Despax, R. (1940). Bulletin de la Société d’Histoire Naturelle de Toulouse, 75:296. Vinçon, G. (1996). Bulletin de la Société Entomologique Suisse , 69:72 Dictyogenus fontium Two New Species Of Dictyogenus Klapálek, 1904 (Plecoptera: Perlodidae) From The Jura Mountains Of France And Switzerland, And From The French Vercors And Chartreuse Massifs Reding, Jean-Paul G. Launay, Bertrand Doaré, Jacques Le Ruffoni, Alexandre Vinçon, Gilles Illiesia 2019 15 2 27 64 8N8LS 2603979 Vincon, Launay, Le Doare, Ruffoni & Reding Vincon, Launay, Le Doare, Ruffoni & Reding 2019 Insecta Perlodidae Dictyogenus Animalia Plecoptera 13 40 Arthropoda species muranyii sp. nov.  http://lsid.speciesfile.org/urn:lsid: Plecoptera.speciesfile.org: TaxonName:506376 ( Figs. 27–51)    Dictyogenus fontium– Despax, R. (1940). Bulletin de la Société d’Histoire Naturelle de Toulouse, 75:296.    Dictyogenus fontium– Vinçon, G. (1996). Bulletin de la Société Entomologique Suisse, 69:72.    Dictyogenus fontiumgr sp 3-GV sensuGilles Vinçon (early-release DNA sequence on: www.boldsystems.org, unpublished)   Materials examined.   Holotypemale: FRANCE, Vercors Massif, Isèredepartment (38), Karstic Spring of Bruyantriver, above Engins, Lans-en- Vercors(38250), 45° 8.798123'N, 5° 37.049358'E,  982ma.s.l.,  09.06.2017, leg. G. Vinçon, deposited in the MZL(catalogue number: GBIFCH00652534).   Paratypes:same locality, same date, 3♂, 1♀, leg. G. Vinçon, deposited in the MZL(catalogue number: GBIFCH00652525, GBIFCH00652519); same locality, same date, 3L, leg. J.-P.G. Reding, deposited in the MZL(catalogue number: GBIFCH00652514).  Additional specimens. We examined many other specimens. These are stored in the collections of Bertrand Launay ( BLC), Gilles Vinçon ( GVC), Jean-Paul G. Reding ( RC), Dávid Murányi ( MC),   Fig. 31.  Dictyogenus muranyii  sp. n., male, hindwing. Karstic spring of Bruyant, Isère dpt, France. Photo B. Launay. Jacques Le Doaré ( JLDC), Alexandre Ruffoni ( ARC) and MZL.   FRANCE Vercors Massif Drôme department (26):   Archiane torrent, Drôme tributary, NE Châtillon-en-Diois, Menée, Cirque d’Archiane, 44° 44.770828'N, 5° 30.214097'E, 760m, 31.07.1990, 1♂;   16.09.1990, 2♀(leg. G. Vinçon; GVC).    Springat Brudour cave, Brudour River, Bournetributary, Bouvante(26190), 44° 55.67493'N, 5° 19.257459'E,  1182m,  13.04.2016, 3L;  02.06.2016, 2♂, 2♀, 1L, 7E (leg. B. Launay; BLC; used for molecular studies by IRSTEA, numbers B079 and B080);  21.05.2017, 1E (leg. G. Vinçon; GVC);   09.06.2017, 9E (leg. J.-P.G. Reding; RC).    Adouin river, near its spring, Vernaisonand Bournetributary, Tourtre, Saint-Martin-en- Vercors(26420), 45° 0.120928'N, 5° 27.550379'E,  793m,  21.05.2017, 1♂, 1♀(leg. G. Vinçon, GVC);   09.06.2017, 11♀, 1E (leg. G. Vinçon; RC; 1♀used for molecular studies by SwissBOL, MZL, catalogue number GBIFCH00280854); 2♂, 17♀(leg. G. Vinçon; GVC).    Cholet river, Lyonneand Bournetributary, Combe Laval, Saint-Laurent-en-Royans (26190), 44° 59.859883'N, 5° 20.75296'E,  355m,  21.05.2017, 2♂, 6E (leg. G. Vinçon; GVC). Isère department (38):   Bruyant river, Furon and Isère tributary, above Engins, Lans-en-Vercors (38250), 45° 8.798123'N, 5° 37.049358'E, 982m, 06.10.1991, 1E;  25.06.1995, 7♂, 4♀;  10.05.1998, 1♂;  10.06.2007, 1♂;  22.06.2008, 1♂;   22.06.2009, 32♂, 6♀( 1♂used for molecular studies by A. Reding);  07.07.2012, 7♂, 2♀, 1L (leg. G. Vinçon; GVC);  07.06.2015, 3♂, 2♀, 6L, 5E (leg. B. Launay; BLC);  09.06.2017, 2♂, 1♀, 17E (leg. G. Vinçon; GVC); 7♂, 1♀, 7L, 3E (leg. J.