taxonID	type	description	language	source
2722A163FFD1FFC0256FFD3D25510525.taxon	description	(Figure 2)	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFD1FFC0256FFD3D25510525.taxon	materials_examined	Neotype. MNHN-IC- 2024 - 1386, 46.2 mm SL, male, France, Lez River, Mediterranean Basin, Hérault Department, no coordinates, 1977, Quignard, J. P. Material examined. MNHN-IC- 1977 - 0134, 2, 37.9 – 44.4 mm SL, topotypes, collected with neotype. MNHN-IC 2010 - 1064, 6 of 7 alc. (1 adult measured, 39.8 mm SL) + 1 mitogenome (voucher tag FFFtag 4260, 41.7 mm SL, GenBank Accession Number MW 288293), collected in the type locality of B. quignardi, France, Lez River at Prades-le-Lez, Dept. Hérault, 43 Ǫ 420000 N, 3 Ǫ 51057.600 E, 24 Nov 2010, Denys, G. and Office National de l'Eau et des Milieux Aquatiques (ONEMA). MNHN-IC- 2021 - 0358, 1, France, Dept. Hérault, 2021, Denys, G. MNCN-ICTIO 17741 – 17742, 2, France, Isle, Sorgue River, tributary of the Rhône River, 44 Ǫ 1305900 N, 5 Ǫ 9026.5000 E (approx. coordinates), 30 Apr 1968, unknown collector.	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFD1FFC0256FFD3D25510525.taxon	diagnosis	Diagnosis. Barbatula barbatula is a member of a monophyletic group of species, herein designated as Western Europe clade (including B. hispanica, B. leoparda, B. fluvicola n. sp. and B. ommata n. sp.), which is diagnosed from all its valid congeners by having a longer dorsal head length 23.2 % – 25.8 % SL (vs. 19 % – 21.8 % in B. hispanica, 20 % – 21.5 % in B. leoparda, 18.4 % – 21.7 % in B. fluvicola n. sp. and 19.3 % – 21.4 % SL in B. ommata n. sp.), longer lateral head length 25.9 % – 29.6 % SL (vs. 22 % – 24.9 %, 22.5 % – 25 %, 21.8 % – 24.5 % and 22.4 % – 26.2 % SL, respectively), longer predorsal length 55.8 % – 60.4 % SL (vs. 49.8 % – 53.3 %, 53.5 % – 54.4 %, 50.5 % – 55.8 % and 50.4 % – 56.8 % SL, respectively), longer prepectoral length 26.2 % – 28.7 % SL (vs. 20.8 % – 25 %, 22.2 % – 24.7 %, 20.9 % – 25.1 % and 21.3 % – 25.8 % SL, respectively), longer dorsal fin 24.1 % – 25.8 % SL (vs. 21.2 % – 23.8 %, 20.4 % – 23.4 %, 18.4 % – 24.2 % and 18.4 % – 23.3 % SL, respectively), shorter caudal peduncle 11.7 % – 12.7 % SL (vs. 13.7 % – 17.4 %, 12 % – 14.4 %, 13.9 % – 17.3 % and 12.7 % – 16.2 % SL, respectively), and protuberant slope present from preceding area of anterior nares (vs. slope preceding anterior nares absent or weakly developed). It is distinguished from B. leoparda, B. fluvicola n. sp. and B. ommata n. sp. by having longer dorsal-fin base 14.5 – 18.6 % SL (vs. 12.8 % – 14.4 % in B. leoparda, 11.2 % – 13.6 % SL in B. fluvicola n. sp. and 11.1 % – 14.3 % in B. ommata n. sp.) and from B. hispanica, B. fluvicola n. sp. and B. ommata n. sp. by having deeper head 51.7 % – 57.3 % HL (vs. 40.2 % – 49.3 %, 46.1 % – 51.7 % and 38.4 % – 46 % HL, respectively). Barbatula barbatula is further distinguished from B. fluvicola n. sp. and B. ommata n. sp. by having the adpressed anal-fin tip slightly anterior to end of caudal peduncle (vs. adpressed anal-fin tip reaching about middle of caudal peduncle) and larger suborbital depth 33.1 % – 36.3 % HL (vs. 26.5 % – 32.4 % and 23.7 % – 31.2 % HL). It further differs from B. leoparda by having ventral surface of head and area between pectoral to pelvic-fin origin without pigmentation (vs. ventral surface of such areas pigmented with brown to dark irregular blotches); and from B. ommata n. sp. by having a longer first dorsal-fin ray 20 % – 24.3 % SL (vs. 13.5 % – 19.8 % SL) and a longer snout 50.8 % – 52.6 % HL (vs. 37.2 % – 46.1 % HL).	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFD1FFC0256FFD3D25510525.taxon	description	Redescription. Morphometric data are shown in Table 1. General appearance is shown in Figure 2. Complementary description details are shown in the original description of Barbatula b. quignardi (B ̆ acescu-Meşter, 1967). Body short and relatively thickened, roughly cylindrical in cross-section at trunk, gradually compressed towards caudal peduncle. Dorsal profile of head straight rising from snout tip to posterior border of supraoccipital; conspicuous hump demarking posterior border of supraoccipital. Ventral profile of trunk slightly convex. Body depth slightly decreasing from end of dorsal fin towards caudal-fin base. Greatest body depth at predorsal area. Greatest body width at opercle or preceding pectoral-fin origin. Head relatively deep, cylindrical in dorsal view. Dorsal head profile with protuberant slope present from preceding area of anterior nares, larger than congeners. Anterior border of snout rounded in dorsal view, tapering towards its tip; anterior portion truncated. Eyes laterodorsally positioned; positioned dorsal half on head. Orbital rim free. Nostrils small. Anterior nostril surrounded by elongate fleshy flap-like tube integument; fleshy flap elongated. Posterior nostril without fleshy flap and located close to anterior border of eye. Anterior and posterior nostril closely set, positioned almost continuously and connected by thin skin ridge. Gill openings united to isthmus laterally approximately at line of inner portion of pectoral-fin base origin. Mouth ventral, arched in a semicircular shape. Upper lip thin; vertical median incision (notch) not evident. Contralateral lower lips separated at midline by a deep notch. Mesial portion of lower lip with well-developed mental lobe forming a fleshy fold, but relatively short; conical protrusion of mental lobe absent. Lateral lobe or projection of lateral portion of lower lips absent. Maxillary barbel longer than premaxillary barbel pairs, surpassing vertical of middle of eye. Two pairs of premaxillary barbel. Mesial premaxillary barbel tip usually not reaching end of mental lobe of lower lip. Lateral premaxillary barbel positioned laterally on snout, longest than mesial barbel; its tip reaching middle of posterior naris. Pectoral-fin rays i, 10 * (1), i, 11 * (1) or 12 * (1); distal margin acute, mesial border somewhat convex. Epithelial tubercles present in pectoral-fin pterygiophores and increase in size towards pectoral-fin tip. Pelvic-fin distal border ellipsoid; pectoral origin situated midbody length, at vertical of second branched dorsal-fin ray. Pelvic-fin rays i, 6 (1) or I, 7 * (2). Axillary pelvic lobe present and conspicuous. Dorsal fin long compared to congeners; its origin located slightly posteriorly midbody; fin tip truncated. Dorsal-fin rays i, 7 ½ * (3). Anal fin i, 5 ½ * (3) rays. Anal fin with distal margin truncated. Anal-fin origin located at third half of body length, posterior to end of ultimate branched dorsal-fin ray. Caudal fin emarginate; upper and lower lobes similar in length. Caudal-fin rays i, 8 + 8, i * (3). Caudal-fin peduncle short and deep, approximately same depth as the remain part of body and head. Procurrent rays pronounced and embedded in the skin forming a dorsal and ventral dermal keel extended two thirds or more of caudal peduncle. Body covered by embedded, tiny, roundish scales, not overlapping each other. Lateral line incomplete and continuous, reaching anterior portion of anal-fin base. Coloration (in alcohol). Background colour yellowish pale- to light brown. Head surface mottled, sparsely set and fainted towards cheek and lateral part of snout. Dorsolateral portion of body with conspicuous, irregular set five dark-brown blotches (Figure 2). Flank with conspicuous, large, dark-brown, irregular rounded blotches, usually separated from each other. Pigmentation fading below lateral line. Ventral surface of body and head mostly without pigmentation, except for caudal-fin peduncle with few sparsely blotches. Dorsal surface of pectoral-fin base and dorsal-fin base densely dark-brown pigmented. Pigmentation on barbels and lips restricted to few spots and small blotches. Pectoral-fin pterygiophores with elongate dark-brown blotches forming two to three elliptical lines. Dorsal fin with pigmentation concentrated in its base; rays with small, somewhat rectangular dark-brown blotches on all rays, usually arranged in rows. Pelvic-fin rays hyaline or with very faint and scattered pigmentation in outermost rays. Anal fin hyaline. Caudal-fin base with dark-brown, slightly m-shaped bar in middle and comma-shaped blotch on upper caudal-fin base. Caudal fin covered by small, dark-brown, elongate blotches forming vertical series or are randomly distributed all over caudal fin. Sexual dimorphism. Barbatula barbatula shows pectoral fin elongated, acute in adult males (vs. elliptical to rounded in females and juveniles). In males, second branched ray longest than remaining (vs. third branched ray longest than remaining in females). Well-developed epidermical tubercles in the pectoral-fin rays of adult males (absent in females and not evident in juvenile males). Tubercles present in the principal unbranched ray and 1 – 6 innermost branched rays, rare in eighth ray (Figure 3 in B ̆ acescu-Meşter, 1967). Urogenital papilla of males small and conical, and urogenital opening in females rounded; in both sexes, smaller in size when compared to its congeners; more difficult to distinguish the sexes.	