identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03A04148B133FE040BC5F8ECFE1FF5E9.text	03A04148B133FE040BC5F8ECFE1FF5E9.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Trichodrilus Claparede 1862	<div><p>Genus Trichodrilus Claparède,1862</p><p>Lumbriculids with two pairs of testes and one pair of ovaries. One pair of male ducts open typically in X, with a semiprosoporous organization, i.e. each atrium receives one pair of vasa deferentia which run independently from two sperm funnels serving separately testes in the pre-atrial and atrial segments. One pair of spermathecae opening in the ovarian segment; some species also have one additional pair in the post ovarian segment and one species with spermathecae in 4 postatrial segments.</p></div>	https://treatment.plazi.org/id/03A04148B133FE040BC5F8ECFE1FF5E9	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Rodríguez, Pilar;Fend, Steven	Rodríguez, Pilar, Fend, Steven (2025): Contributions to the knowledge of the genus Trichodrilus Claparède (Lumbriculidae, Clitellata) with the description of new groundwater species from Spain. Zootaxa 5711 (1): 57-78, DOI: 10.11646/zootaxa.5711.1.2, URL: https://doi.org/10.11646/zootaxa.5711.1.2
03A04148B132FE010BC5FF75F82EF0E2.text	03A04148B132FE010BC5FF75F82EF0E2.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Trichodrilus benati Rodríguez & Fend 2025	<div><p>Trichodrilus benati sp. nov.</p><p>(Figures 1, 2, Table 2)</p><p>Holotype: MNCN 16.03 /3201, 1 mature, mated individual, dissected, stained in haematoxylin and mounted in Canada balsam. 8 December 1987.</p><p>Paratypes: MNCN 16.03 /3202, 1 mature, dissected individual (sperm in sperm funnels, but spermathecal ampullae filled with undetermined mass) . MNCN 16.03 /3203, one mated and dissected individual . MNCN 16.03 /3204, one mated, whole-mounted individual. All from the type locality. 8 December, 1987 .</p><p>Type locality: <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-2.756745&amp;materialsCitation.latitude=43.022705" title="Search Plazi for locations around (long -2.756745/lat 43.022705)">Mairulegorreta Cave</a>, Gorbeia (Itxina Karst), Zigoitia, Alava, Spain (P. Rodriguez leg.). Coordinates N43.022706, W2.756745, 900 m altitude. Sampled at the headwaters of the cave river (Table 1) .</p><p>Etymology: For Beñat Zaldibar, in appreciation for his expert collaboration on several occasions in the histology of the species.</p><p>Other material (in first author’s collection):</p><p>— Mairulegorreta Cave, one mature, mated, whole-mounted individual, from type locality. 8 December 1987, at the resurgence .</p><p>—Azoleta Cave, 1 mature, mated individual, dissected. 10 March 1985. (P. Rodriguez leg.).</p><p>— Arandilla R. source, 5 individuals, mated, whole mounts, 10 September 2023 (P. Rodriguez leg.) .</p><p>Description (based on type locality population): Specimens incomplete, without posterior end, 15–25 segments, body diameter 0.26–0.35 mm in segment X. Prostomium conical. Secondary annulation from segment III. Anterior ventral chaetae shorter in II (70–89 µm), and gradually longer (to 96–128 µm), most posterior chaetae down to 90 µm; nodulus at 0.36–0.44 distance from the tip relative to total chaeta length. Anterior dorsal chaetae 0.8–1 the length of ventral chaetae in the same segment. Clitellum from (posterior part of IX) X to XII (or anterior part of XIII); cells do not show any particular order (Fig. 1B). Male pores open in conical porophores, located very posterior in segment X, in line with ventral chaetae and directed backwards (Fig. 1 D, E). Spermathecal pores in segments XI and XII, behind and in line with ventral chaetae. Female pores in 11/12.</p><p>Pharynx well developed dorsally in II and III (Fig. 1A). Pharyngeal glands from IV (only dorsally) to VII or VIII (only ventrally). First nephridia in segment VII, absent in IX to XI and present again from segment XII (Fig. 1C). Sperm sac back to XII (no egg sac formed).</p><p>Paired testes in IX and X, one pair of ovaries in XI. Semiprosoporous male duct. Atrium oval, gradually narrowing toward the pore, continuous with the conical duct, which, however, is well separated histologically from the ampulla (Figs. 1E, 2). Total length of atrium 120–144 µm (c. 0.5 the body diameter), maximum diameter 56–68 µm, atrial duct 55–58 µm long and 16–24 µm wide, forming a short penis with the ectal end lined by cuticle (Fig. 1D, E). Muscular layer of atrial ampulla 5–10 µm thick; glandular atrial epithelium compact (12–18 µm) with nuclei, and a narrow lumen; prostate cells organized in clusters to 54–73 µm high. One pair of vasa deferentia (10–22 µm Ø) joins the apical part of each atrial ampulla; the posterior one crosses into segment XI.