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Enchytraeus andrasi sp. nov. urn:lsid:zoobank.org:act: 5E2BFD02-E8F8-4689-95F1-57E3A7B01A61 Figs 1C–D, 4, Tables 1–2 Diagnosis (1) Body length 11–14 mm (in vivo), segment number 29–34; (2) chaetae maximum 3 per bundle, straight with ental hook; (3) clitellum in XII–XIII, hyalocytes and granulocytes irregularly arranged dorsally and laterally, ventrally absent between the male lip-like folds; (4) four pairs of nephridia preclitellarly; (5) all pharyngeal glands connected dorsally with ventral lobes, the third pair may be free; (6) dorsal blood vessel origin from XIV, blood colourless; (7) sperm funnel cylindrical, 380–580 μm long, 3–5 × as long as wide (in vivo); (8) vasa deferentia uniform, 24–25 μm wide with 8 μm thick wall (fixed), ciliated, not extending beyond XIII; (9) male glands multiple: one large rounded primary bulb, diameter 50–70 μm (fixed), and many smaller secondary glands; (10) spermathecal ectal duct (50–85 μm long, fixed) covered with gland cells, ampulla irregularly sac-like, about 100–106 μm wide with one large diverticulum (80–100 μm long, fixed), connecting with oesophagus; (11) more small mature eggs present at a time. Etymology Named in the honour of the brother of Klára Dózsa-Farkas, András Dózsa-Farkas, who collected the sample with this species. Material examined Holotype ITALY • last 14 segments, 3.4 mm used for DNA analysis (No. 1386, ID number); Ligurian Sea, Castiglione seashore, decaying seagrass debris; 42°45′56.0″ N, 10°52′51.0″ E; 13 Dec. 2019; András and Kinga Dózsa-Farkas leg.; slide 2871, En.3; ELTE. Paratypes ITALY • 1 spec., last 9 segments, 1.65 mm used for DNA analysis (No. 1390, ID number); same collection data as for holotype; slide 2870, P.144.1; ELTE • 1 spec., last 9 segments, 1.65 mm used for DNA analysis (No. 1391, ID number); same collection data as for holotype; slide 2873, P.144.2; ELTE. Description MEASUREMENTS. Medium-sized to large specimens. Holotype 14.3 mm long, 470 μm wide at VIII and 600 μm at clitellum in vivo, 9.3 mm long, 330 μm wide at VIII and 420 μm at clitellum, when fixed, 41 segments. Body length of paratypes 11–12 mm, width 430–450 μm at VIII and 550–680 μm at clitellum, in vivo. Length of fixed specimens 6.7–8.1 mm, width 420 μm at VIII and 470–580 μm at clitellum. Length of the first 12 segments 2.5–2.6 mm, after fixation. Segment number 29–34. CHAETAE. Chaetal formula: 3–2,3:3–2,3. Chaetae straight with ental hook, about equal in size within bundle, 60–62 μm × 4 μm preclitellarly and 67–75 μm × 4–5 μm posteriorly. Often 2–3 surplus chaetae near bundles (Fig. 4B). Chaetae in XII absent ventrally, but present laterally, 2 per bundle. HEAD PORE. At 0/I. EPIDERMAL GLANDS. Inconspicuous. CLITELLUM. In XII–XIII, hyalocytes and granulocytes irregularly arranged dorsally and laterally (Fig. 4C), ventrally absent between male lip-like folds (Fig. 4E). Before and behind some glands but few. BRAIN (Fig. 4A). About 1.8–2 × as long as wide, truncate posteriorly, sides slightly merging anteriad. OESOPHAGEAL APPENDAGES. Pair of blind-ending tubes. In two specimens all primary pharyngeal glands connected dorsally, with ventral lobes (Fig. 4D), but in third specimen last pair free dorsally. DORSAL BLOOD VESSEL. From XIV, blood colourless. Anterior bifurcation in I. NEPHRIDIA. Four pairs of preclitellar nephridia from 6/ 7–9 /10, anteseptale funnel only, postseptale bulged, short efferent duct, originating postero-ventrally (Fig. 4F). COELOMOCYTES. Oval, texture granulated, about 18–25 μm long, in vivo. No shining, hyaline corpuscles. SUBNEURAL GLANDS. Absent. SPERM SACS. Two paired large lobes of sperm sacs, filling the coelom of X–XI. Testes and sperm funnels in XI, ovaries, male pores and glands in XII. SPERM FUNNELS. Large, 380–580 μm long, 3–5 × as long as wide in vivo (235–330 μm long and 2–2.5 × as