (Fig. 5 A,G) absent except mid-ventrally between chaetal bundles; here 2���3 large, transversely arranged gland cells with finely granular content (fix), present in all segments, XII included; cells 2 x as high as epidermis, larger posteriorly than anteriorly. Clitellum (Fig. 5 C) saddle-shaped, i.e. absent ventrally. Cells in ca. 31 separate (viv) or dense (fix) rows; hyalocytes not in longitudinal rows, present only in dorsal half, extending further ventrally in the middle than at the ends; cells roughly rectangular in living specimens, with hyalocytes larger than granulocytes (diameters ca. 11 ��m vs. 8 ��m (viv) or 10���12 ��m vs. 6 ��m (fix)); ventro-laterally only granulocytes. Cells about as high as wide. A few cells present between male copulatory organs, of same type as in other segments between ventral chaetal bundles. Prostomium (Fig. 5 A) with head pore in mid-dorsal position, slit transverse. Frontal prostomial epithelium (Fig. 5 A,B) thickened, with a recess or cleft at the frontal tip. Dorso-laterally six inner prostomial papillae in bilateral-symmetrical order. Prostomial musculature well-developed. Body wall thin, between 3 and 12 ��m thick, cuticle Septa slightly thickened in preclitellar segments (fix), difference inconspicuous in vivo. Brain incised posteriorly, sides converging anteriad. A pair of bilobed prostomial ganglia present on prostomial nerves. Perikarya of II���IV fused into suboesophageal ganglion, segmental ganglia present from V on, no perikarya at the level of the septa except in hindmost segments; here segmental ganglia interconnected by a thin strand of perikarya at level of septa. Pharyngeal glands (Fig. 5 A,B) with three unpaired dorsal lobes in IV���VI, thickest in the middle, all alike; two pairs of primary ventral lobes, in V and VI; three pairs of secondary ventral lobes, in V, VI, VII, increasing in size posteriad. Oesophageal appendages and intestinal diverticula absent. Gut widening abruptly in VII (Fig. 5 A), gradually tapering posteriorly. Epithelium thickened in VII and VIII, cells enlarged, with fine pale vesicular content. Chloragocytes from V, a dense and flat layer (cell height ca. 5 ��m) from VII, inconspicuous. Dorsal vessel from XIII or XIV; anterior bifurcation prostomial. Pars tumida of midgut over 1���2 segments, from 1 / 2 XVII ��� 1 / 2 X, epithelium strongly proliferated. Preclitellar nephridia (Fig. 5 G) two pairs in preclitellar segments, at 7 / 8 and 8 / 9, not constricted at septum. Anteseptale with parts of nephridial body, longer than wide, funnel attached obliquely. Postseptale larger than anteseptale, length ratio 3: 2, with a dorsal bump in mid-section, no vesicle here; nephridial canal apparently with an up- and down zig-zag course; efferent duct rising subterminally, nephroporus inconspicuous, situated anteriorly of ventral chaetal bundles, no terminal vesicle. Nephridia in postclitellar segments in varying positions, altogether 8���10 in number. Example, holotype: 13 / 14 (paired), 18 / 19 (paired), 20 / 21 (unpaired left side), 24 / 25 (unpaired right side), 25 / 26 (paired). Nephridia elongate, no dorsal bump in mid-section of postseptale, no vesicle, efferent duct short, rising terminally. Coelomocytes (Fig. 5 F) distinctly brownish in vivo, with small spherical and refractile vesicles; vesicles clearly separate from each other, like dots, conspicuous. Cells slightly longer than wide, almost spherical, ca. 25 ��m long. In fixed and dehydrated material cytoplasma pale and homogeneously vesicular, nucleus central, conspicuous. Seminal vesicle absent. Spermatozoa conspicuous on top of sperm funnel when present, ca. 39 ��m long, heads ca. 14 ��m long. Sperm funnel (Fig. 5 H) small, 1 / 4 ��� 1 / 3 as long as body diameter, 1.5���2 as