</p><p>Paired spermathecae in segments XI and XII, sac-like ampulla (90–280 µm long x 45–74 Ø) and a short duct (40–92 µm long x 10–21 µm Ø) with a small ectal vestibule (17–34 µm Ø) (Fig. 1F). First spermathecal ampulla larger than the second one; ampullae can cross into following segment.</p><p>The Azoleta Cave specimen differs from those of the type collection mainly by the larger dimensions of the worm and, in particular, of the atrium (see Table 2). The Arandilla River population has been provisionally ascribed to T. benati sp. nov. due to the general morphology of the male duct (Fig. 1G, H), the body diameter (0.24–0.27 mm), and the atrium total dimensions (105–135 µm long x 54–72 µm Ø), although the duct is shorter. The main differences are: smaller chaetae (37–67 µm long) and a thinner atrial musculature (2–4 µm).</p><p>Remarks: The apical junction of the vasa deferentia to the atrium is one of the most distinct characteristics of the new species. This character is also known in T. intermedius (Fauvel, 1903) from a well in northwestern France, a species that is much larger than T. benati n.sp., with posterior lateral blood vessels, a tubular atrium and large, partially muscular vasa deferentia (for comparison, see Table 2). Hrabě (1937) re-examined a specimen of Trichodrilus intermedius, probably a syntype from Fauvel’s type collection; it was histologically sectioned and deposited in the National Museum of Prague.</p><p>The species T. benati sp. nov. is most similar to T. medius Hrabě, 1960 . Atria of both species have a similar shape, and in both cases the ampulla gradually narrows toward the duct, i.e., the atrium is not petiolate, but the duct is well distinguished histologically from the ampulla. The most peculiar character in the description of T. medius was the very apical junction of the posterior vas deferens, while the anterior one joins at “ a quite considerable distance ” (sic). The study of the type species from the NMP has allowed us to do new measurements (Table 2) and photos. The transverse section in Hrabě´s (1960) fig. 24 (see Fig. 3A) is found on the slide Hr 1517-30 (#9), and we interpret it as misleading, since that atrial ampulla section does not correspond with the middle section of the atrium, but it is somewhat lateral. In fact, the sagittal sections in the slide Hr1517-9 (#1, #2)) show the vasa deferentia joining subapically to the atrial ampulla (Fig. 3B, E). The duct in T. medius is longer than the ampulla, and some individuals have a salient penis, formed by the inner cells of the duct (type-2 penis: Rodriguez &amp; Giani, 1994) (Fig. 3C, D) while T. benati sp. nov. the duct is shorter and the ampulla and has a type-1 penis, i.e., formed by the protrusion of the atrial duct. Spermathecal ampullae in T. medius are larger than in T. benati (Fig. 3F).</p><p>Another five Trichodrilus species with 2 pairs of spermathecae and without lateral posterior blood vessels have been described as having apical or subapical vasa deferentia junctions. Among these species, T. tacensis Hrabě, 1963 clearly differs from T. benati sp. nov. by the tubular atrial ampulla, and the posterior vasa deferentia not penetrating the post-atrial segment. In the other four species, the atrial ampulla is oval or spherical, but they are mainly distinguished from T. benati sp. nov. by the following characters: T. angelieri Giani &amp; Rodriguez, 1994 has very distinct epidermal glands and the atrium has a simple, cylindrical ejaculatory duct; T. claparedei Hrabě, 1938 and T. seirei Timm, 1979 (based on the holotype) are mainly distinguished by the spherical shape of the atrial ampulla and by the thick atrial musculature (up to more than 23 µm), and T. hrabei Cook, 1967 has a small (80 µm long) atrium, with an unusual, laterally-compressed shape, and without a well-defined atrial duct.</p><p>Habitat and distribution. Found in cave streams and karst springs in northern Spain.</p></div>	https://treatment.plazi.org/id/03A04148B132FE010BC5FF75F82EF0E2	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Rodríguez, Pilar;Fend, Steven	Rodríguez, Pilar, Fend, Steven (2025): Contributions to the knowledge of the genus Trichodrilus Claparède (Lumbriculidae, Clitellata) with the description of new groundwater species from Spain. Zootaxa 5711 (1): 57-78, DOI: 10.11646/zootaxa.5711.1.2, URL: https://doi.org/10.11646/zootaxa.5711.1.2