long as wide, fixed). Collar slightly narrower than funnel body (Fig. 4H). Vasa deferentia long, irregularly coiled in XII–XIII, about 24–25 μm wide with 8 μm thick wall (fixed), ciliated (Fig. 4G, I). SPERMATOZOA. Not measurable, heads about 21–36 μm, in vivo. MALE COPULATORY ORGANS. Male glands multiple: one larger rounded primary bulb (diameter 50–70 μm, fixed) near male pore, and many smaller (32–78 μm long and 18–24 μm wide, fixed) secondary glands. Glands arranged roughly in semicircle around male pore and primary bulb (diameter 180–250 μm, fixed) (Fig. 4J). Male pores covered by lip-like folds (about 80 μm wide), recess present. SPERMATHECAE (Figs 1C–D, 4K). With short ectal duct (50–85 μm long, fixed), covered with gland cells (50–75 μm widely in vivo and fixed equally), canal 4–6 μm wide. Ampulla irregularly sac-like, about 100–106 μm wide, heads of spermatozoa embedded in wall of ampulla. Ampulla with one large diverticulum (80–140 μm long, 40–68 μm wide, fixed) (Fig. 1C), in one case ampulla also with small diverticula-like protrusions (Figs 1D, 4K). Ental duct short and opens into oesophagus separately. More small mature eggs present at a time (Fig. 4I). Differential diagnosis In the new species, like as in four of the species in the E. albidus complex (E. albidus s. str., E. moebii, E. polatdemiri and E. irregularis of which sequences are available, the vasa deferentia approximately uniform and ciliated throughout. The principal differences are that the spermathecal ampullae have diverticula in the new species but neither E. moebii nor E. polatdemiri have it. Besides E. moebii is much larger than the new species (body length 25–35 mm with segment number 60–74 vs 6.7–9.3 mm 29–41) and has more chaetae in ventral bundle (4–5 vs 3). Although in E. irregularis the spermatheca has also diverticulum and the maximum number of chaetae is 3, but E. irregularis is larger than the new species (length 9–13 mm with segment number 47–76), and the blood is pink (in E. andrasi sp. nov. colourless) and has extra copulatory glands. Morphologically, E. albidus s. str. is most similar to the new species taking, the form of the spermatheca into consideration, but differs from it in size (10– 13 mm with 58–69 segments vs 6.7–9.3 mm and 29–41 segments) and in the maximum number of chaetae (4–5 vs 3) and the sperm duct extends into XIV–XX (vs only in XII–XIII). The new species is also similar to E. andrasiformis sp. nov., with the morphological differences listed in the differential diagnosis of the latter species.As shown below, E. andrasi is clearly distinguished as separate from all other species by genetic data, which also give strong support for E. andrasi being most closely related to E. andrasiformis, described below. For similarities and differences of the species of Enchytraeus studied by us, see Table 2. Distribution and habitat Castiglione seashore, Ligurian Sea, Italy, decaying seagrass debris., Published as part of Nagy, Hajnalka, Dózsa-Farkas, Klára & Felföldi, Tamás, 2023, New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia, pp. 107-145 in European Journal of Taxonomy 870 on pages 13-19, DOI: 10.5852/ejt.2023.870.2123, http://zenodo.org/record/7986633, {"references":["Erseus C., Klinth M. J., Rota E., Wit P. D., Gustafsson D. R. & Martinsson S. 2019. The popular model annelid Enchytraeus albidus is only one species in a complex of seashore white worms (Clitellata, Enchytraeidae). Organisms Diversity & Evolution 19: 105 - 133. https: // doi. org / 10.1007 / s 13127 - 019 - 00402 - 6","Arslan N., Timm T., Rojo V., Vizcaino A. & Schmelz R. M. 2018. A new species of Enchytraeus (Enchytraeidae, Oligochaeta) from the profundal of Lake Van, the world's largest soda lake (Turkey, East Anatolia). Zootaxa 4382 (2): 367 - 380. https: // doi. org / 10.11646 / zootaxa. 