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Fig. 12. Enchytraeus Henle, 1837 in New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia

Fig. 12. Enchytraeus Henle, 1837 species tree, based on COI, H3 and 16S rRNA gene (Bayes analysis, *BEAST). Posterior probabilities greater than 0.5 are shown at the nodes. Accession codes of sequences with collection information are given in Table 1. Abbreviation: n = number of included specimens for each species. Scale bar = 0.07 substitutions per nucleotide.

Fig. 8 in New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia

Fig. 8. Micrograph of Enchytraeus krumbachi (Čejka, 1913). A–B, D, F. In vivo. C, E, G–H. Fixed, stained. A–B. Sperm ducts (thick-wall portion marked with black arrow, thin-wall portions marked with white arrows). C. Large sperm sacs with more lobes (slide 2868, ELTE). D–E. Male copulatory apparates (many accessory glands marked with arrows). E. Slide 2868 (ELTE). F–H. Spermathecae (ampulla rounded without diverticula). G. Slide 2867 (ELTE). H. Slide 2872 (ELTE). Scale bars = 50 μm. Abbreviation: pbc = primer gland.

Fig. 11 in New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia

Fig. 11. Maximum likelihood (ML) tree of the H3 gene for specimens of the Enchytraeus albidus species complex, based on 225 nucleotide positions using the Kimura 2-parameter substitution model. Bootstrap values greater than 50 are shown at the nodes. Accession codes of sequences with collection information are given in Table 1. Scale bar: 0.01 substitutions per nucleotide.

Fig. 10 in New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia

Fig. 10. Maximum likelihood (ML) tree of the COI gene for specimens of the Enchytraeus albidus species complex, based on 568 nucleotide positions using the General Time Reversible substitution model. Bootstrap values greater than 50 are shown at the nodes. Accession codes of sequences with collection information are given in Table 1. Scale bar: 0.05 substitutions per nucleotide.

Fig. 9 in New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia

Fig. 9. Micrograph of Enchytraeus irregularis Nielsen & Christensen, 1961. A–H, J, K. In vivo. I, L. Fixed, stained. I. Slide 2110 (ELTE). L. Slide 574 (ELTE). A. Brain (refractile globules marked with arrows). B. Clitellar glands, dorsolateral view. C. Coelomocytes. D. Head, anterior bifurcation of blood vessel with red blood (head pore marked with arrow). E. Chloragogen cells. F. Male copulatory organ (lateral view). G. Sperm funnels. H. Segments XII–XIII (male copulatory organ marked with black arrow, extra sexual gland marked with white arrow). I. Multiple copulatory glands (lateral view, primary gland marked with arrow). J. Extra sexual glands in XIII, ventro-lateral view. K–L. Spermathecae (diverticula marked with arrows). Scale bars: A–B, D–L = 50 μm; C = 20 μm. Abbreviation: s = sperm sacs.

Fig. 7 in New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia

Fig. 7. Micrograph of Enchytraeus krumbachi (Čejka, 1913). A–B, D–F, H–I. In vivo. C, G. Fixed, stained. A. Chaetae in ventral bundles preclitellarly (surplus chaetae marked with arrow). B. Coelomocytes. C. Clitellar glands, dorsal view (slide 2868, ELTE). D. Clitellar glands, lateral view (near to male copulatory apparates more granular glands). E. Between the two male copulatory apparates (marked with arrows) and middle ventrally only granulocytes. F–G. Pharyngeal glands (in F first and secondary glands connected dorsally, in G these glands near to each other but not connected dorsally, the spermathecae marked with arrows). G. Slide 2867 (ELTE). H. Sperm sacs in X–XI. I. Sperm funnel. Scale bars = 50 μm. Abbreviation: sf = sperm funnel.

Fig. 6 in New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia

Fig. 6. Micrograph of Enchytraeus albidus Henle, 1837 s. str. A–C, E–H, K–L. In vivo. D, I. Fixed, stained. J. Fixed, not stained. A. Chaetae in ventral bundles preclitellarly. B. Clitellar glands, dorsal view. C–D. Ventral recess between the lip-like folds in XII. D. Slide 3047 (ELTE). E–F Sperm ducts (In F the ciliated canal marked with arrow). G. Nephridium. H. Sperm funnel. I. Multiple male glands (larger primary bulb marked with arrows) (slide 3048, ELTE). J. Lip-like fold (slide 3028, ELTE). K–L. Spermathecae (diverticula marked with black arrow, glands at ectal duct marked with white arrow). Scale bars = 50 μm.

