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Rhyacophila septentrionis McLachlan 1865 Description of the larva Colour: Live specimen golden-olive (Hickin 1954), pale ventrally. Specimens preserved in ethanol reddish brown, dorsum dark, ventrally pale; on abdomen, semicircular-triangular pale anterior dorsolateral areas, dorsocentral area of segments with darker circular pale spots, connected caudally with anterior dorsolateral areas through pale lines, forming W-shape. Sclerotized areas of head and thorax dark brown with pale areas, black in some places (Fig 22a). Head (Figs 14, 22b). Maximum width in posterior third of head, lateral margins almost parallel (tapering slightly towards anterior end, but not constricted at level of eyes, Mackereth 1954, fig 1S; Hickin 1967, fig 40; tapering anteriorly, Edington & Hildrew 1981). Cephalic capsule (Figs 14a, 14b), dorsally and dorsolaterally dark brown, anterior third and areas adjacent to frontoclypeal and anterior coronal sutures and anterior dorsolateral areas paler (dorsal posterior region of head with conspicuous pattern of dark spots on lighter background, Edington & Hildrew 1981, Fig 33, arrow a); ventral side pale, with central and slightly darker U-shape area from middle to posterior end of protogula covered with pale spots (Fig 14c), aboral margins of genae just before union with small black fold (Hickin 1954, fig 4B, 1967, fig 54); in lateral view anterior edge and submentum reddish brown, posterior edge black; oval brown area with dark spots extending laterally from anterior edge of occipital foramen to anterior third of head (Fig 14b). Frontoclypeal apotome (Fig 14d) pale; 2 oval dark brown submesal muscle attachment spots in anterior third between setae #4; dark brown pigmentation in posterior third heart-shaped (crescent shaped, Mackereth 1954, fig 1S; Hickin 1967, fig 40), including pits #4 but not setae #6, with pair of oval lateral and 2–3 circular mesal paler brown muscle attachment spots with dark borders, anterocentral area pale with two oval pale brown spots in anteromesal position (Figs 14a, 22a, 22b). Two-thirds of posterior dorsal area of head brown, marked with darker and paler muscle attachment spots; submesally one pair of dark brown triangular patches with paler spots with dark borders outside posterior edges of frontal sutures; light triangular area on juncture of frontal and coronal sutures, small pale area around pit #9; three pale spots around setae #15, #16, and #19–21, last one connected with posterior pale areas; dark brown dots in posterior area of genae (Mackereth 1954, fig 1S; Hickin 1967, fig 40); dorsal head pits with dark brown borders, posteromesal edge of occipital foramen black. Mandibles (Fig 15) asymmetrical, as in other rhyacophilids; left one bigger than right one. Subapicomesal blade of left mandible slightly concave, right mandible with small tooth in middle, both with darker apices (Figs 15, 22a). Labrum and maxillolabium as in other Rhyacophila species. Mentum rectangular, its anterior half slightly sclerotized. Labrum with small transparent area in central anterior edge. Thorax (Figs 16, 22c): Anterior half of pronotum pale or very light coloured, posterior half with three darker areas, one median and two lateral (Edington & Hildrew 1981, fig 35; Hickin 1954, fig 5B, 1967 fig 56), in darker specimens posterior half brown, with mesal area darker. Sinuous posterior margin bordered by black band, posteriorly and posterolaterally light brown submesally; posterior half of pronotum (Fig 16a) dark, with narrow semicircular area extending anterolaterally and small area laterally, lateral area pale in each pronotal plate; central dark region triangular, tapered posteriorly and reaching posterior edge of sclerite, anteriorly truncate and wider than posteriorly, pale brown spots inside triangular region reaching posterior edge; each side of pronotal plate with two or three lines of pale brown spots with dark border of different shapes and parallel to posterior border; pale areas around setae 2–3#, 5# and 6#, four–six brown spots around setae #5; curved row of two or three dark dots extending from setae #5 toward anterolateral corner, two pale brown spots with dark borders anterolateral and posterolateral of seta 6#, circular and elongate respectively (Figs 16b, 22c); with narrow black band in anterolateral angles, extending posterad laterally above lateral margins to posterolateral corners (Fig 16b); anterior margin light brown, with a line of black spots near anterior angle; mesal area of posterior marginal band black (Fig 16b, 22c). Legs: Similar to those of other Rhyacophila species. Colour pale yellowish, each with tarsus and claw darker, proximal parts of coxa, femur and distal parts of trochanters of fore- and midlegs black or dark brown. Narrow pale dark brown band in proximal dorsolateral area of tibia and distal dorsolateral area of femur of each pair of legs, each inner and outer face of each femur with band of dark brown circular spots. Forelegs each ventrally with spicules, extending from distal half of trochanter, all of femur, tibia and tarsus. Setae black or dark brown, some transparent in ventral area. Abdomen (Figs 17, 18, 22d, 22e): Similar to other Rhyacophila species. Colour as for meso- and metathoracic segments, in preserved larvae semicircular-triangular pale anterior dorsolateral areas, dorsocentral area of each segment with darker circular pale spots, connected caudally with anterior dorsolateral areas through pale lines, forming W-shape, pale yellow ventrally. Widths of anterior abdominal segments similar, narrower in posterior segments. Lateral abdominal gills in tufts of 25–30 filaments. Dorsal sclerite of segment IX (Figs 17, 22d) light coloured, darker in central area, with continuous black band on anterior edge and two discontinuous black bands on posterior edge, posterior to setae #2; band of anterior edge narrower in center, surrounding anteriorly pit #1; pale spots inside central area, two pale areas laterally. Anal prolegs typical of group, each with long sword process (Figs 18a, 18b, 22e). Anal claws each with three short teeth on ventral edge (Figs 18a, 18c, 22e), basoventral hook black basally and reddish brown apically (Figs 18a, 18b). Description of the pupa Biometrics of pupa and cocoon. Male pupa: 10.00 mm (1 specimen); female pupa: 11.60 mm (1 specimen). Cocoon length: 14.40 mm (1 specimen). Head: Distribution of setae on dorsal head similar to R. fasciata. Mandibles dark brown, left one with two large teeth and right one with three, anterior tooth bigger, with numerous fine teeth on inner blade. Antennae of variable length, reaching posterior edge of segment VII and posterior edge of segment IV in male and female studied, respectively. Thorax: Tubercles of prothorax each with 6 setae on male and 4 setae on female studied. Hind wing pads reaching posterior edge of abdominal segment III of male and posterior edge of segment IV of female studied. Abdomen: Paired anterior hook plates pedunculated, almost circular and present from abdominal terga IV to VII (Fig 11), more elongated on segment VII, each with more than 30 spines; also tergum III with pair of small flat hook plates each with 10–15 spines. Paired posterior hook plates sessile on terga III–V, almost circular, each with more than 30 spines. Description of the imago Male body length: 7.70 mm (1 specimen) each forewing: 12.20 mm, each hind wing: 10.60 mm; female, length: 8 mm (1 specimen), each forewing: 11.50 mm, each hind wing: 10.30 mm. Colour: Specimens preserved in ethanol dark brown dorsally, yellowish ventrally, spurs yellowish brown, setae brown, females generally darker than males; small black or dark brown spots dorsally. Forewings dark with small pale spots, pale in preserved specimens, darker in females, hind wings pale; each with dark area on leading edge between subcostal (Sc) and middle space between first radial (R 1) and second radial (R 2) veins in fore- and hind wings, with some darker stripes; each forewing hyaline in circular-oval zone, connecting medial (M) and medio-cubital (m-cu) veins and in distal area of second cubital vein (Cu 2) (Holzenthal et al. 2007); in each hind wing proximal segment of medial veins and connection area (M 1-4), not conspicuous, second cubital (Cu 2) vein not conspicuous either totally or partially. Male genitalia (Figs 19, 24, 31b, 31c, 34): Apical segment of each inferior appendage (Figs 19A, 24a, 34c) with basal and distal edges diverging, dorsal and ventral edges slightly diverging, posterior edge of 2nd segment of each inferior appendage slightly concave, ventral edge slightly convex, 2.5 times longer that dorsal edge (Figs 19A, 24a). Apicodorsal vertex angular, apicoventral angle rounded, projecting as thick lobe narrowing progressively to apex. Parameres in ventral view curved posteromesad in apical half (Figs 19 BV p, 24c, 34d). In lateral view (Fig 19 BL) each constricted at short distance beyond base, dilated at middle, with rounded dorsal margin, without spines in midventral margin, few long and thick spines on lateroventral area; midlateral surface covered by fine spicules or setae from middle to ventral edge, absent in middle anteroventral and posterodorsal edges. Aedeagus (phallicata) in lateral view (Fig 19 CL) with dorsal margin concave and posterior corner of concavity hooked anterad, posterior edge of aedeagus straight, projected posteroventrad in small ventral apex (Fig 19 CL); ventral lobe of aedeagus semicircular (Fig 19 BV vl). Lateroventral lobes of phallus straight, posterior edges strongly convex, apicolateral margins semicircular (Fig 19 CD lvl). Apicodorsal lobe of segment IX dilated subapicolaterally (Figs 19 DD al, 24b, 31b, 34b), preanal appendages (Fig 19 DD pa, 24b, 31b, 34b) round, almost as long as apicodorsal lobe, covered partially by it, mainly posteriorly; in ventral view apical band (Fig 19 DV ab, 31c) V-shaped, longer than wide, posterior edges rounded, almost parallel, non-sclerotized inside area (Fig 19 DV va) straight, anal sclerites triangular (Fig 19 DV as). Female genitalia (Figs 20, 26, 32a, 35): In lateral view (Figs 20L, 26L, 32a, 35c), posterodorsal margin of segment VIII valves on each side round, posterolateral margin with small excision in middle, ventral edge convex, dorsal edge with dorsolateral triangular projection (Fig 20L). In dorsal view (Figs 20D, 26D, 35b), with indentation between segment VIII valves, projections triangular with rounded corners, delimiting oval space (Fig 20D). In ventral view (Fig 20V, 26V, 35d) segment VIII valves forming two elongate sclerites, distal 2/3 of proximal edges diverging, distal edges straight or slightly concave; intersegmental membrane with pair of deep-dark, slightly elongate sclerites. Morphological characters diagnosing larvae of R. fasciata and R. septentrionis Head: In R. fasciata ventral side of head pale, in some specimens one or two rows of low conspicuous pale spots from posterior edge to halfway up the head. In R. septentrionis central area with slightly darker U-shape area from middle to posterior end of protogula, covered with pale spots. In R. fasciata the pair of dark brown patches outside posterior edge of frontal suture