long as wide; collar distinct, not or only little widened and ca. 15 ��m high, contributing to almost 1 / 3 of total sperm funnel length; funnel body variable, either barrel-shaped, not wider than collar, or almost globular, canal opening funnel-like towards proximal end. Vas deferens in dense coils ventro-laterally, ca. 4 ��m thick, wider immediately behind sperm funnel (ca. 6 ��m). Male copulatory organ (Fig. 5 H): male pores on body surface in longitudinal line of ventral chaetae, at half distance between chaetae of XI and XIII; each pore surrounded by a small glandular bulb interspersed with musculature, bulb small, roughly spherical, diameter ca. 20 ��m, outline irregular, occasionally with small extra-bulbs anteriorly and posteriorly; bursa apparently absent, transverse copulatory body muscles sparse, three strands in front of male pore, posteriorly inconspicuous or absent. Accessory glands absent. Spermatheca (Fig. 5 D) a small and proximally blind-ending tube, not attached to oesophagus, confined to V; ectal pore in lateral line, at half-length between chaetae of IV and V; ectal duct and ampulla of about equal length; ectal duct with smooth outline, isodiametric, ampulla not wider than ectal duct but with irregular outline, thin-walled. Small amount of sperm in ampullar distal part. Eggs. One or two mature eggs at a time. Habitat. X. maacki was found at all successional stages of forest regeneration and also in pastures and at agroforestry sites. Remarks. Key characters of Xetadrilus maacki are: (1) Brown coelomocytes; (2) gut widening abruptly in VII, no intestinal diverticula; (3) chaetae in terminal segments large, thick, sigmoid; (4) segmental mid-ventral epidermal gland cells. For a comparison with other congeneric species, see Tables 2 and 3., Published as part of Schmelz, R��diger M., Collado, Rut & R��mbke, J��rg, 2011, Mata Atl��ntica enchytraeids (Paran��, Brazil): A new genus, Xetadrilus gen. nov., with three new species, and four new species of Guaranidrilus ��ernosvitov (Enchytraeidae, Oligochaeta), pp. 1-29 in Zootaxa 2838 on pages 16-18, DOI: 10.5281/zenodo.203260, {"references":["Maack, R. (1968) Geografia fisica do Estado do Parana. Universidade Federal do Parana, Curitiba, 350 pp.","Fabry, R. (2006) Reinhard Maack. Pionier des Umweltschutzes in Sudbrasilien (Parana). Zeitschrift fur Umweltchemie und Okotoxikologie, 18, 37 - 40."]}

Xetadrilus fabryi sp. nov. (Figs 7, 8 G, Table 3) Holotype. UFPR OL- 23, adult specimen with 22 segments, posterior end regenerated, stained whole mount, Antonina, Rio Pequeno, 25 �� 15 ' 56.8 ''S, 48 �� 44 ' 23.5 ''W, 25 m a.s.l., agroforestry system [site 48], Jan 2008, leg. J. R��mbke, R. M. Schmelz. Paratypes. MZUSP, 4 specimens, stained whole mounts, Antonina, Cachoeira: MZUSP 1215, 3 specimens, 25 �� 15 ' 2 ''S, 48 �� 40 ' 24 '' and 25 �� 15 ' 13 ''S, 48 �� 40 ' 34 ''W, 30 and 40 m a.s.l., respectively, pasture, partly abandoned, on Cambisol [sites 1, 4], May 2003, leg. J R��mbke, R. M. Schmelz. MZUSP 1216, 1 specimen, 25 �� 18 ' 25 "S, 48 �� 40 ' 24 "W, 70 m a.s.l., pasture on Cambisol [site 2], March 2004, leg. B. F��rster, R. M. Schmelz. UFPR, 8 specimens, stained whole mounts: UFPR OL-24, 5 specimens, Antonina, Rio Pequeno, 25 �� 15 ' 56.8 ''��� 16 ' 23.4 ''S, 48 �� 43 ' 41.1 ''��� 44 ' 23.5 ''W, 15 and 25 m a.s.l., respectively, agroforestry system [sites 47, 48], Jan 2008, leg. J. R��mbke, R. M. Schmelz. UFPR OL-25, 3 specimens, Curitiba, UFPR University Campus, Agr��rias, 25 �� 24 ' 37 ''S, 49 �� 14 ' 56 ''W, 909 m a.s.l., grassland, Feb 2008, leg. R. M. Schmelz. Additional material. One specimen from UFPR campus, Agrarias, examined in vivo, not preserved. Etymology. Named in honour of Rainer Fabry, versatile German on-site coordinator of