taxonID	type	description	language	source
A50C0231FFAFFF9AFF14FABAFC82F986.taxon	description	(Figs 1, 2)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFAFFF9AFF14FABAFC82F986.taxon	materials_examined	Holotype. MHNUSC _ 10146: mature worm, incomplete (broken at segment XIV), unstained and whole-mounted in Canada balsam. Leg. Ana I. Camacho. Type locality. CAN 002 De los Santos cave, Cantabria, Spain, – 3.58453, 43.19491 (4 June 2002). Paratypes. MHNUSC _ 10147 _ 1 – 2: 1 mature and 1 immature specimen, unstained and whole-mounted in Canada balsam, from type locality (4 June 2002); MHNUSC _ 10147 _ 3 – 5: 2 mature specimens (1 stained in haematoxylin and 1 unstained) and 1 immature stained in haematoxylin, whole-mounted in Canada balsam, CAN 004 Covalagua cave (cueva del Agua), Cantabria, – 3.61173, 43.22852 (7 June 2002); MHNUSC _ 10147 _ 6 – 10: 2 mature specimens (1 stained in haematoxylin and 1 unstained) and 3 immature unstained, whole-mounted in Canada balsam, CAN 022 Upwelling of the Bustablado stream, Cantabria, – 3.61717, 43.27797 (14 Mars 2003); MHNUSC _ 10147 _ 11: 1 mature specimen stained in haematoxylin and whole-mounted in Canada balsam, CAN 034 Bustablado stream, Arredondo, Cantabria, – 3.60441, 43.27404 (8 June 2002); MHNUSC _ 10147 _ 12: 1 mature specimen stained in haematoxylin and whole-mounted in Canada balsam, CAN 008 Arduengo cave (Cuviobramante), Cantabria, – 3.62032, 43.27576 (14 Mars 2003); MHNUSC _ 10147 _ 13: 1 mature specimen stained in haematoxylin and whole-mounted in Canada balsam, CAN 047 Asón stream, La Cubera, Cantabria, – 3.60577, 43.25108 (20 January 2002). All leg. Ana I. Camacho.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFAFFF9AFF14FABAFC82F986.taxon	etymology	Etymology. Named in honor of Dr. Ana Isabel Camacho (MNCN, Madrid, Spain), the leader of the PASCALIS project in Spain, who contributes greatly to the knowledge and development of the biospeleology.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFAFFF9AFF14FABAFC82F986.taxon	description	Description. More than 2.6 mm long (all mature specimens are broken and lack posterior segments). Maximum number of segments (observed in immature specimens), 31. Diameter at segment V (slightly compressed specimens) 90 – 130 μm, at segment XI 100 – 180 μm. Clitellum 1 / 2 X – XII. Prostomium rounded, 42 – 58 μm long and 40 – 70 μm wide. Secondary annulation indistinct or slightly marked, up to VI. Somatic setae bifid, 26 – 35 μm long, with upper tooth shorter and thinner than lower (Fig. 2 A); (2) 3 – 5 setae per bundle anteriorly and 2 – 3 in the clitellar and postclitellar segments. Ventral setae of XI modified into bifid penial setae with short teeth, in number of 6 – 13 per bundle, 28 – 40 μm long and 1 – 1.5 μm thick, arranged comb-like, forming an open arc towards the sagittal plane and placed close to the male pores on their inner side (Figs 1 A, B, 2 D [ps]). One pair of male pores located in the posterior third of segment XI, in line with somatic ventral setae (Fig. 1 A). One pair of spermathecal pores in segment XI; four of the ten specimens studied have an additional pair in XII (Fig. 2 H), and one (from upwelling of Bustablado) has even additional spermathecal pores in X and XIII. All spermathecal pores are placed in the most anterior part of their respective segments, in line with somatic ventral setae (Figs 1 A, 2 H [sp]). Pharyngeal glands in segments IV – VI, poorly developed. Chloragogen cells beginning in VI. Oesophageal appendages and diverticula absent. One pair of testes in segment X. One pair of ovaries in segment XI. Sperm sacs extending by segments (IX) X – XI (XII); mature eggs in segments XI – XII (XIII). Vasa deferentia twisted, shorter than atria, 45 – 60 μm long and around 3 μm wide, entering the apical (ental) end of the atria (Fig. 1 A vd). Atria tubular to spindle-shaped, curved, 78 – 95 μm long and 12 – 16 μm maximum diameter, with a thin muscular outer layer (1 – 2 μm thick); atrial lumen frequently filled by spermatozoids; ectally, each atrium ends in a dilated spherical ampulla which is surrounded by a rather thin muscular layer (atrial bulb), 16 – 26 μm in diameter, acting as a pseudopenis in extrusion (Figs 1 A, 2 B – D, F – H [a, b, pp]). Two prostate glands per atrium, the anterior one large and placed near the entrance of vas deferens, and the posterior one small and placed close to the atrial bulb (Figs 1 A, 2 C, D [pr 1, pr 2]). Spermathecae of XI and XII filled with spermatozoids (Fig. 1 A, D); spermathecae of X and XIII empty (Fig. 1 C). Spermathecal duct muscular, conical and long (up to 30 µm) in XI, globular and short in XII and XIII, tubular and short in X; spermathecal ampulla tubular (up to 80 µm long, 10 – 20 µm in diameter) and directed forward (Figs 1 A, C – D, 2 B, E – H [sd, sa]).	