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFD1FFC0256FFD3D25510525.taxon	distribution	Distribution. Barbatula barbatula is currently confirmed to occur in the Lez River in France (see Nomenclature section) and the Sorgue River (based on morphological identification; Figure 3). The presence of the species, confirmed by coI barcode in the lowermost part of the Rhône drainage, is likely to indicate its presence in other surrounding areas. The range of the species may therefore be wider than currently known (Figure 3).	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFD1FFC0256FFD3D25510525.taxon	discussion	Remarks. Populations from the Doubs and Allaine rivers in the upper portion of Rhône in Switzerland were recovered nested in the B. barbatula clade based on coI gene, but exhibited significant morphological differences compared to B. barbatula populations from the Lez and Sorgue rivers. Additionally, part of the populations from the rivers Le Bied, Broye and Areuse Travor, disconnected streams from the remaining Aare catchment in Switzerland, were also recovered in the coI tree nested within B. barbatula population from the Lez river, although we were unable to check their morphology. Additional material is required to determine whether they are conspecific and to precisely define the geographical range of B. barbatula.	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFD3FFD6256FF92E20B20395.taxon	description	(Figures 4 and 5) urn: lsid: zoobank. org: act: 5 AB 6 B 45 C-C 3 BD- 409 B-A 927 - D 564515 4 D 2 AB	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFD3FFD6256FF92E20B20395.taxon	materials_examined	Holotype. NMBE 1105500, 83.8 mm SL, female, Glane River, appr. 900 m upstream of the confluence between Glane and Saane rivers, Aare catchment, Rhine drainage, Canton of Fribourg, Switzerland, 590 m alt., 46 Ǫ 4701.500 N, 7 Ǫ 705.700 E, 5 Oct 2022, Wegscheider, B., Josi, D., Waldock, C. & Canton of Fribourg (Figure 6 a). Paratype. All from Switzerland: Rhine drainage: Aare River: Canton of Fribourg: NMBE 1105486 – 1105489, NMBE 1105493 – 1105497, NMBE 1105499, 10, 51.2 – 91.2 mm SL (HRXCT of NMBE 1105486), ZFMK-ICH 135303 – 135304, 2, 81.0 – 81.2 mm SL, ZSM 49670, 2, 65.4 – 67.3 mm SL, collected with the holotype. Canton of Bern: — NMBE 1105472 – 1105485, 14 + 1 C & S, 54.0 – 76.7 mm SL, Lyssbach River, at bridge 500 m upstream of confluence with Alte Aare, 440 m alt., 47 Ǫ 5028.100 N, 7 Ǫ 18040.800 E, 18 Oct 2022, Wegscheider, B., Josi, D., Waldock, C, Moreau, S. — NMBE 1105339 – 1105342, 4, 73.7 – 82.4 mm SL, Limpach River, approx. 1.7 km upstream from Sout-West Golf Limpach on road from Unterramsern to Limpach, 466 m alt., 47 Ǫ 702.800 N, 7 Ǫ 29034.300 E, 27 Jan 2022, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C. — NMBE 1105350, NMBE 1105353 – 1105358, NMBE 1105360 – 1105361, NMBE 1105363 – 1105365, 10 + 4 C & S, 64.8 – 87.3 mm SL, ZFMK-ICH 135305 – 135306, 2, 76.6 – 80.6 mm SL, NMW 101168, 2, 78.7 – 81.2 mm SL, Limpach River, 50 m upstream from the bridge on the road Bernstrasse from Wengi to Schnottwil, 471 m alt., 47 Ǫ 5027.300 N, 7 Ǫ 23059.200 E, 23 Feb 2022, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C. — NMBE 1105368 – 1105369, 2, 72.9 – 81.6 mm SL, Limpach River, at Bätterkinden 1 km upstream from the confluence with Emme River, 411 m alt., 47 Ǫ 901000 N, 7 Ǫ 32019.100 E, 2 Mar 2022, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C. — NMBE 1111089 – 1111090, NMBE 1111848 – 1111849, 4, 84.2 – 87.5 mm SL, Urtene River, village Schalunen approx. 85 m upstream where Alpstrasse street cross the river, 486 m alt., 47 Ǫ 6045.100 N, 7 Ǫ 31053.100 E, 1 Mar 2022, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C. — NMBE 1105370 – 1,105,378, NMBE 1105380, 10, NMBE 1105383 (1 C & S), 66.6 – 86.4 mm SL, NMB 6480 – 6481, 2, 72.0 – 72.7 mm SL, MNHN 2024 – 1384, 2, 69.3 – 71.3 mm SL, Emme River, at Wiler bei Utzenstorf next to the Emme Forstbaumschule, 478 m alt., 47 Ǫ 9054.400 N, 7 Ǫ 33018.100 E, 2 Mar 2022, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C. Canton of Solothurn: — NMBE 1105385 – 1105387, NMBE 1105389 – 1105391, NMBE 1105393 – 1105394, 8, 54.4 – 71.8 mm SL, MNCN-ICTIO 298.402 and MNCN-ICTIO 298.403, 2, 67.1 – 69.5 mm SL, Emme River, 1.7 km downstream of the dam of Biberist between Biberist and Derendingen, 486 m alt., 47 Ǫ 11033.900 N, 7 Ǫ 34048.700 E, 4 Mar 2022, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C. Canton of Vaud: — NMBE 1105424, NMBE 1105426 – 1105433, 11, 39.8 – 68.3 mm SL, ZFMK-ICH 135307 – 135308, 2, 53.6 – 69.9 mm SL, Petite Glane River, 1 km east from Villars-legrand, 430 m alt., 46 Ǫ 54022.900 N, 7 Ǫ 0027.700 E, 9 Sep 2022, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C., Sudasinghe, H. Genseq- 2 coI. NMBE 1105371 (tissue tag 230097); Genbank Acession Number PQ 488083. Genseq- 2 coI. NMBE 1105372 (tissue tag 230098); Genbank Acession Number PQ 488084. Genseq- 2 coI. NMBE 1105487 (tissue tag 240565); Genbank Acession Number PQ 488093. Genseq- 2 coI. ZFMK-ICH 135303 (tissue tag 240568); Genbank Acession Number PQ 488094. Genseq- 2 coI. NMBE 1105472 (tissue tag 241024); Genbank Acession Number PQ 488169. Genseq- 2 coI. NMBE 1111089 (tissue tag 229970); Genbank Acession Number PQ 488191. Genseq- 2 coI. NMBE 1111090 (tissue tag 229971); Genbank Acession Number PQ 488192. Genseq- 2 coI. NMBE 1105424 (tissue tag 240249); Genbank Acession Number PQ 488148. Genseq- 2 coI. NMBE 1105426 (tissue tag 240251); Genbank Acession Number PQ 488149. Genseq- 2 coI. NMBE 1105340 (tissue tag 229770); Genbank Acession Number PQ 488130. Genseq- 2 coI. NMW 101168 (tissue tag 229857); Genbank Acession Number PQ 488131. Genseq- 2 coI. NMW 101168 (tissue tag 229858); Genbank Acession Number PQ 488132. Genseq- 2 coI. NMBE 1105369 (tissue tag 230083); Genbank Acession Number PQ 488133. Additional material (non-types). Switzerland: Rhine drainage: Canton of Aargau: NMBE 1085028 – 1085033, 6, 62.1 – 79.0 mm SL, Aabach River, tributary of the Aare River, Niederlenz, 363 m alt., 47 Ǫ 24030.100 N, 8 Ǫ 10010.100 E, 14 Sep 2015, NAWA (National Surface Water Quality Monitoring Program; Switzerland). — NMBE 1085259 – 1085272, 14, 35.5 – 84.8 mm SL, NMBE 1106080 – 1106085, 6 (entire vouchers preserved in tissue collection), 35.0 – 39.0 mm SL, Surb River, tributary of the Aare River, Döttingen, 333 m alt., 47 Ǫ 33052.200 N, 8 Ǫ 15052.600 E, 16 Sep 2015, NAWA. — NMBE 1085103, 1, 70.6 mm SL, Bünz River, tributary of the Aare drainage, Möriken, 374 m alt., 47 Ǫ 24035.600 N, 8 Ǫ 11010.100 E, 15 Sep 2015, NAWA. Canton of Appenzell: — NMBE 1083178, tissue (identified by photo), Urnäsch River, Matzingen, 589 m alt., 47 Ǫ 2404.400 N, 9 Ǫ 19034.800 E, 12 Aug 2015, NAWA. Canton of Basel: — NMBE 1085464 – 1085481, 17, 41.6 – 71.0 mm SL, Birs River, Basel, 249 m alt., 47 Ǫ 33024.200 N, 7 Ǫ 3703.800 E, 24 Sep 2015, NAWA. Canton of Bern: — NMBE 1105343 – 1105349, 7, 36.6 – 53.2 mm SL, Limpach River, 1.7 km upstream from Sout-West Golf Limpach on road direction Unterramsern to Limpach, board between Solothurn and Bern cantons, 466 m alt., 47 Ǫ 702.800 N, 7 Ǫ 29034.300 E, 27 Jan 2022, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C. — NMBE 1105395 – 1,105,401, NMBE 1105403 – 1105406, 11 + 1 C & S, 64.4 – 92.4 mm SL, Kalte Sense River, tributary of the Aare River, downstream of the Hoflanderebrügg, 957 m alt., 46 Ǫ 42059.200 N, 7 Ǫ 19040.700 E, 6 Sep 2022, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C., Sudasinghe, H., Canton of Bern and Canton of Fribourg. — NMBE 1105407 – 1105422, 16, 39.1 – 83.1 mm SL, Sense River, at Sodbachbrügg, 702 m alt., 46 Ǫ 49036.100 N, 7 Ǫ 19020.700 E, 7 Sep 2022, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C., Sudasinghe, H., Canton of Bern and Canton of Fribourg. — NMBE 1105501 – 1105515, 15, 57.6 – 76.2 mm SL, Schwarzwasser River, tributary of the Aare River, near to Rossgraben bridge, 681 m alt., 46 Ǫ 49037.400 N, 7 Ǫ 23055.500 E, 5 Sep 2022, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C., Sudasinghe, H., Canton of Bern and Canton of Fribourg. — NMBE 1083400 – 1083425, 25, 59.2 – 77.9 mm SL, Gürbe River, tributary of the Aare drainage, Kehrsatz, 507 m alt., 46 Ǫ 550000 N, 7 Ǫ 29029.400 E, 21 Aug 2015, NAWA. — NMBE 1080648, NMBE 1080650 – 1080651, 3, 78.9 – 91.7 mm SL, Schwarzwasser River, tributary of the Aare River, Schwarzwasserbrücke, 587 m alt., 46 Ǫ 51051.500 N, 7 Ǫ 24030.900 E, 17 Sep 2013, unknown collector. — NMBE 1080626, 1, 90.3 mm SL, main course of Schwarzwasser River, at Rüschegggraben, 724 m alt., 46 Ǫ 48011.700 N, 7 Ǫ 24030.900 E, 17 Sep 2013, unknown collector. — NMBE 1080692, 1, 80.0 mm SL, NMBE 1080693 (only tissue), Sense River, tributary of the Aare River, Guggersbachbrücke, 763 m alt., 46 Ǫ 45024.500 N, 7 Ǫ 1801300 E, 16 Sep 2013, unknown collector. — NMBE 1080749 – 1080750, 2, 35.7 – 45.7 mm SL, NMBE 1080751 (only tissue), Sense River, tributary of the Aare River, Ruchmühlebrücke, 609 m alt., 46 Ǫ 50046.700 N, 7 Ǫ 20013.900 E, 17 Sep 2013, unknown collector. Canton of Fribourg: — NMBE 1105449 – 1105454, 5, 46.9 – 89.8 mm SL, Saane River, tributary of the Aare River, 0.6 km upstream of the confluence between Saane and La Gérine, 584 m alt., 46 Ǫ 4603200 N, 7 Ǫ 702000 E, 16 Sep 