the SOLOBIOMAproject. Xetadrilus maacki Xetadrilus aphanus Xetadrilus fabryi Segment number 21���28 21���28 33 Description. Body dimensions. Living adult specimens ca. 3 mm long and 0.11���0.12 mm wide, whole mounts 2.5���3.5 mm long and 0.1���0.12 mm wide, up to 0.16 at XII. Segment number 30���33 (N= 3), holotype 22 segments, with regenerated posterior end. Chaetae two per bundle, formula 2,0���0: 2 ��� 2. Lateral chaetae present in II���VII, absent from VIII on. Ventral chaetae from II on, absent in XII. Chaetae distally straight, or slightly bent to same side as proximal bend. Terminal chaetae enlarged, almost 2 x as large as largest anterior chaetae, not sigmoid. Ventral anterior chaetae ca. 2 ��m thick, in II 12 ��m long, gradually increasing in size posteriad to about 28 ��m in IX. Ventral chaetae in hindmost segments 45���55 ��m long and 2.5���3 ��m thick. Epidermal gland cells (Fig. 7 B) variable; entire epidermis glandular to varying degrees, gland cells pale and evenly distributed, or no epidermal gland cells distinguishable in living and preserved material. In specimens with glandular epidermis body surface often covered with foreign particles. Clitellum saddle-shaped, cells in ca. 25 transverse rows, separate (viv) or dense (fix). Hyalocytes on dorsal half, larger than granulocytes, isolated; in ventral half only granulocytes, mid-ventral interruption as wide as distance of male pores. Prostomium with head pore in mid-dorsal position. Frontal prostomial epithelium (Fig. 7 A,E) thickened, with a vesicle-like recess or cleft at the frontal tip. Dorso-laterally several inner papillae in bilateral-symmetrical order, inner papillae also on peristomium. Prostomial musculature present. Body wall medium-thick, cuticle thin. Preclitellar septa not thickened. Brain (Fig. 7 E) in I���III, 2���2.5 x as long as wide, incised posteriorly; prostomial ganglia conspicuous but small; ventral nerve cord with suboesophageal ganglion in II���IV and segmental ganglia from V. Oesophageal appendages absent. Pharyngeal glands (Fig. 7 A,B) as unpaired dorsal lobes in IV���VI, voluminous in IV and V, primary ventral lobes in V and VI, largest in VI; small and spherical secondary ventral lobes in V, VI and VII. Intestinal diverticula (Fig. 7 A,B) one pair in VII, elongate, oval, thick-walled, with distinct lumen; placed laterally of intestine, arranged in "V"- shape (dorsal view), slightly diverging anteriad; diverticula laterally flattened, higher than wide. Chloragocytes often black-grey in posterior segments. Dorsal blood vessel from 1 / 2 XIV in the two mature and complete specimens, from 1 / 4 XIV in holotype; the origin was difficult to see in living specimens. Preclitellar nephridia (Fig. 7 C) 3 pairs, from 7 / 8 to 9 / 10, postseptale without dorsal vesicle. Postclitellar nephridia elongate. Coelomocytes pale, not hyaline, without conspicuous texture. Seminal vesicle absent. Spermatozoa distinct on top of sperm funnel, length not measured in vivo. Sperm funnel (Fig. 7 F) barrel-shaped, ca. 1 / 3 as long as body diameter (40���50 ��m long, fix), collar not wider than funnel body. Vas deferens (Fig. 7 F) quite long, coiled densely in numerous loops in XII, sometimes filling entire dorsal half of segment XII; diameter 4 ��m throughout except near sperm funnel, here ca. 5 ��m (fix). Male copulatory organ (Fig. 7 F) small, circular in top view, not more than a body wall thickening in side view; bursa and bursal slit apparently absent; male pore on body surface, sometimes surrounded by folds. Glandular bulb tripartite, an eversible central bulb, diameter ca. 20 ��m (fix), pierced centrally by vas deferens, and two smaller bulbs attached anteriorly and posteriorly. Copulatory muscles well-developed, encompassing bulbs and