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFAFFF9AFF14FABAFC82F986.taxon	distribution	Distribution and habitat. Caves, springs, phreatic and hyporheic habitats at the upper and middle parts of the Asón River basin, Cantabria, Spain. Alt. 161 – 710 m.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFAFFF9AFF14FABAFC82F986.taxon	discussion	Remarks. Following Erséus (1992 a), the presence of elongated and curved atria, each with two prostate glands attached to proximal and distal ends, respectively, vasa deferentia entering apical ends of atria, slender spermathecae with short ducts, and small straight penial setae allocate the new species described above in the genus Gianius Erséus. From the twelve species of Gianius currently known, Gianius camachoae sp. nov. is clearly distinguished by the presence of spermathecae in the atrial segment (segment XI), a character only shared with the marine species G. eximius Erséus, 1997 and G. urgorrii Martínez-Ansemil & Caramelo, 2025. Among other characters, G. camachoae sp. nov. clearly differs from these two species by the shape and structure of the spermathecae, ampullae elongate and ducts simple in the former, and ampullae globular and ducts bipartite in the two latter. G. crypticus (Rodriguez & Giani, 1989), described from the same geographical area (a cave in the upper part of the Clarin stream basin), appears as its most closely related species, although having exclusively a typical pair of spermathecae in the preatrial segment X. The variability in the number and location of the spermathecae observed in G. camachoae sp. nov. is high, but all specimens have, at least, one functional pair of spermathecae in the atrial segment. Furthermore, G. camachoae sp. nov. clearly differs from G. crypticus for at least two other characters, the atrial muscular layer (thin in the former; very thick in the later), and the atrial bulb (with a large lumen and surrounded by a relatively thin muscular layer in the former; heavily muscular in the later).	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFAAFF99FF14F946FAA4FE7A.taxon	description	(Figs 3, 4)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFAAFF99FF14F946FAA4FE7A.taxon	materials_examined	Holotype. MHNUSC _ 10149: mature worm, incomplete (broken at segment XVI), stained in paracarmine and whole-mounted in Canada balsam. Leg. Fabio Stoch. Type locality. LES 050 Bus delle Piatte cave (441 V / VR), Rocca, Lessinian Massif, Italy, 10.98458, 45.61771 (20 June 2006). Paratypes. MHNUSC _ 10150 _ 1: 1 mature specimen stained in paracarmine and whole mounted in Canada balsam, from type locality (20 June 2006). MHNUSC _ 10150 _ 2: 1 mature specimen stained in paracarmine and whole-mounted in Canada balsam, LES 107 Regosse cave (161 V / VR), Regosse, Lessinian Massif, 11.04847, 45.60577 (22 August 2001); MHNUSC _ 10150 _ 3: 1 mature specimen stained in haematoxylin and whole-mounted in Canada balsam, LES 036 Fumane-Molina interstitial, Lessinian Massif, 10.91867, 45.61550 (August 2001); MHNUSC _ 10150 _ 4: 1 mature specimen stained in haematoxylin and whole-mounted in Canada balsam, LES 124 Verona – Montorio via del Lanificio interstitial, Lessinian Massif, 11.06650, 45.45950 (August 2001). Leg. Fabio Stoch and Diana Galassi.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFAAFF99FF14F946FAA4FE7A.taxon	etymology	Etymology. The specific name refers to the lack of penial setae (Latin inermis, unarmed).	