4382.2.8","Cejka B. 1913. Litorea krumbachi n. spec. n. gen. Ein Beitrag zur Systematik der Enchytraeiden. Zoologischer Anzeiger 17: 145 - 151.","Nielsen C. O. & Christensen B. 1961. The Enchytraeidae. Critical revision and taxonomy of European species. Supplement 1. Natura Jutlandica 10: 1 - 23.","Dozsa-Farkas K. 2019. Enchytraeids of Hungary (Annelida: Clitellata: Enchytraeidae). Pedozoologica Hungarica 7. Eotvos University Press, Budapest.","Henle F. G. J. 1837. Ueber Enchytraeus, eine neue Anneliden-Gattung. Archiv fur Anatomie, Physiologie und Wissenschaftliche Medicin 74 - 90.","Michaelsen W. 1885. Vorlaufige Mitteilungen uber Archenchytraeus Mobii n. sp. Zoologischer Anzeiger 8: 237 - 239."]}

Enchytraeus sp.2 from Svalbard Material examined SVALBARD • 7 specs; Kongsfjorden; ID 264 to 265, slide No. 2140 to 2141, 3039, 3217 to 3218; ELTE. Description Short description of fixed specimens: MEASUREMENTS. Body length 11–16 mm, segment number 55–66, 650–800 μm wide at clitellum. CHAETAE. Chaetal formula: 3,2–2:3(2)–2. Preclitellar chaetae 60–85 μm × 7 μm ventrally, 55–58 μm × 6 μm laterally, postclitellar chaetae 75–85 μm × 6 μm ventrally, and 58–75 μm × 6 μm laterally. DORSAL VESSEL. From XIII–XIV. SPERM SACS. Large sperm sacs, located in X–XI. SPERM FUNNEL. 376 μm long, about 3 × as long as wide, fixed. Sperm duct about the same wide in all length (20–25 μm wide). MALE COPULATORY ORGANS. Male glands multiple: one larger rounded primary bulb (diameter 75–85 μm) and numerous smaller glands. SPERMATHECAE. Ectal duct of spermatheca 62–90 μm long with large glands around orifice of duct. Ampulla spherical (diameter 120–140 μm) with one diverticulum (about 270 μm long), or 2–3 smaller diverticula (80–110 μm long). Additional specimens would be required for accurate description. Distribution and habitat Kongsfjorden, Svalbard, under a bird cliff ‘compost’ of bird excrements, mosses, vascular plants and nest material., Published as part of Nagy, Hajnalka, Dózsa-Farkas, Klára & Felföldi, Tamás, 2023, New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia, pp. 107-145 in European Journal of Taxonomy 870 on page 31, DOI: 10.5852/ejt.2023.870.2123, http://zenodo.org/record/7986633

Enchytraeus sp.3 from Kerguelen Islands Material examined KERGUELEN ISLANDS • 5 specs; Ile Guillou Island; ID 260 to 261, slide No. 3035 to 3037; ELTE. Description Based on the few specimens in imperfect condition, Echytraeus sp.3 from Kerguelen Islands is very similar morphologically to E. albidus s. str. MEASUREMENTS. Body length 10–13 mm, segment number 44–54, 610–660 μm wide at clitellum, fixed. CHAETAE. Chaetal formula: 3–2,3:3,4–2,3. 87–95 μm × 8 μm in ventral bundles, 77–80 μm × 7.5 μm in lateral bundles preclitellarly and 77–80 μm × 7 μm in ventral bundles and 62–70 μm × 5 μm in lateral bundles posteriorly. DORSAL VESSEL. from XIII–XIV. COELOMOCYTES. 31–42 μm long. SPERM SACS. Large sperm sacs, located in X–XI. SPERM FUNNEL. 375–550 μm long, about 4 × as long as wide, fixed. SPERMATOZOA. About 60 μm long, heads 18 μm. Sperm duct about same wide in all length (23–30 μm wide). MALE COPULATORY ORGANS. Male glands multiple: one larger rounded primary bulb (80–100 μm, fixed) and numerous smaller glands. SPERMATHECAE. Ectal duct of spermatheca with glands around duct, not only at the orifice. Ampulla with one diverticulum (about 210 μm long, fixed). Distribution and habitat Guillou Island, Kerguelen Islands, under a small clump of Poa kerguelensis. Results of molecular analysis Results of molecular analysis corroborated that E. adrianensis sp. nov., E. andrasi sp. nov. and E. andrasiformis sp. nov. are new, distinct species, since the sequences obtained from the examined individuals clearly separated in the phylogenetic trees (Figs 10–12). Our study also provided new reference sequences for E. albidus s. str., E. irregularis, E. krumbachi, and it provided sequences for