Revision of the Dicranotropishamata group (Auchenorrhyncha, Delphacidae) and remarks on the implication of chiral dimorphism in its history Author Guglielmino, Adalgisa Author D'Urso, Vera Author Bueckle, Christoph text Deutsche Entomologische Zeitschrift 2016 63 1 89 108 http://dx.doi.org/10.3897/dez.63.6625 journal article http://dx.doi.org/10.3897/dez.63.6625 1860-1324-1-89 59C70D2324BD4D07BD17D7736A26A0EC Dicranotropis remaniaca sp. n. Figs 9-16, 17-26, 44-46 Measurements. Body length: 2.45-2.90 mm (brachypterous males), 3.85-4.20 mm (macropterous males), 2.90-3.40 mm (brachypterous females), 4.30-4.50 mm (macropterous females). Head length: 0.20-0.28 mm (males), 0.26-0.30 mm (females). Head width including eyes: 0.78-0.92 mm (males), 0.88-0.96 mm (females). Pronotum length: 0.20-0.25 mm (males), 0.22-0.26 mm (females). Mesonotum length: 0.40-0.50 mm (brachypterous males), 0.58-0.64 mm (macropterous males), 0.44-0.52 mm (brachypterous females), 0.66-0.80 mm (macropterous females). Length of fore wings from shoulder to wing tip: 1.15-1.48 mm (brachypterous males), 3.15-3.50 mm (macropterous males), 1.24-1.48 mm (brachypterous females), 3.65-3.75 mm (macropterous females). Description. In size, coloration and shape very similar to D. hamata : Median carina of frons forked below junction with vertex (Figs 15, 16); lateral carinae of pronotum not reaching hind margin; wings of brachypterous specimens between 1.5 and 2 x longer than wide, apically rounded (Figs 9, 14). Figures 1-8. Dicranotropis hamata (Boheman), male. 1: Aedeagus, left lateral view. 2: Aedeagus, right lateral view. 3: Aedeagus, dorsal view. 4: Aedeagus, ventral view. 5: Right genital style from inside. 6: Pygofer, ventral view. 7: Pygofer, left lateral view. 8: Pygofer, caudal view. (1-5: Germany, Baden-Wuerttemberg , Freudenstadt. 6-8: Germany, Bayern, Berchtesgaden). Figures 9-16. Dicranotropis remaniaca sp. n. (Italy, Latium, Rieti). 9: Brachypterous male, dorsal view. 10: Brachypterous male, dorsolateral view. 11: Brachypterous male, lateral view. 12: Macropterous male, dorsal view. 13: Macropterous female, dorsal view. 14: Brachypterous female, dorsal view. 15: Brachypterous male, frontal view. 16: Brachypterous female, frontal view. Coloration . Males (Figs 9-12, 15): Face with carinae white and areas between carinae black or light brown bordered with black; vertex light brown, pronotum light brown with carinae white; mesonotum light brown or more or less dark with white central longitudinal stripe extending onto scutellum; upper side of abdomen black, often with central part and some spots on lateral parts more or less light brown; pygofer black with more or less extended light brown areas; anal tube white; anal style black; forewings (brachypterous) hyaline brown, in brachypterous specimens apical half of suture black with adjacent area of wing dark, basal half of suture and hind margin with adjacent veins white; in macropterous specimens forewings hyaline with apical half of clavus and narrow adjacent area dark; underside mostly black; legs black with knees, tibiae and tarsi light or dark brown, third tarsomere generally dark. Females (Figs 13, 14, 16): similar to males but generally lighter: areas between frontal carinae light brown narrowly bordered with