-P.G. Reding; RC; 1♂used for molecular studies by SwissBOL, MZL, catalogue number GBIFCH00280811).    Cuves de Sassenagestrong rheocrene spring, Germebrook, Furonand Isèretributary, Sassenage(38474), 45° 12.516243'N, 5° 39.078175'E,  330m,  15.06.1991, 1♂, 2♀strongly brachypterous (leg. G. Vinçon; MC).   Drevenne river, Isère tributary, Pont Chabert, Saint-Gervais (38470), 45° 10.643943'N, 5° 29.885486'E,  885m,  15.06.1991, 1L (leg. G. Vinçon; GVC);  31.05.2012, 2♂;  17.07.2012, 2L;   11.07.2015, 3L (leg. J. Le Doaré; JLDC);   06.06.2015, 1E (leg. B. Launay; BLC).    Font Noire river, Bournetributary, Villard-de- Lans(38250), 45° 4.472474'N, 5° 33.970966'E,  1010m,  01.06.2016, 1♂, 1E (leg. B. Launay; BLC; used for molecular studies by IRSTEA, number B120).    Fauge river, Bournetributary, Villard-de-Lans(38250), 45° 3.373805'N, 5° 34.664105'E,  1230m,  01.06.2016, 1E (leg. B. Launay; BLC).   Figs. 32-35.  Dictyogenus muranyii  sp. n., male terminalia. 32.Hemitergal lobes, dorsal view. Karstic spring of Font Noire, Isère dpt, France. Photo B. Launay. 33.Epiproct, lateral view. Karstic spring of Font Noire, Isère dpt, France. Photo B. Launay. 34.Lateral stylet, lateral view. Karstic spring of Brudour, Drôme dpt, France. Photo B. Launay. 35.Epiproct, dorso-caudal view. Karstic spring of Font Noire, Isère dpt, France. Photo B. Launay.   Fig. 36.  Dictyogenus muranyii  sp. n., female, subgenital plate. Karstic spring of Brudour, Drôme dpt, France. Photo B. Launay.    Spring of Furonriver, Lans-en-Vercors(38250), 45° 6.304551'N, 5° 36.221224'E,  1294m,  12.07.2015, 6L (leg. J. Le Doaré; JLDC). Chartreuse Massif Isère (38) and Savoie (73) departments:   Sarcenas river at Sarcenas, Isère tributary, (38700), 45° 16.829887'N, 5° 45.251584'E, 1093m, 20.04.1986, 5L; 01.06.1986, 4L;   05.10.1986, 1♂(leg. G. Vinçon; GVC).   Spring of Guiers-Vif river, Rhône tributary, Cirque de Saint-Même, Saint-Pierre-D’Entremont (73670), 45° 23.491629'N, 5° 53.464859'E, 1050m, 21.06.1992, 3♀;   12.09.1992, 4♀, 1E;   21.06.1992, 3♀(leg. G. Vinçon; GVC).   Karstic spring of Guiers Mort, Rhône tributary, Saint-Pierre-de-Chartreuse (38380), 45° 19.571976'N, 5° 51.452039'E, 1360m, 11.09.1938, stage and numbers not specified (leg. Ms. Daudin, fide Despax 1940).   Diagnosis.General color dark brown with tawny and yellow spots ( Figs. 27, 28). Males and females macropterous; only two femaleswere found strongly brachypterous. Apex of the frontal sclerite of the epiproct of adult males very slightly turned downwards, in lateral view ( Fig. 33). Female subgenital plate covering half of the abdominal sternum 9 ( Fig. 36). Body length of males 17.2 to 20.9 mm; females 18 to 23.6 mm. Anterior wings of males 15 to 19.3 mm; females 17.7 to 21.5 mm. Posterior wings of males 12.5 to 15.8 mm; females 15.3 to 17.4 mm.   