further unnamed taxa within the E. albidus group (E. sp.2 and E. sp.3). Enchytraeus andrasi sp. nov. and E. andrasiformis formed distinct lineages, they were placed outside the clade of the E. albidus complex in the phylogenetic trees. The p-distances between the COI sequences of E. andrasi and E. andrasiformis are 16.8–17.6%, the H3 distances between them are 3%. The COI distances between E. andrasi and the members of E. albidus complex (including E. adrianensis) are 15.9–20.6%, the H3 distances between them are 5.4–8.5%. The intraspecific COI distances between specimens of E. andrasi are 0–1.8%. There are no intraspecific differences between the H3 sequences of E. andrasi individuals. The COI distances between E. andrasiformis and the species of E. albidus complex (including E. adrianensis sp. nov.) are 15.6–20.3%, the H3 distances between them are 5.2–8.1%. Enchytraeus adrianensis and additional species missing from the study of Erséus et al. (2019), namely E. irregularis, E. sp.2 and E. sp.3 were placed inside the clade of the E. albidus complex. The COI distances between E. adrianensis and members of E. albidus complex are 12.8–17.8%, the H3 distances between them are 0.4–5.5%. There are no intraspecific differences between the COI and between the H3 sequences of specimens of E. adrianensis. Enchytraeus sp.2 and E. sp.3 formed one group on the gene trees. It is worth noting that the COI distance between E. sp.2 and E. sp.3 is very small (0.2%), while the H3 distance between them is bigger (2.5%), and comparable to interspecific differences in the rest of the H3 gene tree (Fig. 11). Enchytraeus adrianensis appeared as the sister group of E. krumbachi in the COI and H3 gene trees, and E. irregularis appeared as the sister group of E. sp.2 / sp.3 clade in both of the gene trees (Figs 10– 11). The COI distances between E. adrianensis and E. krumbachi are 12.9–13.4%, the H3 distances between them are low, 0.4–1.6%. The COI distances between E. irregularis and E. sp.2 / sp.3 clade are 12.2–12.9%, the H3 distances between them are 2.1–3.8%. According to the molecular analysis, our E. krumbachi individuals and E. cf. krumbachi CE1684 and CE1689 specimens belong to the same species. The intraspecific COI distances between our E. krumbachi and E. cf. krumbachi individuals are 0.7–3%, the intraspecific H3 distances between them are 0–0.8%. On the species tree (Fig. 12), the E. albidus complex formed a distinct clade with maximum posterior probability support and contains E. adrianensis, E. irregularis and E. sp.2 / sp.3, as in the gene trees. E. adrianensis sp. nov. appeared as the sister group of E. krumbachi, E. sp.2 / sp.3 appeared as the sister group of E. irregularis. Enchytraeus andrasi and E. andrasiformis were placed outside the E. albidus complex and they appeared there as sister groups of each other. E. andrasi and E. andrasiformis clustered with E. norvegicus Abrahamsen, 1969 on the species tree (Fig. 12) but with only low posterior probability support, and the latter species differs from the new species in many traits., Published as part of Nagy, Hajnalka, Dózsa-Farkas, Klára & Felföldi, Tamás, 2023, New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia, pp. 107-145 in European Journal of Taxonomy 870 on pages 31-32, DOI: 10.5852/ejt.2023.870.2123, http://zenodo.org/record/7986633, {"references":["Erseus C., Klinth M. J., Rota E., Wit P. D., Gustafsson D. R. & Martinsson S. 2019. The popular model annelid Enchytraeus albidus is only one species in a complex of seashore white worms (Clitellata, Enchytraeidae). Organisms Diversity & Evolution 19: 105 - 133. https: // doi. org / 10.1007 / s 13127 - 019 - 00402 - 6"]}