black; dark spot on wing suture small; upper side of abdomen in great part light brown; ovipositor sheath light; femura often in part light brown. Genital morphology. Males (Figs 17-26): Pygofer with distinctly protruding dorsocaudal protuberance on each side; protuberances apically with small and short spine in medioventral position (Figs 22-24); anal tube on each side with small tooth of variable size near the base in subbasal position (Figs 25, 26); styles subbasally on the mediocaudal side with scabrous surface and acute spine shaped process, in the middle distinctly curved mediocaudad and provided with preapical tooth (Fig. 21); aedeagus laterally depressed, ventrally bent, with phallotreme on the right side, only in rare exceptions on the left side; on its dorsal margin in central position with carina comprised of varying number of fused teeth and in preapical position with large single tooth, both bent towards right side; on right side, close to ventral margin, with group of about three small teeth in preapical position and, basally of them, single large tooth curved somewhat dorsad; on left side very close to ventral margin with one or more series of small teeth, varying in size and number, and with group of about three teeth more apically and quite distant from each other and from ventral aedeagus margin (Figs 17-20). Females: Gonocoxae VIII wide, median margin equally convex (Fig. 46); genital scale distinct, +/- triangular, with narrow deep apical incision reaching about half length of genital scale (Figs 44, 45). Figures 17-26. Dicranotropis remaniaca sp. n., male. 17: Aedeagus, left lateral view. 18: Aedeagus, right lateral view. 19: Aedeagus, dorsal view. 20: Aedeagus, ventral view. 21: Right genital style from inside. 22: Pygofer, ventral view. 23: Pygofer, left lateral view. 24: Pygofer, caudal view. 25: Anal tube, lateral view. 26: Anal tube, caudal view. (17-21: Italy, Emilia Romagna, Sologno. 22-24: Italy, Abruzzo, Campotosto. 25, 26: Italy, Abruzzo, Sella di Corno). Remarks. The pygofer and aedeagus morphology (in males), e.g. width of the aedeagus, number of the aedeagal spines (Figs 159-188), and shape of pygofer protuberances (Figs 201-206, 219-224, 249-256), and the morphology of the genital scale (in females) is to some degree variable, and apparently there are also slight regional differences (e.g. aedeagi of specimens from northern Italy, southern Switzerland and Slovenia (Figs 167-180) are particularly slender). For the variability of the genital styles see Figs 84-97. Figures 27-32. Transitional forms between Dicranotropis hamata (Boheman) and D. remaniaca sp. n. (hybrids?). 27: Aedeagus, left lateral view. 28: Aedeagus, right lateral view. 29: Right genital style from inside. 30: Aedeagus, left lateral view. 31: Aedeagus, right lateral view. 32: Right genital style from inside. (27-29: France, Haute-Garonne, Luchon. 30-32: France, Haute-Garonne, Luchon (other specimen)). Figures 33-39. Dicranotropis zenata Logv. (Georgia, Goderdsi). 33: Aedeagus, left lateral view. 34: Aedeagus, right lateral view. 35: Aedeagus, dorsal view. 36: Right genital style from inside. 37: Pygofer, ventral view. 38: Pygofer, left lateral view. 39: Pygofer, caudal view. 40-42. Dicranotropis sagata Logv. (Georgia, Sasaredneo). 