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFAAFF99FF14F946FAA4FE7A.taxon	description	Description. Mature worms up to 5 mm long. Number of segments 28 – 33. Diameter at segment V (slightly compressed specimens) 105 – 135 μm, at segment XI 100 – 160 μm. Clitellum 1 / 2 X – XII. Prostomium rounded, 45 – 65 μm long and 60 – 82 μm wide. Secondary annulation unnoticeable or slightly marked, up to V. Somatic setae bifid, 30 – 45 μm long, with upper tooth shorter and thinner than lower (Fig. 4 A); (2) 3 – 4 somatic setae per bundle anteriorly and 2 (3) in the clitellar and postclitellar segments (ventral setae absent in XI). Penial setae absent. One pair of male pores located in the posterior third of segment XI, in line with somatic ventral setae. One pair of spermathecal pores in segment X, placed in the most anterior part of the segment, in line with somatic ventral setae. Pharyngeal glands in segments IV – VI. Chloragogen cells beginning in VI. Oesophageal appendages and diverticula absent. One pair of testes in segment X. One pair of ovaries in segment XI. Sperm sacs extending by segments X – 1 / 2 XII. Egg sac extending to 13 / 14. Vasa deferentia about as long as atria, around 5 μm wide, entering the apical end of the atria. (Figs 3 A, 4 C [vd]). Atria tubular and curved, 115 – 140 μm long and 13 – 20 μm maximum diameter, with a thin muscular outer layer (1 – 2 μm thick); atrial lumen frequently containing spermatozoids; ectally, the atrium ends directly in a simple pore, surrounded by muscular strands, and can extrude somewhat as a small pseudopenis (Figs 3 A, 4 E [mp, ms]). Two roundish compact prostate glands per atrium, similar in size, the anterior one placed near the entrance of vas deferens, and the posterior one near the ectal end (Figs 3 A, 4 B – C). Spermathecae composed by a short conical muscular duct (13 – 20 μm long) dilated in a roundish vestibulum (16 – 30 μm in diameter) and followed by a long tubular ampulla (90 – 110 µm long, 13 – 22 µm wide) generally directed backwards in the segment (Figs 3 B, 4 D).	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFAAFF99FF14F946FAA4FE7A.taxon	distribution	Distribution and habitat. Caves and interstitial habitats in the western and southern parts of the Lessinian Massif (Progno di Fumane basin, Progno di Valpantena basin and Vaio di Squaranto basin), southernmost part of the Venetian pre-Alps, Italy. Alt. 71 – 700 m.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFAAFF99FF14F946FAA4FE7A.taxon	discussion	Remarks. Following Erséus (1992 a), the presence of elongated and curved atria, each with two prostate glands attached to proximal and distal ends, respectively, vasa deferentia entering apical ends of atria, slender spermathecae with short ducts, and small straight penial setae allocate the new species described above in the genus Gianius Erséus. The species most closed relative to Gianius inermis sp. nov. appears to be G. labouichensis (Rodriguez & Giani, 1989), described from the underground stream of Labouiche (Ariège, France). G. labouichensis is the only species of Gianius known until now with somatic setae in the ventral bundles of XI at maturity, instead of differentiated penial setae; the mature specimens of G. inermis sp. nov. are completely devoid of ventral setae in XI. The male ducts of G. labouichensis and G. inermis sp. nov. are similar in shape, but clearly smaller in the former (87 μm long, 7.5 μm in diameter). The location of the male pores in about the middle of the segment in G. labouichensis and in the posterior third of the segment in G. inermis sp. nov., and the absence in G. inermis sp. nov. of a dorsal appendage in the gut, present in G. labouichensis, are two other characters that clearly separate these two species.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA9FF97FF14FE1AFABFFDEB.taxon	description	(Figs 5, 6)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA9FF97FF14FE1AFABFFDEB.taxon	materials_examined	Holotype. MHNUSC _ 10148: mature worm stained in paracarmine and whole-mounted in Canada balsam. Leg. Ana I. Camacho. Type locality. CAN 072 Upwelling of the Clarión stream, Cantabria, Spain, – 3,54130, 43.34713 (13 Mars 2003).	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA9FF97FF14FE1AFABFFDEB.taxon	etymology	Etymology. The specific name refers to the small (Latin parvus) size of the genital organs (Latin genitalia).	