O. Merkl szerk.: Folia Entomologica Hungarica 68. (Budapest, 2007)
DEFINITION AND DIVERSITY OF DENIVIA The genus Denivia was erected by JOHNSON (1992) including the type species Thecla deniva HEWITSON, 1874 plus Papilio hemon CRAMER, 1775 and Denivia maggae JOHNSON, 1992. It was described as an "outgroup" of the diverse and primarily Andean "elfin butterflies" (= Micandra section sensu ROBBINS, 2004) based on characters in genital structures and wing pattern elements. The ventral hindwing androconia was not taken into consideration. The genus Denivia is best defined by the ventral hindwing androconia, which is a characteristic pocket situated along the cubital vein on the subbasal area of the wing membrane (GODMAN & SALVIN 1887: pi. 49, fig. 7a; BÁLINT 2006: fig. 5). The fold opens towards the anal margin of the wing and in the pocket there are specialized, presumably scent producing or disseminating scales of butter-yellow colour (BÁLINT, in prep.). This character is unique in the eumaeine lycaenids and we consider this character state as an apomorphy for the genus Denivia. Certain species, which were originally placed in Denivia, do not possess this character, or their fold is not a homology, and because of this homoplasy they were clustered together (CONSTANTINO et al. 1993, BÁLINT & JOHSNON 1997; BÁLINT et al. 2007). These taxa are in need of revision. Similarly, we doubt that Papilio Usus STOLL, 1790 and its close relatives belong to Denivia, because they lack this special ventral hindwing androconia. Probably this group of lycaenid butterflies without androconia is the basal sister group of Denivia. How this organ is working that is not yet clear for us. One hypothesis is that during copulation the female wings cover the male ones, and via rubbing the fold can be opened by the female; by this the scent can be disseminated from the pocket. This is supported by the observation that generally in Lycaenidae the females flap in the male wings during copulation. Another hypothesis that Denivia males are able to open this fold by themselves via a fan-like folding or wrinkling of the wing membrane. Species of Denivia can be arranged into three groups (see Table 1 ) : ( 1 ) the Mesoamerican-Andean-Atlantic theocritus species group, whose taxa possess a medial pattern in their ventral wing surfaces comprised by characteristic intercellular gleaming dots; (2) the Amazonian-Atlantic-Guyananphegeus species group, which is probably an isolate of the previous group as indicated by their similary formed hindwing tornai lobe and female genital terminalia with large posterior ductus bursae (with length one half to nearly equal to that of anterior ductus bursae), with asymmetric lamella postvaginal spines; and (3) the Panamerican hemon species group, whose males possess a large visual brand