2022, Wegscheider, B.; Josi, D.; Sudasinghe, H.; Talbi, M., Canton of Fribourg. — NMBE 1080702 – 1080703, 2, 70.8 – 77.7 mm SL, Sense River, tributary of the Aare River, Guggersbachbrücke, 764 m alt., 46 Ǫ 45024.800 N, 7 Ǫ 18010.500 E, 16 Sep 2013, unknown collector. — NMBE 1083482 – 1083483, 2, 69.4 – 89.1 mm SL, NMBE 1106139 – 1106140, 2 (entire vouchers preserved in tissue collection), 43.0 – 44.0 mm SL, Sionge River, tributary of the Aare River, Vuippens, 690 m alt., 46 Ǫ 39022.900 N, 7 Ǫ 4041.800 E, 28 Aug 2015, NAWA. Canton of St. Gallen: — NMBE 1105434 – 1105448, 14 + 1 C & S, 53.7 – 84.8 mm SL, Jona River, tributary of the Limmat River, upstream of Rapperswil-Jona at the bridge, 422 m alt., 47 Ǫ 14012.600 N, 8 Ǫ 50012.800 E, 13 Sep 2022, Calegari, B. B., Wegscheider, B.; Josi, D., Sudasinghe, H. — NMBE 1111834 – 1111837, NMBE 1111057 – 1111067, and NMBE 1109967 – 1109971, 19 + 1 C & S, 64.4 – 81.4 mm SL, Thur River, tributary of the Aare River, upstream of Thur bridge near to Niederbüren, 705 m alt., 47 Ǫ 28011.500 N, 9 Ǫ 11037.900 E, 14 Oct 2023, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C. — NMBE 1084866 – 1084886, 19 + 2 C & S, 43.3 – 76.5 mm SL, Thur River, Golfplatz, 476 m alt., 47 Ǫ 2801300 N, 9 Ǫ 11041.700 E, 9 Sep 2015, NAWA. — NMBE 1085366 – 1085367, NMBE 1085369 – 1085374, NMBE 1085376 – 1085377, NMBE 1085380 – 1085382, NMBE 1085384 – 1085389, 19, 57.1 – 82.2 mm SL, Glatt River, Buechental, 496 m alt., 47 Ǫ 26038.400 N, 9 Ǫ 901700 E, 22 Sep 2015, NAWA. Canton of Schwyz: — NMBE 1105455 – 1105470, 16, 63.3 – 100.5 mm SL, Chli-Aa River, tributary of the Limmat River, at Lachen about 400 m upstream of confluence to Lake Zurich, 409 m alt., 47 Ǫ 11052.600 N, 8 Ǫ 51032.500 E, 14 Oct 2022, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C., Schmid, A., Moreau, S. Canton of Thurgau: — NMBE 1083709 – 1083710, NMBE 1083715 – 1083722, 10, 71.0 – 85.0 mm SL, Lauche River, Kubel, 441 m alt., 47 Ǫ 31011.900 N, 8 Ǫ 55050.400 E, 26 Aug 2015, NAWA. — NMBE 1083628 – 1083631, 4, 40.6 – 50.5 mm SL, Chemibach River, Märstetten, 419 m alt., 47 Ǫ 35050.700 N, 9 Ǫ 302000 E, 25 Aug 2015, NAWA. Canton of Zürich: — NMBE 1105531 – 1105545, 15, 57.8 – 77.9 mm SL, Sihl River, tributary of the Limmat River, downstream of bridge near to the train station of Leimbach, 435 m alt., 47 Ǫ 2000.700 N, 8 Ǫ 31011.500 E, 12 Sep 2022, Calegari, B. B., Wegscheider, B., Josi, D. — NMBE 1105546 – 1105547, 2, 85 – 88.3 mm SL, Sihl River, tributary of the Limmat River, downstream of hydroelectric dam Sihlwehr, 705 m alt., 47 Ǫ 10017.700 N, 8 Ǫ 39051.900 E, 12 Sep 2022, Calegari, B. B., Wegscheider, B., Josi, D. — NMBE 1084662 – 1084678, 16, 49.6 – 77.8 mm SL, Mönchaltorfer Aa River, Mönchaltorf, 441 m alt., 47 Ǫ 18043.900 N, 8 Ǫ 43010.500 E, 1 Sep 2015, NAWA. — NMBE 1084301 – 1084307, 7, 47.5 – 67.1 mm SL, Ellikerbach River, downstream at wastewater treatment plant Ellikon, 377 m alt., 47 Ǫ 34020.500 N, 8 Ǫ 49013.400 E, 15 Oct 2015, NAWA. — NMBE 1084369 – 1084383 and NMBE 1106708 – 1106709, 17, 57.1 – 91.8 mm SL, Eulach River, downstream at ARA Elgg, 497 m alt., 47 Ǫ 3004.100 N, 8 Ǫ 5103.200 E, 16 Oct 2015, NAWA. — NMBE 1083144 and NMBE 1083147, 2, 62.6 – 71.0 mm SL, and NMBE 1083143 (only tissue), Furtbach River, tributary of the Limmat River, Otelfingen, 419 m alt., 47 Ǫ 26058.600 N, 8 Ǫ 23012.900 E, 11 Aug 2015, NAWA. Germany: Danube drainage: Bavaria State: — ZSM 35660, 14, 62.8 – 81.9 mm SL, stream Kessel, at Bergmühle, Unterbissingen, West to Donauwörth, Swabia, 48 Ǫ 42028.6200 N, 10 Ǫ 3806.7500 E, 11 Jun 2007, Schubert, M. — ZSM 34592, 20, 80.2 – 46.9 mm SL, stream tributary of the Isar River, Pförreraugraben at Freising, Upper Bavaria, 48 Ǫ 210 N, 11 Ǫ 440 E, 17 Jun 2006, Miller, M. — ZSM 33502, 12, 80.7 – 64.9 mm SL, stream Geltnach, tributary of the Wertach River, below Bertoldshofen near to Marktoberdorf, Upper Bavaria, 47 Ǫ 47.50 N, 10 Ǫ 39.40 E, 6 Jul 2005, A. Kolbinger and B. Ott. — ZSM 32918, 1, 78.9 mm SL, Swabia, 48 Ǫ 19040.500 N, 11 Ǫ 007.800 E, 13 Apr 2005, Schubert, M. — ZMS 31550, 1, 50.6 mm SL, stream Westliche Günz, tributary of the Günz River, upstream of Westerheim near Memmingen, Allgäu region, Swabia, 47 Ǫ 59.70 N, 10 Ǫ 17.50 E, 17 Nov 2004, Schubert, M. and Neumann, D. — ZSM 31620, 2, 57.0 – 58.1 mm SL, Grosse Vils upstream of Lebertshausen, Lower Bavaria, 48 Ǫ 28.80 N, 12 Ǫ 24.10 E, 6 Dec 2004, Kolbinger, A., and Ott, B. — ZSM 33459, 1, 56.0 mm SL, Danube River, upstream on left bank below Danube-bridge at public bath, Dillingen, Swabia, 48 Ǫ 34.30 N, 10 Ǫ 30.80 E, 7 Jun 2005, Schubert, M. — ZSM 37270, 5, 58.9 – 76.9 mm SL, Iller River, 5.6 km South of Ulm, Wiblingen, Swabia, 48 Ǫ 21030.200 N, 9 Ǫ 59052.600 E, 23 Jun 2008, Langenargen, F. F. S. — ZSM 36024, 3, 65.0 – 80.0 mm SL, Inn River, at Simbach on the borders between German and Austria, Lower Bavaria, 48 Ǫ 15045.300 N, 13 Ǫ 02011.500 E, 31 Jul 2007, Ott, B., and Bohl, E. Austria: Danube drainage: — NMW 92845, 1, 84.3 mm SL, Fischerwirtsbach, Liefering, Salzburg, 21 Jul 1993, Glechner. — NMW 93066, 4, 26.4 – 64.2 mm SL, Statzenbach, Neumarkt at Wallersee, Salzburg, 28 Jul 1995, Glechner.	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFD3FFD6256FF92E20B20395.taxon	diagnosis	Diagnosis. Barbatula fluvicola is a member of a monophyletic group of species, herein designated as the Western Europe clade (including B. barbatula, B. hispanica, B. leoparda and B. ommata n. sp.), which is diagnosed by the following combination of characters. Barbatula fluvicola differs from B. leoparda and B. barbatula by having a shallower head 46.1 % – 51.7 % HL (vs. 51.5 % – 58.3 % HL in B. leoparda and 51.7 – 57.3 % HL in B. barbatula), and smaller suborbital depth 26.5 % – 32.4 % HL (vs. 32.2 % – 36.3 % and 33.1 – 36.3 % HL, respectively). It is further distinguished from B. leoparda and B. hispanica by possessing lower caudal-peduncle depth 8.7 % – 11.8 % SL (vs. 12.1 % – 13.6 % and 12.1 – 13.9 % SL, respectively); from B. leoparda by having upper lip covering entirely premaxilla (vs. premaxilla visible at mid-line portion), ventral surface of head and area between pectoral to pelvic-fin origin without pigmentation (vs. ventral surface of such areas pigmented with brown to dark irregular blotches), a shorter posterior nareal distance 21.3 – 26.5 % HL (vs. 26.8 % – 30 % HL), and a shorter lower lip 13.7 – 17.2 % HL (vs. 17.4 % – 21.2 % HL), and from B. hispanica by having a shorter maxillary barbel, reaching to middle of eye (vs. long maxillary barbel, reaching to posterior border of eye). It is further distinguished from B. barbatula by having a shorter dorsal head length 18.4 % – 21.7 % SL (vs. 23.2 % – 25.8 % SL), a smaller dorsal-fin base length 11.2 % – 13.6 % SL (vs. 14.5 % – 18.6 % SL), slope preceding anterior nares absent or weakly developed (vs. protuberant slope present from preceding area of anterior nares) and adpressed anal-fin tip reaching about middle of caudal peduncle (vs. adpressed anal-fin tip slightly anterior to end of caudal peduncle). Additionally, the new species distinguished from the sympatric congener B. ommata n. sp. by having smaller orbit 16.1 % – 19.8 % HL (vs. 21.1 % – 28.3 % HL), deeper head 46.1 % – 51.7 % HL (vs. 38.4 – 46 % HL), a longer coracoid, occupying at least two thirds of cleithrum length, usually almost reaching its tip (vs. coracoid short, half-length of cleithrum; Figure 7), ventral border of dentary curved (vs. ventral border of dentary straight; Figure 8), a smaller swimbladder bony capsule, not reaching half of head depth (Figure 9) (vs. swimbladder bony capsule large, occupying slightly more than half of head depth), posterior border of swimbladder bony capsule straight in ventral view (Figure 9) (vs. posterior border of swimbladder bony capsule rounded) and condyle of quadrate positioned at vertical line passing through posterior naris (Figure 9) (vs. condyle of quadrate positioned at vertical passing through anterior border of orbit).	