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA9FF97FF14FE1AFABFFDEB.taxon	description	Description. Small worm, 1.8 mm long. Number of segments 26. Diameter at segment V 125 μm, at segment XI 175 μm. Clitellum 1 / 2 X – XII. Prostomium rounded, 45 μm long. Secondary annulation up to VI. Somatic setae bifid, 18 – 32 μm long, with upper tooth equal or slightly thinner and longer than lower (Fig. 6 B); 3 – 5 setae per bundle in segments II – VII and 2 thereafter. Penial setae bifid, with a minute upper tooth, 2 – 3 setae per bundle, about 22 μm long and 1.5 μm thick (Figs 5 A – B, 6 D [ps]). One pair of male pores located in the posterior third of segment XI, in line with somatic ventral setae. One pair of spermathecal pores in segment X, placed anteriorly in the segment, slightly lateral with respect to the alignment of the ventral somatic setae. Pharynx with a well-developed pharyngeal pad. Pharyngeal glands in segments IV – VII. Chloragogen cells beginning in VI. Oesophageal appendages and diverticula absent. One pair of testes in segment X. One pair of ovaries in segment XI. Sperm sacs not observed. Egg sac extending to 12 / 13. Vasa deferentia coiled, 4 – 5 μm wide, about as long as the atria (Figs 5 A, 6 C, E [vd]). Atria cylindrical and curved (C-shaped), about 50 μm long and 7 – 8 μm in diameter, with a thin muscular outer layer (about 1 μm thick); ectally, the atrium ends directly in the tegument, surrounded by a thin circular muscular layer (Figs 5 A, 6 C – E [a]). Two prostate glands per atrium, the anterior one placed near the entrance of the vas deferens, and the posterior one at the ectal part of the atrium, close to the penial setae (Figs 5 A, 6 C – E [pr 1, pr 2]). Spermathecae composed of a short duct (22 μm long, 11 μm wide) and a thin-walled, pyriform, slender ampulla (48 µm long, up to 14 µm wide), filled with spermatozoa (Figs 5 A, 6 F [sd, sa]).	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA9FF97FF14FE1AFABFFDEB.taxon	distribution	Distribution and habitat. Upwelling of the Clarión stream, Cantabria, Spain. Alt. 57 m.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA9FF97FF14FE1AFABFFDEB.taxon	discussion	Remarks. Following Erséus (1992 a), the presence of elongated and curved atria, each with two prostate glands attached to proximal and distal ends, respectively, vasa deferentia entering apical ends of atria, slender spermathecae with short ducts, and small straight penial setae allocate the new species described above in the genus Gianius Erséus. The narrow cylindrical C-shaped atria, only slightly tapering towards the ends, its small size (only 50 μm long and 7 – 8 μm in diameter), and the presence of bifid penial setae in low number (only 2 – 3 per bundle) are distinctive characters of Gianius parvogenitalis sp. nov. with respect to all the other known species in the genus.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA7FF97FF14FD6BFA2BFC1E.taxon	type_taxon	Type species. Cantabrodrilus pilarae Martínez-Ansemil sp. nov.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA7FF97FF14FD6BFA2BFC1E.taxon	diagnosis	Diagnosis. Freshwater subterranean phallodrilines. Somatic setae bifid. Penial setae present, single pointed hook-shaped. Coelomocytes absent. Spermathecal pores in segment X. Vasa deferentia narrow, entering apical ends of atria. Atria tripartite, elongate; ental part strongly muscular, middle part tubular U-shaped, ectal part thin-walled. Two large prostate glands per atrium; the anterior one attached near entrance of vas deferens and the posterior one at ectal end of atrium.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA7FF97FF14FD6BFA2BFC1E.taxon	etymology	Etymology. The name Cantabrodrilus means that the type species is a “ worm ” (Greek drilos) from Cantabria.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA7FF92FF14FBFFFBF2FE06.taxon	description	(Figs 7, 8)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA7FF92FF14FBFFFBF2FE06.taxon	materials_examined	Holotype. MHNUSC _ 10151: mature worm, incomplete (broken after the setae of segment XII), unstained and whole-mounted in Canada balsam. Leg. Ana I. Camacho. Type locality. CAN 177 Latarmá stream near Toyu cave, Venta Fresnedo, Cantabria, Spain, – 4.48694, 43.28030 (3 September 2002).	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA7FF92FF14FBFFFBF2FE06.taxon	etymology	Etymology. Named after Professor Pilar Rodriguez (University of the Basque Country, Spain), who contributes greatly to the knowledge of the oligochaetes in inland waters.