in the shape of open wings, with dark spots inside. In R. septentrionis the pair of dark brown patches more nearly triangular, with paler spots with dark borders inside. In R. fasciata mandibles dark reddish brown, uniformly coloured, left mandible with inner blade slightly convex. In R. septentrionis mandibles reddish brown, with darker apices, left mandible with inner blade slightly concave. Thorax: In R. fasciata on posterior half three dark areas, one median and two lateral, two lines of dark spots inside median area, circular spot on anterior half; posterior area with two to three rows of brown spots with dark border in circular-oval shape and parallel to posterior border. In R. septentrionis on posterior half three dark areas (in Hickin 1954) or dark base with darker mesal stripe with two lines of pale spots inside, two pale brown spots with dark borders anterolateral and posteriolaterally to seta 6#, circular and elongate respectively; posterior area with two-three lines of darkly-bordered pale brown spots of different shapes and parallel to posterior border. In R. fasciata anterior margin with black spot near anterolateral angle; mesal area of posterior marginal band of sclerite dark brown or black posteriorly. In R. septentrionis anterior margin with a line of black spots near anterolateral angle, light brown in middle; mesal area of posterior marginal band of each sclerite black posteriorly. Abdomen: In R. fasciata dorsal sclerite of segment IX uniformly coloured, brown spots in posteromesal area, band of anterior edge with two extensions surrounding pit #1, connected with dark stripes in darker larvae. In R. septentrionis sclerite of segment IX light coloured, mesal area slightly darker, with paler spots inside. Morphological characters diagnosing pupae of R. fasciata and R. septentrionis Abdomen: In R. fasciata paired anterior hook plates palmate, pair of small flat anterior plates without spines on tergum III; paired posterior hook plates slightly oblong transversally. In R. septentrionis paired anterior hook plates almost circular, pair of small flat anterior plates on tergum III with 10–15 spines; paired posterior hook plates almost circular. Morphological characters diagnosing males of R. fasciata and R. septentrionis Genitalia: In R. fasciata, posterior edge of 2nd segment of each inferior appendage slightly concave dorsally and convex ventrally, ventral edge slightly concave and 2.0 times longer than dorsal edge, apicodorsal vertex slightly angular. In R. septentrionis, posterior edge of 2nd segment of each inferior appendage completely concave, ventral edge slightly convex and 2.5 times longer than dorsal edge, apicodorsal vertex more conspicuously angular. In R. fasciata parameres in lateral view each slender near base, dilated in middle, with almost parallel dorsal and ventral margins in central area (S-shape), with two rows of long and thick spines on midventral margin, pointing outward; midlateral surface with sparse fine spicules or setae, from middle posteroventral to posterodorsal edge, covering most of apex, absent on middle anteroventral and dorsal edges. In R. septentrionis, parameres each slender near base, dilated in middle, with rounded dorsal margin, without spines in midventral margin, few long and thick spines on lateroventral area; midlateral surface covered by fine spicules or setae, from middle to ventral edge, absent on middle anteroventral and posterodorsal edges. In R. fasciata posterior edge of aedeagus straight, projected posteroventrad and rounded at posteroventral apex; ventral lobe of aedeagus triangular, lateral edges nearly straight; lateroventral lobes of phallus straight, posterior edges convex, apicolateral margins rounded. In R. septentrionis posterior edge of aedeagus straight, projected posteroventrad in small posteroventral protrusion; ventral lobe of aedeagus semicircular; lateroventral lobes of phallus straight, posterior edges convex, apicolateral margins semicircular. In R. fasciata apicodorsal lobe of segment IX slightly dilated subapicolaterally, almost round, with small apicomesal excision in some specimens; preanal appendages round, shorter than apicodorsal lobe, almost covered by it; in ventral view, apical band V-shaped and 1.8 times as long as wide, posterior arms widely separated, nonsclerotized area between arms of apical band heart-shaped and with posteromesal excision and posterior edges rounded. In R. septentrionis apicodorsal lobe of segment IX strongly dilated subapicolaterally; preanal appendages round, almost as long as apicodorsal lobe, covered partially by it, mainly posteriorly; in ventral view, apical band V-shaped and 1.2 times as long as wide, posterior arms close together and almost parallel, non-sclerotized area between arms of apical band straight and narrow and without posteromesal excision. Morphological characters diagnosing females of R. fasciata and R. septentrionis Genitalia: In R. fasciata in lateral view with dorsal margin of segment VIII 3/4 as long as venter and with prominent dorsolateral projection, posterolateral margin of valve on each side irregular, with 1 or 2 posterior projections, posteroventral projection more conspicuous, ventral margin slightly concave near base and straight or convex distally, in some specimens slightly indented posteriorly. In R. septentrionis in lateral view, with dorsal margin of segment VIII half as long as venter and with prominent dorsolateral projection, posterolateral margin of valve on each side round and with small excision in middle, ventral margin convex. In R. fasciata in dorsal view with indentation between segment VIII valves nearly enclosed by pair of thick posterolateral projections, with rounded apical and subapical corners, small projection on apical corner of some specimens. In R. septentrionis with indentation between segment VIII valves nearly enclosed by pair of triangular projections, with more-nearly acute apical corners. In R. fasciata in ventral view segment VIII valves forming two elongate sclerites, basal third of proximal edges parallel, distal edges irregular, straight, diverging; intersegmental membrane with pair of dark ovoid or slightly elongate sclerites. In R. septentrionis in ventral view segment VIII valves forming two elongate sclerites, basal 1/3 of proximal edges converging, distal edges straight or slightly concave, diverging; intersegmental membrane with pair of deep-dark slightly elongate sclerites. Genetic analysis Rhyacophila fasciata has a wide area of distribution in northern and central Europe and there is no GeneBank entry from the type locality (Elberfeld, Germany), so we have included some sequences from localities around it (Belgium and Germany). In the first analysis, we used these sequences, together with others from localities that previously have been checked as R. fasciata from Bosnia and Herzegovina, Belgium, Croatia, Czech Republic, Estonia, Finland, Germany, Kosovo, Norway, Poland, Republic of Macedonia, Russia, Slovakia and Sweden, together with one sequence of R. cf. nubila from Russia and three of R. cf. obliterata from Poland and Russia. In the second analysis, to build the tree, we selected a total of 52 sequences: a maximum of two sequences of R. fasciata per country (25 in total), as well as 3 sequences of R. septentrionis, 3 sequences of R. denticulata, 4 sequences of R. sociata from Spain and France (promoted to species by Valladolid et al. 2018), 3 sequences of R. kykladica (promoted to species by Valladolid et al. 2019), 7 sequences of R. fasciata delici Kučinić & Valladolid (in Valladolid et al. 2020) and 3 sequences of R. fasciata viteceki Valladolid & Kučinić 2020 (in Valladolid et al. 2020), 1 sequence of R. cf. nubila from Russia and 3 of R, Published as part of Valladolid, María, Arauzo, Mercedes, Chertoprud, Mikhail V., Chvojka, Pavel, Czachorowski, Stanisław, Dorda, Beatriz A., Hinić, Jelena, Ibrahimi, Halil, Karaouzas, Ioannis, Krpač, Vladimir, Kučinić, Mladen, Lodovici, Omar, Salokannel, Juha, Stamenković, Valentina Slavevska & Stojanović, Katarina, 2021, The Rhyacophila fasciata Group in Europe: Rhyacophila fasciata Hagen 1859 and formerly synonymized species (Trichoptera: Rhyacophilidae), with new description of Rhyacophila fasciata and Rhyacophila septentrionis McLachlan 1865 (stat. prom.)., pp. 1-57 in Zootaxa 4975 (1) on pages 23-33, DOI: 10.11646/zootaxa.4975.1.1, http://zenodo.org/record/4804044, {"references":["McLachlan, R. (1865) Trichoptera Britannica: A monograph of the British species of caddis-flies. Transactions of the Entomological Society of London, 3 rd Series, 5, 1 - 183. https: // doi. org / 10.1111 / j. 1365 - 2311.1967. tb 01433. x","Hickin, N. E. (1954) Larvae of the British Trichoptera. 42. Rhyacophila septentrionis McLachlan (Rhyacophilidae). Proceedings of the Royal Entomologica Society of London, Series A, General Entomology, 29 (4 - 6), 59 - 61. https: // doi. org / 10.1111 / j. 1365 - 3032.1954. tb 01199. x","Mackereth, J. C. (1954) Taxonomy of the larvae of the British species of the genus Rhyacophila (Trichoptera). Proceedings of the Royal Entomologica Society of London, Series A, General Entomology, 29 (10 - 12), 147 - 152. https: // doi. org / 10.1111 / j. 1365 - 3032.1954. tb 01189. x","Hickin, N. E. (1967) Caddis larvae. Larvae of the British Trichoptera. London, Hutchinson, 467 pp.","Edington, J. M. & Hildrew, A. G. (1981) A key to the caseless caddis larvae of the British Isles. Freshwater Biological Association, Scientific Publication No. 43, 91 pp.","Hagen, H. (1859) Die Phryganiden Pictet's. Entomologische Zeitung, 20 (4 - 6), 131 - 170.","Holzenthal, R. W., Blahnik, R. J., Prather, A. L. & Kjer, K. M. (2007) Order Trichoptera Kirby, 1813 (Insecta), Caddisflies. Zootaxa, 1668 (1), 639 - 698. https: // doi. org / 10.11646 / zootaxa. 1668.1.29","McLachlan, R. (1863) Notes on British Trichoptera, with description of a new species of Rhyacophila. The Entomologist's Annual, 1863, 129 - 136","Valladolid, M., Kucinic, M., Arauzo, M., Cerjanec, D., Cuk, R., Dorda, B. A., Lodovici, O., Stanic Kostroman, S., Vuckovic, I. & Rey, I. (2020) The Rhyacophila fasciata Group in Croatia and Bosnia and Herzegovina: Rhyacophila f. fa sciata Hagen 1859 and the description of two new subspecies, Rhyacophila fasciata delici Kucinic & Valladolid (ssp. nov.) from Croatia and Bosnia and Herzegovina and Rhyacophila fasciata viteceki Valladolid & Kucinic (ssp. nov.) from Bosnia and Herzegovina (Trichoptera: Rhyacophilidae). Zootaxa, 4885 (1), 051 - 075. https: // doi. org / 10.11646 / zootaxa. 4885.1.3","McLachlan, R. (1879) A Monographic Revision and Synopsis of the Trichoptera of the European Fauna, Part VIII. John Van Voorst, London, 1874 - 1880, pp. 429 - 500, plates 45 - 51. https: // doi. org / 10.5962 / bhl. title. 28556","Navas, L. (1916) Tricopteros nuevos de Espana. 1 ª Serie. Broteria, Serie Zoologica, 14, 5 - 11.","Malicky, H. & Sipahiler, F. (1993) Kocherfliegen (Trichoptera) aus der Turkei, mit Bemerkungen zu weiteren mediterranean Kocherfliegen. Bulletin de la Societe Entomologique Suisse, 66, 457 - 468.","Zetterstedt, J. W. (1840) Insecta lapponica. Voss, Lipsiae, 1068 pp.","Valladolid, M., Arauzo, M., Basaguren, A., Dorda, B. A. & Rey, I. (2018) The Rhyacophila fasciata Group in Western Europe: Confirmation of Rhyacophila denticulata McLachlan 1879 (stat. prom.) and Rhyacophila sociata Navas 1916 (stat. res.), based on morphological and molecular genetic evidence (Trichoptera: Rhyacophilidae). Zootaxa, 4418 (6), 526 - 544. https: // doi. org / 10.11646 / zootaxa. 4418.6.2","Valladolid, M., Karaouzas, I., Arauzo, M., Dorda, B. A. & Rey, I. (2019) The Rhyacophila fasciata Group in Greece: Rhyacophila kykladica Malicky & Sipahiler 1993 (stat. prom.) (Trichoptera: Rhyacophilidae). Morphological description, genetic and ecological features. Zootaxa, 4657 (3), 503 - 522. http: // dx. doi. org / 10.11646 / zootaxa. 4657.3.5"]}