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFD3FFD6256FF92E20B20395.taxon	description	Description. Morphometric data are provided in Table 2. Body robust elongate, roughly cylindrical in cross-section at trunk, gradually compressed towards caudal peduncle. Dorsal profile straight to slightly convex from snout tip to dorsal-fin origin. Dorsal and ventral profile of trunk approximately straight. Dorsal head profile slightly concave from snout tip to anterior border of eye and relatively straight from that point to posterior border of supraoccipital. Some individuals with shallow or conspicuous hump demarking posterior border of supraoccipital. Body depth decreasing posterior to dorsal-fin base to middle caudal-fin peduncle. Greatest body depth slightly anterior to dorsal-fin origin or at half distance between pectoral-and pelvic-fin origins at abdominal area. Greatest body width in half distance between pectoral-and pelvic-fin origins. Head relatively deep, cylindrical in dorsal view, longer than wide. Eyes laterodorsally positioned on dorsal half of head; located at middle between snout tip and posterior opercle margin. Ocular capsule slightly ellipsoid formed by thin and transparent skin. Nostrils small; anterior naris opening slightly smaller than posterior one. Anterior nostril with an elongate fleshy flap-like tube. Posterior naris in a drop shape. Anterior and posterior nostril close to each other, connected by a thin skin ridge. Ventral border of gill opening attached to body at inner pectoral-fin base. Mouth ventral, arched in a semicircular shape. Mouth wide, almost equal to head width. Upper lip thick, covering all premaxilla area. Mid-portion of upper lip larger than lateral portion. Short median incision (notch) in upper lip reaching to half lip depth; poorly developed in some individuals. Upper lip papillose in some individuals, smooth in others. Anterolateral portion of upper lip more depressed than middle portion. Outer skin of upper lip with three protuberances behind interspace of barbels. Lower lip separately at midline by a deep notch. Mesial portion of lower lip well-developed into a mental lobe vertically elongated formed by a fleshy fold in inverted U-shaped. Mental lobes usually in close contact at mid-line; without protrusion, leaving just a small V-shaped space of dentary uncovered at mid portion. Lateral portion of lower lip positioned inclined in 45 Ǫ angle to ventral midline, with deep furrows, posteriormost tip without projection. Distal parts of furrows on lower lip with small papillae, proximal parts usually smooth. Tip of mesial premaxillary barbel reaching approximately to base of maxillary barbel. Lateral premaxillary barbel reaching anterior border of posterior naris, when folded upwards. Maxillary barbel reaching or slightly surpassing vertical of anterior margin of eye, never surpassing eye. Pectoral-fin rays i, 10 (15), i, 11 * (17) or i, 12 (4). Innermost ray unbranched and very small, usually visible only in C & S specimens, with a single process base attached to first radial. Distal margin of pectoral fin truncated or convex. Principal unbranched pectoral-fin ray two thirds of longest pectoral-fin ray. Pelvic fin rays i, 6 (3) or i, 7 * (33), plus one additional, tiny, unbranched ray (sometimes absent). Pelvic fin ellipsoid, its origin at vertical through dorsal-fin origin or at principal, unbranched dorsal-fin ray. Pelvic fin with distal margin convex, its tip slightly anterior to a point below end of adpressed dorsal fin. Axillary pelvic lobe present, conspicuous fully attached to body. Dorsal-fin origin situated approximately at middle of body, straight or slightly convex. Proximal radial component of first dorsal-fin pterygiophore inserted anteriorly to neural spine of 13 th vertebrae (11 C & S). Dorsal fin with 3 – 4 small (11 C & S), unbranched rays completely covered by skin, followed by i, 7 ½ * (30) or i, 8 ½ rays (6). Dorsal fin with 8 (10 C & S) to 9 (1 C & S) pterygiophores and one additional posterior smaller pterygiophore with its radial element (sometimes radial absent) not bearing any rays. First dorsal-fin pterygiophore composed of likely supraneural and first proximal radials fused to each other in single complex structure supporting 3 – 4 shorter unbranched rays + principal unbranched ray. Remaining pterygiophores bearing posterior laminar bony keel supporting 7 – 8 ½ branched rays, each one associated individually to one of those pterygiophores, last 1 ½ rays supported by penultimate pterygiophore, and one additional last pterygiophore smaller in size lacking any correspondent ray. Dorsal-fin principal unbranched ray shorter than subsequent posteriormost four branched rays, reaching two thirds of its length. Distal anal-fin margin straight and slightly inclined anteriorly. Anal fin with three simple rays plus i, 5 ½ * (36) rays (one specimen with first two principal rays unbranched; morphological aberration). First simple rays and first unbranched principal ray supported by composed structure of at least three pterygiophores fused into single element, and five pterygiophores supporting remaining branched rays and last radial element much smaller at end of fin without rays attached; ultimate pterygiophore supporting 1 ½ rays. First pterygiophore inserted anteriorly to hemal spine of 24 th or 25 th vertebrae (11 C & S). Caudal fin truncated or slightly emarginate. Upper and lower lobe of caudal fin equal in length. Caudal-fin rays i, 8 + 7, i * (18), i, 7 + 8, i (5), i, 8 + 8, i (12), or i, 9 + 8, i (1 C & S). Procurrent rays in dorsal lobe 7 – 10, and ventral lobe with 7 rays. Procurrent rays pronounced and embedded in skin of shallow dorsal and ventral adipose keels extended on at least half of caudal peduncle. Ventral adipose keel slightly elevated extending to tip of adpressed anal fin. Epural long articulated to base of vertebrae. Upper hypural plate as single element with fused hypurals 3, 4 and 5. Lower hypural plate with co-ossified parhypural and hypurals 1 and 2 separated, forming the compound caudal centrum. Total vertebrae, 39 (8 C & S) or 40 (3 C & S); 14 – 15 (11 C & S) pairs of ribs. Body covered by embedded, tiny, rounded, not overlapping scales, irregularly set along trunk. Scales present on flank from about middle of pectoral fin to caudal-fin base. No scales on pre-dorsal region, except small patch before dorsal-fin origin. Scales on ventral surface of body present from end of pelvic-fin girdle to urogenital opening. Dermal tubercles present in some individuals, dense on head in adult males and females, and on dorsal surface of pectoral-fin rays. Lateral line continuous and complete, reaching caudal-fin base, with 64 – 67 pores (11 C & S), first three pores visibly larger than remaining. Cephalic lateral sensorial system with canals only partially ossified. Infraorbital canal with 11 – 12 pores (formed by two pores + posterior-half pore of antorbital segment, and anterior-half pore + nine pores + posterior-half pore of suborbital segment). Infraorbital canal composed of antorbital branch bearing two pores (anterior and dorsomedial), posteriorly ending in half-pore close but separately from anterior-half pore of adjacent suborbital branch, followed by nine pores and ending in posterior-half pore attached to distinct postorbital canal. Postorbital canal turned backwards with anterior-half pore, plus three pores and posterior-half pore in its end connected to both supratemporal canal and lateral line canal (or trunk canal). Supratemporal canal with half pore connected to both posterior-half pore of postorbital and half pore of lateral line canal, and two pores dorsally positioned. Trunk canal with three larger pores. Nasal canal with three pores weakly ossified (anterior, lateral and posterior). Frontal canal reaching near to anterior orbital border, but truncated at mid-portion splitting in two separate branches, each one bearing three pores (anterior, lateral and posterior). Preopercular canal with eight pores, anterior part located ventrally to articular and posteriorly to dentary. Colouration (in alcohol). Background colouration yellowish, head, back and flank with dark-brown pattern (Figure 10). Head very densely mottled. Back and flank mottled or marmorated, body and head almost plain brown in some individuals. Juveniles smaller than 40 mm SL often with inconspicuous, irregular shaped, blotches on back, most constant below dorsal-fin origin and at end of dorsal-fin base. Inconspicuous series of blotches along lateral midline in some individuals. In individuals with plain-brown flank, pigmentation dissociating and fading, below a line between origin of pectoral and pelvic fins and abdominal cavity. Ventral portion of head and abdomen area without pigmentation. Lateral part of head with dense pigmentation on cheek, lacking melanophores on ventral portion of the opercle. Presence of thin stripe from snout tip to anterior portion of eye, limited dorsally by nasal opening. Pigmentation absent on maxillary barbel, except when present, only on base. Premaxillary barbel usually without pigmentation, but when present more frequently in lateral pair of barbel with irregular dark markings on dorsal surface, most concentrated on proximal half. Lips completely deprived of pigmentation except by anterior border of upper lips. Pectoral fin with elongate dark blotches on rays, distal margin unpigmented. Blotches larger, more concentrate and conspicuous along anterior half of fin (first six rays). Dorsal-fin rays with small, somewhat rectangular dark blotches on rays, distributed irregularly, but sometimes organised in three bands, distal dorsal fin border without pigmentation. Pelvic fin hyaline. Anal fin hyaline, in few individuals with a faint vertical row of pigmentation on base. Caudal fin with small, elongated blotches on rays, forming three to four bands in most individuals, without bands in others. Distal margin of caudal fin unpigmented. Colouration in life. Similar to that observed in preserved specimens, but with fins and maxillary barbels orangish yellow to orange, more evident in paired fins.	