Fig. 5 in New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia

Fig. 5. Micrograph of Enchytraeus andrasiformis sp. nov. A–C, F–G, I, K. Holotype (slide 3132, En. 4; ELTE). A–C, F–G, K. In vivo. D–E, H–J, L. Fixed, stained. A. Brain. B. Clitellar glands, dorsal view. C. Clitellar glands, lateral view (three eggs visible). D. Paratype (slide 3094, P.145.2, ELTE), chaetae in ventral bundles with surplus chaetae, preclitellarly. E. Paratype (slide 3093, P.145.1, ELTE), pharyngeal glands. F. Coelomocytes. G. Sperm funnel. H. Paratype (slide 3094, P.145.2). J. Paratype (slide 3094, P.145.2, ELTE). L. Paratype (slide 3093, P.145.1, ELTE). H–I. Multiple male glands (large primary bulb marked with black arrows, extra sexual glands marked with white arrows). J–L. Spermathecae. Scale bars = 50 μm.

Fig. 1 in New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia

Fig. 1. Schematic drawing of spermathecae of the specimens studied.A. Enchytraeus adrianensis sp. nov. B. E. krumbachi (Čejka, 1913). C–D. E. andrasi sp. nov. E. E. andrasiformis sp. nov. F. E. albidus Henle, 1837. G. E. irregularis Nielsen & Christensen, 1961.

Fig. 4 in New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia

Fig. 4. Micrograph of Enchytraeus andrasi sp. nov. A–C, E, G–I, K. In vivo. D, F, J. Fixed, stained. A. Brain. B. Chaetae in ventral bundles preclitellarly (surplus chaetae marked with arrow). C. Clitellar glands of holotype, dorsal view. D. Paratype (slide 2870, P.144.1, ELTE), pharyngeal glands. E. Clitellar glands of holotype, ventral view (male lip-like folds marked with arrows, between the folds the glands absent: marked with *). F. Paratype (slide 2870, P. 144.1, ELTE), nephridium at 7/. G. Sperm duct. 8 H. Sperm funnel. I. Sperm duct and many small eggs. J. Holotype (slide 2871, En.3, ELTE), multiple male glands of holotype (large primary bulb marked with arrow). K. Spermatheca (large diverticulum marked with short black arrow, small diverticula like protrusions marked with white arrows, ectal duct covered with glands marked with long, black arrow). Scale bars = 50 μm.

Fig. 2 in New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia

Fig. 2. Micrograph of Enchytraeus adrianensis sp. nov. A–D, I–J. In vivo. E–H, K–L. Fixed, stained. A. Brain (small aggregations of refractile globules marked with arrows). B. Chaetae in a ventral bundles with 3 surplus chaetae. C. Clitellar glands in regular rows. D. Clitellar glands in reticulate pattern. E. Paratype (slide 2748, P.143.3, ELTE). F. Slide 2684 (ELTE). H. Slide 2761 (ELTE). K. Paratype (slide 2851, P.143.10, ELTE). L. Paratype (slide 2760, P.143.6). E–F. Clitellar glands, ventral view (male copulatory organs marked with black arrow, between these organs the granulocytes marked with white arrows). G. Paratype (slide 2856, P.143.15, ELTE), first pharyngeal glands free dorsally. H. All pharyngeal glands connected dorsally (spermathecae marked with arrows). I–K. Coelomocytes (in J the small hyaline, refracting corpuscules also present). L. Large paired lobes of sperm sacs. Scale bars: A–I, K–L = 50 μm; J = 20 μm.