Rhyacophila fasciata Hagen 1859 Description of the larva (Figs 4–9, 21). The description of the specimens examined will be complemented with descriptions of previous authors, adding or confirming features. Biometrics of last instar larva and prepupa: Length: 14.25–19.00 mm ( = 16.09, n = 13). Maximum width: 2.98–3.90 mm ( = 3.28, n = 13) (Lepneva 1964, 5 th instar larval length: 23.00– 23.40 mm; Nielsen 1942, 5 th instar larval length: 12.00– 23.40 mm; Ulmer 1909, larval length: 22.00–24.00 mm, larval width: 3.00– 3.50 mm). Maximum width of larva at metathorax. Colour: Live specimen greenish, dorsally with dark areas and light stripes, pale ventrally. Specimens preserved in ethanol reddish brown, dorsum dark, ventrally pale; on abdomen, semicircular pale anterior dorsolateral areas, dorsocentral area of segments with irregular pale spots, connected with anterior dorsolateral areas. Sclerotized areas of head and thorax brown with pale areas and stripes, black in some places (Fig 21a). Head (Figs 4, 21b). Length: 1.49–2.20 mm ( = 1.90, n = 13). Width: 1.14–1.50 mm ( = 1.30, n = 13). Maximum width in posterior third of head, lateral margins almost parallel. Cephalic capsule (Figs 4a, 4b), dorsally and dorsolaterally light brown in anterior third, areas adjacent to frontoclypeal and anterior coronal sutures and anterior dorsolateral areas paler; ventral side variable (Silfvenius 1905), pale (Pitsch 1993), or with dark spots in middle central area (Sedlák 1985, p. 111, table 3, fig 10); anterior edge and submentum reddish brown, posterior edge black (Nielsen 1942); oval brown area with dark spots extending laterally from anterior edge of occipital foramen to anterior third of head (Fig 4b) (Klapálek 1893; Lepneva 1964, p. 223, figs 181, 182B). Frontoclypeal apotome (Fig 4c) pale; 2 oval dark brown muscle attachment spots in central anterior third, between setae #4; dark pigmentation in posterior third heart-shaped, including pits #4 but not setae #6, with 4–6 oval brown muscle attachment spots with dark borders (Klapálek 1893; Silfvenius 1905; Nielsen 1942; Lepneva 1964; Pitsch 1993). Two-thirds of posterior dorsal area of head light brown, marked with darker muscle attachment spots; centrally one pair of dark brown patches, wing-shaped, outside posterior edge of frontal suture; light circular area on juncture of frontal and coronal sutures; three pale spots around setae #15, #16, and #19–21; dorsal head pits with dark border, central posterior occipital foramen black. Mandibles (Fig 5) asymmetrical, as in other rhyacophilids; left one bigger than right one. Inner blade of left mandible irregular, convex, right mandible with small tooth in middle (Nielsen 1942, pl. 2, fig 13; Lepneva 1964, p. 223, fig 184). Labrum and maxillolabium as in other Rhyacophila species (Nielsen 1942, pl. 2, figs 12A, 12B, 14A, 14B; Lepneva 1964, p. 223, figs 183A, 183B, 185). Mentum rectangular, its anterior half slightly sclerotized. Labrum with small transparent area in central anterior edge. Thorax (Figs 6, 21c): Anterior half of pronotum pale or very light coloured. Sinuous posterior margin bordered by dark band, brown in middle of each half, black posteromesally and posterolaterally, light brown submesally; posterior half of pronotum (Fig 6a) with three dark areas separated by light zones, central dark region tapered posteriorly and reaching posterior edge of sclerite, anteriorly truncate, wider that posteriorly, two lines of dark brown spots inside central dark region and reaching posterior edge, several dark brown spots along posterior edge; each pronotal plate with lateral dark area almost triangular, from anterior submesal region to posterolateral edge of sclerite, posterior area with two–three rows of brown spots with dark borders distributed in circular-oval shape and parallel to posterior border. Four–six brown spots around setae #5; curved row of dark dots from setae #5 toward anterior submesal corner, curving around circular spot on anterior half (Fig 6a); pale posterior and median areas connected (Fig 6b); with narrow black band in anterolateral angles, extending posterad laterally to posterolateral margins, inner to margins (Fig 6b; Lepneva 1964); anterior margin light brown, with black spot near anterolateal angle; central area of posterior marginal band of each sclerite dark brown or black (Fig 6b) (Nielsen 1942, pl.3, fig 17A; Lepneva 1964, p. 223, fig 188). Legs (Fig 9): Similar to those other Rhyacophila species. Colour yellowish, proximal regions of coxa, femur and distal region of trochanter of fore- and midlegs black or dark brown. Narrow pale reddish-brown band in proximal dorsolateral area of tibia and distal dorsolateral area of femur of each pair of legs, each inner and outer face of each femur with band of circular spots. Forelegs each ventrally with spicules, extending from distal half of trochanter, all of femur, and distal two-thirds of tibia and tarsus. Setae black or dark brown, some transparent in ventral area. Abdomen (Figs 7, 8, 21d, 21e): Similar to other Rhyacophila species. Colour as for meso- and metathoracic segments, greenish with dark dorsal areas and light stripes in living larvae, in preserved specimens semicircular pale anterior dorsolateral areas, dorsocentral area of segments with irregular pale spots, connected with anterior dorsolateral areas, pale yellow ventrally. Widths of anterior abdominal segments similar, narrower in posterior segments. Lateral abdominal gills in tufts of 25–30 filaments (20–30 filaments, Lepneva 1964). Dorsal sclerite of segment IX (Figs 7, 21d) brown, with continuous black band on anterior edge and two small and widely separated black bands on posterior edge, under setae #2 (Lepneva 1964); band of anterior edge narrower in center, with pair of narrow extensions backward beyond and surrounding pit #1; with some brown spots submesally near posterior margin; in darker larvae, extensions of anterior edge and spots of posterocentral area connected by dark stripes, creating X-shape figure; three pale areas along posterior margin, on either side of two black bands and central area. Anal prolegs typical of group, each with long sword process (Figs 8a, 8b, 21e). Anal claws each with three short teeth on ventral edge (Figs 8a, 8c, 21e), basoventral hook black basally and reddish brown apically (Figs 8a, 8b). Description of the pupa Biometrics of pupa and cocoon. Pupal length (total): 9.70–15.70 mm ( = 12.20, n = 28); male pupa: 10.07– 12.28 mm ( = 11.06, n = 11) (length: 11.20–12.20, Lepneva 1964; Nielsen 1942); female pupa: 9.70–15.70 mm ( = 12.94, n = 17) (length: 12.40–15.50, Lepneva 1964; Nielsen 1942; up to 16.00 mm, Silfvenius 1905). Cocoon length (total): 12.37–18.35 mm ( = 15.11, n = 68) (length: 12.50–18.00 mm, Nielsen 1942); male cocoon length: 12.39–16.27 mm ( = 14.43, n = 10); female cocoon length: 13.43–17.73 mm ( = 15.99, n = 16). Head: Distribution of dorsal setae: four before, two between and two in middle of antennal pads, three on posterior edge of each cephalic tubercle. Mandibles brown, left one with two large teeth and right one with three, with numerous fine teeth on inner blade (Klapálek 1893; Nielsen 1942; Ulmer 1909). Antennae of variable length, reaching from middle of abdominal segment IV to posterior edge of segment VI (to anterior edge of segment IV in females and posterior edge of segment VI in males, Klapálek 1893; to abdominal segment VIII in males and V in females, Lepneva 1964; Nielsen 1942). Thorax: Tubercles of prothorax each with 5 setae, directed backwards, two central setae directed forwards. Hind wing pads variously reaching from anterior edge of abdominal segment III to posterior edge of segment IV (forewing pads reaching anterior edge of abdominal segment IV, Klapálek 1893; forewing pads reaching abdominal segment V in males and IV in females, Lepneva 1964; hind wing pads reaching end of segment IV in females, Silfvenius 1905). Abdomen: Paired anterior hook plates pedunculated, straight basally and dilated posteriorly, palmate, and present on abdominal terga IV to VII (Fig 10) (Lepneva 1964; Ulmer 1909), slightly bigger on segments V and VI, each with 15–20 spines; also, pair of inconspicuous small flat plates without spines on tergum III. Paired posterior hook plates sessile on terga III–V, slightly oblong tranversally (length: width = 1:1.3) (Klapálek 1893: paired anterior hook plates present on abdominal terga III to VIII, paired posterior hook plates on terga III to V; Silfvenius 1905: paired anterior hook plates present on abdominal terga IV to VII, sometimes without plates on segment III, paired posterior hook plates present on terga III to V), each with variable number of spines, more than 25. Description of the imago Male body length: 7.04–10.46 mm ( = 9.23, n = 36) each forewing: 10.16–13.54 ( = 11.67, n = 36), each hind wing: 8.52–11.99 mm ( = 10.25, n =36); females, length: 8.73–12.19 mm ( = 10.46, n = 19), each forewing: 10.05–13.90 mm ( = 12.14, n = 19), each hind wing: 8.24–12.03 mm ( = 10.68, n = 19) [Ulmer 1909: male length (R. fasciata) = 10.00 mm, wingspan = 27.00 mm; male length (R. septentrionis) = 6.50–9.00 mm, wingspan = 22.50– 29.00 mm; Malicky 2004: male forewing length = 10.00–14.00 mm, female forewing length = 11.00–14.00 mm]. Colour: Specimens preserved in ethanol reddish brown dorsally, yellowish ventrally, spurs reddish, setae golden brown, females generally darker than males; small black or dark brown spots in dorsal area; in lateral view, some specimens with large black spots marking border between dorsal and ventral areas. Forewings dark with small pale spots, three transversal dark bands (fascias) in dry specimens (Fig 30a), pale in preserved specimens, darker in females, hind wings pale: each with dark area on leading edge between subcostal (Sc) and second radial (R 2) veins in fore- and hind wings; each forewing hyaline in irregular zone connecting medial (M) and medio-cubital (m-cu) veins and in distal area of second cubital vein (Cu 2) (Holzenthal et al. 2007); in each hind wing proximal segment of medial veins and connection area (M 1-4) not conspicuous, second cubital (Cu 2) and first anal (A 1) veins not conspicuous either totally or partially, proximal portion of cubital vein (Cu) and distal portions of first cubital vein (Cu 1a-1b) thicker. Male genitalia (Figs 2b, 12, 23, 30b): Apical segment of each inferior appendage (Figs 2b, 12A, 23a) with basal and distal edges diverging, dorsal and ventral edges diverging, posterior edge of 2nd segment of each inferior