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA7FF92FF14FBFFFBF2FE06.taxon	description	Description. Small worm, about 80 µm in diameter at segments V and XI (specimen broken after the setae of segment XII). Clitellum unnoticeable. Prostomium short (28 µm long), rounded. Secondary annulation not observed. Somatic setae bifid, 20 – 30 μm long, with short teeth, upper slightly shorter than lower, 2 – 3 somatic setae per bundle (Fig. 8 A); 5 single pointed hook-shaped penial setae per bundle in segment XI, about 20 μm long (Figs 7 B, 8 E [ps]). One pair of male pores located in the posterior third of segment XI, in line with somatic ventral setae. One pair of spermathecal pores in segment X, surrounded by a glandular area, in the lateral anterior part of the segment (Fig. 8 B, D [ga]). Pharyngeal glands in segments III – V. Chloragogen cells beginning in VI. One pair of testes in segment X. One pair of ovaries in segment XI. Sperm sacs entering the posterior part of segment IX. Vasa deferentia short and twisted, 4 – 5 μm wide, entering apical ends of atria (Fig. 7 A [vd]). Atria elongated, divided in three clearly differentiated parts. The ental part consist in a long, sigmoid, spindle shaped and heavily muscular structure (75 – 85 μm long, 13 μm in maximum diameter, muscular wall up to 4 μm thick) (Figs 7 A, 8 F [ma, m]), the middle part is a simple short curved U-tube (about 5 μm in diameter) connecting the ental and the ectal part (Fig. 7 A [ta]), and the latter is a pyriform, thin-walled and granulated ampulla (60 – 75 μm long, 20 – 22 μm in maximum diameter, muscular wall about 2 μm thick), which ends in a slightly curved and gradually narrowing ejaculatory duct (about 15 μm long) provided with a thin layer of circular muscles (Figs 7 A, 8 C, E [pa, ad]). Two large prostate glands per atrium, one attached to the anterior end of its muscular ental part, and the other to the posterior end of the ectal ampulla (Figs 7 A, 8 B – C, F [pr 1, pr 2]). Spermathecae composed by a tubular duct (36 – 40 μm long), and a very long, thin-walled ampulla (115 – 130 μm long, 20 μm in maximum diameter) directed backwards all along the segment X (Figs 7 C, 8 B [sd, sa]).	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA7FF92FF14FBFFFBF2FE06.taxon	distribution	Distribution and habitat. Hyporheic flow of the Latarmá stream, a tributary of the Lamasón stream at the left side of the Nansa stream basin, Cantabria, Spain. Alt. 180 m.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA7FF92FF14FBFFFBF2FE06.taxon	discussion	Remarks. The new species described here is a typical member of the Phallodrilinae, a characteristic subfamily of Naididae that, according to Erséus (1992 a) has two important autapomorphies: the absence of coelomocytes and the presence of two compact prostate glands attached to each atrium, one to anterior face, another to posterior face of latter. Cantabrodrilus pilarae gen. nov., sp. nov. is the only phallodriline with tripartite atria. The highest degree of complexity observed until know in the atria of the Phallodrilinae was the bipartite condition of the atria of Nootkadrilus Baker, 1982 (atria elongate, bipartite and folded at middle), with which C. pilarae sp. nov. also shares the shape of the penial setae. At first glance, the well set-off U-tube connecting the ental and the ectal part of the atria in Cantabrodrilus could be interpreted as an evolved character with respect to the bipartite atria of Nootkadrilus, but the latter is an atypical genus among the Phallodrilinae, living in the Northwest Pacific Ocean, and defined not only by the bipartite condition of the atria but also by the following apomorphies: absence of anterior prostate gland in the atria, presence of thickened and heavily granulated walls, resembling (and functioning as?) prostate glands, or diffuse prostate-like cells in the ental part of the vasa deferentia (see Erséus 1992 a).	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA2FF90FF14FDC7FD6AF96A.taxon	description	(Figs 9, 10) Abyssidrilus sp. 1 in Giani et al. (2011)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA2FF90FF14FDC7FD6AF96A.taxon	materials_examined	Holotype. MHNUSC _ 10144: mature worm, incomplete (broken at segment XVII), stained in paracarmine and whole-mounted in Canada balsam. Leg. Boris Sket. Type locality. Cave 4101, Brje pri Koprivi, Sežana, Slovenia, 13.86220, 45.77766 (December 1979). Paratypes. MHNUSC _ 10145 _ 1 – 4: 3 mature specimens, unstained and whole-mounted in Canada balsam; 1 immature specimen, stained in paracarmine and whole-mouted in Canada balsam. All from type locality (December 1979). Leg. Boris Sket.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA2FF90FF14FDC7FD6AF96A.taxon	etymology	Etymology. Named in honor of the late Professor Boris Sket (University of Ljubljana, Slovenia), who made outstanding contributions to the knowledge and development of biospeleology.