Indalia pygmaeola pallifrons (Zeller, 1847) comb. nov.

Original combination: Lithosia pallifrons Zeller, 1847. Stettiner Entomologische Zeitung 8: 339.

T. L.: Festungsglacis (Glogau), Poland.

Material examined. SPAIN. ARABA: 1 ♀, Pto. de Azáceta, 800 m, 25.viii.1984, J. Gastón leg. ÁVILA: 1 ♂ and 1 ♀, Puerto Castilla, 1185 m, 17.viii.2001, J. Gastón leg. BARCELONA: 11 ♂ and 9 ♀, Coll d´Heures, Collsuspina, 950 m, 15.vii.2006, R. Macià leg.; 5 ♂ and 2 ♀, Parc Natural Castell de Montesquiu, Montesquiu, 637 m, 31TDG36, 11.viii.2007, R. Macià & J. Ylla leg.; 2 ♂, Balenyà, 587 m, 29.vii.1945, J. Vilarrubia leg. in coll. MCNB.; BURGOS: 1 ♀, Obarenes, 715 m, 18.vii.2003, J. Gastón leg.; 2 ♂, El Ribero, 750 m, 17.vii.1982, J. Gastón leg.; 1 ♂ and 1 ♀, Herrera de Ircío, 500 m, 5.viii.1994, J. Gastón leg.; 1 ♂ and 1 ♀, Páramo de Masa, 1050 m, 24.vii.2001, J. Gastón leg.; 7 ♂ and 6 ♀, El Cerro, Gredilla la Polera, 980 m, 31TUN40, 14.vi.2017, R. Macià leg.; 9 ♂ and 5 ♀, Quintanilla Sobresierra, 1060 m, 3.vii.2000, R. Macià & J. Ylla leg.; 5 ♂ and 3 ♀, Toca de Cuevaburgos, Covarrubias, 993 m, 30TVM65, 17.vii.2007, 13.vii.2012, R. Macià & J. Ylla leg.; 8 ♂ and 2 ♀, Paramos de Lora, Sargentes de Lora, 992 m, 30 TVN33, 7.vii.2015, R. Macià leg.; 2 ♂, San Pedro del Páramo, 975 m, 1.viii.1998, R. Macià leg.; CASTELLÓN: 2 ♂ and 1 ♀, Corral de Gracià, Alto Palancia, Barracas, 1029 m, 30TXK93, 16.vi.2009, R. Macià & J. Ylla leg.; CUENCA: 3 ♂ and 4 ♀, Sierra de Valdemeca, Collado Bajo, 1500 m, 31.vii.1999, R. Macià & J. Ylla leg.; 3 ♂ and 1 ♀, Herreria de los Chorros, Tragacete, 1510 m, 30.vii.2009, R. Macià & J. Ylla leg.; GIRONA: 1 ♂ and 3 ♀, Font Home Mort, Queralbs, 1750 m, 29.vii.2006, R. Macià & J. Ylla leg.; 4 ♂, Viladrau, 821 m, 8.ix.2019, 27.vii.1920, S. Novellas leg. in coll. MCNB.; GRANADA: 1 ♂, Sª Guillimona, Arroyo Montano, Huéscar, 1480 m, 11.viii.2010, J. Gastón leg.; 1 ♂, La Losa, Huéscar, 1300 m, 14.viii.1998, J. Gastón leg.; 1 ♂, Sª de Dúrcal, 1900 m, 2.viii.1998, J. Gastón leg. HUESCA: 1 ♀, Villanúa, 1100 m, 6.viii.2018, J. Gastón leg. LA RIOJA: 1 ♂, Villanueva de Cameros, 1150 m, 1.ix.1984, J. Gastón leg. LEÓN: 2 ♂ and 1 ♀, Mirantes de Luna, 1600 m, 2.viii.2008, J. Gastón leg.; 2 ♂ and 1 ♀, Las Médulas, 725 m, 21.vii.2005, R. Macià leg.; 7 ♂ and 9 ♀, Caldas de Luna, 1140 m, 23.vii.2006, R. Macià leg.; LLEIDA: 1 ♀, Pto. del Portillón, Val d’Arán, 1300 m, 2.viii.2018, J. Gastón leg.; 2 ♂, Parc Natural Cadí-Moixeró, Gòsol, 1500 m, 13.viii.1995, R. Macià leg.; 2 ♂, Sarroca de Bellera, 1050 m, 12.viii.2010, F. Vallhonrat leg. in coll. MCNB.; PALENCIA: 1 ♀, Velilla del Río Carrión, 1200 m, 24.vii.2009. J. Gastón leg. SEGOVIA: 1 ♂, Pto. de Pasapán, Sª Guadarrama, 1700 m, 18.viii.2001, J. Gastón leg.; 1 ♀, Casla, Sª Arcones, 1165 m, 17.vii.2009, J. Gastón leg. TARRAGONA: 4 ♂, El Tillar, Vimbodí-Poblet, 891 m, 31TCF37, 10.viii.2013, R. Macià.; TERUEL: 6 ♂ and 4 ♀, Camino Forestal, Calomarde, 1550 m, 30TXK26, 2.viii.2017, R. Macià leg.; 8 ♂ and 6 ♀, Valle de Valdevecar, Albarracín, 1125 m, 30TXK37, 28.vii.2009, 11.viii.2018, R. Macià leg.; 3 ♂ and 1 ♀, La Pajarera, El Vallecillo, 1483 m, 30TXK25, 18.vii.2018, R. Macià leg.; 8 ♂ and 4 f #, Pino Gordo, Moscardón, 1413 m, 21.vii.2001, 29.vii.2009, R. Macià leg.; 3 ♂, Torrecilla del Rebollar, 1200 m, 24.vii.1999, J. Gastón leg.; 2 ♂, Olalla, 1100 m, 28.vii.2006, J. Gastón leg. ZARAGOZA: 2 ♀, Torralba de los Frailes, 1050 m, 11.vii.1997, J. Gastón leg.