40: Aedeagus, left lateral view. 41: Aedeagus, right lateral view. 42: Right genital style from inside. Figures 43-49. Dicranotropis hamata group, females. 43: Dicranotropis hamata (Boheman) (Bulgaria, Western Rhodopes, Eleshnica), genital scale. 44: Dicranotropis remaniaca sp. n. (Slovenia, Nova Gorica), genital scale. 45: Dicranotropis remaniaca sp. n. (Italy, Umbria, Lago di Piediluco), genital scale. 46: Dicranotropis remaniaca sp. n. (Italy, Basilicata, M. Sirino), abdomen, ventral view. 47: Dicranotropis zenata Logv. (Georgia, Goderdsi, Advigeki), genital scale. 48: Dicranotropis sagata Logv. (Georgia, Sasaredneo), genital scale. 49: Dicranotropis sagata Logv. (Georgia, Sasaredneo), abdomen, ventral view. Figures 50-73. Dicranotropis hamata (Boheman), right genital style from inside. 50: France, Herault . 51: France, Nantes. 52: France, Jura. 53: France, Ardeche . 54: France, Vaucluse. 55: Finland. 56: Denmark. 57: Switzerland, Jura. 58: Germany, Baden-Wuerttemberg , Tuebingen . 59: Germany, Bayern, Berchtesgaden. 60: Austria, Kaernten , Woerthersee. 61: Austria, Steiermark, Leibnitz. 62: Austria, Niederoesterreich , Himberg. 63: Czech Republic, Moravia, Mikulov. 64: Hungary, Velence. 65: Slovenia, Zgornje Jezersko. 66: Serbia, Dzep . 67: Macedonia, Strumica distr.. 68: Bulgaria, Goce Delchev. 69: Bulgaria, Western Rhodopes, Eleshnica. 70: Greece, Katara region, Pindos. 71: Greece, Katara region, Pindos (other specimen). 72: Ukraine, Crimea. 73: Russia, Krasnodar. Figures 74-97. Dicranotropis hamata group, right genital style from inside. Dicranotropis hamata (?). 74: Poland, Zdory. 75-83. Transitional forms between D. hamata (Boheman) and D. remaniaca sp. n. (hybrids?). 75: France, Gard. 76: France, Saone-et-Loire. 77: France, Ariege . 78: France, Aude, Laroque de Fa. 79: France, Toulouse. 80: France, Haute-Garonne, St. Beat . 81: France, Tarn, Albi. 82: France, Pyrenees Orientales, Mt. Louis. 83: France, Haute-Garonne, Arlos. 84-97. Dicranotropis remaniaca sp. n.. 84: Spain, Teruel. 85: Spain, Lerida. 86: Spain, Girona, Ripoll. 87: Germany, Bayern, Fuessen , Trauchberg. 88: Switzerland, Canton Ticino. 89: Switzerland, Canton Ticino, Monteggio. 90: Switzerland, Canton Ticino, Monteggio (other specimen). 91: Slovenia, Bovec. 92: Slovenia, Nova Gorica. 93: Slovenia, Vogrsko. 94: Italy, Valle d'Aosta , Monte Avic. 95: Italy, Abruzzo, Sella di Corno. 96: Italy, Abruzzo, Campotosto. 97: Italy, Basilicata, Monte Sirino. Figures 98-129. Dicranotropis hamata (Boheman), aedeagus, left and right lateral view. 98, 99: France, Jura. 100, 101: France, Nantes. 102, 103: France, Herault . 104, 105: France, Ardeche . 106, 107: France, Vaucluse. 108, 109: Denmark. 110, 111: Switzerland, Jura. 112, 113: Germany, Baden-Wuerttemberg , Tuebingen . 114, 115: Germany, Bayern, Berchtesgaden. 116, 117: Austria, Niederoesterreich , Himberg. 118, 119: Austria, Steiermark, Leibnitz. 120, 121: Austria, Kaernten , Woerthersee. 122, 123: Czech Republik, Moravia, Mikulov. 124, 125: Hungary, Velence. 126, 127: Slovenia, Zgornje Jezersko. 128, 129: Serbia, Dzep . Figures 130-157. Dicranotropis hamata group, aedeagus, left and right lateral view. 130-143. Dicranotropis hamata (Boheman). 130, 131: Macedonia, Strumica distr.. 132, 133: Bulgaria, Western Rhodopes, Eleshnica. 