Adults ( Figs. 27–36).Upper side of the head brown, with large yellow spots ( Fig. 28). M-line yellow, not interrupted in its middle ( Fig. 28). Between the lateral ocelli, a large, ovoid, yellow area delimitated posteriorly by the epicranial suture ( Fig. 28). Presence of a large, yellow, median band extending from the anterior margin of the pronotum to its posterior margin ( Fig. 28). Band slightly constricted in the middle and then steadily widening toward the posterior margin ( Fig. 28). A tawny area on each side of the pronotum, with dark, sculpted rugosities ( Fig. 28). Anterior and posterior angles of pronotum almost rectangular ( Fig. 28). Abdominal sterna 1 to 6 pale yellow, with two dark patches. Antennae dark brown; cerci light brown with basal part pale yellow ( Figs. 29, 30). Wing venation as typical for the genus ( Fig. 31; cf. Fig. 8). Forewing with the two cross-veins “ra- rp” and “rp- ma”   Fig. 37.  Dictyogenus muranyii  sp. n., larva. Karstic spring of Brudour, Drôme dpt, France. Photo A. Ruffoni. nearly aligned (like in Fig. 8). Numerous crossveins forming a reticulated area between RA and RP ( Fig. 31; cf. Fig. 8). Cross-vein “ra- rp” and subcostal area faintly infuscate (as in Fig. 8).  Male terminalia ( Figs. 29, 30, 32–35).Epiproct flanked by flat and spatulate lateral stylets ( Figs. 33, 34). Abdominal tergum 10 divided into hemiterga whose lobes are covered with a bunch of 20 to 25 long setae in which 3 to 6 stronger and longer spines (half of the length of the hemitergal lobes) are embedded ( Figs. 29, 32). Hemitergal lobes bulb-shaped with a slight distal knob ( Fig. 30), both pointing rearwards ( Fig. 30) and almost horizontally toward each other ( Figs. 29, 32). Apex of frontal sclerite of epiproct slightly turned downwards, in lateral view ( Fig. 33). Lateral stylets long, only slightly enlarged at apex, in lateral view ( Figs. 33, 34). Abdominal sternum 7 composed of multiple plates (as in Figs. 7, 59, 82).  Females ( Fig. 36).Females not formally identifiable to species level. Female subgenital plate ( Fig. 36) covering at most half of sternum 9. Its general shape is semi-circular with a shallow Vshaped median notch.  Mature larvae ( Figs. 37–45).Interocellar area with a narrow yellow patch nearly reaching lateral ocelli ( Figs. 37–39). Lateral ocelli with small lateral circum-ocellar yellow patch ( Fig. 38). A large, elliptical yellow patch above each lateral ocellus ( Figs. 37–39). M-line well visible only in its medial, crescent-shaped, section ( Figs. 37–39). Pronotum wider than long ( Figs. 38, 39). Medio-dorsal setae on pronotum long, but not compacted, sometimes covering only the posterior half of the pronotum ( Figs. 40, 41). Medio-dorsal row of setae present on mesonotum and metanotum ( Fig. 40). Abdominal terga with a row of short and sparse medio-dorsal setae ( Fig. 42). Paragenital plates, in ventral view, without spines, or with only 1 spine on one of the paragenital plates, but with numerous empty spine insertion points ( Fig. 44). Medio-dorsal row of swimming-hairs of caudal setae sparse, slightly longer than the diameter of the cerci ( Fig. 43). General aspect as in Fig. 37.  Egg characteristics ( Figs. 46–51).General shape triangular or trilateral in cross section. Posterior pole of egg regularly rounded; ridges protruding ( Figs. 46, 47). Chorionic surface spring of Brudour, Drôme dpt, France. Photo B. Launay. 43.Medio-dorsal setae on cerci. Karstic spring of Brudour, Drôme dpt, France. Photo A. Ruffoni. 44.Paragenital plates, ventral view. Karstic spring of Brudour, Drôme dpt, France. Photo B. Launay. 45.Stipe. Karstic spring of Brudour, Drôme dpt, France. Photo B. Launay.   Figs. 38-45.  Dictyogenus muranyii  sp. n., larva. 38.Head, pronotum and mesonotum, dorsal view. Karstic spring of Brudour, Drôme dpt, France. Photo B. Launay. 