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA2FF90FF14FDC7FD6AF96A.taxon	description	Description. Small worms, 3.2 mm long and 32 segments (complete paratype specimen). Diameter at segment V (slightly compressed specimens) 150 – 185 μm, at segment XI 175 – 195 μm. Clitellum hardly noticeable, extending over 1 / 2 X – XII. Prostomium rounded, 60 – 62 μm long. Secondary annulation at segments II – V. Somatic setae 32 – 50 μm long, 3 – 5 (6) in the preclitellar bundles and 2 – 5 per bundle in clitellar and postclitellar ones; ventral setae absent in XI in mature specimens; preclitellar dorsal setae and ventral setae until about segment XXV bifid, with upper tooth slightly longer than or as long as lower (Fig. 10 A, C – D); a mixture of single-pointed and bifid setae with upper tooth reduced in the following one or two segments; thereafter the bundles are composed exclusively of single pointed sigmoid setae (Fig. 10 B). Penial setae absent. One pair of male pores located in two conical protuberances (porophores) in the posterior third of segment XI, in line with somatic ventral setae (Fig. 10 E – F, H [mp, po]); each porophore has a large ring of glandular cells surrounding the male pore (Fig. 10 F g). One pair of spermathecal pores in segment XII, surrounded by a small circle of glandular cells, and placed anteriorly in the segment, in a lateral position slightly ventral with respect to the lateral lines (Figs 9, 10 H – G [sp, g]). Pharyngeal glands in (III) IV – VI. Chloragogen cells beginning in VI. One pair of testes in X. Anterior and posterior sperm sacs extending to IX and XI, respectively. One pair of ovaries in XI. Egg sac not developed, one mature egg present in the ovarian segment. Vasa deferentia shorter than atria, 5.5 – 7 μm in diameter, entering apical end of atria (Figs 9, 10 J [vd]). Atria tubular, comma-shaped, 205 – 215 μm long, 16 – 20 μm wide, strongly muscular (muscular layer 5 – 7 μm thick) (Figs 9, 10 H, K [a]). Two prostate glands per atrium. The anterior prostate is large, compact and lobed and is attached to the apical end of the atrium (Figs 9, 10 J [pr 1]), and the posterior prostate consists in a small group of cells, hardly visible, connected by a few strands with the ectal end of the atrium (or with the porophore?) near the male pore (Figs 9, 10 I [pr 2]). Spermathecae elongated, confined to XII, each consisting of a short conical duct (about 45 μm long), curved at base, and dilating in a club-shaped ampulla filled with spermatozoids; length of the ampulla about 75 μm long, maximum width 33 μm (Figs 9, 10 E [sd, sa]).	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA2FF90FF14FDC7FD6AF96A.taxon	distribution	Distribution and habitat. Cave in the Slovenian coastal karst, Sežana, Slovenia. Alt. 290 m.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFA2FF90FF14FDC7FD6AF96A.taxon	discussion	Remarks. The genus Aberrantidrilus Martin was erected to include a new groundwater species of Phallodrilinae, A. stephaniae Martin, 2015, from the Mercantour National Park, France, and the only two groundwater species of Abyssidrilus Erséus known until then, A. cuspis (Erséus & Dumnicka, 1988), from Umbria, Italy and A. subterraneus (Rodriguez & Giani, 1989), from Cantabria, Spain (Martin et al. 2015). The decision to erect this genus was supported by “ a unique combination of characters rather than formally proposed apomorphies ”. Thus, these three species share the “ aberrant ” position of the spermathecal pores (in the postatrial segment XII), the presence of a single penial seta per bundle oriented in anterior direction, the atria covered by a thick muscular layer and opening to the exterior on conical male protuberances (porophores), and the small size of the posterior prostate glands. A fourth species of Aberrantidrilus, A. mihaljevici Vučković, Rodriguez & Kerovec, 2020 was recently described from the tufa barrier of Roški slap, Croatia (Rodriguez et al. 2020); the authors remark that this species shows all the abovementioned characters except the thick muscular layer of the atria and, perhaps also, the presence of posterior prostate glands (“ not seen, probably due to its small size ”). The new species described here is tentatively ascribed to the genus Aberrantidrilus. The only major character by which A. sketi sp. nov. stands out in the genus is the lack of penial setae, but this is probably an unstable character in the genus as the absence of penial setae was observed also