03A04148B136FE0D0BC5FCA0F969F1B6.text	03A04148B136FE0D0BC5FCA0F969F1B6.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Trichodrilus tenuis , Hrabe 1960	<div><p>Trichodrilus tenuis Hrabě, 1960</p><p>(Figures 4, 5; Table 3)</p><p>Trichodrilus tenuis Hrabě, 1960: 271, figs. 26–28. Wachs 1967: 234; Juget &amp; Dumnicka 1986: 240; Rodriguez &amp; Giani 1994: 36, table 2; Juget &amp; des Chatelliers 2001: 23, fig. 4; Achurra &amp; Rodriguez 2008: 165, table 1; Achurra et al. 2015: 159, table 1; Des Chatelliers et al. 2009: 685; Giani et al. 2011: 91, table 1; Rodriguez-Noriega 2013: 166, table 5.3.6.1.c; Dumnicka 2014: table 2; Camacho &amp; Puch 2021: 208.</p><p>Trichodrilus cf. tenuis Hrabě: Martin et al. 2015: 564.</p><p>Material from Spain examined (see Table 1 for locality data) (first author’s collection):</p><p>—Argatxa spring, Bizkaia (18 December 1984, P. Rodriguez leg.) 1 mated, dissected individual.</p><p>—Goikoetxe Cave, Bizkaia (31 January, 2016, P. Rodriguez leg.) 1 mated, whole-mounted individual.</p><p>— Escalón Cave, Cantabria (18 September, 2008, A. Achurra leg.) 1 partly mature, dissected individual .</p><p>— La Cullalvera Cave, Cantabria (28 March 1988, A. Camacho leg.) 1 mated, dissected individual .</p><p>— Source of the Bustablado River, Cantabria, (14 March, 2003, Ana Camacho leg., donated by E. Martinez-Ansemil) 2 mated, dissected individuals .</p><p>— El Molino Spring, Bustablado, Cantabria (14 March, 2003, Ana Camacho leg., donated by E. Martinez-Ansemil) 1 mated, whole-mounted individual .</p><p>— La Coventosa Cave, Cantabria (1 April, 2003, Ana Camacho leg., donated by E. Martinez-Ansemil) 2 mated, whole mounted individuals .</p><p>—La Cubilla Cave (15 March, 2003, Ana Camacho leg., donated by E. Martinez-Ansemil) 1 whole-mounted individual with eggs.</p><p>— Rio Chico Cave, source of the Gandara River, Cantabria (5 June, 2003, Ana Camacho leg., donated by E. Martinez-Ansemil) 2 mated, whole-mounted individuals .</p><p>—Los Santos Cave (4 June 2003, Ana Camacho leg., donated by E. Martinez-Ansemil) 1 unmated, whole-mounted individual.</p><p>—Ojo Guareña Cave, Burgos (Pilar Rodriguez and Ana Camacho leg.) 5 dissected individuals, four of them mated and one with eggs, sampled in 18 March 1995, 3 November 1995, 30 August 2002, 22 November 2002, 24 April 2004.</p><p>— Well in Velamazan, Soria (13 September 1976, R. Rouch leg.) 1 whole-mounted and 2 dissected, mated individuals .</p><p>— Ginel R. source, La Magdalena Spring, Zaragoza (5 September 2024, Pilar Rodriguez leg.) 3 whole-mounted and 2 dissected, mated individuals .</p><p>—Well in Los Picos, Alicante (5 July 1986, A. Camacho leg.) 1 mated, dissected individual.</p><p>—Well in Zafra-Rio Bodión, Badajoz (3 November, 1988, A. Camacho leg.) 1 mated, whole-mounted individual.</p><p>—Well in Ermita San Isidro, Gibraleón, Huelva (4 November, 1988, A. Camacho leg.) 1 mated, dissected individual.</p><p>Description (measurements from Spanish mated individuals; those data marked with (a) are from a very small individual in Rio Chico Cave).</p><p>Number of segments 44 [Velamazan] and 60 [Ginel R. source] of the only two individuals found complete. Body diameter in X (0.14 (a)) 0.20–0.35 mm. Prostomium 84–150 µm long. Anterior secondary annulus beginning in segment III or IV (rarely in V) and extending throughout the body. Chaetae with nodulus slightly distal; in preclitellar segments chaetae 41–103 µm long in dorsal bundles, 61–107 µm long in ventral bundles, shorter in II, and dorsals about 0.7–0.9 times the length of ventrals of their respective segments; chaetae become shorter in posterior segments (to 80–90 µm) (measurements taken from two individuals).</p><p>All genital pores in the line of ventral chaetae. One pair of male pores in the posterior part of segment X, close to the septum 10/11, open on short, round to conical porophores (c. 25 µm high) (Fig. 4A–C). Two pairs of simple spermathecal pores in segments XI and XII, which open in the line of ventral chaetae, behind the ventral bundles. One pair of female pores, in 11/12.</p><p>Clitellum from X to XII (sometimes also in the anterior part of XIII, or extending through XIII). The pharyngeal glands present from IV or V to VII or the anterior part of segment VIII. First nephridium seen in 6/7, although fixation and orientation in whole mounts or damage during dissection makes it difficult to see this character in some individuals.</p><p>Paired testes in IX and X, one pair of ovaries in XI. Sperm sacs extend to XI– XIII; egg sacs to XII–XV. Male funnels on 9/10 and 10/11. Free portion of both anterior and posterior vasa deferentia 6–14 µm diameter, posterior vas deferens forms a loop in XI before passing to X; junction to the atrial ampulla median to subapical (Fig 4A–D). The ratio of total length of atrium to the body diameter in X is 0.22–0.43 (mean=0.27). Total length of atrium (50 (a)) 63–112 μm, and 32–75 μm wide; atrial ampulla spherical to oval; a short, broadly-conical atrial duct (23–42 µm long) narrows gradually to the male pore, and is about the length of the porophore. In some individuals, the atrial duct forms a short penis by the extrusion of the inner cells (type-2 penis: Rodriguez &amp; Giani 1994) that may protrude through the male pore (to 50 µm) (Fig. 4D). Epithelial cells of the ampulla of variable height (4–22 µm); atrial musculature very narrow (c. 2–3 µm). Apical part of the atrial ampulla covered by long prostatic cells (34) 54–104 µm high), usually forming long, tightly packed clusters.