39.Head, pronotum, dorsal view. Karstic spring of Brudour, Drôme dpt, France. Photo A. Ruffoni. 40.Head, pronotum, mesonotum and metanotum, lateral view. Karstic spring of Bruyant, Isère dpt, France. Photo A. Ruffoni. 41.Pronotum, lateral view. Karstic spring of Bruyant, Isère dpt, France. Photo J.-P.G. Reding. 42.Abdominal segments, lateral view. Karstic   Figs. 46-51.  Dictyogenus muranyii  sp. n., egg characteristics. 46.Egg, upper view from one ridge with two faces and the micropylar region visible. Bruyant, Vercors Massif, Isère dpt, France. 47.Entire egg. Archiane, Vercors Massif, Isère dpt, France. 48.Egg collar. Archiane, Vercors Massif, Isère dpt, France. 49.Egg collar. Bruyant, Vercors Massif, Isère dpt, France. 50.Detail of micropyles. Bruyant, Vercors Massif, Isère dpt, France. 51.Detail of chorion and micropyles. Archiane, Vercors Massif, Isère dpt, France.   Figs. 52-53.  Dictyogenus muranyii  sp. n., biotopes. 52.Cave and Spring of Brudour river, Drôme dpt, France. Photo A. Ruffoni. 53.Spring of Adouin River, Vercors Massif, Isère dpt, France, in summer. Photo J. Le Doaré. with polygonal follicle cell impressions ( Figs. 48, 50, 51). Anchor papillate, donut shaped apically with central depression ( Figs. 48, 49). Micropyles not protruding and arranged regularly in a line in the middle of the egg ( Figs. 47, 50). Eclosion line absent ( Figs. 46, 47).  Comparison to Congeners.  Adults.In the adult male of  Dictyogenus muranyii  sp. n., a wide, V-shaped membranous area between the hemitergal lobe and the inner anterior corner of the hemitergum is present ( Figs. 29, 32), whereas this area is sclerotized and much narrower in  D. alpinum( Fig. 56). The lateral stylets in  D. muranyii  sp. n.are slightly enlarged apically ( Figs. 33, 34), intermediate in form between those of  D. alpinum( Fig. 57) and those of the  D. fontiumspecies complex ( Fig. 80). In the adult female of  D. muranyii  sp. n., the subgenital plate ( Fig. 36) covers at most the upper half of sternum 9, as is also the case for specimens of the  D. fontiumspecies complex ( Figs. 83, 84), whereas the subgenital plate of  D. alpinumcovers ¾ of sternum 9 ( Figs. 60, 61). The anterior and posterior angles of the pronotum of  Dictyogenus muranyii  sp. n., D. jurassicum  sp. n.and  D. fontiumarealmost rectangular ( Figs. 2, 28, 79), whereas those of  D. alpinumare rounded ( Fig. 54). Adult males and females of  Dictyogenus muranyii  sp. n.hence have more affinities with those of the  D. fontiumspecies complex than with those of  D. alpinum.  Mature larvae.  Dictyogenus muranyii  sp. n.differs from  D. fontiumby the presence of medio-dorsal setae on the pronotum ( Fig. 41compared to Fig. 86). Medio-dorsal setae on pronotum are long and sparse in  D. muranyii  sp. n.( Figs. 40, 41), whereas they are long and dense in  D. alpinum( Fig. 71), and short and scattered, arranged as two loosely demarcated rows in  D. jurassicum  sp. n.( Fig. 13).   Distribution and ecology.  Dictyogenus muranyii  sp. n.inhabits karstic springs (some of them intermittent; Fig. 52) in the French Vercors and Chartreuse massifs ( Fig. 92). The localities occurring on the south western part of the Vercors Massif belong to the Drôme watershed (main Rhône tributary); those occurring on the northern and eastern part of the Vercors Massif, and on the southern part of the Chartreuse Massif belong to the Isère watershed (main Rhône tributary), and those occurring on the western   Figs. 54-55.  