in one mature specimen of A. stephaniae (Martin et al. 2015) and “ maybe ” also in the specimens of A. cf. cuspis cited from Trieste, Italy (Sambugar et al. 1999). The tubular, narrow, comma-shaped and strongly muscular atria, the characteristically lobed anterior prostate glands and the thin and long spermathecal ducts clearly separate A. sketi sp. nov. from all the other species of the genus; the presence of glandular cells surrounding the spermathecal pores is also noteworthy, as similar structures or true glandular tissues associated or close to spermathecal pores are also present in Abyssidrilus opulentus Erséus, 1992 and Abyssidrilus potens Erséus, 1992 (Erséus 1992 a). As pointed out by Martin (in Martin et al. 2015), the specimens tentatively identify by Sambugar et al. (1999) as A. cuspis or A. cf. cuspis from Italy, France and Slovenia can be considered as a complex of cryptic species. Aberrantidrilus can be regarded now as a diversified groundwater genus of Phallodrilinae from the karst of southern Europe.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBAFF8AFF44FE3EFA2CFDCE.taxon	distribution	Romana Cave, Venezia Giulia, Italy and Škocjanske Cave, Slovenia (Sambugar et al. 1999) [see comments above]	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBAFF8AFF44FEAEFD2CFE06.taxon	description	Umbrian wells near Perugia, Italy (Erséus & Dumnicka 1988; Dumnicka 1990); Bocca Lupara Cave, La Spezia, Liguria, Italy, Papipano Cave, Udine, Friuli, Italy and Aven de la Cuisinière Cave, Calcomier, Aveyron, France (Sambugar et al. 1999) [see comments above]	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBAFF8AFF44FDC6FC93FDB6.taxon	distribution	Tufa barrier of Roški slap, Croatia (Rodriguez et al. 2020)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBAFF8AFF44FD8EFC95FD7E.taxon	distribution	Cave 4101, Brje pri Koprivi, Sežana, Slovenia (this work)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBAFF8AFF44FD56FA20FD25.taxon	distribution	Hyporheic zone in alluvial aquifer at Bévéra River, Sospel, Mercantour National Park, France (Martin et al. 2015)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBAFF8AFF44FD1EFDF3FC89.taxon	distribution	Jivero 1 Cave at Ozana, Matienzo, Cantabria, Spain (Rodríguez & Giani 1989); Alisas – Ramales karst, Matienzo, Cantabria, Spain (Achurra et al. 2015)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBAFF8AFF44FC4AFB91FBC5.taxon	description	Karstic spring of Argatxa, Bizkaia, Spain (Giani & Rodriguez 1988); Santa Eufemia – Ereñozar, Bizkaia, Spain, Llanes – Ribadesella and Gijón – Villaviciosa, Asturias, Spain (Achurra et al. 2015)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBAFF8AFF44FBFEFC92FBAA.taxon	distribution	Mineral spring in Charco Verde, Isla de la Palma, Canary Islands (Erséus, 1992 b). A. cuneus is a species commonly found in marine intertidal environments around the world.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBAFF8AFF44FB62FF6FFB3E.taxon	distribution	Budjur Mar Cave (pool area of a stream, 25 m asl), Leeuwin – Naturaliste Ridge, Western Australia (Pinder et al. 2006)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBAFF8AFF44FB16FD53FAAE.taxon	distribution	Anchyaline karst window about 500 m from the sea at Cape Range, Western Australia (Pinder et al. 2006). A. parvithecatus is a circumtropical species, also found in brackish anchialine waters in the Cabo Verde Islands (Erséus 1992 b) and frequent in littoral marine environments.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBAFF8AFF44FA66FEC7FA32.taxon	distribution	Tombstone Rocks Cave (deep pool of freshwater about 9 km from the ocean), Nambung region, Western Australia (Pinder et al. 2006)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBAFF8AFF44FA0AFCE5F9FA.taxon	distribution	Spring of the river Tione, Verona, Italy (Sambugar et al. 1999)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBAFF8AFF44F9D2FC69F9A2.taxon	distribution	Exsurgence Aïn Maarouf, El Hajeb, Morocco (Martin & Boughrous, 2012)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBAFF8AFF44F922FC48F8D2.taxon	description	Latarmá stream near Toyu cave, Venta Fresnedo, Cantabria, Spain (this work)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBAFF8AFF44F8B2FCCEF842.taxon	description	Göksu stream, West-Central Anatolia, Turkey (Arslan et al. 2007)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBBFF8BFF44FF5EFCCEFF2F.taxon	