134, 135: Bulgaria, Goce Delchev. 136, 137: Greece, Katara region, Pindos. 138, 139: Greece, Katara region, Pindos (other specimen). 140, 141: Ukraine, Crimea. 142, 143: Russia, Krasnodar. 144-157. Transitional forms between D. hamata (Boheman) and D. remaniaca sp. n. (hybrids?). 144, 145: France, Ariege . 146, 147: France, Gard. 148, 149: France, Saone-et-Loire. 150, 151: France, Aude, Laroque de Fa. 152, 153: France, Haute-Garonne, St. Beat . 154, 155: France, Tarn, Albi. 156, 157: France, Haute-Garonne, Arlos. 158: Transitional forms between D. hamata (Boheman) and D. remaniaca sp. n. (hybrids?), aedeagus, left lateral view, France, Toulouse. Figures 159-188. Dicranotropis remaniaca sp. n., aedeagus, left and right lateral view. 159, 160: Spain, Girona, Ripoll. 161, 162: Spain, Teruel. 163, 164: Spain, Lerida. 165, 166: Germany, Bayern, Fuessen , Trauchberg. 167, 168: Switzerland, Canton Ticino, Monteggio. 169, 170: Switzerland, Canton Ticino, Monteggio. 171, 172: Switzerland, Canton Ticino. 173, 174: Slovenia, Bovec. 175, 176: Slovenia, Nova Gorica. 177, 178: Slovenia, Vogrsko. 179, 180: Italy, Valle d'Aosta , Monte Avic. 181, 182: Italy, Abruzzo, Campotosto. 183, 184: Italy, Abruzzo, Sella di Corno. 185, 186: Italy, Abruzzo, Sella di Corno (other specimen). 187, 188: Italy, Basilicata, Monte Sirino. Figures 189-206. Dicranotropis hamata group, pygofer, ventral view. 189-197: Dicranotropis hamata (Boheman). 189: France, Nantes. 190: Finland. 191: Switzerland, Jura. 192: Germany, Baden-Wuerttemberg , Kaiserstuhl. 193: Germany, Bayern, Berchtesgaden. 194: Austria, Niederoesterreich , Himberg. 195: Slovenia, Zgornje Jezersko. 196: Macedonia, Strumica distr.. 197: Russia, Krasnodar. 198-200: Transitional forms between D. hamata (Boheman) and D. remaniaca sp. n. (hybrids?). 198: France, Haute-Garonne, St. Beat . 199: France, Aude, Laroque de Fa. 200: France, Haute-Garonne, Luchon. 201-206. Dicranotropis remaniaca sp. n.. 201: Switzerland, Canton Ticino. 202: Switzerland, Canton Ticino, Monteggio. 203: Slovenia, Vogrsko. 204: Italy, Basilicata, Monte Sirino. 205: Germany, Bayern, Fuessen , Trauchberg. 206: Spain, Girona, Ripoll. Figures 207-224. Dicranotropis hamata group, pygofer, caudal view. 207-215: Dicranotropis hamata (Boheman). 207: France, Nantes. 208: Finland. 209: Switzerland, Jura. 210: Germany, Baden-Wuerttemberg , Kaiserstuhl. 211: Germany, Bayern, Berchtesgaden. 212: Austria, Niederoesterreich , Himberg. 213: Slovenia, Zgornje Jezersko. 214: Macedonia, Strumica distr.. 215: Russia, Krasnodar. 216-218: Transitional forms between D. hamata (Boheman) and D. remaniaca sp. n. (hybrids?). 216: France, Haute-Garonne, Luchon. 217: France, Aude, Laroque de Fa. 218: France, Haute-Garonne, St. Beat . 219-224: Dicranotropis remaniaca sp. n.. 219: Switzerland, Canton Ticino. 220: Switzerland, Canton Ticino, Monteggio. 221: Slovenia, Vogrsko. 222: Italy, Basilicata, Monte Sirino. 223: Germany, Bayern, Fuessen , Trauchberg. 224: Spain, Girona, Ripoll. (the arrows in Figures 210, 213-215 indicate the position of the preapical tooth) Figures 225-256. Dicranotropis hamata group, pygofer, left lateral view. 225-242: Dicranotropis hamata (Boheman). 225: France, Nantes. 226: France, Herault . 227: France, Ardeche . 228: France, Vaucluse. 