appendage slightly concave dorsally and slightly convex ventrally, ventral edge slightly concave (Figs 12A, 23a). Apicodorsal vertex slightly angular, apicoventral angle rounded, projecting as thick lobe narrowing progressively to apex. Parameres in ventral view curved posteromesad in apical half (Figs 12 BV p, 23c). In lateral view (Fig 12 BL) each constricted at short distance beyond base, dilated at middle, with almost parallel dorsal and ventral margins in central area (S-shape), two files of long and thick spines on midventral margin, pointing outward; midlateral surface covered by sparse thin spicules or setae, from middle posteroventral to posterodorsal edge and covering most of apex, absents in middle anteroventral and dorsal edges. Aedeagus (phallicata) in lateral view (Fig 12 CL) with dorsal margin concave and posterior corner of concavity hooked anterad, posterior edge of aedeagus straight, projected posteroventrad, subapically rounded ventrally (Fig 12 CL); ventral lobe of aedeagus triangular (Fig 12 BV vl), lateral edges slightly rounded; lateroventral lobes of phallus straight, posterior edges strongly convex, apicolateral margins rounded (Fig 12 CD lvl). Apicodorsal lobe of segment IX slightly dilated subapicolaterally, almost round in shape (Figs 12 DD al, 23b), with small apicomesal excision in some specimens; preanal appendages (Fig 12 DD pa) round, shorter than apicodorsal lobe, almost covered by it; in ventral view, apical band V-shaped (Fig 12 DV ab) nearly as wide as long, non-sclerotized inside area with posteromesal excision, posterior edges rounded (Fig 12 DV va), anal sclerites triangular (Fig 12 DV as). Female genitalia (Figs 13, 25): In lateral view (Figs 13 La, 13Lb, 25L), posterodorsal margin of segment VIII valves on each side irregular posteriorly, convex on ventral edge, with dorsolateral projection, in some specimens slightly indented posteriorly, posterolateral margin irregular, with 1–2 posterior projections, ventral projection more conspicuous (Fig 13 Lb 1–3). In dorsal view (Figs 13 Da, 13Db, 25D), with indentation between segment VIII valves, with rounded apical and subapical corners, delimiting oval space, small projection in apical corner in some specimens (Fig 13 Db). In ventral view (Fig 13V, 25V) segment VIII valves forming two elongate sclerites, basal 1/4 of mesal edges parallel, distal 3/4 of mesal edges irregular, slightly concave, diverging; intersegmental membrane with pair of dark, ovoid or slightly elongate sclerites., Published as part of Valladolid, María, Arauzo, Mercedes, Chertoprud, Mikhail V., Chvojka, Pavel, Czachorowski, Stanisław, Dorda, Beatriz A., Hinić, Jelena, Ibrahimi, Halil, Karaouzas, Ioannis, Krpač, Vladimir, Kučinić, Mladen, Lodovici, Omar, Salokannel, Juha, Stamenković, Valentina Slavevska & Stojanović, Katarina, 2021, The Rhyacophila fasciata Group in Europe: Rhyacophila fasciata Hagen 1859 and formerly synonymized species (Trichoptera: Rhyacophilidae), with new description of Rhyacophila fasciata and Rhyacophila septentrionis McLachlan 1865 (stat. prom.)., pp. 1-57 in Zootaxa 4975 (1) on pages 17-23, DOI: 10.11646/zootaxa.4975.1.1, http://zenodo.org/record/4804044, {"references":["Hagen, H. (1859) Die Phryganiden Pictet's. Entomologische Zeitung, 20 (4 - 6), 131 - 170.","Lepneva, S. G. (1964) Fauna of the USSR. Trichoptera. 1. Larvae and pupae of Annulipalpia. Zoological Institute of the Academy of Science of the USSR, New Series, 88, 1 - 638 [Jerusalem: Israel Program for Scientific Translations (1970)].","Nielsen, A. (1942) Uber die entwicklung und biologie der Trichoptera mit besonderer Berucksichtgung der queltrichopteren Himmerlands [About the development and biology of the Trichoptera with special consideration of spring Trichoptera from Himmerlands]. Archiv fur Hydrobiologie, Suppl. 17, (3 - 4), 337 - 358.","Ulmer, G. (1909) Die Susswasserfauna Deutschlands, Heft 5, 6: Trichoptera. Verlag, Gustav Fischer, Jena, 326 pp.","Silfvenius, A. J. (1905) Beitrage zur Metamorphose der Trichopteren. Acta Societatis pro Fauna et Flora Fennica, 27 (6), 1 - 168.","Pitsch, T. (1993) Zur Larvaltaxonomie, Faunistik und Okologie Mitteleuropaischer Fliessgewasser-Kocherfliegen (Insecta: Trichoptera) [Larval Taxonomy, Faunistics and Ecology of Central European River Caddisflies (Insecta: Trichoptera)]. Landschaftsentwicklung und Umweltforschung, Schriftenreihe des Fachbereichs Landschaftsentwicklung Sonderheft 8, TU Berlin, 322 pp. https: // depositonce. tu-berlin. de / handle / 11303 / 5106","Sedlak, E. (1985) Bestimmungsschlussel fur mitteleuropaische Kocherfliegenlarven (Insecta, Trichoptera). [Identification key for Central European caddis larvae (Insecta, Trichoptera)]. Wasser und Abwasser Vienna, 29, 1 - 146. (Translation from Czech and edition: J. Waringer). https: // www. zobodat. at / pdf / WasserAbwasser _ 1985 _ 0001 - 0146. pdf","Klapalek, F. (1893) Untersuchungen uber die Fauna der Gewasser Bohmens. I. Metamorphose der Trichopteren, II serie. Archiv fur die Naturwissenschaftliche Landesdurchforschung von Bohmen, 8 (6), 1 - 142.","Malicky, H. (2004) Atlas of European Trichoptera. Springer, Dordrecht, The Netherlands, 359 pp. https: // doi. org / 10.1007 / 978 - 1 - 4020 - 3026 - 0 _ 1","Holzenthal, R. W., Blahnik, R. J., Prather, A. L. & Kjer, K. M. (2007) Order Trichoptera Kirby, 1813 (Insecta), Caddisflies. Zootaxa, 1668 (1), 639 - 698. https: // doi. org / 10.11646 / zootaxa. 1668.1.29","McLachlan, R. (1865) Trichoptera Britannica: A monograph of the British species of caddis-flies. Transactions of the Entomological Society of London, 3 rd Series, 5, 1 - 183. https: // doi. org / 10.1111 / j. 1365 - 2311.1967. tb 01433. x"]}

Rhyacophila ferruginea (Scopoli), nomen dubium Phryganea ferruginea Scopoli, 1763: 266, fig. 691. Rhyacophila ferruginea (Scopoli): Hagen, 1859: 153. The original description of this species, as Phryganea ferruginea (Figs 1a, 1b) was: “ Phryganea ferruginea. Length of 6 1/3 lines [supposedly French lines, that gives approximately 14.287 mm of length. M.A. Alonso Zarazaga, personal communication]. Diagnosis. All rust colored; forewings reticulated dark; black eyes. In forests by the waters; month of May. Abdomen with lateral black dots, as many as segments of it. Antennae 6 lines long, rusty. The other sex with a round white common spot, and white spots (5–6) on the outer margin of the forewings.” Hagen (1854), in his study of the species of Phryganea included in Scopoli’s Entomologia Carniolica stated that the description of P. ferruginea does not match the drawing, even with other Phryganea. Following the “reticulate wings” and “white spots” of wings, he proposed three options that could be one of the following: Chaetopteryx villosa (Fabricius, 1798) (based on body contour and reticulation of wings), one of two Hydropsyche Pictet 1834, and one Rhyacophila (reticulation of wings); the color (all rust) only matched with the mentioned Rhyacophila (maybe R. nebulosa Stephen), and, as he had only the female, he couldn’t check if males also had the white spots on the wings. In 1859 Hagen cited this species and R. fasciata, together with R. vulgaris, R. dorsalis and R. paupera: “4. R. ferruginea Scop. Anal appendage rounded, almost completely covering the lateral appendages. The smallest species. Carniola, Austria. Maybe dark colored females of my collection from Zurich belong to it.” “5. R. fasciata Hag. The appendages are very similar to the previous species, but the size of the animal is much more significant and the upper wings have three dark cross-ties. A single male from Elberfeld”. McLachlan (1865), at the end of his description of Rhyacophila septentrionis wrote: “I have some doubt in announcing this as a new species, since the form of the appendices agrees very well with the figure of those of R. ferruginea, sent to me by Dr. Hagen. I submitted an example to that gentleman, and he says that it is larger and darker than his continental types of that species and may be distinct. R. ferruginea is described as ‘die kleinste Art’ of the group (the true genus Rhyacophila) to which it belongs”. In his publication of 1868, McLachlan said: “ R. ferruginea, (Scopoli) Hagen. I have not seen this insect, which, according to Hagen (in litt.) is smaller than septentrionis, but with similar lobes and appendices. Scopoli’s name cannot be said to apply to this species with certainty.” Finally, in his publication of 1879, McLachlan concluded at the end of the paragraph about Rhyacophila septentrionis: “I do not feel justified in adopting Scopoli’s name ferruginea (as was done by Hagen for the example from Carniola). Even supposing that Scopoli had a Rhyacophila before him (which is just possible), the particular species must remain uncertain.” We agree with this statement, due to the difficulty of identification from the original species described, so we propose to change the status of Rhyacophila ferruginea (Scopoli 1763) to nomen dubium., Published as part of Valladolid, María, Arauzo, Mercedes, Chertoprud, Mikhail V., Chvojka, Pavel, Czachorowski, Stanisław, Dorda, Beatriz A., Hinić, Jelena, Ibrahimi, Halil, Karaouzas, Ioannis, Krpač, Vladimir, Kučinić, Mladen, Lodovici, Omar, Salokannel, Juha, Stamenković, Valentina Slavevska & Stojanović, Katarina, 2021, The Rhyacophila fasciata Group in Europe: Rhyacophila fasciata Hagen 1859 and formerly synonymized species (Trichoptera: Rhyacophilidae), with new description of Rhyacophila fasciata and Rhyacophila septentrionis McLachlan 1865 (stat. prom.)., pp. 1-57 in Zootaxa 4975 (1) on pages 13-14, DOI: 10.11646/zootaxa.4975.1.1, http://zenodo.org/record/4804044, {"references":["Scopoli, I. A. (1763) Entomologia Carniolica Exhibens Insecta Carnioliae Indigena et Distributa in Ordines, Genera, Species, Varietates. Methodo Linnaeana. I. T. Trattner, Vindobonae [Vienna], xxxvi + 420 pp. https: // doi. org / 10.5962 / bhl. title. 119976","Hagen, H. (1859) Die Phryganiden Pictet's. Entomologische Zeitung, 20 (4 - 6), 131 - 170.","Hagen, H. (1854) Die Kupfer zu Scopoli's Entomologia Carniolica. Entomologische Zeitung, 15 (3), 81 - 91.","Fabricius, J. C. (1798) Entomologia systematica emandata et aucta. Secundum classes, ordines, genera, species adjectis synonimis, locis, observationibus descriptionibus. Supplementum Entomologiae Systematicae. Hafniae [Copenhagen], apud Proft et Storch.","Pictet, F. J. (1834) Recherches pour Servir a l'Histoire et a l'Anatomie des Phryganides. A. Cherbuliez, Geneve [Geneva]. J. - B. Bailliere, Paris, iii + 235 pp., 20 coloured plates. https: // doi. org / 10.5962 / bhl. title. 66017","McLachlan, R. (1865) Trichoptera Britannica: A monograph of the British species of caddis-flies. Transactions of the Entomological Society of London, 3 rd Series, 5, 1 - 183. https: // doi. org / 10.1111 / j. 1365 - 2311.1967. tb 01433. x"]}

Valladolid, María, Arauzo, Mercedes, Chertoprud, Mikhail V., Chvojka, Pavel, Czachorowski, Stanisław, Dorda, Beatriz A., Hinić, Jelena, Ibrahimi, Halil, Karaouzas, Ioannis, Krpač, Vladimir, Kučinić, Mladen, Lodovici, Omar, Salokannel, Juha, Stamenković, Valentina Slavevska, Stojanović, Katarina (2021): The Rhyacophila fasciata Group in Europe: Rhyacophila fasciata Hagen 1859 and formerly synonymized species (Trichoptera: Rhyacophilidae), with new description of Rhyacophila fasciata and Rhyacophila septentrionis McLachlan 1865 (stat. prom.). Zootaxa 4975 (1): 1-57, DOI: https://doi.org/10.11646/zootaxa.4975.1.1