</p><p>Spermathecae paired in XI and XII, filled with unordered sperm. Spermathecal ampulla usually in the same segment as the corresponding spermathecal pore, although in several individuals some or all spermathecae may penetrate the following segment. The ental region of the ampulla is usually vacuolated and in some cases the sperm is clearly visible inside the vacuoles; it is usually narrower than the ectal section and it may be in the process of degenerating. Spermathecal ducts short (28–72 µm long with a narrow, ental neck (9–12 µm diameter), widening ectally to 21–27 µm (Fig. 4E). Spermathecal ampulla irregular, sac-like or tubular, well separated from the duct and variable in size (88–255 µm long, 30–120 µm wide); folded backwards or straight, in a dorsoventral orientation.</p><p>Remarks. Among the Trichodrilus species with 2 pairs of spermathecae, T. tenuis Hrabě, 1960 is one of the smallest worms, with a small atrium (usually about one third the body diameter or less), a thin atrial musculature (c. 2 µm) and a cover of high prostatic cells. In the present study, data on the morphology of the Iberian metapopulation are given for the first time, and the general morphological characteristics are compared with those in the type collection of the species (from the NMP) (Table 3). All characters in the Spanish metapopulation fit the dimensions of the type collection, except for the position of the pharyngeal glands. In the original description of T. tenuis, Hrabě discussed the difference of this species from similar Trichodrilus species ( T. moravicus Hrabě, 1937 and T. allobrogum Claparède, 1862). In addition to the small size, Hrabě highlighted that in T. tenuis the pharyngeal glands (“chromophile cells”) are restricted to segments IV– VI, while they extend to VIII in the other two species (Table 3). In the Spanish collection of T. tenuis, the position of the pharyngeal glands extends to VII or VIII. The examination of Hrabě’s collections of T. tenuis confirms that the position of pharyngeal glands is always in segments IV– VI. Cook (1967: 363) seemed not to be convinced of the taxonomic importance of this character and pointed that this is a variable character in other lumbriculids. In the same way, Rodriguez &amp; Giani (1994) reported some Trichodrilus species showing variability in the extension of the pharyngeal glands, e.g. T. macroporophorus Hrabě, 1954 (back to VII or VIII) or T. pragensis Vejdovský, 1876 (to VIII or IX). It is, however, consistent in the Czech populations of T. tenuis, and may be a useful morphological character in future taxonomic analyses of the species, based on molecular studies.</p><p>For the present identification of Trichodrilus tenuis, we emphasize characters of the reproductive system as seen in the type series: an oval, small atrial ampulla (Fig. 5A–C) where the vasa deferentia join in a median to subapical position (Fig. 5B–D), and a short, conical atrial duct, not petiolated (that is, duct is not well separated by a constriction from the ampulla), which opens in a small, round porophore (Fig. 5A). In some individuals a type-2 penis extends through the male pore (Fig. 5C, E), perhaps depending on the strength of the fixation, since it was also observed in an unmated individual. Spermathecae have a short duct, bulbous at the ectal end, and ampullae may be straight in the same segment as the pore, or bent and passing to the next segment (Fig. 5F).</p><p>The spermathecae of T. tenuis were described in detail by Juget &amp; des Chatelliers (2001) from a large collection of several hundred individuals from groundwaters in the region of Lyon, East France. These authors described “ a pseudovestibule with an inner epithelium of conspicuous high cells arranged in a circle around the zone of junction between the duct and the ampulla ”. In our collection, that part of the ampulla close to the duct has a columnar epithelium, and we don’t see anything resembling a vestibule. The more strongly stained (most recent) sperm cells are usually restricted to the ectal section of the spermathecal ampulla, while in French worms the sperm appears distributed along the odd ampulla, which has a thick wall with many constrictions (see Juget &amp; des Chatelliers 2001: fig. 4). In their description, Juget &amp; des Chatelliers also stated that the spermathecal ampullae were always in the same segment as the pore. However, in both Hrabě’s and the Spanish collection, the ampulla may cross to the following segment or not, and the spermathecal ampullae appear either standing up to the dorsal region of the body, as Juget &amp; des Chatelliers (2001) described, or folded on the ventral side toward the posterior part of the segment. All of these differences suggest that the French population of T. tenuis may be a different species.