Enchytraeus irregularis Nielsen & Christensen, 1961 Figs 1G, 9, Tables 1–2 Enchytraeus irregularis Nielsen & Christensen, 1961: 13. Enchytraeus irregularis – Hong & Dozsa-Farkas 2018: 81–85, figs. 2–4. Enchytraeus irregularis – Dózsa-Farkas K. 2019: 81–82, fig. 6.39.1 Material examined KOREA • 5 specs; Jeollabuk-do, Songcheon-dong; ID: 262, 906, slide 573 to 575; ELTE. HUNGARY • 5 specs; Érd; ID 946, slide 3029 to 3031, 3034 • 2 specs; Tiszalök, slide 2110 to 2111; ELTE. Description The species was described by Nielsen & Christensen (1961) from compost heap in the Botanical Garden of Copenhagen, Denmark. In 1987, Dózsa-Farkas found it in an earthworm-farm in Tiszalök (Hungary) and in 2016, in a sewage-sludge compost-bed in Érd (Hungary). In 2007, individuals of E. irregularis were collected from an earthworm breeding box in house (Korea, Jeollabuk-do) and a culture was made from these specimens. The results of the latter study were published in Hong & Dózsa-Farkas (2018). Schmelz & Collado (2010) proposed its synonymy with E. capitatus von Bülow, 1957, but Dózsa-Farkas (2019) disagreed with it. Now we were able to study these specimens using DNA sequencing, so all earlier specimens and some new specimens from Korea and Hungary were studied again morphologically. On the basis of the result of this new study, we give an additional description of this species. MEASUREMENTS. Body length 14–20 mm, width 540–700 μm at clitellum in vivo (length 9–13 mm and 530–880 μm wide, if fixed), segment number 47–66 (segment number 41–49 in Nielsen & Christensen 1961). Length of the first 12 segments 2.3–3.0 mm, after fixation. CHAETAE. Straight or slightly curved, formula of 3–2,3: 3–2,3,(4). CLITELLUM. Saddle-shaped, over XII–XIII (XIV), gland cells small, irregularly scattered (Fig. 9B). HEAD PORE. At 0/1. BRAIN. About 150 μm long, 1.6 × as long as wide in vivo, with one or two pairs of aggregations of refractile globules (Fig. 9A). OESOPHAGEAL APPENDAGES. Short, outline irregular, thin-walled with single central lumen. All three pairs of primary pharyngeal glands with connection dorsally and ventral lobes. DORSAL VESSEL. From XIV–XVI, blood faintly red, anterior bifurcation in peristomium (Fig. 9D). COELOMOCYTES. One type, only mucocytes, small, oval or elongate, pear-shaped, 20–29 μm long in vivo, filled with light granules (Fig. 9C). Shining hyaline corpuscles also occur but in few numbers. NEPHRIDIA. Four pairs of nephridia preclitellarly. SPERM SACS. Paired, well developed and lobed, located in IX–XI (Fig. 9G). SPERM FUNNELS (Fig. 9G). 415–730 μm long, 4–7 × as long as wide, about as long as the body diameter in vivo, (270–600 μm long, 2–4 × as long as wide, if fixed), collars slightly wider than funnel. SPERMATOZOA. About 80–100 μm long, heads about 30 μm in vivo, sperm ducts uniform, about 20 μm wide. MALE COPULATORY ORGANS. Male glands multiple with larger primary bulb (70–100 μm × 70–100 μm) and 6–8 narrower, long, glandular secondary glands (50–90 μm × 12–14 μm) around male pores (Fig. 9F–G). After male copulatory organs pair of oval or squarish glandular structure visible with subdivision (‘accessory sexual glands’) (150–250 μm long, 110–225 μm wide, about 100 μm high, fixed), without external orifice (Fig. 9F, H–I). Usually in XIII, directly before the ventral chaetal bundles. In one specimen, visible immediately before male copulatory organs, and in another specimen, where clitellum was from XII to XIV, this organ was found in XV and only on one side (slide no. 573). Now revised again under microscope, so Hong & Dózsa-Farkas (2018): 85 is not correct: “clitellum extended over XII–XV and accessory sexual gland found in XIV” (but correct caption of fig. 3g). In Nielsen & Christensen (1961), location of this organ in front of penial bulb and also in XIII ventrally. These organs mostly larger than male copulatory organ. SPERMATHECAE. Spermathecal ectal ducts (87–110 μm long), with distinct canals (5–8 μm wide) and ring of glands around ectal orifices. Length of glands about 55–75 μm (fixed). Length of duct without glands proximally 13–25 μm. Duct widens into ampulla proximally (diameter of ampullae about 75–130 μm), which again gradually tapers into short ental ducts, communicating separately with oesophagus. From ampullae anteriorly or laterally arises one large diverticulum (170–220 μm long in vivo, 100–200 μm fixed) filled with spermatozoa (Fig. 9K–L). 