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFD3FFD6256FF92E20B20395.taxon	discussion	Remarks (variability). All specimens identified as Barbatula fluvicola were readily identifiable by their overall body and head shape and proportions. In some geographically restricted areas, phenotypes of some individuals did not match the species assignment based on genetic barcode (see in Molecular analyses and species delimitation Section). This was the case in the streams Jona and Chli-Aa, both in the Limmat drainage (listed as non-type material), near to the confluence with Lake Zürich, where a few individuals are intermediate in phenotype and colour pattern between B. fluvicola and B. ommata n. sp., suggesting hybridisation in a secondary contact zone. Two individuals from Lake Zürich, near to the inflow of the stream Jona (NMBE 1071779 and NMBE 1071775), were molecular assigned to B. fluvicola, corroborating this hypothesis. On the other hand, the loach population of Constance and Geneva lakes recovered as nested in the B. fluvicola clade in coI tree were differentiated in several traits from B. fluvicola, such as having larger snout length and eyes, and a narrower head in adults, but recovered these populations nested. Since we are uncertain if these populations from the Constance and Geneva lakes represent hybrid populations in a contact zone or a distinct species closely related to B. fluvicola, which has recently diverged (see discussion); we are herein eventually assigning those populations as Barbatula aff. fluvicola until further work can resolve their status (see Comparative Material section). The molecular analysis further assigned part of the population from Schwarzwasser stream in the Sense River catchment (NMBE 1080648, NMBE 1080650, NMBE 1080651), Sense River itself (NMBE 1080702, NMBE 1080703, NMBE 1080692, NMBE 1080750, NMBE 1080751, NMBE 1105471 and NMBE 1105408), Saane River (NMBE 1086859 and NMBE 1105452) and some individuals of Furtbach in the Limmat River catchment (NMBE 1083146, NMBE 1083148, NMBE 1083143, and NMBE 1083144) to B. ommata n. sp. However, these individuals clearly share the overall morphology and diagnostic characters of B. fluvicola, and they are herein assigned as such. We hypothesised an indicative of past introgression from the latter species (see discussion). Further clarification of the status of these populations will require a population genomic and phylogenomic analyses (Calegari et al. in prep.). For now, the few individuals with mismatch between phenotype and mitochondrial haplotype are listed as non-type material under B. fluvicola.	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFD3FFD6256FF92E20B20395.taxon	distribution	Distribution. Barbatula fluvicola is known from several tributaries of the upper Rhine drainage (here defined to include also high Rhine and Alpine Rhine drainage sections), including rivers in the Aare, Reuss and Limmat catchments. The species is absent from lakes in the Aare catchment. The species is also found in the upper Danube drainage in Germany and in the Inn drainage in Austria (vouchers listed in non-type series were morphologically and genetically confirmed). The overall distribution of the species comprises the upper Rhine drainage in Switzerland and Germany, and upper Danube River drainage in Germany, Austria and Switzerland (Figure 3).	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFD3FFD6256FF92E20B20395.taxon	description	Sexual dimorphism. Epidermical tubercles in pectoral fin present mostly in 1 st – 4 th branched rays in males (vs. tubercles absent or not evident in females and juveniles). Differences in pectoral-fin shape are easily noticed in adults, with males showing pointed pectoral-fin tip and enlargement of outermost branched-fin rays (vs. rounded in females); juveniles and some subadults showed no conspicuous differences in pectoral fin shape. Urogenital papilla of males relatively small and conical (vs. rounded in females).	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFD3FFD6256FF92E20B20395.taxon	etymology	Etymology. The name fluvicola is from Latin meaning inhabitant of rivers, alluding to the stream preference habitats where this species has been collected. A noun in apposition.	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFD3FFD6256FF92E20B20395.taxon	vernacular_names	Vernacular name. North-Prealpine Stone Loach (English), Nord-Voralpine Bartgrundel (German), Loche Préalpes du Nord (French).	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFD3FFD6256FF92E20B20395.taxon	discussion	Ecological notes. Specimens were collected in small to medium size rivers, clear water, with swiftly flowing water. This species inhabits at the bottom of rivers under stones and among pebbles and sometimes close to larger rocks when in more rapid deeper waters. Juveniles are often found associated with greater water moss (Fontinalis antipyretica) and algae, as well as aquatic plants, which seems to be an important habitat providing protection against predators. Water parameter ranges from sampled sites overall all seasons inhabited by B. fluvicola: temperature from 4.8 to 18.2 ǪC; pH between 7.57 and 8.4; conductivity from 350 μs / cm to 640 μs / cm. The species seems to be primarily insectivorous based on few individuals dissected for clearing and staining technique (listed in type material).	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFD3FFD6256FF92E20B20395.taxon	conservation	Conservation status. Barbatula fluvicola has a wide distribution and it is known in Switzerland from the Aare, Reuss and Limmat drainages, tributaries of the Rhine River, and from the upper Danube and its tributaries in Germany, and the Inn drainage in Switzerland and Austria. The extent of occurrence (EOO) is estimated in 47,420 km 2 by the Minimum Convex Polygon, and the area of occupancy (AOO) is about 188 km 2 (calculated using GeoCAT tool). However, it is expected that the species also occurs in similar habitats along other portions of its distribution in Upper Rhine and Danube drainages in Germany and Austria. Part of the distribution of the species in Switzerland (mainly subpopulations from Aare catchment around Bern and Fribourg cities, and Limmat catchment) is situated in large urban and agricultural areas that is experience impacts on its habitat quality due to urbanisation and pesticides. Additionally, almost all rivers inhabit by the species in Switzerland are fragmented by several artificial small to median barriers; however, this species is not migratory and may not represent a direct impact that could threaten the population long-term persistence. Barbatula fluvicola is relatively widely distributed, and despite it suffering continued negative impacts in its occupancy area due to decreasing habitat quality, those threats do not affect a majority of localities, where the species seems to be abundant. Thus, Barbatula fluvicola is preliminary categorised as Least Concern (LC) according to IUCN criteria (IUCN Standards and Petitions Subcommittee, 2022). However, it is recommended that monitoring of the species subpopulations and its area of occupancy be conducted for better understanding of its abundance and population stability.	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFC5FFD22697FF7E25E703B7.taxon	description	(Figures 11 and 12) urn: lsid: zoobank. org: act: 4 A 1 BCF 03 - B 054 - 41 B 6 - 8020 - F 34 B 01 B 13 ADE	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFC5FFD22697FF7E25E703B7.taxon	materials_examined	Holotype. NMBE 1105555, 74.1 mm SL, female, Corcelles-près-Concise beach, Lake Neuchatel, Aare catchment, Rhine drainage, Canton of Vaud, Switzerland, 46 Ǫ 5003300 N, 6 Ǫ 4204200 E, 20 Jul 2023, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C. Paratype. All from Switzerland: Rhine drainage: Aare catchment: Canton of Vaud: NMBE 1105553 – 1105554, 2, 69.1 – 73.7 mm SL, collected with the holotype. — NMBE 1105282 – 1105290, 9, 53.1 – 70.9 mm SL, Lake Neuchatel, 46 Ǫ 5104700 N, 6 Ǫ 5103800 E, 19 Jul 2023, Calegari, B. B., Wegscheider, B., Waldock, C., Josi, D. — NMBE 1105291 – 1105293, 3, 56.9 – 70.7 mm SL, Lake Neuchatel, Corcelles-près-Concise beach, 46 Ǫ 5004800 N, 6 Ǫ 4205900 E, 20 Jul 2023, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C. Canton of Neuchâtel: NMBE 1105309 – 1105311, NMBE 1105313, NMBE 1105315 – 1105326, NMBE 1105328, 17, 46.8 – 57.6 mm SL (HRXCT of NMBE 1105313), NMB 6482 – 6483, 2, 52.7 – 54.5 mm SL, ZFMK-ICH 135312, 1 C & S, 54.6 mm SL, Lake Neuchatel, littoral zone, 367 m alt., 47 Ǫ 008.500 N, 7 Ǫ 004000 E, 17 Jul 2022, Calegari, B. B., Wegscheider, B., Josi, D. — MHNN 89 - 2491 and MHNN 89 - 2492, 2, 64.1 – 67.2 mm SL, Lake Neuchatel, littoral zone at camping Paradis-Plage, 46 Ǫ 5800.400 N, 6 Ǫ 52025.500 E, 16 Oct 2024, Calegari, B. B., Sudasinghe, H. Canton of Fribourg: NMBE 1105261 – 1105264, NMBE 1105266 – 1105269, NMBE 1105272 – 1,105,278, NMBE 1105280, 48.7 – 66.2 mm SL, ZFMK-ICH 135309 – 135,311, 3, 62.9 – 65.6 mm SL, ZSM 49671, 2, 53.7 – 56.4 mm SL, NMW 101169, 2, 52.4 – 65.9 mm SL, Lake Neuchatel at Estavayer harbour, 46 Ǫ 4903100 N, 6 Ǫ 4006000 E, 28 Jun 2023, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C. Canton of St. Gallen: NMBE 1105245 – 1105249, 5, 49.1 – 75.7 mm SL, Lake Walen, near to Quinten, 47 Ǫ 705700 N, 9 Ǫ 1103200 E, 28 Jun 2023, Calegari, B. B., Wegscheider, B., Josi, D., Reichlin, P. — NMBE 1105237 – 1105238, NMBE 1105241 – 1,105,242, NMBE 1105244, 6, 51.3 – 82.3 