</p><p>Habitat and distribution. Groundwaters in the Iberian Peninsula host several populations which have been attributed in present study to the lumbriculid species T. tenuis Hrabě. These sites are up to 700 km distant within Spain and more than 1500 km from the southern and western regions of Germany, where the species was first reported (see Fig. 6). The morphospecies T. tenuis is Euro-Mediterranean. In Central Europe, the species distribution covers the High Rhine and Ruhr rivers in Germany (Juget &amp; Dumnicka 1986). In the Mediterranean region, it has already been reported in groundwaters of eastern France (Juget &amp; des Chatelliers 2001), Slovenia (Giani et al 2011), and Spain (wells in Velamazan, Soria: Rodriguez &amp; Giani 1994; Ojo Guareña Cave, Burgos: Rodriguez-Noriega 2013; cave rivers and springs in Bizkaia and Cantabria: Achurra et al. 2015). Unpublished records were also reported from wells in the Rif (Morocco) (Juget &amp; des Chatelliers 2001), although these were not included in the guide on the aquatic oligochaetes of Maghreb by Martin &amp; Aït Boughrous (2012). The present study extends the range of distribution of T. tenuis to practically the entire karstic carbonate region in the Iberian Peninsula (Fig. 6), and its habitat covers all types of groundwaters: caves, hyporheos and phreatic waters in wells. However, we are aware that future molecular studies may demonstrate the existence of cryptic speciation in this groundwater species.</p></div>	https://treatment.plazi.org/id/03A04148B136FE0D0BC5FCA0F969F1B6	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Rodríguez, Pilar;Fend, Steven	Rodríguez, Pilar, Fend, Steven (2025): Contributions to the knowledge of the genus Trichodrilus Claparède (Lumbriculidae, Clitellata) with the description of new groundwater species from Spain. Zootaxa 5711 (1): 57-78, DOI: 10.11646/zootaxa.5711.1.2, URL: https://doi.org/10.11646/zootaxa.5711.1.2
03A04148B13AFE0B0BC5FC14FE1FF68A.text	03A04148B13AFE0B0BC5FC14FE1FF68A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Trichodrilus stygodytes Rodríguez & Fend 2025	<div><p>Trichodrilus stygodytes sp. nov.</p><p>(Figures 7, 8D–H, Table 3)</p><p>Trichodrilus moravicus Hrabě, 1937 . Hrabě 1938: 73, fig. 1–8 (partim)</p><p>Holotype. MNCN 16.03 /3206, one mature, mated individual, stained and whole-mounted in Canada balsam.</p><p>Type locality. <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-2.59979&amp;materialsCitation.latitude=43.32556" title="Search Plazi for locations around (long -2.59979/lat 43.32556)">Omaerreka River</a> headwaters (below a resurgence), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=-2.59979&amp;materialsCitation.latitude=43.32556" title="Search Plazi for locations around (long -2.59979/lat 43.32556)">Kortezubi</a>, Bizkaia, Spain. 13 March 1985. Coordinates: N 43.32556, W 2.59979, 150m altitude .</p><p>Paratypes. MNCN 16.03 /3207, Omaerreka River source, 13 March 1985, 1 mature, mated individual, dissected, stained and mounted in Canada balsam . MNCN 16.03 /3208, Okamika Cave, at the cave resurgence, 27 December 2023, 1 mature, mated individual, dissected, stained and mounted in Canada balsam . MNCN 16.03 /3209, Goiketxe / Lamina Cave, 30 July 2025, 1 mature, mated individual, stained and whole-mounted in Canada balsam. (All sampled by Pilar Rodriguez) .</p><p>Other material in first author’s collection. One mature, mated individual from Goiketxe/ Lamina Cave (30 July 2025), stained and whole-mounted in Canada balsam. One mature individual from Lamiñak/Las Lamiñas Cave River (5 January 1986), stained and whole-mounted in Canada balsam.</p><p>Other material ascribed to the species. Hrabě’s collection in NMP under the name of Trichodrilus moravicus: Býčí Skála Cave, Blansko, Moravian Karst, Czech Republic (https://mapy.cz/s/robekerede): Hr 1007-1 a whole mount. Hr 1007–2 (I– VII) sagittal histological sections (# 88–14). Hr 1026 transverse sections (#25, 27, 31) and sagittal sections (#24, 28, 30), and one whole mounted immature individual (#29). Hr 1028 sagittal sections (#36) and 5 whole-mounted individuals (#38).</p><p>Etymology. The Latin species epitheton stygodytes is derived from ancient Greek and it can be interpreted as dweller (dytes) of subterranean river (stygos, underground river known as the boundary between the worlds of the living and the dead, in Greek mythology). Noun in apposition.