2–8 mature eggs at a time. Differential diagnosis The main differences between this species and the other species investigated by us are the pink blood, and the characteristic large extra sexual glands without external orifice. For similarities and differences of this species compared to other species of the E. albidus complex, see Table 2. Distribution and habitat Songcheon-dong, Jeollabuk-do, Korea, earthworm breeding box in house.Tiszalök, Hungary, earthworm-farm. Érd, Hungary, sewage-sludge compost bed. This species has unknown origin, only appeared in culture, it has not yet been found in the wild., Published as part of Nagy, Hajnalka, Dózsa-Farkas, Klára & Felföldi, Tamás, 2023, New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia, pp. 107-145 in European Journal of Taxonomy 870 on pages 27-30, DOI: 10.5852/ejt.2023.870.2123, http://zenodo.org/record/7986633, {"references":["Nielsen C. O. & Christensen B. 1961. The Enchytraeidae. Critical revision and taxonomy of European species. Supplement 1. Natura Jutlandica 10: 1 - 23.","Hong Y. & Dozsa-Farkas K. 2018. New description of enchytraeid species (Clitellata: Enchytraeidae) from Korea. Journal of Species Research 7: 80 - 91. https: // doi. org / 10.12651 / JSR. 2018.7.1.080","Schmelz R. M. & Collado R. 2010. A guide to European terrestrial and freshwater species of Enchytraeidae (Oligochaeta). Soil Organisms 82: 1 - 176.","von Bulow T. 1957. Systematisch-autokologische Studien an eulitoralen Oligochaeten der Kimbrischen Halbinsel. Kieler Meeresforschungen 13: 96 - 116.","Dozsa-Farkas K. 2019. Enchytraeids of Hungary (Annelida: Clitellata: Enchytraeidae). Pedozoologica Hungarica 7. Eotvos University Press, Budapest."]}

Enchytraeus albidus Henle, 1837 s. str. Figs 1F, 6, Tables 1–2 Material examined HUNGARY • 16 specs; Budapest; ID 1300 to 1301, 1461 to 1462, slide 3012, 3013, 3016 to 3018, 3024, 3028, 3044 to 3048; ELTE. Description of new material We investigated a living population of E. albidus s. str. from the laboratory of Szent István University, Hungary, which was derived from the culture of ECT Oekotoxikologie GmbH, Flörsheim, Germany. From this culture the neotype of E. albidus has been selected (Erséus et al. 2019). In this way our morphological characters substantially agree with those described in Erséus et al. (2019). MEASUREMENTS. Segment number 58–65, body length 15.5–20.8 mm, 850–900 μm wide at clitellum in vivo (700–980 μm, when fixed). CHAETAE. In ventral bundles mostly 3–5 (Fig. 6A). BRAIN. 125–165 μm long, 1.2–1.8 × as long as wide (fixed). PHARYNGEAL GLANDS. Variable, all converging or connected dorsally, or not converging (in Erséus et al. 2019: all converging). COELOMOCYTES. 17–32 μm long, mostly oval or spindle-shaped, granulated (in Erséus et al. 2019: 10– 20 μm). In the coelomocytes mostly shining, hyaline, round or tetragonal corpuscles (diameter 9–15 μm) are also present, similar to E. adrianensis sp. nov. CLITELLUM. Hyalocytes and granulocytes of clitellum irregularly arranged dorsally (Fig. 6B). Ventral surface of XII with a recess between lip-like folds (Fig. 6C–D, J). NEPHRIDIA. Four pairs of nephridia preclitellarly (Fig. 6G). SPERM FUNNELS. 