mm SL, ZFMK-ICH 135313 – 135314, 2, 53.6 – 71.5 mm SL, Lake Walen, near to Quinten, 47 Ǫ 705200 N, 9 Ǫ 120200 E, 28 Jun 2023, Calegari, B. B., Wegscheider, B., Josi, D., Reichlin, P. — NMBE 1105250 – 1105258, 8 + 1 C & S, 56.7 – 71.7 mm SL, Lake Walen, near to Mols, 47 Ǫ 6051.400 N, 9 Ǫ 16055.800 E, 28 Jun 2023, Calegari, B. B., Wegscheider, B., Josi, D., Reichlin, P. Canton of Lucerne: NMBE 1105219 – 1105221, NMBE 1105224 – 1105226, NMBE 1105228 – 1105229, NMBE 1105231 – 1105235, 12 + 1 C & S, 50.6 – 71.1 mm SL, MNHN 2024 – 1385, 2, 55.0 – 63.8 mm SL, MNCN-ICTIO 298.404 and MNCN-ICTIO 298.405, 2, 56.5 – 68.2 mm SL, Lake Lucerne at Vitznau, 430 m alt., 47 Ǫ 001400 N, 9 Ǫ 290400 E, 27 Jun 2023, Calegari, B. B., Wegscheider, B., Josi, D., Reichlin, P. Canton of Nidwalden: NMBE 1105236, 1, 58.5 mm SL, Lake Lucerne, Beckenried, 440 m alt., 46 Ǫ 5801800 N, 8 Ǫ 2605700 E, 27 Jun 2023, Calegari, B. B., Wegscheider, B., Josi, D., Reichlin, P. Genseq- 2 coI. NMBE 1105553 (tissue tag 241420); Genbank Acession Number PQ 488228. Genseq- 2 coI. NMBE 1105282 (tissue tag 241424); Genbank Acession Number PQ 488229. Genseq- 2 coI. NMBE 1105291 (tissue tag 241441); Genbank Acession Number PQ 488230. Genseq- 2 coI. NMBE 1105310 (tissue tag 239503); Genbank Acession Number PQ 488120. Genseq- 2 coI. NMBE 1105245 (tissue tag 241280); Genbank Acession Number PQ 488223. Genseq- 2 coI. NMBE 1105248 (tissue tag 241283); Genbank Acession Number PQ 488224. Genseq- 2 coI. NMBE 1105250 (tissue tag 241286); Genbank Acession Number PQ 488225. Genseq- 2 coI. NMBE 1105261 (tissue tag 241392); Genbank Acession Number PQ 488227. Genseq- 2 coI. NMW 101169 (tissue tag 241410); Genbank Acession Number PQ 488226. Genseq- 2 coI. NMBE 1105237 (tissue tag 241240); Genbank Acession Number PQ 488221. Genseq- 2 coI. NMBE 1105220 (tissue tag 241212); Genbank Acession Number PQ 488217. Genseq- 2 coI. MNCN-ICTIO 298.404 (tissue tag 241214); Genbank Acession Number PQ 488218. Genseq- 2 coI. NMBE 1105226 (tissue tag 241218); Genbank Acession Number PQ 488219. Genseq- 2 coI. NMBE 1105236 (tissue tag 241228); Genbank Acession Number PQ 488220. Additional material (non-types). All from Switzerland. Canton of Bern: NMBE 1120429 and NMBE 1120489, 2, Lake Biel, 57.4 – 58.7 mm SL, 47 Ǫ 0404800 N, 7 Ǫ 11046.700 E, 20 Sep 2017, Décourcière, H. — NMBE 1120476 and NMBE 1120485, 2, 53.5 – 59 mm SL, Lake Biel, 47 Ǫ 03039.500 N, 7 Ǫ 10029.300 E, 20 Sep 2017, Décourcière, H. Canton of Fribourg: NMBE 1061957, NMBE 1061974, NMBE 1061976 – 1061978, NMBE 1061993 – 1061995, NMBE 1062005, 7 + 1 C & S, 53.2 – 76.3 mm SL, Lake Neuchatel, 46 Ǫ 52014.900 N, 6 Ǫ 52016.500 E, 3 Oct 2011, unknown collector. — NMBE 1061958, 1, 64.2 mm SL, Lake Neuchatel, at Sous la Corbière, 46 Ǫ 51056.900 N, 6 Ǫ 51050.300 E, 3 Oct 2011, Project Lac. — NMBE 1074762, 1, 52.3 mm SL, Lake Neuchatel, at Sous la Corbière, 46 Ǫ 5205.900 N, 6 Ǫ 51011.100 E, 7 Oct 2011, unknown collector. Canton of Neuchâtel: NMBE 1074763, 1, 53.6 mm SL, Lake Neuchatel, at La Grande Béroche, 46 Ǫ 5506.900 N, 6 Ǫ 48055.600 E, 7 Oct 2011, Project Lac. Canton of Lucerne: NMBE 1069200 – 1069202, 3, 54.0 – 60.3 mm SL, Lake Lucerne at Vitznau, 430 m alt., 47 Ǫ 001600 N, 8 Ǫ 290100 E, 20 Aug 2014, Vonlanthen, P. — NMBE 1066769 – 1066772, 4, 36.3 – 40.3 mm SL, Lake Zug, near to Immensee, 47 Ǫ 0603000 N, 8 Ǫ 2806000 E, 22 Aug 2013, Tourreau, G. — NMBE 1080978, 1, 59.9 mm SL, Steinibach, 47 Ǫ 1014.500 N, 8 Ǫ 17059.900 E, 9 Sep 2013, Vonlanthen, P. — NMBE 1080958 – 1080971, 14, 58.2 – 76.0 mm SL, Steinibach, Ringstrasse, 47 Ǫ 0054.500 N, 8 Ǫ 18035.500 E, 9 Sep 2013, Vonlanthen, P. Canton of St. Gallen: NMBE 1074613, 1, 69.3 mm SL, Lake Walen, near to Sargans, 47 Ǫ 0704900 N, 9 Ǫ 12016.300 E, 23 Oct 2012, Brodersee, J. — NMBE 1111831 – 111833, NMBE 1109966, 4, 32.1 – 76.1 mm SL, Rufibach River, Limmat drainage, northwest of the airport Schänis between Maseltrangen and Ussbühl, 413 m alt., 47 Ǫ 10043.700 N, 9 Ǫ 1044.800 E, 13 Sep 2022, Calegari, B. B., Wegscheider, B.; Josi, D., Sudasinghe, H. Canton of Schwyz: NMBE 1105516 – 1105530, 15, 54.9 – 88.8 mm SL, Canton of Schwyz, Hoggibach River, at side channel of Hintergraben / Linthkanal, Aare River, Reichenburg, 406 m alt., 47 Ǫ 10032.700 N, 8 Ǫ 59021.900 E, 14 Oct 2022, Calegari, B. B., Wegscheider, B., Josi, D., Waldock, C., Schmid, A., Moreau, S. Canton of Vaud: NMBE 1061953, NMBE 1061974 – 1061975, NMBE 1061992, NMBE 1062001, 4 + 1 C & S, 52.2 – 57.6 mm SL, Lake Neuchatel, at Corcelles-près-Concise beach, 46 Ǫ 50016.900 N, 6 Ǫ 42027.900 E, 3 Oct 2011, Project Lac. — NMBE 1061963 and NMBE 1061989, 2, 54.4 – 55.3 mm SL, Lake Neuchatel, at Corcelles-près-Concise beach, 46 Ǫ 5008.700 N, 6 Ǫ 42012.300 E, 3 Oct 2011, Project Lac. — NMBE 1062004, 1, 59.5 mm SL, Lake Neuchatel, near to La Poissine, 46 Ǫ 49035.100 N, 6 Ǫ 4109.100 E, 3 Oct 2011, Project Lac. Canton of Zürich: NMBE 1105329 – 1105338, 9 + 1 C & S, 36.9 – 51.6 mm SL, Lake Zurich at Linthkanal, 394 m alt., 47 Ǫ 1306.200 N, 8 Ǫ 56028.900 E, 11 Jul 2022, Calegari, B. B., Wegscheider, B., Josi, D. — NMBE 1071771, NMBE 1071774, NMBE 1071787, NMBE 1071789, NMBE 1071793 – 1071794, NMBE 1071797, 7, 36.6 – 49.8 mm SL, Lake Zurich, Richterswill, 47 Ǫ 1301000 N, 8 Ǫ 41033.300 E, 1 Oct 2014, Vonlanthen, P.	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFC5FFD22697FF7E25E703B7.taxon	diagnosis	Diagnosis. Barbatula ommata is a member of a monophyletic group of species, herein designated as Western Europe clade (including B. barbatula, B. hispanica, B. leoparda and B. fluvicola), which is diagnosed from all clade congeners, except by B. hispanica, by having a shallower head 38.4 % – 46 % HL (vs. 46.1 – 51.7 % in B. fluvicola, 51.7 – 57.3 % in B. barbatula and 51.5 – 58.3 % HL in B. leoparda). The new species differs from B. hispanica and B. fluvicola by having a larger eye 21.1 % – 28.3 % HL (vs. 16.8 % – 22.5 % and 16.1 % – 19.8 % HL). It is distinguished from B. leoparda and B. hispanica by having a shallower caudal peduncle 7.7 % – 10.9 % SL (vs. 12.1 % – 13.6 % and 12.1 % – 13.9 % SL). It is further distinguished from B. leoparda and B. barbatula by having a smaller suborbital depth 23.7 % – 31.2 % HL (vs. 32.2 % – 36.3 % and 33.1 % – 36.3 % HL). It further differs from B. leoparda by having a shorter lower lip 12.7 % – 17.1 % HL (vs. 17.4 % – 21.2 % HL), and ventral surface of head and area between pectoral to pelvic-fin origins without pigmentation (vs. ventral surfaces with brown to dark irregular blotches). B. ommata is further distinguished from B. barbatula by having a shorter dorsal head length 19.3 % – 21.4 % SL (vs. 23.2 % – 25.8 % SL), a shorter first dorsal-fin ray 13.5 % – 19.8 % SL (vs. 20 % – 24.3 % SL), a shorter dorsal-fin base 11.1 % – 14.3 % SL (vs. 14.5 % – 18.6 % SL), a shorter dorsal fin length 18.4 % – 23.3 % SL (vs. 24.1 % – 25.8 % SL), a shorter prepectoral length 21.3 % – 25.8 % SL (vs. 26.2 % – 28.7 % SL), a longer caudal peduncle 12.7 % – 16.2 % SL (vs. 11.7 % – 12.7 % SL), a shorter snout length 37.2 % – 46.1 % HL (vs. 50.8 % – 52.6 % HL), a weekly developed slope present between anterior nares (vs. protuberant slope from preceding area of anterior nares) and adpressed anal-fin tip reaching about middle of caudal peduncle (vs. adpressed anal-fin tip slightly anterior to end of caudal peduncle). Barbatula ommata differs from B. fluvicola, which have widely overlapping distribution ranges, by having ventral border of dentary straight (vs. ventral border of dentary curved; Figure 8), shorter coracoid, half-length of cleithrum (vs. coracoid long, occupying at least two thirds of cleithrum length; Figure 7), larger swimbladder bony capsule, occupying slightly more than half of head depth (Figure 13) (vs. smaller swimbladder bony capsule, not reaching half of head depth), and posterior border of swimbladder bony capsule rounded in ventral view (Figure 13) (vs. posterior border of swimbladder bony capsule straight in ventral view).	