Pierinae

Pierinae: Pierini

The well­known genus Pieris Schrank has been extensively studied and revised (Mariani 1937; Bernardi 1947, 1958; Kudrna 1974; Robbins & Henson 1986). Klots (1933) recognised four subgenera (Pieris, Pontia Fabricius, Synchloe Hübner, Glennia Klots) and, although he narrowed the concept of Pieris substantially to include a much smaller representation of taxa, the genus has been split further and ‘his’ subgenera are now usually recognised as distinct genera on phylogenetic grounds (Mariani 1937; Bernardi 1947; Geiger & Scholl 1985; Robbins & Henson 1986). Klots (1933), for instance, erected Glennia as a monotypic subgenus of Pieris to accommodate the species pylotis Godart from South America. Robbins and Henson (1986), however, advocated full generic status for this taxon on the basis of significant differences in the female genitalia. Bernardi (1947) revised the systematics of Pontia, and Geiger and Scholl (1985) provided biochemical evidence for the separation of Pontia from Pieris, but recommended that the concept of Synchloe be restricted to include a smaller number of species and treated as a subgenus of Pontia. That is, they divided Pontia into two species­groups or subgenera: the callidice group (Synchloe) and the daplidice group (Pontia). Bernardi (1958) described the genus Talbotia Bernardi to accommodate the single species naganum Moore from Asia previously included in Pieris. He considered Talbotia to be closely allied to Pieris, Ganyra and Belenois Hübner. Verity (1947) proposed Artogeia Verity as a subgenus of Pieris, which Kudrna (1974) and others subsequently elevated to generic rank. However, several recent studies (Geiger 1981; Geiger & Scholl 1985; Robbins & Henson 1986) have rejected this proposal on either biochemical or phylogenetic grounds. Robbins and Henson (1986) recommended that Artogeia be considered a synonym of Pieris.

Klots (1933, p 224) proposed the taxon Pieriballia Klots as subgenus of Itaballia Kaye, and included two species in it; he also considered that Pieriballia and Itaballia were closely related to Perrhybris Hübner. Lamas (2005), however, tentatively treated Pieriballia as a monobasic genus (with type species viardi Boisduval).

Klots (1933, p 224) treated Ganyra Billberg as a subgenus of Ascia Scopoli because the included species were more closely related to Ascia than anything else, but noted that “They also show many points of similarity to the African Belenois.” Bernardi (1958) and Robbins and Henson (1986) treated Ganyra as a distinct genus; the latter authors provided morphological evidence based on study of the female genitalia that Ganyra, together with Itaballia and Perrhybris, were probably the closest relatives of Pieris.

The Tatochila group of pierine genera from South America has been extensively revised and expanded, from three (Klots 1933) to eight (Ureta 1955; Field 1958; Herrera & Field 1959; Ackery 1975; Field & Herrera 1977; Shapiro 1991, 1994). The five new genera are Hypsochila Ureta, Theochila Field, Pierphulia Field, Infraphulia Field, and Reliquia Ackery. Two of the new taxa (Pierphulia, Infraphulia) were initially treated as subgenera of Piercolias Grote and Phulia Herrich­Schäffer, respectively (Field 1958), but Field and Herrera (1977) later raised them to full generic status. Chionanema Ureta was described as a subgenus of Hypsochila by Ureta (1955), but Field (1958) seriously questioned its validity and subsequently synonymised it (Field & Herrera 1977). The genus Tatochila, comprising 11 species (Lamas 2005), is possibly polyphyletic, comprising two or more genera (Shapiro 1991; pers. comm. 2004), however, a phylogenetic analysis of the entire Tatochila group is needed to verify this.