229: France, Jura. 230: Finland. 231: Denmark. 232: Switzerland, Jura. 233: Germany, Baden-Wuerttemberg , Kaiserstuhl. 234: Germany, Bayern, Berchesgaden. 235: Austria, Niederoesterreich , Himberg. 236: Hungary, Velence. 237: Slovenia, Zgornje Jezersko. 238: Serbia, Dzep . 239: Macedonia, Strumica distr.. 240: Greece, Katara region, Pindos. 241: Greece, Katara region, Pindos (other specimen). 242: Russia, Krasnodar. 243-248: Transitional forms between D. hamata (Boheman) and D. remaniaca sp. n. (hybrids?). 243: France, Haute-Garonne, St. Beat , 244: France, Aude, Laroque de Fa. 245: France, Haute-Garonne, Luchon. 246: France, Ariege . 247: France, Saone-et-Loire. 248: France, Gard. 249-256: Dicranotropis remaniaca sp. n.. 249: Switzerland, Canton Ticino. 250: Switzerland, Canton Ticino, Monteggio. 251: Slovenia, Vogrsko. 252: Italy, Basilicata, Monte Sirino. 253: Germany, Bayern, Fuessen , Trauchberg. 254: Spain, Girona, Ripoll. 255: Spain, Lerida. 256: Spain, Teruel. Diagnosis. Main differences to D. hamata consist in the shape of the genital styles and the aedeagus. The genital styles are stout, curved and provided with a preapical tooth in D. remaniaca while they are slender, straight, devoid of preapical tooth in D. hamata (Fig. 5). The aedeagus has its phallotreme on the right side, only in rare exceptions on the left side, while it is typically on the left side in D. hamata , and also in all other characters of the aedeagus D. remaniaca is the mirror image to D. hamata (Figs 1-4). Other differences lie in the shape of the pygofer which is in D. remaniaca generally with a less protruding dorsocaudal portion and further caudally and dorsally located preapical teeth, therefore these are often visible in lateral view (Figs 22-24), while D. hamata has a more protruding dorsal portion of the pygofer and the preapical teeth are not visible in lateral view (Figs 6-8). However, the pygofer characters are rather variable and can be misleading in some cases. Distribution (Fig. 257). Spain (Figs 84-86, 159-164); Switzerland south of the main Alpine chain (Canton Ticino) (Figs 88-90, 167-172); Italy except for Sicily and Sardinia and a small part in the northeastern Alpine region (Figs 94-97, 179-188); western Slovenia (Figs 91-93, 173-178); and some regions in Germany (southeastern Baden-Wuerttemberg , southwestern Bavaria) (Figs 87, 165, 166). Figure 257. Outline map of the distribution of the species of the Dicranotropis hamata -group. 1 = Dicranotropis remaniaca sp. n.; 2 = Dicranotropis hamata (Boheman); 1/2 = transitional forms between D. remaniaca and D. hamata (hybrids?); 2! = D. hamata specimens from northern Poland with particular style shape; 3 = D. zenata Logv.; (2) D. hamata specimen from northern Friuli, checked by Manfred Asche; (2!) D. hamata specimen figured by Ossiannilsson (1978) . Ecology. D. remaniaca shares its ecological characteristics with D. hamata and is found generally on not too dry meadows, often near forest margins or groups of bushes, from low to medium high altitude until about 1600m. Host plants are different species of Poaceae . Biology. The species was mostly found from beginning of June until end of August, but one record from April (340m) indicates that the taxon may be bivoltine in lowlands. In