also extending dorsad as transverse copulatory muscles. Accessory glands absent. Spermathecae (Fig. 7 A) not attached to oesophagus, a simple tube. Ectal duct short (length 20 ��m, diameter 10 ��m, fix) with wide lumen, often filled with sperm, gradually widening into distal part of ampulla, here sperm arranged in parallel, or no ampulla distinguishable. The following tube narrow (diameter ca. 5 ��m, fix), widening into thin-walled ental reservoir in VI or VII; reservoir not always developed. Sperm may be present in any part of the spermatheca. One mature egg at a time. Habitat. X. fabryi was found in grazed and abandoned pastures and at agroforestry sites. It was not found at forest sites. Remarks. This is the only species of the new genus with intestinal diverticula. Diverticula are similar in shape to the ones in Guaranidrilus joanae Christoffersen, 1977 and G. hoeferi sp. nov. (see above). The two Guaranidrilus species are much larger and with very large sperm-related sexual organs. X. fabryi has been assigned to Xetadrilus (and not to Guaranidrilus) by virtue of the following characters: (1) Lateral chaeta absent from VIII, (2) epidermal gland cells oval, not transversely elongate, (3) prostomium with inner papillae and ganglia, (4) nephridia without dorsal vesicle, (5) pharyngeal glands with secondary lobes not only in V, VI, but also in VII, (6) post-clitellar origin of dorsal blood vessel., Published as part of Schmelz, R��diger M., Collado, Rut & R��mbke, J��rg, 2011, Mata Atl��ntica enchytraeids (Paran��, Brazil): A new genus, Xetadrilus gen. nov., with three new species, and four new species of Guaranidrilus ��ernosvitov (Enchytraeidae, Oligochaeta), pp. 1-29 in Zootaxa 2838 on pages 20-23, DOI: 10.5281/zenodo.203260, {"references":["Christoffersen, M. L. (1977) New species of Guaranidrilus Cernosv. (Enchytraeidae, Oligochaeta) from Serra do Mar, Sao Paulo, Brazil. Studies on Neotropical Fauna and Environment, 12, 187 - 206."]}

Xetadrilus aphanus sp. nov. (Figs 6, 8 F, Table 3) Holotype. MZUSP 1209, adult specimen, stained and whole-mounted in Canada-Balsam, Cachoeira, Antonina, 25 �� 18 ' 25 "S, 48 �� 40 ' 24 "W, 70 m a.s.l., pasture on Cambisol [site 2], Mar 2004, leg. B. F��rster, R. M. Schmelz. Paratypes. MZUSP, 32 specimens, stained whole mounts: MZUSP 1210, 2 specimens, Antonina, Cachoeira, 25 �� 15 ' 22 ''S, 48 �� 40 ' 14 ''W, 40 m a.s.l., young secondary forest on Cambisol [site 7], May 2003, leg. J. R��mbke, R. M. Schmelz. MZUSP 1211, 11 specimens, same data as holotype. MZUSP 1212, 10 specimens, Antonina, Cachoeira, 25 �� 18 ' 56 "S, 48 �� 39 ' 29 "W, 20 m a.s.l., pasture on Gleysol [site 20], Mar 2004, leg. B. F��rster, R. M. Schmelz. MZUSP 1213, 7 specimens, Antonina, Cachoeira, 25 �� 18 ' 32 "S, 48 �� 39 ' 38 "W, 50 m a.s.l., pasture on Cambisol [site 3], Oct 2004, leg. B. F��rster, R. M. Schmelz. MZUSP 1214, 2 specimens, Curitiba, UFPR University Campus, Agrarias, 25 �� 24 ' 37 ''S, 49 �� 14 ' 56 ''W, 909 m a.s.l., grassland, Mar 2004, leg. R. M. Schmelz. UFPR, 30 specimens, stained whole mounts, Guareque��aba, Itaqui: UFPR OL-20, 8 specimens, 25 �� 14 ' 16.44 ''S, 48 �� 29 ' 52.1 ''W and 25 �� 18 ' 49.5 ''S, 48 �� 27 ' 10.4 ''W, 8 and 13 m a.s.l., respectively, abandoned ("herbaceous") pasture on Cambisol [sites 31, 33], Oct 2007, leg. P. Heine, R. M. Schmelz. UFPR OL-21, 6 specimens, 25 �� 18 ' 31.8 ''S, 48 �� 27 ' 2.3 ''W, 28 m a.s.l., medium old secondary forest on Cambisol [site 37], Sep 2007, leg. P. Heine, R. M. Schmelz. UFPR OL-22, 16 specimens, appr. 25 �� 13 ' 30 ''S, 48 �� 27 ' 40 ''W, 4 m a.s.l., pasture near SPVS guesthouse in Taga��aba, Jan 2008, leg. J. R��mbke, R. M. Schmelz. Additional material. Ca. 15 specimens, several sites in Cachoeira and Itaqui, examined in vivo, not preserved. Etymology. 