FRANCE: 5 ♂ and 3 ♀, Col de Serenne, Isère, 1800 m, 8.vii.2004, R. Macià & J. Ylla leg.; 3 ♂, Col de la Madone, Provence, Alpes Maritimes, 800 m, 24.vii.2000, R. Macià & J. Ylla leg.; 3 ♂, Mont Ventoux, Provence, 1020 m, 29.vii.1994, R. Macià & J. Ylla leg.; 3 ♂, Saint Crépin, Alpes-Côte d´Azur, 1025 m, 8.vii.2005, R. Macià leg.

Diagnosis. Indalia pygmaeola pallifrons is the subspecies present in most of Europe. In Britain, it is present in one locality in Kent (Dungeness). This subspecies is characterized by its larger size and yellow-ochre colour, the males with pale grey brush strokes on the forewings. In Indalia pygmaeola pygmaeola the forewings are a lighter straw-yellow and the hindwings are ivory coloured.

Re-description. Imago (Figs. 31–32). Average wingspan males 28.6 mm (n=10; 24–31 mm); average wingspan females 23.0 mm (n=10; 21–26 mm).

Genitalia (Fig. 64). Male genitalia: Similar to those of nominotypical Indalia pygmaeola, from which it differs in two details: the apical saccular process does not have a uniform curvature, but as it approaches the cucullus, it opens outwards; in the aedeagus, one of the cornuti is larger and stronger than the other two. Female genitalia: Identical to those of the nominotypical subspecies.

Immature stages (Fig. 87). The final instar larva is 13 mm long. Cephalic capsule well developed, with black upper area. Body with very short setae, with grey to light brown verrucae; dorsal line black, sometimes broken, on each side two light brown spots in front of D1 and behind D2; sides with scattered black spots; anal shield with 4 scattered small verrucae. Pupa thick, with spots on the dorsal area; clypeus indistinct. Pupae of both sexes very similar. The cocoon among the lichens and detritus, with little silk.

Molecular data. The four samples analysed for this species cluster together in a well-supported clade (PP=1) originating close to the base of the Indalia genus. RESL Cluster Analysis returned a single cluster and all sequences matched the single BIN defined for Indalia pygmaeola (BOLD: AAE8202). This species is related to the Indalia lutarella and Setema cereola pair and, despite the poor BI support for this grouping (PP=0.55) the COI distances among these three species are quite low (TN-dist between 3.03% and 5.05%).

Biology. Univoltine, flying in a single generation throughout July and August to mid-September. In open and sunny biotopes, favouring a calcareous substratum, at elevations between 550 and 1800 m. Adults fly by day and at dusk, and are attracted to bait and artificial light. The larvae feed on lichens, in captivity accepting dry leaves and lettuce (Ylla et al. 2010) and also artificial diet.

Distribution (Fig. 110). Mediterranean-Asiatic. The area of distribution extends from North Africa and central/ southern Europe to Armenia, Russia, Mongolia, Asia Minor and Iran. There are stable colonies in an extensive area throughout the northern third of the Iberian Peninsula including Portugal, and is also present in mountainous areas of the south and east. In the Balearic Islands there is a record of Indalia pymaeola attributed to the grey form (Petra locality in Mallorca), which could well be a misidentification for Indalia marcida (Pérez De-Gregorio & Vallhonrat, 1995).