</p><p>Description (based on material collected in Spain; for comparison with material from the Hrabě collection, see below). Very small worms, body diameter in X 0.17–0.30 mm; all incomplete (up to 40 segments), with missing posterior part of the body. Clitellum from X–XII (indistinct in some individuals). Secondary annulation from III or IV. One pair of male pores in X, behind and in line with ventral chaetae; one pair of female funnels in 11/12; two pairs of spermathecal pores, in XI and XII, behind and in line with the ventral chaetae. Male pores open on round porophores (15–20 µm high, 45–50 µm wide). One pair of chaetae per bundle, simple pointed; in the anterior body segments, dorsal chaetae c. 50–84 µm, ventral chaetae (55) 62–100 µm, dorsals 0.8–1 times smaller than ventral chaetae, nodulus distal, at 0.4 times the total length from the distal end.</p><p>Pharynx in segments II and III, with a well-developed dorsal pad (Fig. 7B). Pharyngeal glands to VIII. Sperm sacs back to XI or XII, eggs sac to XII or XIII. Two pairs of testes (segments IX and X) and one pair of ovaries (segment XI). Semiprosoporous male duct. Atrium petiolate, ampulla well separated from the duct (Fig. 7A), its total length 0.3–0.5 the body diameter. Atrium total length 80–121 µm; round to oval ampulla (50–76 µm long, 50–80 µm Ø). Atrial ampulla epithelium 4–16 µm high; ampullar musculature 2–8 µm thick, up to 10 µm at the vasa deferentia junction. Prostate glands are packed clusters of glandular cells to 47–80 µm high. The atrial duct (30–40 µm long, 17–30 µm Ø) can protrude a little through the male pore forming a penis (Fig. 7E). One pair of vasa deferentia (8–13 µm Ø) join the basal part of the atrial ampulla, close to the atrial duct (Fig. 7A, C, D); the posterior vas deferens crosses into segment XI.</p><p>Spermathecal ampullae round or elongate (c. 50–120 µm long, c. 50–70 µm Ø), remaining in the same segment or crossing to the next one. Short spermathecal ducts (40–70 µm long), narrow close to the ampulla (10–15 µm Ø), and with a distal widening forming a vestibule (21–30 µm Ø) (Fig. 7F).</p><p>Remarks. Among the Trichodrilus species with two pairs of spermathecae, Trichodrilus stygodytes sp. nov. is morphologically similar to other species of small size with a small, spherical to oval atrial ampulla, covered by a thin muscular layer (&lt;10 µm), with a short duct (see Annex). In particular, the new species resembles T. moravicus (see below). The main character that separates T. stygodytes from this group of species is the basal/ectal junction of the vasa deferentia to the atrial ampulla.</p><p>The comparison of the measurements of the Spanish population of T. stygodytes sp. nov. with the Czech collection from Cave Býčí Skála is shown in Table 3. The Spanish worms are smaller and with smaller chaetae than the Býčí Skála population. Both populations have similar characteristics of the male duct and share the distinctive basal junction of the vasa deferentia to the atrial ampulla (compare Figs. 7A, C, D and 8D–G). The spermathecae in the Czech population are somewhat larger than in the Spanish worms, and also show an ectal vestibule in the spermathecal duct (Fig. 8H).</p><p>Habitat and distribution. The habitat of the species includes cave streams and headwaters of karstic rivers. Spain and Czech Republic. Spanish populations are at more than 1600 km distance from the sites in the Czech Republic, and future molecular studies could demonstrate that they belong to different species. Among the relatively large collection from subterranean waters that we have studied in the Iberian Peninsula, we have found T. stygodytes sp. nov. only in a single karst area, while T. tenuis is a widespread species in subterranean waters, and T. moravicus (see below) has not been found, so far.</p></div>	https://treatment.plazi.org/id/03A04148B13AFE0B0BC5FC14FE1FF68A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Rodríguez, Pilar;Fend, Steven	Rodríguez, Pilar, Fend, Steven (2025): Contributions to the knowledge of the genus Trichodrilus Claparède (Lumbriculidae, Clitellata) with the description of new groundwater species from Spain. Zootaxa 5711 (1): 57-78, DOI: 10.11646/zootaxa.5711.1.2, URL: https://doi.org/10.11646/zootaxa.5711.1.2