'Aphanus' is a latinized form of the Greek 'aphanes', meaning inconspicuous. The name refers to the small body size of the worms. Description. Specimens inconspicuous, among the smallest enchytraeids in all samples. Body dimensions. living adult specimens ca. 2���3 mm long and 0.07���0.1 mm wide; fixed adult specimens 2���2.2 mm long and ca. 0.1 mm wide, 0.08���0.12 mm at V, 0.1���0.14 mm at XII. Segment number of adult individuals (19)- 23-28 (N = 35). Chaetae 2 per bundle, formula 2,0���0: 2 ��� 2. Lateral chaetae present in II���VII, absent from VIII on. Ventral chaetae from II on, absent in XII. Chaetae distally straight, or slightly bent to same side as proximal bend. Terminal chaetae enlarged, twice as large as anterior chaetae, not sigmoid. Ventral anterior chaetae ca. 2 ��m thick, in II 10���16 ��m long, gradually increasing in size posteriad to about 20���26 ��m. Ventral chaetae in hindmost segments 34���40 ��m long and 2.5���3 ��m thick. Epidermal gland cells (Fig. 6 G) variable; entire epidermis glandular to varying degrees, gland cells pale and evenly distributed, or epidermal gland cells indistinguishable in living and preserved material. In specimens with glandular epidermis body surface often covered with foreign substance (humus particles). Clitellum saddle-shaped, not developed ventrally; cells in separate (viv) or dense (fix) rows; dorsally in ca. 30 transverse rows; hyalocytes and granuloctyes alternating, granulocytes isolated; ventro-laterally only granulocytes. Clitellum extending posteriorly to chaetae of XIII. Prostomium (Fig. 6 A,C,D) with head pore in mid-dorsal position. Frontal prostomial epithelium thickened, with a vesicle-like recess or cleft at the frontal tip. Dorso-laterally several inner papillae in bilateral-symmetrical order, papillae also on peristomium. Prostomial musculature present. Body wall less than 5 ��m thick (viv); fix: 4��� 9 ��m. Cuticle thin, indistinguishable (viv), or Septa 4 / 5���6 / 7 (- 9 / 10) thicker than the rest. Brain in I���III, 2���2.5 x as long as wide, incised posteriorly, sides converging anteriad (Fig. 6 D). A pair of ganglia on prostomial nerves. Suboesophageal ganglion in III���IV, perikarya of ventral nerve cord in segmental ganglia from V on. Pharyngeal glands (Fig. 6 A,B) with unpaired dorsal lobes in IV and V; primary ventral lobes in V, elongate; secondary ventral lobes in V and VI, spherical, smaller than primary ventral lobes. In VI���VII a pair of separate elongate lobes, consisting of an anterior dorsal part in VI and a posterior ventral part in VI���VII; both parts broadly connected in Z-like fashion, ventral portion extending into VII to varying degrees, without constriction at septum. Oesophageal appendages and intestinal diverticula absent, intestine from VII, here epithelium thickened in VII and parts of VIII (Fig. 6 B); transition from oesophagus to intestine at 6 / 7, widening abrupt here, or gradual over several following segments. Chloragocytes sparse, inconspicuous, small, first cells from VI. Dorsal vessel from 1 / 2 XII ��� XIII, difficult to see in living specimens. Pars tumida of midgut from 1 / 2 XVI ��� XXIII, extending over 1.5���4 segment lengths. Preclitellar nephridia (Fig. 6 B) two pairs, at 7 / 8 and 8 / 9, not constricted at septum, ca. 50���70 ��m long (fix). Anteseptale with parts of nephridial body, longer than wide, funnel attached in obliquely upright position. Postseptale larger than anteseptale, length ratio ca. 2: 1, with a dorsal bump in mid-section, usually without vescile, gradually tapering into short efferent duct; nephridial canal apparently with an up- and down zig-zag course; nephroporus inconspicuous, anterior to ventral chaetal bundles, no terminal vesicle. A