mountain regions it has apparently only one generation. Type series. Holotype, male: Lazio (Frosinone), Monti Ernici, road Collepardo-Veroli , east of Civita; N41°45.596' , E13°24.384' ; 735m; 09/08/2012; St. 679; dry open area with Poaceae , thistles, Thymus , Satureja and shadowy path near dry brook with Acer , Corylus etc.; Guglielmino & Bueckle leg.. Paratypes: Same data as holotype, 6♂♂, 2♀♀. - Emilia-Romagna (Parma), SP 81 3,9km east (direction Bedonia) of Passo Tomarlo (km 11,4); ~ 1200m; 09/06/2007; St.385; dry meadow and moderately moist meadow near beech wood, Salix myrsinifolia , Urtica , Poaceae ; Guglielmino & Bueckle leg.; 4♂♂, 6♀♀. - Same locality; 22/08/2008; St.444; Guglielmino & Bueckle leg.; 12♂♂, 10♀♀. - Emilia-Romagna (Parma), road from Ponteceno to Anzola, 2,4km east of Anzola; ~ 850m; 21/08/2008; St.443; meadow with Dactylis surrounded by Quercus , Acer , Corylus , Clematis ; Guglielmino & Bueckle leg.; 7♂♂, 6♀♀. - Toscana (Massa), Alpi Apuane, ca. 3km south of Vinca; ~ 1000m; 05/06/2008; St.420; mixed forest with Alnus cordata (?), and undergrowth with Rubus , ferns, Poaceae ; Guglielmino & Bueckle leg.; 4♂♂, 2♀♀. - Same locality; 17/08/2008; St .435; Guglielmino & Bueckle leg.; 5♂♂, 4♀♀. - Lazio (Rieti), Monti Reatini, M. Terminillo, S.P. 10, 4 Km from Leonessa; ~ 1200m; 22/8/1999; St. 36; vegetation along a brook; Guglielmino & Bueckle leg.; 6♂♂, 15♀♀. - Lazio (Rieti), East of Lago di Piediluco, Madonna della Luce, SS 79 near fork Labro, Km 29,5; N42°31 '15.0" , E12°46 '38.2" ; 372m; 21/8/2000; St. 86; herbaceous vegetation with Equisetum , Phragmites , Carex , Cyperaceae , between Ulmus , Salix , Quercus ; Guglielmino & Bueckle leg.; 8♂♂, 5♀♀. - Lazio (Rieti), Amatrice, ca. 1km south of Preta, Tronto river; ~ 1150m; 18/6/2005; St.154; undergrowth of mixed forest with Quercus cerris , few Ulmus , Salix , Populus tremula ; Guglielmino & Bueckle leg.; 11♂♂, 12♀♀. - Lazio (Rieti), Rieti, Riserva Ripasottile-Lago Lungo, st. 3, 22/7/2009; St.454; Guglielmino & Bueckle leg.; 8♂♂, 2♀♀. - Abruzzo ( L'Aquila ), slope south of Sella di Corno; ~ 1100m; 26/8/1999; St. 46; dry meadows with Ostrya carpinifolia Scop., Quercus , Acer ; Guglielmino & Bueckle leg.; 7♂♂, 1♀. - Abruzzo ( L'Aquila ), slope south of Sella di Corno; ~ 1200m; 26/8/1999; St. 47; meadows on the borders of a mixed forest; Guglielmino & Bueckle leg.; 8♂♂, 23♀♀. - Campania (Caserta), St.320: Strada da Gallo Matese a Fontegreca, prima del passo ~ 1,5 km a ovest di Gallo Matese, 850 m, 27/8/2006, prati fra siepi di Acer monspessulanum , Rosa , Prunus spinosa , Crataegus con Poaceae , Fabaceae ecc.; Guglielmino & Bueckle leg.; 6♂♂, 8♀♀. - Basilicata (Potenza), Monte Sirino, road to Lauria , fountain 7,5km south of fork to Moliterno; ~ 1000m; 02/08/2009; St.470; forest with Quercus , Crataegus , Alnus cordata , Spartium , Rubus and small open pasture with Poaceae , Lamiaceae , Holcus ; Guglielmino & Bueckle leg.; 2♂♂, 3♀♀. Type material deposited in Department of Agricultural and Forestry Sciences (DAFNE), University of Tuscia, Viterbo, Italy ( Guglielmino's collection) (CG), two male and two female paratypes in Senckenberg Naturhistorische Sammlungen Dresden, Museum fuer Tierkunde, Dresden, Germany. (For further material of this taxon see Suppl. material 1).