700–820 μm long in vivo, 5–6 × as long as wide, the collar as wide as funnel body (Fig. 6H). Head of spermatozoa 30–60 μm long. Sperm duct ciliated (Fig. 6E–F), elongate, about the same width throughout (30–44 μm wide, with 9–16 μm thick wall), coiled, extending into XIV–XVII. MALE COPULATORY ORGANS. Male glands multiple: one larger rounded primary bulb (65–80 μm, fixed) and numerous smaller glands (40–50 μm, fixed) (Fig. 6I). SPERMATHECAE. Ectal duct of spermatheca 105–200 μm long, diameter of glands around duct 90–130 μm in vivo. Diverticulum of ampulla 180–237 μm long and 70–125 μm wide in vivo (Figs 1F, 6K–L). For similarities and differences of the species of Enchytraeus studied by us, see Table 2. Distribution and habitat Laboratory culture. For additional references, including synonymies and further distributional data, see Erséus et al. (2019)., Published as part of Nagy, Hajnalka, Dózsa-Farkas, Klára & Felföldi, Tamás, 2023, New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia, pp. 107-145 in European Journal of Taxonomy 870 on pages 22-24, DOI: 10.5852/ejt.2023.870.2123, http://zenodo.org/record/7986633, {"references":["Henle F. G. J. 1837. Ueber Enchytraeus, eine neue Anneliden-Gattung. Archiv fur Anatomie, Physiologie und Wissenschaftliche Medicin 74 - 90.","Erseus C., Klinth M. J., Rota E., Wit P. D., Gustafsson D. R. & Martinsson S. 2019. The popular model annelid Enchytraeus albidus is only one species in a complex of seashore white worms (Clitellata, Enchytraeidae). Organisms Diversity & Evolution 19: 105 - 133. https: // doi. org / 10.1007 / s 13127 - 019 - 00402 - 6"]}

Enchytraeus krumbachi (Čejka, 1913) Figs 1B, 7–8, Tables 1–2 Enchytraeus krumbachi Cejka, 1913: 145–151, figs 1–10. Enchytraeus krumbachi – Erséus et al. 2019: 126–129, fig. 12. Emended diagnosis As a result of our comparisons, we suggest the following diagnosis of the species: (1) Body length 15–25 mm (in vivo), segment number 40–66; (2) chaetae maximum 4 (5, 6) per bundle; (3) granulocytes and hyalocytes irregularly arranged dorsally in clitellum; (4) four pairs of nephridia preclitellarly; (5) the pharyngeal glands are connected dorsally; (6) dorsal blood vessel origin from XV, blood colourless; (7) sperm funnel 1.2–3 × as long as wide (fixed); (8) vasa deferentia tripartite, extending into XIV–XIX, the middle part thicker, unciliated, the thinner part ciliated; (9) male copulatory organs: one large rounded primary bulb, slightly longer than wide (80–100 μm × 70–90 μm), many smaller secondary glands; (10) ampulla of spermatheca rounded, large, without diverticula, ectal duct with large glands covered; (11) 4–10 mature eggs present at a time. For similarities and differences of the species of Enchytraeus studied by us, see Table 2. Material examined ITALY • 11 specs (+ 12 specs only in vivo); Castiglione seashore; ID 1383, 1385, slide 2867 to 2869, 2872, 3004, 3006 to 3007, 3014 to 3015; ELTE • 3 specs; Punta Ala Grosseto; slide 3174, 3180 to 3181; ELTE. Description of new material and comparison with previous accounts Morphological characters identical with the description by Čejka (1913) and Erséus et al. (2019). Unfortunately, we did not have the opportunity to study the type material, but the original description is quite detailed and can be compared with our own data. MEASUREMENTS. Segment number 40–66. In vivo, body length 15–25 mm, width at clitellum 700– 950 μm; when fixed, length 7–14 mm (with first 12 segments 2.7–3.8 mm), width 500–860 μm. Čejka (1913) originally reported 66–75 segments, and body ca 15 mm long, and Erséus et al. (2019), studying fixed worms, noted length of 2.9–3.8 mm for first 12 segments, and width of 400–620 um at clitellum. Thus, our specimens are roughly identical in general size to earlier studied material, although slightly wider than individuals examined by Erséus et al. (2019). CHAETAE. Chaetal formula: 2,3–2,3:2,3,4,(5,6)–2,3. In ventral preclitellar bundles of some specimens 3 chaetae, and only one or two bundles with 4 chaetae, in other specimens 4 chaetae in all or most bundles; sometimes 2–3 surplus chaetae can be found. In Erséus et al. (2019), chaetal formula 3–2:3,4–2 and there are always 2 chaetae per bundle postclitellarly. Some of our individuals with 3 chaetae per bundle postclitellarly. In one specimen, 5 and 6 chaetae occured in some preclitellar ventral bundle. Čejka mentioned 3 chaetae in lateral and 4 chaetae in ventral bundles. Size of chaetae similar to size given by Erséus et al. (2019) (Table 2). 