Fig. 3 in New insights into the Enchytraeus albidus complex (Annelida, Enchytraeidae), with the description of three new species from seashores in Italy and Croatia

Fig. 3. Micrograph of Enchytraeus adrianensis sp. nov. A, C, G–H. In vivo. B, D–F, I. Fixed, stained. A. Sperm funnels. B. Paratype (slide 2748, P. 143.3, ELTE), sperm ducts (slender part marked with white arrow, wider part with black arrows). C–F. Multiple male glands (E–F primary bulb marked with arrow). D. Paratype (slide 2748, P.143.3, ELTE). E. Paratype (slide 2748, P.143.3, ELTE). F. Holotype (En.2. slide 2842, ELTE). I. Paratype (slide 2827, P. 143.8, ELTE). F. Holotype (En.2. slide 2842, ELTE). G–I. Spermathecae (ectal duct covered with glands marked with white arrows, ampullae filled with sperm in it marked with black arrows).Scale bars = 50 μm.

Fig. 6 Enchytraeus albidus s. str. a Male apparatus viewed dorsally. b in The popular model annelid Enchytraeus albidus is only one species in a complex of seashore white worms (Clitellata, Enchytraeidae)

Fig. 6 Enchytraeus albidus s. str. a Male apparatus viewed dorsally. b Schematic illustration of the male apparatus in cross section. ag accessory glands, pb penial bulb, vd vas deferens. Scale bar 100 μm

Fig. 4 Enchytraeus albidus s in The popular model annelid Enchytraeus albidus is only one species in a complex of seashore white worms (Clitellata, Enchytraeidae)

Fig. 4 Enchytraeus albidus s. str., live specimen, from the same locality as CE20518 (see Table 1). Photo: C. Erséus

Fig. 7 Enchytraeus albidus s. str. a–d in The popular model annelid Enchytraeus albidus is only one species in a complex of seashore white worms (Clitellata, Enchytraeidae)

Fig. 7 Enchytraeus albidus s. str. a–d Spermathecae. ed ectal duct, eg ectal gland, sa spermathecal ampulla, sd spermathecal diverticulum, sm sperm. Scale bars 100 μm

Fig. 9 Enchytraeus moebii a male copulatory apparatus. b–c in The popular model annelid Enchytraeus albidus is only one species in a complex of seashore white worms (Clitellata, Enchytraeidae)

Fig. 9 Enchytraeus moebii a male copulatory apparatus. b–c Spermathecae. ag accessory glands, ed ectal duct, eg ectal gland, mu musculature, pb penial bulb, sa spermathecal ampulla, sm sperm, vd vas deferens. Scale bar 100 μm

Fig. 2 Species tree for the E in The popular model annelid Enchytraeus albidus is only one species in a complex of seashore white worms (Clitellata, Enchytraeidae)

Fig. 2 Species tree for the E. albidus species complex (shaded), estimated using the multi-locus coalescent model. The number of included speci- mens for each species is given after the species name. Values above branches are posterior probability (PP) support values, values under terminal branches are mean PP from species delim- itation analyses in BPP, written as: mean analysis A/mean analysis B/ mean analysis C. Scale measures the branch lengths in proportion to total tree length

Fig. 12 in The popular model annelid Enchytraeus albidus is only one species in a complex of seashore white worms (Clitellata, Enchytraeidae)

Fig. 12 Enchytraeus cf. krumbachi a vas deferens, showing the transition between the thin- and thick-walled portions; note the lack of cilia inside the thick-walled part. b Male copulatory apparatus. c Spermatheca. ag accessory gland, ed ectal duct, eg ectal gland, pb penial bulb, sa spermathecal ampulla, sm sperm, vd vas deferens. Scale bars 100 μm

Fig. 3 in The popular model annelid Enchytraeus albidus is only one species in a complex of seashore white worms (Clitellata, Enchytraeidae)

Fig. 3 Gene trees for the Enchytraeus albidus species complex (shaded), estimated by using Bayesian Inference. a COI. b H3 (specimens marked with B-1^ or B-2^ after the species name are heterozygous and the

Fig. 11 in The popular model annelid Enchytraeus albidus is only one species in a complex of seashore white worms (Clitellata, Enchytraeidae)

Fig. 11 Enchytraeus albellus Klinth, Erséus and Rota, sp. nov., holotype. a Sperm funnel. b Transverse section comparing thin- and thick-walled portions of vas deferens. c Longitudinal sec- tion comparing thin- and thick- walled portions of vas deferens. d Male apparatus. e Spermatheca. ag accessory glands, ed ectal duct, eg ectal gland, mu musculature, pb penial bulb, sa spermathecal ampulla, sf sperm funnel, sm sperm, vd vas deferens. Scale bars 100 μm