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFC5FFD22697FF7E25E703B7.taxon	description	Description. Morphometric data and counts are provided in Table 2. Body robust and elongate, cylindrical in cross-section at trunk. Dorsal profile of trunk approximately straight descending from dorsal-fin origin to middle of caudal-fin peduncle; ascending from that point to caudal-fin origin. Ventral profile of trunk straight, except from anal fin descending towards caudal-fin end. Dorsal head profile slightly concave from snout tip to posterior border of supraoccipital. Trunk depth similar along its length slightly decreasing posteriorly dorsal-fin base towards middle of caudal-fin peduncle. Greatest body depth slightly anterior to dorsal-fin origin. Body width decreasing and gradually compressed from pectoral-fin origin towards caudal fin. Greatest body width at dorsal-fin origin. Head cylindrical in dorsal view, longer than wide. Eyes laterodorsally positioned, located at middle of head length. Ocular capsule rounded to slightly ellipsoid. Nostrils small; anterior naris opening slightly smaller than posterior one. Anterior nostril with an elongate fleshy flap-like tube oriented laterodorsally. Posterior naris in oval shape with anterior border slightly pointed. Anterior and posterior nostril close to each other and connected by thin skin ridge. Ventral border of gill opening attached to body at line of pectoral-fin base. Mouth ventral, arched in a semicircular shape. Mouth wide, almost equal to head width. Gape twice times longer than deep. Upper lip somewhat slim, leaving exposed middle portion of premaxilla area. Median incision usually deep, occupying approximately entire lip depth; poorly developed in some individuals. Lower lip separately at midline by a deep notch. Mesial portion of lower lip developed in a fleshy fold in inverted U-shape forming a mental lobe. Mental lobes usually separate each other at mid-line along entire length; without protrusion. Anterior border of lower lips in majority of specimens not covering anterior border of dentary, leaving this portion visible. Lateral portion of lower lip positioned inclined in 45 Ǫ angle to vertical midline, not bearing evident furrows; posterior tip without projection. Lower and upper lips rugous covered with small rounded papillae in its entire length. Barbels tapering gradually towards tips. Two pairs of premaxillary barbel of similar width. Tip of mesial premaxillary barbel reaching or almost reaching to base of maxillary barbel. Lateral premaxillary barbel tip surpassing middle of posterior naris usually reaching anterior border of eye. Maxillary barbel slightly surpassing vertical of middle of eye. Pectoral-fin rays i, 11 * (31). Innermost soft ray unbranched and tiny, usually visible only in C & S specimens. Distal margin of pectoral fin truncated or convex. Principal unbranched pectoral-fin ray slightly longer than two-thirds of longest pectoral-fin ray. Pelvic-fin rays i, 7 * (31). Pelvic fin ellipsoid, its origin at vertical through second branched dorsal-fin ray. Pelvic fin with distal margin convex, its tip slightly anterior to vertical of end of adpressed dorsal fin. Axillary pelvic lobe present, conspicuous attached to body, in oblate ellipsoid shape. Dorsal-fin origin located slightly after middle of body; distal margin slightly convex to truncated. Dorsal fin with 3 small (6 C & S), unbranched rays, and i, 7 ½ * (30) or i, 8 ½ (1). Dorsal fin with 9 (4 C & S) or 10 (2 C & S) pterygiophores. First dorsal-fin pterygiophore composed by what seems a supraneural and first proximal radials fused each other in a single complex structure supporting 3 shorter simple rays + a principal longest unbranched ray. Remaining 8 or 9 pterygiophores with posterior laminar bony keel, supporting their respectively branched ray, except last pterygiophore lacking any correspondent ray, and smaller in size. Ultimate 1 ½ dorsal-fin rays supported by the penultimate pterygiophore. Principal dorsal-fin unbranched ray shorter than the subsequent outermost three branched rays, reaching two-thirds of its length. First dorsal-fin pterygiophore inserted anterior to neural spine of 13 – 14 th vertebrae (6 C & S). Distal margin of anal fin slightly concave due to longer mid-rays. Anal fin with 2 – 3 shorter simple rays (6 C & S) plus i, 5 ½ * (25) rays. Two to 3 simple rays + first unbranched principal ray supported by composed structure of at least 3 proximal radials fused in a single element; remaining branched rays supported by 5 pterygiophores; ultimate pterygiophore supporting 1 ½ rays. Anal-fin origin located posterior to vertical of dorsal-fin end. First pterygiophore inserted anteriorly to hemal spine of 25 th or 26 th vertebra (6 C & S). Caudal fin weakly emarginate (rarely truncated). Caudal-fin rays i, 7 + 6, i (1), i, 7 + 7, i (2), i, 8 + 7, i * (23), or i, 8 + 8, i (5 C & S). Procurrent rays in dorsal lobe 7, and ventral lobe with 6 rays. Procurrent rays pronounced and embedded in skin forming a dorsal and ventral adipose keels extended on up to half of caudal-peduncle length. Ventral adipose keel slightly elevated, shorter than dorsal keel. Ventral margin of caudal fin sometimes somewhat convex, while dorsal margin approximately straight in most of specimens; rarely straight in both margins. Epural reaching half-length of caudal skeleton. Upper hypural plate as a single element with hypurals 3, 4 and 5 articulated, but not fused, separate each other along entire length. Lower hypural plate with co-ossified parhypural, and hypurals 1 and 2 separated each other, forming the compound caudal centrum. Total vertebrae, 39 (3 C & S) or 40 (3 C & S); 13 pairs of ribs (6 C & S) gradually decreasing length towards caudal fin. Body covered by embedded tiny, rounded, not overlapping scales, separated by a distance larger than scale diameter. Scales irregularly set along trunk. Scales present on flank from anterior vertical of dorsal-fin origin to caudal-fin base. Ventral portion of body with scales present only between end of pelvic-fin girdle to anal-fin origin. Scales completely absent on predorsal region and head. No evident developed dermal tubercles in any mature or juvenile individuals, even at reproductive season. Lateral line incomplete, ending before caudal-fin base approximately at vertical of end of anal fin. Lateral line quite variable with 32 – 55 (6 C & S) canals; first three to four canals well ossified. Lateral line interrupted usually at vertical through dorsal-fin origin continuing approximately at vertical of end of anal-fin base. Lateral line canals drastically decreasing size after first half of trunk and more spaced each other towards caudal fin. Cephalic lateral sensorial system with canals only partially ossified, between pores, sometimes weakly ossified. Infraorbital canal 11 – 12 pores (two pores + posterior-half pore of antorbital segment, and anterior-half pore + nine pores + posterior-half pore of suborbital segment). Infraorbital canal composed by the antorbital branch bearing two pores (anterior and dorsomedial) and posteriorly half-pore, close but separately from anterior-half pore of adjacent suborbital branch, followed by nine to 10 pores and ending in posterior-half pore. Posterior-half pore of suborbital segment connected to postorbital canal with anterior-half pore, plus three pores and ending in posterior-half pore connected to half pore of both supratemporal and lateral line canal. Supratemporal canal with two and half pores. Nasal canal with three pores (anterior, mid-lateral, and posterior) usually unossified (rarely weakly ossified). Frontal canal bordering orbital border, but truncated at mid-portion splitting in two branches, each one bearing three pores (anterior, lateral and posterior). Preoperculomandibular canal with eight to nine pores with anterior part located ventrally to articular and posteriorly to dentary. Trunk canal short with two pores limited by pectoral-fin girdle. Nasal canal present with three pores (Figure 1). Colouration (in alcohol). Overall background colouration pale yellow. Body trunk covered by light to medium brown pigmentation forming irregular roundish blotches, randomly distributed (Figure 11). Pigmentation more concentrated on dorsal surface of body, with three to four brown blotches on predorsal region, and three blotches on postdorsal region, less evident in most individuals. Blotches mostly separately from each other, but coalescent. Lateral line portion usually bearing a faint brown stripe formed by coalescent blotches of pigmentation, sometimes more conspicuous and continuous at middle to end of trunk. Pigmentation fading abruptly in ventral portion of head and body, between pectoral-and-pelvic fins origins, and abdominal region. Head with more uniform dark-brown pigmentation along dorsal profile by concentration of pigmentation. Suborbital area and distal margin of opercle pale yellow to whitish with small, sparse pigmentation. Dark-brown, narrow stripe present on prenasal area, extending from snout tip to anterior portion of eye, but interrupted by nasal opening. Specimens varying in pigmentation over axillary pelvic-fin keel and at dorsal surface of pectoral-fin base, from barely present to conspicuous darker brown colour. Pigmentation absent on maxillary barbel, but present and faded in premaxillary barbels, usually restricted to base of barbel, rarely reaching its half length: mesial premaxillary barbel more pigmented than lateral pair. Upper and lower lips deprived of any pigmentation, except by the anterior area preceding upper lips. Pectoral fin with elongate dark brown blotches over rays reaching almost entire length, except by posterior margin. Outermost six pectoral-fin rays with larger blotches, more concentrate along anterior half of fin length; fainting towards mesial portion. Concentrated dark-brown pigmentation present on dorsal-fin base and at distal portion of fin; fading at middle portion of rays. Pigmentation on dorsal-fin rays forming or not rectangular dark bars over all rays, distributed irregularly; more conspicuous on first unbranched ray. Pelvic fin with no pigmentation or rarely showing few faint pigments concentrated only on distal portion of fin. Pigmentation on anal fin mostly absent, except by distal portion with faded small pigmentation. All fins lack pigmentation on ventral surface. Caudal-fin rays covered by small irregular pigmentation over entire length, except on distal margin. Elongated dark-brown to black blotch on caudal-fin base. Colouration in life. Similar to that observed in preserved specimens but with ground colour of body more pale yellow and mid-portion of lateral body with gold iridescent coloration in some individuals.	