2–3 surplus chaetae also occasionally occur (Fig. 7A). COELOMOCYTES. Round or narrowed at one end, texture granulated, 15–27 μm length in vivo (Fig. 7B), (fixed not well measurable) Erséus et al. (2019): 10–15 μm, fixed. Shining, hyaline corpuscles mostly absent, but in some specimens occuring in few numbers. CLITELLUM. Granulocytes and hyalocytes irregularly arranged dorsally (Fig. 7C). In Erséus et al. (2019), ventral surface of XII with invaginations creating two recesses with overhanging lips. In our specimens near to male copulatory glands laterally (Fig. 7D) and ventrally between lips and midventrally only granulocytes (Fig. 7E). Čejka (1913) does not give any description of epidermal gland cells of epidermis. PHARYNGEAL GLANDS. In Erséus et al. (2019), pharyngeal glands are connected dorsally, in our individuals the position is mostly the same (Fig. 7F), but sometimes the first and second pairs of primary pharyngeal glands did not connect or they just contacted (Fig. 7G). In an unusual way, the third pair of pharyngeal glands was completely absent in one specimen. NEPHRIDIA. Four pairs of preclitellar nephridia at 6/ 7–9 / 10. SPERM FUNNELS. Length: width ratio of sperm funnel (Fig. 7I) 1.2–3:1 (fixed), (Erséus et al. 2019: 1.5–2:1, Čejka 1913: 3:1). Vas deferens (Fig. 8A–B) has same structure, tripartite, middle part thicker (55–75 μm with 20–30 μm wall thickness), unciliated, thinner ductus 20–37 μm thick and ciliated. Sizes of thicker part of ductus slightly larger than given by Erséus et al. (2019) (50–55 μm, with 15 or 15–30 μm wall thickness). In Erséus et al. (2019), vas deferens reaches XVIII, in our specimens variable: XIV–XIX, in Čejka (1913), it reaches XVI. SPERM SACS. Well developed, paired sperm sacs (Figs 7H, 8C) in IX–XI. MALE COPULATORY ORGANS. One large rounded primary bulb, slightly longer than wide (80–100 μm × 70– 90 μm), according to Erséus et al. (2019): diameter 85–90 μm and many smaller secondary (accessory) glands (Fig. 8D–E). In Čejka (1913), 6 gland complexes around penial bulb. SPERMATHECAE (Figs 1B, 7F–G, 8F–H). Without diverticula, ampulla round, large [70–140 μm, in Erséus et al. (2019): 95–125 μm], ectal duct with large glands covered (Figs 1B, 8F–H). In Erséus et al. (2019), ectal duct of spermatheca entally opening into small rounded chamber but we did not observe this chamber, and it is not mentioned in Čejka’s original description. Besides, differences in length and diameter of ectal duct (see Table 2). According to the morphological comparisons above, we think that our specimens agree better with the morphological description of Čejka (1913). Distribution and habitat Castiglione seashore, Ligurian Sea, Italy, decaying seagrass debris. Punta Ala Grosseto, Castiglione della Pescaia, Italy, decaying seagrass debris., Published as part of Nagy, Hajnalka, Dózsa-Farkas, Klára & Felföldi, Tamás, 2023, New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia, pp. 107-145 in European Journal of Taxonomy 870 on pages 24-27, DOI: 10.5852/ejt.2023.870.2123, http://zenodo.org/record/7986633, {"references":["Cejka B. 1913. Litorea krumbachi n. spec. n. gen. Ein Beitrag zur Systematik der Enchytraeiden. Zoologischer Anzeiger 17: 145 - 151.","Erseus C., Klinth M. J., Rota E., Wit P. D., Gustafsson D. R. & Martinsson S. 2019. The popular model annelid Enchytraeus albidus is only one species in a complex of seashore white worms (Clitellata, Enchytraeidae). Organisms Diversity & Evolution 19: 105 - 133. https: // doi. org / 10.1007 / s 13127 - 019 - 00402 - 6"]}