Rhyacophila fasciata Hagen Rhyacophila fasciata Hagen, 1859: 153. Lectotype: 1 male. Entomology 11067, Entomology Collection of the Museum of Comparative Zoology, Harvard University. Type locality: Elberfeld (Germany). Other information: (in label) 57; E.-illegible; Hagen; 107; ML; type 11067; (vial); Lectotype Rhyacophila fasciata Hagen. (MCZBase, 2020). (Figs 2a, 2b, 30a, 30b) Ross (1952) included the species in his publication about the lectotypes of Hagen in the collection of the Museum of Comparative Zoology at Harvard University, Cambridge, Massachusetts. Description. This species has been described by several authors over many years. The first publication where this species is cited is that of Hagen (1859), where he provided a very short description: “A. Lower appendages undivided at the top. 1. The tongue-like extention of the last abdominal segment with an enlarged end. 5. R. fasciata Hag. The app. are very similar to the previous species (R. ferruginea Scop.), but the size of the animal is much more significant, and the upper wings have three dark cross bands. A single male from Elberfeld” (Fig 30a).” McLachlan (1868) described it as follows: “ R. fasciata Hagen, may be distinguished almost with certainty by the dark fasciae on the anterior wings. The form of the dorsal lobe is very distinct from the preceding (R. paupera), being broad and rounded, and nearly concealing the app. sup. The second joint of the app. inf. is obliquely truncate (Pl.XIV. fig. 16). I have three ♂ from Carinthia from Zeller.” In 1879, McLachan described R. fasciata and R. septentrionis. In the description of R. fasciata he included the original specimen from Elberfeld, his specimens from Carinthia, saying that they should be referred to septentrionis, and 7 ♂ and 1 ♀ from the Pyrenees (France), writing “it may be that these should form a distinct species”. These specimens from the Pyrenees were described in 1916 by Navás as Rhyacophila sociata (Valladolid et al. 2018). On the other hand, in the description of R. septentrionis McLachlan included his specimens from Scotland, as well as others from different countries of North and Central Europe. Based on results of our studies of morphology and genetics, we can say that most of the specimens cited belong to R. fasciata, keeping only those from Scotland as R. septentrionis. Klapálek (1893) described the larva and the pupa of R. septentrionis in Bohem, but the area matches with those of R. fasciata, so it should be this species. The same situation pertains to Silfvenius (1905), who described larvae and pupae from South Finland as R. septentrionis that should be R. fasciata. Ulmer (1909), based on previous authors, described the imagos and larvae of R. fasciata and R. septentrionis, first one from Elberfeld and the second one from different places in Germany. We consider that these specimens all belong to R. fasciata. There are more recent authors that have described R. fasciata and R. septentrionis specimens: Nielsen (1942) as R. septentrionis from Denmark; Nóvak (1963) as R. fasciata from Czechoslovakia; Hickin (1954) and Mackereth (1954) as R. septentrionis from United Kingdom; Fotius-Jaboulet (1964) as R. septentrionis from France; Lepneva (1964) as R. septentrionis from northwest, west and southwest Russia; Eidel (1974) as R. fasciata from Austria; Buholzer (1978) as R. fasciata from Switzerland; Sedlak (1985) as R. fasciata from Central Europe; Pitsch (1993) as R. fasciata from Central Europe. Of all these descriptions, only those from the United Kingdom could belong to R. septentrionis; the rest, from continental Europe, should be R. fasciata or some related species. Finally, Malicky & Sipahiler (1993) illustrated the differences among male specimens of R. fasciata from Europe, Turkey, and Lebanon that they considered as subspecies of the R. fasciata Group. Some of them have been promoted to species (R. denticulata McLachlan 1879, by Valladolid et al. 2018; R. kykladica Malicky & Sipahiler 1993, by Valladolid et al. 2019), one species has been resurrected (R. sociata Navás 1916, previously a synonym of R. denticulata, by Valladolid et al. 2018), and others could be considered also valid species (Valladolid, unpublished data)., Published as part of Valladolid, María, Arauzo, Mercedes, Chertoprud, Mikhail V., Chvojka, Pavel, Czachorowski, Stanisław, Dorda, Beatriz A., Hinić, Jelena, Ibrahimi, Halil, Karaouzas, Ioannis, Krpač, Vladimir, Kučinić, Mladen, Lodovici, Omar, Salokannel, Juha, Stamenković, Valentina Slavevska & Stojanović, Katarina, 2021, The Rhyacophila fasciata Group in Europe: Rhyacophila fasciata Hagen 1859 and formerly synonymized species (Trichoptera: Rhyacophilidae), with new description of Rhyacophila fasciata and Rhyacophila septentrionis McLachlan 1865 (stat. prom.)., pp. 1-57 in Zootaxa 4975 (1) on pages 14-15, DOI: 10.11646/zootaxa.4975.1.1, http://zenodo.org/record/4804044, {"references":["Hagen, H. (1859) Die Phryganiden Pictet's. Entomologische Zeitung, 20 (4 - 6), 131 - 170.","Ross, H. H. (1952) Lectotypes of Hagen species belonging to certain families of Trichoptera. Psyche: A Journal of Entomology, 59, 31 - 36. https: // doi. org / 10.1155 / 1952 / 52847","McLachlan, R. (1868) Contributions to a knowledge of European Trichoptera. Transactions of the Entomological Society of London, 4 th Series, 1, 289 - 308.","Valladolid, M., Arauzo, M., Basaguren, A., Dorda, B. A. & Rey, I. (2018) The Rhyacophila fasciata Group in Western Europe: Confirmation of Rhyacophila denticulata McLachlan 1879 (stat. prom.) and Rhyacophila sociata Navas 1916 (stat. res.), based on morphological and molecular genetic evidence (Trichoptera: Rhyacophilidae). Zootaxa, 4418 (6), 526 - 544. https: // doi. org / 10.11646 / zootaxa. 4418.6.2","Klapalek, F. (1893) Untersuchungen uber die Fauna der Gewasser Bohmens. I. Metamorphose der Trichopteren, II serie. Archiv fur die Naturwissenschaftliche Landesdurchforschung von Bohmen, 8 (6), 1 - 142.","Silfvenius, A. J. (1905) Beitrage zur Metamorphose der Trichopteren. Acta Societatis pro Fauna et Flora Fennica, 27 (6), 1 - 168.","Ulmer, G. (1909) Die Susswasserfauna Deutschlands, Heft 5, 6: Trichoptera. Verlag, Gustav Fischer, Jena, 326 pp.","Nielsen, A. (1942) Uber die entwicklung und biologie der Trichoptera mit besonderer Berucksichtgung der queltrichopteren Himmerlands [About the development and biology of the Trichoptera with special consideration of spring Trichoptera from Himmerlands]. Archiv fur Hydrobiologie, Suppl. 17, (3 - 4), 337 - 358.","Novak, K. (1963) Beschreibung und bestimmungstabelle der weibchen bohmischer arten der gattung Rhyacophila. Acta Societatis Entomologicae Cechosloveniae, 60 (4), 304 - 311.","Hickin, N. E. (1954) Larvae of the British Trichoptera. 42. Rhyacophila septentrionis McLachlan (Rhyacophilidae). Proceedings of the Royal Entomologica Society of London, Series A, General Entomology, 29 (4 - 6), 59 - 61. https: // doi. org / 10.1111 / j. 1365 - 3032.1954. tb 01199. x","Mackereth, J. C. (1954) Taxonomy of the larvae of the British species of the genus Rhyacophila (Trichoptera). Proceedings of the Royal Entomologica Society of London, Series A, General Entomology, 29 (10 - 12), 147 - 152. https: // doi. org / 10.1111 / j. 1365 - 3032.1954. tb 01189. x","Fotius-Jaboulet, M. C. (1964) Description de la larve de Rhyacophila praemorsa Mcl. et comparaison de cette larve a celle de Rhyacophila septentrionis Mcl. Travaux du Laboratoire de Zoologie et de la Station Aquicole Grimaldi de la Faculte des Sciences de Dijon, 62, 1 - 13.","Lepneva, S. G. (1964) Fauna of the USSR. Trichoptera. 1. Larvae and pupae of Annulipalpia. Zoological Institute of the Academy of Science of the USSR, New Series, 88, 1 - 638 [Jerusalem: Israel Program for Scientific Translations (1970)].","Eidel, K. (1974) Die Kocherfliegen (Trichoptera) des Wutachgebietes. Mitteilungen des Badischen Landesvereins fur Naturkunde und Naturschutz, 11 (2), 181 - 195. https: // www. zobodat. at / pdf / Mitt-Bad-Landesver-Natkde-Natschutz-Freiburg _ NF _ 11 _ 0181 - 0195. pdf","Buholzer, H. (1978) Larvenmorphologie und Verbreitung der Schweizerishen Rhyacophila Arten (Trichoptera, Rhyacophilidae). Ph. D. Dissertation, ETH Nr. 6177, Zurich, 151 pp. https: // www. research-collection. ethz. ch / handle / 20.500.11850 / 135968","Sedlak, E. (1985) Bestimmungsschlussel fur mitteleuropaische Kocherfliegenlarven (Insecta, Trichoptera). [Identification key for Central European caddis larvae (Insecta, Trichoptera)]. Wasser und Abwasser Vienna, 29, 1 - 146. (Translation from Czech and edition: J. Waringer). https: // www. zobodat. at / pdf / WasserAbwasser _ 1985 _ 0001 - 0146. pdf","Pitsch, T. (1993) Zur Larvaltaxonomie, Faunistik und Okologie Mitteleuropaischer Fliessgewasser-Kocherfliegen (Insecta: Trichoptera) [Larval Taxonomy, Faunistics and Ecology of Central European River Caddisflies (Insecta: Trichoptera)]. Landschaftsentwicklung und Umweltforschung, Schriftenreihe des Fachbereichs Landschaftsentwicklung Sonderheft 8, TU Berlin, 322 pp. https: // depositonce. tu-berlin. de / handle / 11303 / 5106","Malicky, H. & Sipahiler, F. (1993) Kocherfliegen (Trichoptera) aus der Turkei, mit Bemerkungen zu weiteren mediterranean Kocherfliegen. Bulletin de la Societe Entomologique Suisse, 66, 457 - 468.","McLachlan, R. (1879) A Monographic Revision and Synopsis of the Trichoptera of the European Fauna, Part VIII. John Van Voorst, London, 1874 - 1880, pp. 429 - 500, plates 45 - 51. https: // doi. org / 10.5962 / bhl. title. 28556","Valladolid, M., Karaouzas, I., Arauzo, M., Dorda, B. A. & Rey, I. (2019) The Rhyacophila fasciata Group in Greece: Rhyacophila kykladica Malicky & Sipahiler 1993 (stat. prom.) (Trichoptera: Rhyacophilidae). Morphological description, genetic and ecological features. Zootaxa, 4657 (3), 503 - 522. http: // dx. doi. org / 10.11646 / zootaxa. 4657.3.5","Navas, L. (1916) Tricopteros nuevos de Espana. 1 ª Serie. Broteria, Serie Zoologica, 14, 5 - 11."]}

Los ríos pueden presentar una biodiversidad alta, en función de su estado de conservación general y de la variabilidad de hábitats que posean. Desde la cabecera, donde nacen, hasta su desembocadura en otro río o en el mar, el cauce de cada río posee numerosos sustratos (grandes rocas, gravas de diversos tamaños, arenas, limos), vegetación (bosques de ribera, plantas semisumergidas y sumergidas, algas filamentosas y microscópicas) y velocidades de la corriente del agua (zonas de corriente rápida y media, remansos, zonas estancadas) que permiten la existencia de todo tipo de animales vertebrados e invertebrados en sus aguas. Por otra parte, la presencia de ciertos factores como la contaminación de las aguas por residuos de todo tipo (químicos, biológicos, físicos) procedentes de agricultura, ganadería, industria o entornos urbanos, así como la modificación del paisaje (por ejemplo, en la extracción de arenas y gravas) hace que esta biodiversidad baje en la zona afectada, llegando incluso a desaparecer los organismos vivos en los casos más extremos.