distribution	Göksu stream, West-Central Anatolia, Turkey (Arslan et al. 2007)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBBFF8BFF44FEE6FC5DFE06.taxon	distribution	Several localities along the lower part of the River Weser, Germany (Hrabĕ 1960); Mairuelegorreta Cave at Zigoitia, Alava, Spain, Jivero 1 Cave at Ozana, Matienzo, Cantabria, Spain and Labouiche Cave, Vernajoul, Ariège, France (Rodriguez & Giani 1989); Tonawanda Channel, Niagara River, North America (Farara & Erséus 1991); Santa Eufemia – Ereñozar, Bizkaia (Achurra & Rodriguez 2008 and Achurra et al. 2015); Itxina, Gorbeia, Bizkaia, Spain, and Alisas – Ramales karst, Matienzo, Cantabria, Spain (Achurra et al. 2015)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBBFF8BFF44FE3EFC37FDCF.taxon	distribution	Several groundwater environments in Cantabria, Spain (this work)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBBFF8BFF44FDC6FAB5FDB7.taxon	distribution	Le Trou des Voleurs Cave, near Poncin, Southern Jura, France (Juget & Creuzé des Chȃtelliers 2001)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBBFF8BFF44FD8EFBB4FD7F.taxon	distribution	Orillon Cave at Ozana, Matienzo, Cantabria, Spain (Rodriguez & Giani 1989)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBBFF8BFF44FD56FC59FD27.taxon	distribution	Several groundwater environments in the Lessinian Massif, Italy (this work)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBBFF8BFF44FD1EFC1CFCEF.taxon	distribution	Labouiche Cave, Vernajoul, Ariège, France (Rodriguez & Giani 1989)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBBFF8BFF44FCA6FCE9FC57.taxon	distribution	Ojo Guareña Cave, Burgos, Spain (Rodriguez & Achurra 2010)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBBFF8BFF44FC6EFCFBFC1F.taxon	distribution	Upwelling of the Clarión stream, Cantabria, Spain (this work)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBBFF8BFF44FC36FF6FFB8F.taxon	distribution	Hyporheic flow of the river Tambre, Galicia, Spain (Giani & Martínez-Ansemil 1981); hyporheic flow of the river Dordogne, France (Lafont 1989); underground water at Ile du Grand Gravier, river Rhône (Lafont & Durbec 1990)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBBFF8BFF44FB43FA24FB33.taxon	distribution	Anchialine groundwater (well about 500 m from the ocean) at Cape Range, Western Australia (Pinder et al. 2006)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBBFF8BFF44FAD7FB6BFAA6.taxon	description	Lake Baikal, southern Siberia, Russia (Martin & Brinkhurst 1998), as Rhyacodriloides gladiiseta.	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBBFF8BFF44FA27FBBCF9D7.taxon	description	Inland anchialine Prospero Cave, near Castle Harbour, Bermuda (Erséus 1986)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBBFF8BFF44F9B7FD4BF92B.taxon	description	Salt spring near Zadar, Dalmatia, Croatia (Hrabĕ 1973); Jama pri ogradi, Gangarol island, Murter, Croatia (Martínez-Ansemil et al. 2016); freshwater (probably influenced by sea water) at the mouth of a small stream, Tjärnö, Sweden (Erséus et al. 1992). S. modricensis is also present in marine littoral sandy habitats, and is qualify as a probably oligohaline brackish-water species (Erséus et al. 1992)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFBBFF8BFF44F8E3FC36F82F.taxon	distribution	Hyporheic and phreatic waters at the alluvial plain of the river Rhône, upstream from Lyon, France (Juget 1987); hyporheic flow of the river Neste d’Aure, Hautes-Pyrénées, middle reach of the river Dordogne, and Labouiche Cave, Vernajoul, Ariège, France (Erséus et al. 1992); hyporheos of the Lachein creek, Alas, Ariège (Route et al. 2004); alluvial aquifer at Forez plain, catchment of the river Loire, France (Paran et al. 2005) [more detailed data of the French sites are given in Artheau & Giani (2006)]; spring v Dragi 1, spring at Resnik, spring Malence and hyporheal of the stream Borovniščica, Slovenia (Giani et al. 2011)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
A50C0231FFB8FF88FF44FF5EFC13FF2E.taxon	description	Saint Lawrence Great Lakes, North America (Cook & Hiltunen 1975)	en	Martínez-Ansemil, Enrique, Sambugar, Beatrice (2025): Inventory of the inland-water Phallodrilinae (Clitellata: Naididae), with descriptions of a new genus and five new species from groundwaters of southern Europe. Zootaxa 5692 (3): 446-468, DOI: 10.11646/zootaxa.5692.3.2, URL: https://doi.org/10.11646/zootaxa.5692.3.2