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFC5FFD22697FF7E25E703B7.taxon	distribution	Distribution. Barbatula ommata is known only from the lakes Neuchatel (Figure 6 b), Lucerne, Walen, Zurich, Zug, Biel and Murten, all systems of the Aare drainage in Switzerland (Figure 3). The species has been further historically reported from lakes Thun and Brienz (Figure 3), but its continued occurrence in these systems remains uncertain due to the lack of recent records, highlighting the urgent need for further surveys to confirm its persistence and presence in these lakes.	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFC5FFD22697FF7E25E703B7.taxon	discussion	Remarks. There are four records of Barbatula in the InfoFauna database (www. infofauna. ch), dated 1988, 1998, 2001 and 2011, from two sites in Lake Thun and one record (1998) from Lake Brienz. These historical records lack accompanying photographs, voucher specimens or precise geographic coordinates, but given the ecological differences between the two new sympatric species, only B. ommata inhabits Aare lake systems. We therefore assign these historical records tentatively to this species. Despite considerable sampling efforts using electrofishing, minnow traps and nets across different seasons and years, no Barbatula specimens were observed from lakes Thun and Brienz after 2011. Given the very limited number of records, each separated by a decade or more (except for 2001), we assume that B. ommata previously inhabited these lakes but that its population has since declined drastically, likely potentially approaching local extinction. Given the limited historical records of the species from lakes Thun and Brienz and the lack of recent confirmed occurrences, a targeted search for these subpopulations is recommended to clarify their potential persistence in these lakes and assess the species' conservation status. Additionally, only three specimens of B. ommata were recorded in Lake Murten (also known as Lac de Morat) in 2010, two of which were detected during Project Lac and one in NAWA monitoring. The InfoFauna database contains just two further records from this lake, in 1985 and 2001. A similar situation applies to Lake Zug, where there are only four specimens recorded in 2013 during project Lac. Data indicate that B. ommata is present at low abundance in Lakes Murten and Zug. Given the declining habitat quality in these lakes, we recommend systematic monitoring to assess the population dynamics and conservation status of this species. Such monitoring is crucial for developing effective conservation strategies to mitigate further habitat degradation and support the survival of B. ommata in these lakes. Sexual dimorphism. Barbatula ommata shows epidemical tubercles in the pectoral-fin rays of adult males. The tubercles are present mostly on the innermost fourth branched pectoral-fin rays (vs. tubercles absent or not evident in females and juveniles). Difference in pectoral-fin shape is easily noticed in adults, with male showing pointed pectoral-fin tip and enlargement of outermost branched-fin rays (vs. rounded in female, and fin rays similar in width). Urogenital papilla of males small and conical (vs. urogenital in females rounded).	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFC5FFD22697FF7E25E703B7.taxon	etymology	Etymology. Barbatula ommata from the Greek ómmata (ὄμμ ατα) for eyes, and is given in reference to the species diagnostic great diameter of its eyes. A noun in apposition.	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFC5FFD22697FF7E25E703B7.taxon	vernacular_names	Vernacular name. Lake Stone Loach (English), Seebartgrundel (German), Loche du Lac (French).	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFC5FFD22697FF7E25E703B7.taxon	discussion	Ecological notes. Specimens were collected in the littoral zone of lakes. Collected on the shore down to 1.2 m water depth, most abundantly found in small pebbles substrate, and among middle-sized stones (~ 20 cm). Barbatula ommata and Cottus sp. are syntopic and were collected together. Few individuals dissected indicate that the species is primarily insectivorous.	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
2722A163FFC5FFD22697FF7E25E703B7.taxon	conservation	Conservation status. Barbatula ommata seems endemic to Switzerland, with its distribution largely confined to lakes within the Aare catchment, including Neuchatel, Biel, Murten, Lucerne, Walen, Zug and Zurich (Figure 3). Despite extensive sampling efforts, the species has not been observed in lakes Thun and Brienz in the past 15 years, despite sporadic historical records. Assessing whether the species is approaching local extinction in these lakes is critical, as such a loss would account for 25 % of its overall distribution and one entire subcatchment of the four within its original range. The Extent of Occurrence (EOO) based on the extant range of the species was estimated at 4302 km 2 by the Minimum Convex Polygon (calculated using GeoCAT tool) meeting the IUCN criteria of Endangered (B 1: EOO <5000 km 2). The area of occupancy (AOO) based on IUCN methodology (2 × 2 km grid) of known records was estimated at 132 km 2 meeting the criteria of Endangered (B 2: AOO <500 km 2). The AOO for the entire lake area for all lakes where the species occurs was also estimated at 490 km 2, which represents an overestimation because this species is confined to shallow littoral zones of the lakes (<1.5 m depth) requiring pebbles and small stones substrate, and it is absent from deeper areas as well as large boulders and rock faces. Barbatula ommata has a restricted distribution driven by its specific habitat requirements. Accordingly, we estimate the extent of suitable habitat within the very restricted range for the species at 26.2 km 2, considering only the shoreline areas of all lakes and excluding deeper zones where the species does not occur. We used a geospatial buffer from the shoreline to make this calculation, assuming all habitat within 50 m of the shoreline was habitable within all lakes where the species has recently been recorded (Zurich, Neuchatel, Murten, Biel, Lucerne, Walen and Zug; note that the median distance from the shoreline for B. ommata records was 13 m). The distribution of B. ommata in Swiss lakes is severely affected by urbanisation. Many areas are impacted by habitat degradation, primarily from urbanisation, pollution and the alteration of littoral zones caused by constructions, harbours, boulders and retaining walls, yet the impact degree of these modifications remains largely unknown. The loss of suitable habitats, already highly restricted (26.2 km 2), poses a significant threat to the species, which is confined to shallow littoral zones, absent from deeper lake areas and dependent on pebble and small substrate for survival. This specific habitat has currently a patchy distribution in the lakes that B. ommata occurs in, which further isolates subpopulations within the lakes of this species whose range is already naturally fragmented across lakes. Additionally, the extensive recreational use of the last natural lake shores, particularly in the summer, has potentially caused a continued decline of habitat quality and area of occupancy due to this species having its reproductive activity, feeding areas and nursery grounds in the littoral zones. Climate change is also an important factor that has been negatively impacting cold-water lakes due to the increase of water temperature in the shallow inshore zones in summer, despite it being difficult to measure the extent of impact. Barbatula ommata is naturally confined to large, oligotrophic, clear water lakes and its range is already naturally severely fragmented. Many of these lakes are surrounded by urban areas undergoing significant habitat degradation, with a notable loss of suitable habitat across multiple locations. Subpopulations in lakes Thun, Brienz, Murten, Biel and Zug are very small and have not been observed for over 12 years despite targeted collection efforts. These lakes together represent approximately 50 % of the species' distribution within the lake system. The prolonged absence of B. ommata suggests a significant reduction in the AOO of the species and the population is assumed to be severely fragmented. Thus, Barbatula ommata is preliminary assessed as Endangered (EN) based on the criteria B 1 ab (ii, iii) + B 2 ab (ii, iii), according to IUCN criteria (IUCN Standards and Petitions Subcommittee, 2022).	en	Calegari, Bárbara B., Freyhof, Jörg, Waldock, Conor, Wegscheider, Bernhard, Josi, Dario, Rüber, Lukas, Seehausen, Ole (2025): Two new species of stone loaches of the genus Barbatula (Cypriniformes: Nemacheilidae) from Europe with a neotype designation of B. barbatula (Teleostei: Nemacheilidae). Journal of Fish Biology 107 (4): 1364-1397, DOI: 10.1111/jfb.70108