In our previous paper (Valladolid et al. 2021) we have detected two errors: in the title, Rhyacophila septentrionis McLachlan 1865 (stat. prom.), the correct reference is Rhyacophila septentrionis McLachlan 1865 (stat. res.). And the correct e-mail address of Pavel Chvojka is: pavel.chvojka@nm.cz. After the publication, some specimens of Rhyacophila fasciata Hagen 1859 from Poland (cited as belonging to the M. Valladolid Collection) were deposited in the Collection of Entomology of the Museo Nacional de Ciencias Naturales (MNCN – CSIC), Madrid, with the following numbers: [Adults]. R. fasciata: 4 ♂, river Pasłęka (NE Poland), 53.618N, 20.326E, 180 m a.s.l., 11/v/85 [MNCN_Ent 296070, MNCN_Ent 296071, MNCN_Ent 296073, MNCN_Ent 296074 (MNCN)]. 1 ♀, river Łyna (NE Poland), 53.441N, 20.413E, 190 m a.s.l., 11/v/91 [MNCN_Ent 296072 (MNCN)]. Leg. S. Czachorowski. [Pupae]. R. fasciata: 2 immature pupae (IP) + 1 ♂P + 1 ♀P, Rurka, river Rurzyca (W Poland), 52.999N, 14.481E, 130 m a.s.l., 07/xii/96 [2 IP, MNCN_Ent 296077, MNCN_Ent 296078; 1 ♂P, MNCN_Ent 296075; 1 ♀P, MNCN_Ent 296076 (MNCN)]. 1 ♂P + 1 ♀P, river Wałsza (NE Poland), 54.236N, 20.128E, 70 m a.s.l., 19/iv/99 [1 ♂P, MNCN_Ent 296062; 1 ♀P, MNCN_Ent 296063 (MNCN)]. 1 ♂P + 1 ♀P, Gaznoty, river Gizela (NE Poland), 53.534N, 19.901E, 190 m a.s.l., 9/vi/99 [1 ♂P, MNCN_Ent 296064; 1 ♀P, MNCN_Ent 296065 (MNCN)]. 1 ♂P + 1 ♀P, Zajączki, river Gizela (NE Poland), 53.560N, 19.882E, 170 m a.s.l., 9/vi/99 [1 ♂P, MNCN_Ent 296066; 1 ♀P, MNCN_Ent 296067 (MNCN)]. 1 ♂P + 1 ♀P, river Mała Wkra (NE Poland), 53.552N, 19.964E, 140 m a.s.l., 17/iv/00 [1 ♂P, MNCN_Ent 296060; 1 ♀P, MNCN_Ent 296061 (MNCN)]. 1 ♂P + 1 ♀P, Elbląg, river Kumiela (N Poland), 54.158N, 19.414E, 15 m a.s.l., 10/vii/03 [1 ♂P, MNCN_Ent 296058; 1 ♀P, MNCN_Ent 296059 (MNCN)]. 1 ♂P + 1 ♀P, brook Srebrny Potok (NE Poland), 54.162N, 19.491E, 135 m a.s.l., 11/vii/03 [1 ♂P, MNCN_Ent 296068; 1 ♀P, MNCN_Ent 296069 (MNCN)]. Leg. S. Czachorowski. [Larvae]. R. fasciata: 1 L, Góry Stołowe, brook Kudowski Potok (SW Poland), 50.482N, 16.337E, 840 m a.s.l., 19/ix/94 [MNCN_Ent 296079 (MNCN)]. Leg. S. Czachorowski.

Rhyacophila fasciata viteceki Valladolid & Kučinić (ssp. nov.) This subspecies has been found in Bosnia and Herzegovina. Etymology. The specific name is the genitive of Vitecek, given in honour of Dr. Simon Vitecek, biologist and trichopterologist of Wasser Cluster Lunz, Lunz am See, Austria. Type material. Holotype ♂: BOSNIA AND HERZEGOVINA, river Neretva, Bačevići (43.254154º N 17.824627º E, 35 m a.s.l.), 30/vi/17 (Stanić-Koštroman & M. Kučinić) [N1, MNCN _ Ent 269367 (MNCN)]. Paratypes: 2 ♂, same locality as holotype, 30/vi/17 (Stanić-Koštroman & M. Kučinić) [N2, no. 4455, (MCK) and N3, no. 1111 (UMBH)]. 2 ♂ and 1 ♀, river Sturba, 30/viii/18 (M. Kučinić) [1 ♂, S1, MNCN _ Ent 269368 (MNCN); 1 ♀, S3, MNCN _ Ent 269369 (MNCN); 1 ♂ S2, no. 4456 (MKC)]. 1 ♂, river Bunica, 04/vi/17 (StanićKoštroman & M. Kučinić) [B1, no. 1112 (UMBH)]. Description of the imago. Holotype (N1): Length from front of head to distal edge of segment IX 8.91 mm, each forewing 12.55 mm, each hind wing 10.54 mm. Males: Length 8.29–9.21 mm (x = 8.72, n = 6), each forewing 9.83–12.55 (x = 11.34, n = 6), each hind wing 8.85–10.54 mm (x = 9.79, n = 6). Female: Length from front of head to distal edge of segment VIII 11.26 mm (n =1), forewing 14.52 mm (n = 1), hind wing 12.15 mm (n = 1). In ethanol-preserved specimens colour generally pale brown, yellowish, with golden brown setae. Head, thorax, and abdomen dorsally pale brown, with small dark brown spots in dorsal area, some specimens in lateral view with long black spots marking borders between dorsal and ventral areas, abdomen ventrally yellowish; female darker than male, anterior 2/3 of abdominal segments pale brown or brown in darker specimens. Legs light, with spurs reddish brown. Forewings brown, spotted; hind wings pale. Male genitalia (Figs 3, 6 a–6e): Apical segment of each inferior appendage (Figs 3A, 6a) with basal and distal edges diverging, posterior edge oblique and slightly convex, ventral edge straight to slightly concave, at least two times longer that dorsal edge. Apicodorsal vertex angular, apicoventral angle projecting as thick lobe narrowing progressively to round apex (Fig 6a). Parameres (Figs 3 BV–3BL, 6c–6d) in ventral view curved posteromesad in apical 2/3 (Figs. 3 BV p, 6c). In lateral view (Figs. 3 BL, 6d) each slender at base, slightly dilated in middle, with rounded dorsal margin before midlength and almost straight ventral margin at midlength, pointed at apex; row of few thick spines on midventral margin (Figs 6 c–6d); midlateral surface covered by very fine spicules or setae, reaching from anteroventral to posterodorsal edges of paramere, absent from dorsal edge. Aedeagus (phallicata) in lateral view (Figs 3 CL, 6e) with dorsal margin deeply concave and posterior corner of concavity hooked anterad; upper posterior edge concave to obliquely truncate apex, ventral edge sinous. Lateroventral lobes of phallus (Fig. 3 CD lvl) convex, narrowing progressively towards apex, apicolateral margins round, posteromesal edge of each lobe concave. Ventral lobe of aedeagus nearly round, tapering and acute caudally (Fig 3 BV vl). Apicodorsal lobe of segment IX (Figs 3 DD, 6b) slightly dilated subapicolaterally, posterior edge semicircular, lateral edges converging anterad; preanal appendages (fig 3DD pa) shorter than apicodorsal lobe (Fig 3 DD al), obliquely subtruncate apicolaterally, with slightly concave posteromesal edges (Figs. 3 DD, 6b). In ventral view, apical band V-shaped, inner and outer edges converging in posterior third, pointed apically, longer than wide (Fig 3 DV ab); posterior edge of non-sclerotized ventral area slightly concave, with small mesal incision (Fig 3 DV va), anal sclerites subtriangular, concave apically (Fig 3 DV as)., Published as part of Valladolid, María, Kučinić, Mladen, Arauzo, Mercedes, Cerjanec, Darko, Ćuk, Renata, Dorda, Beatriz A., Lodovici, Omar, Stanić-Koštroman, Svjetlana, Vučković, Ivan & Rey, Isabel, 2020, The Rhyacophila fasciata Group in Croatia and Bosnia and Herzegovina: Rhyacophila f. fasciata Hagen 1859 and the description of two new subspecies, Rhyacophila fasciata delici Kučinić & Valladolid (ssp. nov.) from Croatia and Bosnia and Herzegovina and Rhyacophila fasciata viteceki Valladolid & Kučinić, pp. 51-75 in Zootaxa 4885 (1) on pages 65-66, DOI: 10.11646/zootaxa.4885.1.3, http://zenodo.org/record/4296372

Rhyacophila sociata Nav��s 1916 (status resurrectus) . In his publication, Nav��s (1916) described one male collected in Solares (Cantabria), and as stated previously, Schmid (1949) did not included it in his study of Nav��s��� collection. A staff person of the University of Zaragoza, where the insect collection of Nav��s was housed years ago, suggested that the holotypes, including that of R. sociata, possibly were sent to the Museo de Ciencias Naturales of Barcelona. The curator of the arthropod collection of this museum informed us that this specimen is not in the Barcelona collection, nor is the specimen described as R. gemella (Nav��s 1923). Consequently, we decided to designate a neotype for this species. In a second step, we tried to collect new specimens from Solares, without success, and researchers working in this area confirmed that they never collected the species there. We decided to designate also a new type locality, where the neotype specimen was captured. Neotype: 1 male (08/06/09). Collection number: MNCN _ Ent 160675. Tissues and DNA Collection Number: MNCN / ADN 57444. Other specimens: 3♂, 30/06/08; 1♂ + 6♀, 06/08/08; 1♂ + 1♀, 08/06/09; 1♂, 3/07/09. Collection numbers, see Table 1. Type locality: Casalarreina, La Rioja (Spain). River Oja. Latitudinal and longitudinal geographic coordinates: 42.546105 N, 2.915515 W. Altitude: 490 m a.s.l. Description of the larva. Biometrics last instar larva and prepupa: Length: 9.43���15.34 mm (x��= 2.28, n = 26). Maximum width of larva at metathorax. Maximum width of abdominal segments II���III: 2.32���4.13 mm (x��= 2.96, n = 26). Color: Thorax and abdomen greenish, dorsum with dark areas delimited by narrow, pale, dorsolateral marks. Sclerotized areas of head and thorax dark brown with pale spots and stripes. Specimens preserved in alcohol generally pale yellow, with sclerotized areas reddish brown to black in some places. Head (Fig. 1). Length: 1.37���2.23 mm (x��= 1.58, n = 26). Width: 1.00���1.84 mm (x��= 1.22, n = 26). Maximum width in posterior third of head, approximately in line with sensory pits #9. Cephalic capsule (Figs 1a, 1b), in dorsal view with yellow background anterior of setae #6 and light brown posterior of them, across full width of dorsal view (Figs. 1a, 1b) and halfway down sides (Fig. 1c), except sometimes lighter posterior occiput and beside frontoclypeal and coronal sutures. Ventral view uniformly yellow, without dark spots. Frontoclypeal apotome (Fig. 1d) with brown pattern in posterior third, posterior of setae #6, this pattern U���shaped in 59% of specimens studied (modified U��� shape in 41% of specimens), and with 3���4 pale muscle attachment spots with dark borders inside pattern. Posterior dorsal area of head with pair of brown patches, marked with darker muscle attachment spots (in 78% of specimens). Three light spots around setae #15, #16 and #19���21, forming pair of light longitudinal stripes along head in 53% of specimens. Light triangular spot on juncture of frontal and coronal sutures in 85% of specimens, sometimes replaced by pair of light spots around sensory pits #9. On posterior occipital zone, pair of light areas connecting with dorsal light stripes of setae #15, #16 and #19���21 in 61% of specimens. Most specimens (73%) with light areas bordering frontal and coronal sutures (n = 100). Mandibles (Fig. 2) asymmetrical, as in other rhyacophilids, left one bigger than right one. Inner blade of left mandible smooth, convex, right mandible with small tooth in middle. Maxillolabium and labrum (Fig. 3) as in other Rhyacophila species. Mentum rectangular (Fig. 3a), its anterior half slightly sclerotized. Labrum (Fig. 3b) with small transparent area in middle of anterior edge. Thorax (Fig. 4): Anterior half of pronotum pale or very light coloured in 88% of specimens studied. Sinuous posterior margin bordered by dark band, black in middle and posterolaterally, light brown submesally. Posterior half of pronotum (Fig. 4a) with three dark triangular areas separated by light zones, central dark region tapered posteriorly and reaching posterior edge of sclerite, its anterior reaching sensory pit #P2. Lateral dark triangular areas separated from posterior margins by light stripes, each delimited by line of small darker points parallel to back edge. Each pronotal plate with curved row of dark dots from anterior edge of lateral dark triangular area toward anterolateral corner, curving around circular spot on anterior half of most pigmented specimens (Fig. 4a). Circle of dark dots also around setae #5. Anterolateral margin of each pronotal sclerite black only in anterolateral angle and extending posterad laterally without contacting posterolateral margin (Fig. 4b). Anterior margin not coloured, sometimes with small, brown spot near anterior angle. Central area of posterior marginal band of each sclerite light brown, not black (Fig. 4b). Abdomen (Figs. 5���7): Similar to other species of Rhyacophila. Colour as for meso- and metathoracic segments, greenish with dark dorsal areas delimited by light bars in living larvae and light reddish brown dorsally and pale yellow ventrally in fixed larvae. Widths of anterior abdominal segments similar, narrower in posterior segments. Lateral abdominal gills in tufts of 21���25 filaments. Dorsal sclerite of the segment IX (Fig. 5) light coloured, with two narrow discontinuous black bands on anterior and posterior edges. Anal prolegs (Figs. 6D, 6V, 7a) typical of group, with long sword process (Figs. 7a, 7c). In this aspect, R. denticulata differing from R. sociata in that, instead of this long sword process, basilateral plate with short and angular process (Despax 1928a, fig. 6). Anal claws each with two ��� three teeth on ventral edge (Figs. 7a, 7b), basoventral hook black basally and reddish brown apically (Figs. 6V, 7c). Legs (Fig. 8): Similar to those of other Rhyacophila species. With light reddish brown colour, dorsoproximal coxa black, with small black areas also in dorsoproximal femur and proximal trochanter (dorsal and ventral). Light- coloured zone in distal dorsolateral area of tibia of each of three pairs of legs. Forelegs each with spicules, extending ventrally from distal part of trochanter to end of tarsus, with different disposition on each segment: On trochanter forming isolated tufts, on femur forming double rows, on tibia tufts ventrally and in rows laterally and on tarsus, similar to trochanter but with tufts more densely distributed. Setae reddish, some thicker and darker: On each foreleg, setae 1,3,4 of coxa and 1,4 of trochanter and 1,2,4,6 of femur; on each midleg, setae 