03A04148B13CFE160BC5FB01F9C9F5B7.text	03A04148B13CFE160BC5FB01F9C9F5B7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Trichodrilus moravicus Hrabe 1937	<div><p>Trichodrilus moravicus Hrabě, 1937</p><p>(Figures 8A–C, Table 3)</p><p>Trichodrilus moravicus Hrabě, 1937: 3, 20. Hrabě 1938: 73, fig. 1–8 (partim); 1942: 28; 1960: 265; 1981: 107, 108, table 19(3). Dumnicka 1995:19; 2000: 359, 385, tables 7–10; 2014:18, tables 1 &amp; 2. Dumnicka &amp; Galas 2017: table 1. Dumnicka et al. 2020: 7, fig. 5, table 3.</p><p>Lectotype: NMP Trichodrilus moravicus Hr 1007 -2(IV) (#11) NMP P6 j-20/89/144, sagittal sections, mature, with egg sac.</p><p>Type locality: <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=16.74944&amp;materialsCitation.latitude=49.24444" title="Search Plazi for locations around (long 16.74944/lat 49.24444)">Říčka River</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=16.74944&amp;materialsCitation.latitude=49.24444" title="Search Plazi for locations around (long 16.74944/lat 49.24444)">Moravian Karst</a>, Czech Republic, about 10 km from <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=16.74944&amp;materialsCitation.latitude=49.24444" title="Search Plazi for locations around (long 16.74944/lat 49.24444)">Brno</a> (https://mapy.cz/ s/narozularu). <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=16.74944&amp;materialsCitation.latitude=49.24444" title="Search Plazi for locations around (long 16.74944/lat 49.24444)">Hrabě’s</a> collection was sampled from several sites in a section of the <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=16.74944&amp;materialsCitation.latitude=49.24444" title="Search Plazi for locations around (long 16.74944/lat 49.24444)">Říčka river</a>, near <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=16.74944&amp;materialsCitation.latitude=49.24444" title="Search Plazi for locations around (long 16.74944/lat 49.24444)">Pekárna</a> (N49.24305, E16.74583), <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=16.74944&amp;materialsCitation.latitude=49.24444" title="Search Plazi for locations around (long 16.74944/lat 49.24444)">Netopýrka</a> and Ochozská Caves (N49.24444, E16.74944), Brno District, separated about 300 m (coordinates from Timm &amp; Abarenkov, gbif.org). The lectotype is from site B described in Hrabě 1938: 80, Dr. P. Dolejs, pers. comm .)</p><p>Paralectotypes: NMP P6 j-20/89/144. Hr 1006: not fully mature, unmated individual in histological sections (#1); two immature, whole-mounted individuals (#2). Hr 1007-2 (#4, #6) whole mount, mature, unmated, no egg sac. Hr 1007-2 (#8, #14) sagittal sections, mature, unmated, no egg sac. Hr 1015 transverse sections, mated individual (#17). Hr 1015-2 (#19) 1 immature, whole-mounted individual. Three sites located 10 km of Brno, Moravian Karst, Czech Republic .</p><p>Description (based on four mature individuals from the NMP type material from Říčka River, mated or unmated but with sperm in male funnels and long sperm sacs). Small worms, body diameter in X 0.21–0.31 mm. Prostomium round. Clitellum not well separated from adjacent segments, but with thicker epidermis: to 14 µm high in X– XIII, vs. 5–10 µm in anterior segments. Secondary annulation from III. One pair of male pores in X (in IX in one of the examined individuals) behind and in line with ventral chaetae, one pair of female funnels in 11/12 and two pairs of spermathecae, in XI and XII, behind in line with the ventral chaetae. One pair of chaetae per bundle, simple pointed; in the anterior body segments, dorsals somewhat smaller than or equal in length to ventrals (dorsals 60–78 µm, ventrals 68–82 µm in Hr 1007-2c (#6); dorsals 83–96 µm, ventrals 85–106 µm in Hr 1015-2 (#19).</p><p>Pharynx in segments II and III, with a well-developed dorsal pad. Pharyngeal glands from V to the anterior part of VIII. Chloragogenous tissue from segment VII. First nephridia in VII. Sperm sacs only developed backwards to XI–XV, no egg sacs. Two pairs of testes (segments IX and X) and one pair of ovaries (segment XI).</p><p>Semiprosoporous male duct. Atrium petiolate, with a round ampulla and a well-defined duct (Fig. 8A), its total length about half the body diameter. Round to oval ampulla (74–122 µm long, 53–91 µm Ø), atrial duct 33–51 µm long, 20–27 µm Ø, roughly cylindrical or slightly conical, narrowed to the pore (to 12–17 µm). Atrial musculature ≤ 2–3 µm thick, epithelium with glandular, nucleated cells, 10–18 µm high. Prostate glands in clusters, to 80–94 µm high. One pair of vasa deferentia joins medially or subapically to atrial ampulla (Fig. 8B), the posterior one crossing septum 10/11 and forming a short loop in segment XI.</p><p>Spermathecal ampullae 100–247 µm long and 60–130 µm Ø, in the same segment as the corresponding pore, or crossing to the following one. Long spermathecal ducts, about 105–160 µm long, 15–30 µm Ø, with a small distal widening, that in mated specimens can form a small vestibule containing sperm (Fig 8C).