Dictyogenus alpinum, male. 54.Head and pronotum. Vorz River, Isère dpt, France. Photo B. Launay. 55.Hemitergal lobes, lateral view. Vorz River, Isère dpt, France. Photo B. Launay.   Fig. 56.  Dictyogenus alpinum, male, hemitergal lobes, dorsal view. Nant Bénin River, Savoie dpt, France. Photo B. Launay.   Fig. 57.  Dictyogenus alpinum, male, epiproct, lateral view. Nant Bénin River, Savoie dpt, France. Photo B. Launay. part of the Chartreuse Massif belong to the Guiers watershed (Rhône tributary). The life cycle of  Dictyogenus muranyii  sp. n.is probably similar to the one of  D. jurassicum  sp. n.The main emergence period of adults is in May and June, although isolated specimens occur until September. We have noted on one occasion a spectacular upstream flight of females in mid-June on the river Adouin ( Fig. 53). Some of the females were ovipositing under large stones right at the spring head. Mature larvae emerge preferentially on small islets located in the middle of the river.   Etymology of  Dictyogenus muranyii sp. n.This species is dedicated to Dr. Dávid Murányi, Hungarian Natural History Museum, Budapest, Hungary, in recognition of his outstanding contributions to the plecopteran taxonomy. 3111831366 2017-09-06 MZL G. Vincon France Vercors 982 45.146637 Vercors Massif 1 5.6174893 Engins 13 40 1 holotype 3111831352 2017-09-06 MZL G. Vincon France Vercors 982 45.146637 Vercors Massif 1 5.6174893 Engins 13 40 4 1 3 paratype 3111831332 2017-09-06 MZL J. - P. G. Reding France Vercors 982 45.146637 Vercors Massif 1 5.6174893 Engins 13 40 1 paratype 3111831354 14 41 3111831307 [189,622,1102,1127] 1990-09-16 GVC G. Vincon 14 41 2 2 3111831355 2016-02-06 2017-05-21 2016-02-06 GVC B. Launay & G. Vincon Bouvante 1182 44.927914 Spring 1 5.3209577 Bourne 14 41 4 2 2 3111831322 [272,737,1354,1379] 2017-09-06 RC J. - P. G. Reding 14 41 1 3111831336 2017-05-21 GVC G. Vincon Vercors 793 45.002014 Adouin river 1 5.459173 Tourtre 14 41 2 1 1 3111831398 2017-09-06 MZL, GVC G. Vincon 14 41 31 29 2 3111831318 2017-05-21 GVC G. Vincon Combe Laval 355 44.997665 Cholet river 1 5.345883 Bourne 14 41 2 2 3111831306 14 41 3111831311 14 41 3111831367 [948,1117,923,947] 14 41 3111831389 [1132,1302,923,947] 14 41 3111831375 14 41 3111831331 2009-06-22 Reding 14 41 39 6 33 3111831323 14 41 3111831345 14 41 3111831386 GVC G. Vincon 14 41 8 1 7 3111831372 MZL J. - P. G. Reding 14 41 1 1 3111831396 1991-06-15 MC de Sassenage & G. Vincon Sassenage 330 45.208603 Cuves de Sassenage 1 5.651303 Isere 14 41 3 2 1 3111831309 1991-06-15 GVC G. Vincon 885 45.1774 1 5.498091 14 41 1 3111831353 [930,1096,1498,1522] 14 41 3111831317 [1108,1272,1498,1522] 14 41 3111831302 2015-11-07 JLDC J. Le Doare 14 41 1 3111831325 2015-06-06 BLC B. Launay 14 41 1 3111831314 2016-01-06 B. Launay Lans 1010 45.074543 Font Noire river 1 5.5661826 Bourne 14 41 1 1 3111831346 2016-01-06 BLC B. Launay Villard-de-Lans 1230 45.05623 Fauge river 1 5.577735 Bourne 14 41 1 3111831388 2015-12-07 JLDC J. Le Doare Lans-en-Vercors 1294 45.105076 Spring of Furon 1 5.6036873 Furon river 16 43 1 3111831347 16 43 3111831344 1986-05-10 GVC G. Vincon 16 43 1 1 3111831377 16 43 3111831391 [377,596,1472,1497] 1992-12-09 true 16 43 4 4 3111831380 1992-06-21 GVC G. Vincon 16 43 3 3 3111831321 16 43