1,4 of coxa and trochanter and 1,2,4,6 of femur; on each hindleg, setae 1,4,5 of coxa and 1,4 of trochanter and 1,2,3,4,6 of femur. Description of the pupa. Biometrics of pupa and cocoon. Pupal length (total): 8.24���14.08 mm (x��= 10.83, n = 39); male pupa: 8.24���11.45 mm (x=�� 9.72, n = 19); female pupa: 8.36���14.08 mm (x��= 11.08, n = 20). Maximum width of young pupa in abdominal segments II and III. Cocoon length (total): 11.00��� 15.50 mm (x=�� 13.32, n = 32); width: 3.00��� 4.64 mm (x��= 3.67 mm, n = 32); male cocoon length: 11.00��� 14.28 mm (x��= 12.68, n = 15), width: 3.18���4.05 mm (x��= 3.55 mm, n = 15); female cocoon length: 11.58���15.30 mm (x��= 13.89, n = 13), width: 3.00���4.37 mm (x��= 3.77, n = 13). Head: Distribution of setae on dorsal head similar to that of R. denticulata (Despax 1928a). Mandibles similar in R. denticulata and R. sociata, left one with two large teeth and right one with three, with numerous fine teeth on inner blade. Labrum similar in shape to that of R. denticulata. Antennae in R. sociata of variable length, reaching from middle of abdominal segment V to anterior part of segment VII. Thorax: Tubercules of prothorax each with 5���6 setae in R. sociata but only 4 in R. denticulata. Wing pads reaching posterior edge of abdominal segment IV in R. sociata, and posterior edge of segment III in R. denticulata (Despax 1928a). Abdomen: Paired hook plates of anterior margins pedunculated and present from abdominal segments IV to VII; anterior hook plates on abdominal segment III not visible in R. sociata (Fig. 9), small and with 3���4 spines developed and 3���4 very small in R. denticulata (Despax 1928a). Paired posterior hook plates sessile and on segments III���V in the two species. Number of hooks per plates varying among these species, being least numerous in R. denticulata (see Despax 1928a), and of intermediate abundance in R. sociata (Fig. 9). Description of the imago. Males length: 9.50���10.80 mm (x��= 10.07, n = 7), each forewing: 10.8���11.8 (x��= 11.29, n = 7), each hind wing: 9.30���10.40 mm (x=�� 9.83, n = 7); females, length: 10.70���12.60 mm (x=�� 11.54, n = 7), each forewing: 11.80���12.80 mm (x��= 12.36, n = 7), each hind wing: 10.70���11.10 mm (x��= 10.96, n = 7). Colour generally pale, yellowish with setae reddish brown. Head and thorax reddish brown, abdomen yellowish, with small black spots in dorsal and lateral areas, sometimes also with dark bands on each side of abdominal segments IV���V to VII. Legs light, with spurs reddish brown. Wings pale, darker in females. Dark area on leading edge, between subcostal (Sc) and radial 1 (R1) veins in forewing and between Sc and radial 2 (R2) in hind wing; in R. fasciata this area present between Sc and R1 in both wings, but in R. denticulata only in forewing, between Sc and R2 together with dark spots in edge of wing, at R1���R5 and medial 1 (M1) veins. Male genitalia (Figs. 10, 11): 2nd segment of each inferior appendage quadrangular, with basal and distal edges nearly parallel, posterior and ventral edges slightly concave. Apicodorsal and especially apicoventral angles rounded, apicoventral angle projecting as lobe narrowing progressively to apex. In R. denticulata and R. fasciata males, 2nd segment of each inferior appendage wider basally than in R. sociata, with basal and distal edges diverging, apicodorsal vertex more angular and ventral lobe thicker; also posterior edge sinuous in these two species, in R. denticulata concave in the upper and lower halves, concavities less conspicuous in R. fasciata (Fig. 10A). Parameres (Figs. 10B, 11) each in lateral view slender at base, dilated in middle, with rounded dorsal and ventral margin, and pointed at apex, with few thick spines on midventral margin and small spicules or chaetae on posterior half of external surface. In dorsal and ventral views parameres curved posteromesad in apical 1/3. In R. denticulata parameres more dilated than in R. sociata, each with midventral and lower half of midlateral areas covered with spines long and thick on ventral margin and decreasing in size dorsally, middorsal half with small spines or chaetae; parameres more strongly curved posteromesad in apical 1/3, apex often sinuous and/ or overlapping. In R. fasciata, parameres in lateral view each with base slightly thicker than for other two species and less dilated in middle, midlateral surface almost covered with spines, thicker spines on ventral margin, decreasing in size dorsally, spines of ventral margin less numerous than in R. denticulata; slightly curved posteromesad in apical 1/3 and often overlapping apically (Figs. 10B, 11). Aedeagus (phallicata) in lateral view (Fig. 10 CL) slender and clearly curved in middle, with mid-dorsal margin concave and midventral margin convex, apically with margins almost parallel. Ventral lobe of phallus with parallel margins basally in dorsal view, abruptly narrower apically, apicolateral margins rounded, aedeagus apically blunt (Fig. 10 CV). Aedeagus of R. denticulata in lateral view with mid-dosal margin concave for short distance but midventral margin not convex, ventral lobe in dorsal view with margins convex and narrowing progressively toward apex, apicolateral margins slightly projecting caudad, and aedeagus slightly pointed (Fig. 10 CV). Aedeagus of R. fasciata in lateral view with mid-dorsal margin concave and posterior corner of concavity hooked anterad, midventral margin with small notch, in dorsal view margins of ventral lobe similar and aedeagus similar to R. sociata (Fig. 10 CV). Apicodorsal lobe of segment IX dilated subapicolaterally, with small apicomesal excision; preanal appendages shorter than that lobe, with round edges and small invagination near contact area between segment IX and preanal appendages (Fig. 10 DD). In ventral view, apical band V-shaped, anal sclerites triangular (Fig. 10 DV). In R. denticulata apicodorsal lobe of segment IX dilated subapicolaterally about as much as for R. sociata, but with apex rounded and without excision, preanal appendages shorter than that lobe, more nearly squared than in R. sociata, in ventral view, apical band U-shaped, anal sclerites triangular. In R. fasciata, apicodorsal lobe more dilated than in R. sociata and R. denticulata, almost from its base, and with broad apicomesal excision, similar to R. sociata. In ventral view, apical band V-shaped, with apices close together, anal sclerites triangular (Fig. 10 DV). Female genitalia (Fig. 12): In lateral view, segment VIII valves of R. sociata with posterior margin shallowly concave apically, with each dorsolateral apex extending farther posterad than ventrolateral apex. In R. denticulata, posterior margin with two small projections on each side, one dorsal and one ventral separated by modest indentation and ventrolateral apex longer than dorsolateral apex. In R. fasciata posterolateal margin on each side triangular and longer than small dorsolateral projection, excision between them corresponding to shallow posterior concavity of R. sociata and dorsolateral indentation of R. denticulata. In dorsal view, R. sociata with indentation between broad segment VIII valves, apicodorsal ends of valves with rounded apical and subapical corners and delimiting subcircular space. Rhyacophila denticulata also with apicodorsal ends of valves broad but with apical and subapical corners more nearly rectilinear than in R. sociata and delimiting transversely oval space. In R. fasciata apicodorsal ends of segment VIII valves small, blunt, with delimited space more nearly triangular, wider anteriorly. In ventral view, R. sociata with segment VIII valves forming two elongated sclerites, fused basally, intersegmental membrane with pair of dark sclerites united anteriorly to form heart-shape structure. In R. denticulata, segment VIII valves not united basally, two intersegmental membrane sclerites coma-shaped, not fused, thicker in anterior 1/3 and more widely separated posteriorly. In R. fasciata, segment VIII valves widely separated and not fused basally, intersegmental membrane sclerites longitudinally oval and more widely separated than in R. denticulata. Genetic analysis. The first approximations at the genetic level for the European samples of R. fasciata, R. denticulata, and R. sociata showed that R. fasciata samples could be separated into at least two different groups: One group of specimens from Austria and a second one from central-northern Europe. The difference was sufficient to considerer these groups to represent at least two species. A study of these two species will be conducted in the future. Since the original description of R. fasciata was based in specimens from Carinthia (Austria), we used the sequences from Austria for the comparison with R. denticulata and R. sociata. The mitochondrial sequences analyzed show clear differences between the R. fasciata specimens from Austria and those of R. denticulata and R. sociata. Based on the divergence of sequences, there are two differentiated branches, one corresponding to R. fasciata and another that includes R. sociata and R. denticulata species (Fig. 13). The specific clades are supported by very high bootstrap values in the three cases (98 for R. fasciata, and 99 for the R. sociata ��� R. denticulata node and for each of the R. sociata and R. denticulata branches). The genetic distances (maximum composite likelihood model) between the species, including an outgroup species (R. relicta McLachlan 1879), were 0.041 (R. sociata ��� R. denticulata), 0.052 (R. sociata ��� R. fasciata) and 0.056 (R. denticulata ��� R. fasciata). The pairwise distances between each of these species and the outgroup were 0.076 (R. relicta ��� R. fasciata), 0.095 (R. relicta-R. sociata), and 0.103 (R. relicta ��� R. denticulata). The maximum intraspecific distances obtained were 0.0062 for R. sociata, 0.0053 for R. denticulata, and 0.0040 for R. fasciata., Published as part of Valladolid, Mar��a, Arauzo, Mercedes, Basaguren, Ana, Dorda, Beatriz A. & Rey, Isabel, 2018, The Rhyacophila fasciata Group in Western Europe: Confirmation of Rhyacophila denticulata McLachlan 1879 (stat. prom.) and Rhyacophila sociata Nav��s 1916 (stat. res.), based on morphological and molecular genetic evidence (Trichoptera: Rhyacophilidae), pp. 526-544 in Zootaxa 4418 (6) on pages 530-534, DOI: 10.11646/zootaxa.4418.6.2, http://zenodo.org/record/1245456

Genus Discophrya Lachmann, 1859 Freshwater suctorians. Evaginative swarmer is dorsoventrally flattened and ciliated on the ventral side. Cup-shaped species without lorica, and when they have lorica, it is a extension of the stalk. Cellular body flattenned. Stalk is strong especially in symphoriont species, with a proximal widening. Contractile vacuole easily distinguishable. (Matthes 1988; Lynn & Small 2000)., Published as part of Fern��ndez-Leborans, Gregorio, Valladolid, Mar��a, Arauzo, Mercedes, Mill��n, Andr��s, Gabilondo, Regina, Gallardo, Manuela, Jim��nez, Luis & Ram��rez-Ballesteros, Mireya, 2017, Epibionts on Hydraena species (Coleoptera: Hydraenidae) from high mountain rivers of Pyrenees (Ordesa and Monte Perdido National Park), with the description of a new species, pp. 79-94 in Zootaxa 4317 (1) on page 85, DOI: 10.11646/zootaxa.4317.1.3, http://zenodo.org/record/880068, {"references":["Matthes, D. (1988) Suctoria. In: Matthes, D., Guhl, W. & Haider, G. (Eds.), Suctoria und Urceolariidae (Peritricha). Gustav Fischer Verlag, Stuttgart, pp. 1 - 226.","Lynn, D. H. & Small, E. B. (2000) Phylum Ciliophora. In: Lee, J. J., Leedale, G. F. & Bradbury, P. (Eds.), An Illustrated Guide to Protozoa. Allen Press, Kansas, pp. 371 - 656."]}

Description of Discophrya ordesae Fernández-Leborans & Valladolid, sp. n. Derivatio nominis: from Ordesa valley (Pyrenees, Comunity of Aragón, Spain). Ciliate suctorians located mainly in ventral areas of abdomen and thorax (in this case between legs), in number of 1–2, rarely 3 individuals per basibiont (Figure 5). Fanlike body (222–238 µm long, and 255–277 µm wide) with distal end uniformly rounded and proximal end strait and connected to the stalk (Figures 6–7). Body very flattened. Distal edge of the lorica double with capitate tentacles inserted (135–148), of variable length (Figures 8–9) as they are retracted or not (5.3–15.5 µm retracted; 27.2–102.2 extended). Stalk short (86.3–97.2 µm) and thick. Distal end of stalk joining to the basibiont by a slightly expanded region. Proximal part of the stalk continued by a lorica (stylotheca) with thick edge and strong surrounding the cellular body. Macronucleus located in the middle of body (Figure 10–11), oval or rounded (100.4–125 µm long, 109.3– 127 µm wide). In several specimens, larval kineties may be observed in the centre of the body (Figure 12), with the cilia protruding laterally, probably constituting the development of the swarmer on brood pouch (Table 4). Evaginative budding. Registration numbers: MNCN 39.04/5-39.04/7. Distribution. It was found attached to ventral area, mainly in abdomen, of Hydraena delia and H. truncata (Coleoptera: Hydraenidae). From Yaga river (Table 2).