small dorsal vesicle seen in the postseptale a few specimens, similar to the one in Guaranidrilus. First postclitellar nephridia at 13 / 14 or 14 / 15, longer than in anterior segments (ca. 70���90 ��m, fix), in following segments only few positions occupied. Coelomocytes elongate, pale, broadly oval, ca. 15���20 ��m long (viv, fix), filled with distinct, pale, spherical vesicles; a few vesicles may be refractile; cells of some specimens dark in aggregations but without colour or tint. Seminal vesicle absent. Spermatozoa ca. 42 ��m long, heads ca. 18 ��m long in vivo. Sperm funnel (Fig. 6 E) small, less than half as long as body diameter and only slightly longer than wide (1.3���1.5 x), collar distinct, thickwalled, slightly narrower than widest funnel body diameter, canal comparatively wide. Fix: length 35 ��m diameter 20���22 ��m, almost cylindrical in cross section (i.e. only slightly flattened). Vas deferens ca. 4 ��m wide (viv, fix), in 2���3 wide coils in XII, of same diameter throughout. Male copulatory organ (Fig. 6 E) small, glandular bulb spherical or oval in top view, longest diameter ca. 20 ��m (viv, fix), not more than a swelling of body wall in side view, pierced centrally by vas deferens, occasionally accompanied by a pair of small extra bulbs (diameter Accessory glands absent. Spermatheca most often small and inconspicuous, often difficult to see in living specimens. Ectal duct ca. 25 ��m long and ca. 8 ��m wide (fix), short and simple, as if absent in living specimens (viv); distal part of ampulla as wide as or slightly wider than ectal duct, spherical or ovoid, diameter 10���15 ��m, thin-walled, with sperm aligned in parallel in longitudinal axis of spermatheca; connecting tube thinner than ectal duct, diameter 4���5 ��m, ental reservoir reaching VII or VIII when fully extended (reaching X in 1 specimen), often coiled either in VI or V; diameter 10��� 12 ��m proximally; ental reservoir not always developed; in these cases the proximal end as narrow as the connecting tube and difficult to distinguish in living and preserved material. Eggs. One or two mature eggs at a time. Habitat. X. aphanus was found in grazed and abandoned pastures and in early and medium-growth stages of forest regeneration. It was absent in old-growth forest and almost absent at the agroforestry sites (1 specimen found). Remarks. Xetadrilus aphanus worms were among the smallest enchytraeids in all samples; adult specimens were often considered as juveniles or hatchlings of other species. The most conspicuous trait is the Z-shape of the posterior pharyngeal gland in VI and VII, unique in the genus and to our knowledge also in the family. Further differentiating traits are listed in Table 3. The species is variable with respect to (1) length of spermatheca, (2) presence and conspicuousness of epidermal gland cells, (3) size or length of pharyngeal glands in VII, (4) dorsal vesicle in the nephridial postseptale (mostly absent, seen in a few specimens), and (5) conspicuousness of male glands and copulatory body muscles., Published as part of Schmelz, R��diger M., Collado, Rut & R��mbke, J��rg, 2011, Mata Atl��ntica enchytraeids (Paran��, Brazil): A new genus, Xetadrilus gen. nov., with three new species, and four new species of Guaranidrilus ��ernosvitov (Enchytraeidae, Oligochaeta), pp. 1-29 in Zootaxa 2838 on pages 18-20, DOI: 10.5281/zenodo.203260

Einleitung: Ziel der vorliegenden Studie ist es, mittels intraartikulärer Drucksensoren an anatomischen Frischpräparaten Belastungsveränderungen nach isolierter Capitatumverkürzungsosteotomie, wie sie zur Behandlung der Lunatumnekrose durchgeführt wird, zu bestimmen. Damit soll[for full text, please go to the a.m. URL], 127. Kongress der Deutschen Gesellschaft für Chirurgie