</p><p>Remarks. The species name was made available in a key to species of Trichodrilus (Hrabě 1937: 20), but the formal and detailed description is in Hrabě (1938). The description of T. moravicus was based on several populations, but mainly from two sites: the Říčka River and in the Býčí Skála Cave, in the Czech Republic. Hrabě (1938) reported differences in size (number of segments and body diameter) and the study of the type series from the NMP showed additional morphological differences between these populations. Most interesting was the difference in the position of the junction of the vasa deferentia to the atrial ampulla, subapical in specimens from Říčka R. vs. basal in specimens from Býčí Skála. This fact has led us to review the diagnosis of T. moravicus and to evaluate two options to deal with this problem:</p><p>1) To amend the description of T. moravicus, considering that these characters are variable for the species;</p><p>2) Not to modify the original description, so that T. moravicus shows the subapical junction of the vasa deferentia, which occurs in the type series from Říčka River. This option implies the proposal of a new identity for Hrabě´s population in the Cave Býčí Skála, mostly based on the different position of the vasa deferentia junction, but also considering the differences already evident in the original description.</p><p>We have taken the second option. This meant the designation of a lectotype for T. moravicus from the Říčka River, since all specimens from Říčka River and Cave Býčí Skála were syntypes, due to the lack of original type designation. We chose a specimen consistent with the original description of the species, with junction of vasa deferentia and atrium clearly visible. The present description, based on the revision of the type material from Říčka River, fits the original descriptions by Hrabě (1938) where the junction of vasa deferentia to the atrial ampulla is described as ental (nahe seinem entalen Ende einmündet). The figures that Hrabě included in his description (1938: figs.7, 8) from the Řicka R. are from transversally sectioned individuals, where we were not able to determine the position of the vasa deferentia junction. However, this character was clearly seen in both sagittal sections and whole-mounts from the Říčka River collection examined by the authors .</p><p>On the other hand, the Trichodrilus population from Cave Býčí Skála is identified here as T. stygodytes sp. nov., conspecific with the population from Bizkaia, Spain (see above), which shares the structure of the reproductive system. In order to facilitate future DNA-based studies, the holotype of this species was selected from the Spanish population and not from the slide collection of the National Museum of Prague.</p><p>In the original description of T. moravicus, there is another population from the Glatzer Schneeberg (near Králický, Czech Republic), a subterranean river in Cave Quarglöcher (Hr 1027), that was not examined here. With the lectotype designation, these specimens are no longer syntypes of T. moravicus .</p><p>As Hrabě (1960) pointed out, T. moravicus is morphologically close to T. tenuis in several characteristics: very small worms, atrium ampulla round to oval, with narrow musculature, the posterior vas deferens passes into segment XI and joins subapically to the atrial ampulla. This species can be separated from T. tenuis by the size and shape of atrium [described by Hrabě (1960) as “slightly different”]. The atrium is petiolate in T. moravicus, i.e. with the cylindrical duct well-separated from the ampulla, while in T. tenuis, the duct gradually narrows and it appears as if contained within the ectal part of the ampulla, or protrudes forming a conspicuous penis. Total length of the atrium in T. moravicus is about half the body diameter, while in T. tenuis it is usually about one third.</p><p>Habitat and distribution. Trichodrilus moravicus has been reported so far in headwaters of karst rivers, interstitial waters and the profundal zone of a lake in Czech Republic, Slovakia and Poland (Fig. 6). Kasprzak (1979: 73–74, fig.14) described a small population of T. moravicus in Poland, with very thick atrial musculature and few tiny prostatic cells, which does not match the original description of T. moravicus . The presence of T. moravicus in several European countries should be confirmed in light of the new taxonomic data.</p><p>......continued on the next page</p></div>	https://treatment.plazi.org/id/03A04148B13CFE160BC5FB01F9C9F5B7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Rodríguez, Pilar;Fend, Steven	Rodríguez, Pilar, Fend, Steven (2025): Contributions to the knowledge of the genus Trichodrilus Claparède (Lumbriculidae, Clitellata) with the description of new groundwater species from Spain. Zootaxa 5711 (1): 57-78, DOI: 10.11646/zootaxa.5711.1.2, URL: https://doi.org/10.11646/zootaxa.5711.1.2