Description of Tokophrya cf. stenostyla Hamilton & Jahn The suctorians analyzed (Figures 2–3) had an oval elongated body (51–53 µm long, 20–42 µm wide). In several individuals there was a cap-shaped region protruding from the front end of the body. The posterior end of the body was truncated just above the junction of the stalk. Tentacles spread at the distal end, not grouped in actinophores. There were 20–25 tentacles, directly attached to the surface of the ciliate. The stalk was long (105–121 µm), and its posterior end was widen slightly in attachment with substrate. The anterior end showed a widening near the body, composed by an unequal double ring. The macronucleus was spherical or oval (19–29 µm long, 17–27 µm wide), located in the middle or slightly towards the anterior half of the body (Table 3). Registration numbers: MNCN 39.04/2-39.04/4 Distribution. They were found attached principally to the dorsal area (Figure 4) of elytra of several species of Hydraena: H. delia, H. subimpressa and H. truncata (Coleoptera: Hydraenidae). Found in rivers Bellós and Yaga. (Table 2)., Published as part of Fernández-Leborans, Gregorio, Valladolid, María, Arauzo, Mercedes, Millán, Andrés, Gabilondo, Regina, Gallardo, Manuela, Jiménez, Luis & Ramírez-Ballesteros, Mireya, 2017, Epibionts on Hydraena species (Coleoptera: Hydraenidae) from high mountain rivers of Pyrenees (Ordesa and Monte Perdido National Park), with the description of a new species, pp. 79-94 in Zootaxa 4317 (1) on page 83, DOI: 10.11646/zootaxa.4317.1.3, http://zenodo.org/record/880068

[EN]: Nitrate pollution from diffuse sources is the main cause of groundwater quality degradation. The Nitrates Directive of the European Union (EU; 91/676/EEC) establishes that water resources should be considered affected by nitrate pollution when their nitrate contents exceed 50 mg L-1. The Directive defines Nitrate Vulnerable Zones (NVZ) as the areas of land draining into waters affected by nitrate pollution. In these areas farmers are required to comply with the measures laid out in Action Programmes designed to improve water quality. However, one major obstacle to a more efficient implementation of EU environmental policies for nitrate pollution control is the lack of consensus on the criteria to be used for designating NVZ. In this regard, the use of inappropriate criteria for the designations can seriously reduce the effectiveness of the Action Programmes implemented in the NVZ. Bearing all this in mind, the new LU-IV procedure (Arauzo, 2017), for assessing and mapping groundwater vulnerability to nitrate pollution from diffuse sources, was applied to the territory of the Community of La Rioja (northern Spain; Fig. 1) in which, several aquifers (Table I) are chronically affected by nitrate pollution. More specifically, the study had the following objectives: (1) to analyse the distribution of nitrate contents in the 14 aquifers that, totally or partly, belong to the Community of La Rioja (Table I; Fig. 1), (2) to generate the thematic maps of the intrinsic groundwater vulnerability and the specific groundwater vulnerability to nitrate pollution, using the LU-IV procedure, and (3) to define the NVZ in the study area. The LU-IV procedure combines a map of intrinsic vulnerability (based on the IV index) with a map of the risks associated with different land uses (using the Over tool from logical toolset of Spatial Analyst Tools in ArcGIS 10.3). The most basic formulation of the IV index uses four environmental parameters that are commonly related to intrinsic groundwater vulnerability: lithology of the vadose zone, depth to groundwater table, topography slope and annual precipitation (Table II). This method stands out as it meets the following requirements: (1) it uses readily available parameters that provide enough data to feed the model, (2) it excludes redundant parameters, (3) it avoids the need to assign insufficiently contrasted weights to parameters, (4) it assesses the entire catchment area that potentially drains N-polluted waters into the receptor aquifer, (5) it is implementable within a GIS, and (6) it provides a multi-scale representation (Arauzo, 2017). The spatial modelling of nitrate contents in the 14 aquifers under study (Fig. 2) showed large nitrate polluted areas in all of the alluvial aquifers, which exhibited the worst conditions. It was observed that the alluvial aquifers located in first- and second-order basins (aquifers no 044, 045 and 047) showed significant negative correlations between nitrate concentration and water table elevation (Table I). This result was consistent with the existence of stagnant groundwater zones in the lower sections of these aquifers (Fig. 2), where nitrates tend to accumulate. The map of the intrinsic groundwater vulnerability (based the IV index; Fig. 3) showed high to extreme levels of intrinsic vulnerability across 12.3% of the territory of La Rioja (617 km2) and moderate levels in 21.6 % (1.084 km2). All the alluvial areas were within the range of high to extreme intrinsic vulnerability. The map of the specific groundwater vulnerability to nitrate pollution (according the LU-IV procedure; Fig. 4) showed high to extreme levels of specific vulnerability across 483 km2 and moderate levels in 527 km2. To date, the Community of La Rioja has officially designated a total of three NVZ within the study area, jointly covering a surface of 124 km2 (Fig. 1). It was estimated that 53.3% of the alluvial areas in La Rioja exhibited high to extreme vulnerability to nitrate pollution, while 23.8% was subject to moderate risk (Fig. 4). This high vulnerability can be attributed to the combined effect of some typically alluvial characteristics, including: the shallow water table in the alluvial areas, the interconnections between rivers and alluvial aquifers, the permeability of the alluvial deposits and the typically high concentration of irrigated agriculture on the lower terraces and floodplains along river banks. These results highlight the need to redefine what have been the officially designed NVZ in the study area until now. First of all, it seems necessary to ensure the effective protection of the alluvial aquifers, by extending the designations of the NVZ to all the alluvial areas (this high level of protection should be generalized for all the alluvial areas in Europe). On the other hand, territories with moderate vulnerability to nitrate pollution (beyond the alluvial areas) should also be monitored and controlled, given their wide extension. In other words, if we want to effectively implement Action Programmes designed to restore and protect water quality, in line with Directive 91/676/EEC, it is first necessary to review and redefine the current criteria for the designation of NVZ., [ES]: El procedimiento LU-IV es una nueva herramienta que permite cartografiar la vulnerabilidad específica de las aguas subterráneas a la contaminación por nitrato y delimitar las Zonas Vulnerables a la Contaminación por Nitrato (ZVN; Directiva 91/676/CEE). Combina un mapa de vulnerabilidad intrínseca (a partir del índice IV, que estima los riesgos asociados a parámetros del medio físico) y un mapa de riesgos asociados a los usos del suelo (LU). El procedimiento ha sido aplicado al territorio de la Comunidad de La Rioja (España). El mapa de vulnerabilidad específica reveló que una superficie de 483 km2 presentaba niveles de vulnerabilidad de alto a extremo, mientras que otros 527 km2 mostraban niveles medios. Centrándonos en los territorios aluviales (los más vulnerables, atendiendo a factores del medio físico), el 53,3% presentó una vulnerabilidad específica de alta a extrema, el 23,8% niveles medios y el 22,9% niveles bajos o inapreciables. De estos resultados se desprende la necesidad de una protección integral de la totalidad de los territorios aluviales de La Rioja mediante la ampliación de las designaciones de ZVN. Las zonas que presentaron niveles medios de vulnerabilidad en territorios no aluviales también deberían ser susceptibles de seguimiento, dada su amplia extensión., Este trabajo ha sido financiado mediante los proyectos AGL2011-29861/AGR y CGL2016-81110-R (AEI/EU). La Confederación Hidrográfica del Ebro aportó gran parte de los datos hidroquímicos y de profundidades del nivel freático.

Ponencia Invitada Inaugural del Congreso de Espeleología "EspeleoMeeting Ciudad de Villacarrillo" (Villacarrillo (Jaén), 13, 14 y 15 de agosto de 2016)., [ES] En este trabajo se describe a través de la citología, histología y secuenciación del gen de la rodopsina, la única población de peces encontrada en una cueva de la Península Ibérica. Esta población pertenece a la especie Squalius alburnoides (Steindachner, 1866) (Cyprinidae) y tiene la particularidad de tener un origen híbrido con individuos y poblaciones de diferentes ploidias (2n=50; 3n=75 y 4n=100) y distintos mecanismos de reproducción. Los peces de la cueva de Peal del Becerro I presentaron una degeneración incipiente de la retina, con menor cantidad de fotorreceptores y pigmentación que los peces de la misma especie situados en el exterior de la cueva (Embalse de La Bolera). Todos resultaron ser tetraploides simétricos AAPP lo que sugiere que este biotipo favorece de algún modo la viabilidad de poblaciones de esta especie permitiendo recuperar la bisexualidad y un genoma balanceado. Sin embargo, es necesario realizar estudios en profundidad desde la genómica y transcriptómica y tener un mejor conocimiento de la biología y ecología de esta población cavernícola., [EN] In this study we described through a multidisciplinary approach (cytology, histology and rhodopsin gene sequencing) the only cave population of freshwater fish found in the Iberian Peninsula. This population belong to the species Squalius alburnoides (Steindachner, 1866) (Cyprinidae), which has the singularity of being a species of hybridogenetic origin with individuals and populations of different ploidies (2n=50; 3n=75 y 4n=100) and diverse mechanisms of reproduction. The individuals of S. alburnoides found in the Peal del Becerro I cave showed an incipient degeneration of the retina, with lower amount of photoreceptor cells and pigmentation relative to individuals outside the cave (de la Bolera reservoir). All individuals from the cave were symmetric tetraploids AAPP, which suggests that this biotope is favouring the viability of this population through the recovery of bisexuality and a balanced genome. Nonetheless, further deeper studies using genomics and transcriptomics approaches are required to achieve a better understanding of the biology and ecology of this cave-dweller population.