Coliadinae

Braby, Michael F., 2005, Provisional checklist of genera of the Pieridae (Lepidoptera: Papilionoidea), Zootaxa 832, pp. 1-16 : 7-9

Coliadinae

The pantropical genus Eurema Hübner has been extensively revised in the Old World ( Yata 1989), and only two subgenera are now recognised: Eurema and Terias Swainson. Klots (1933) treated Nirmula Moore (with type species venata Moore) and Maiva Grose­ Smith & Kirby (with type species brigitta Stoll ) as subgenera of Eurema , but Yata (1989) considered them to be synonyms of Eurema (Eurema) based on similarities of the male and female genitalia. Field (1950) revised Teriocolias Röber from the New World and treated it as a monobasic subgenus (with type species zelia Lucas ) of Eurema . Lamas (2005), however, provisionally treated Teriocolias , together with the two other New World subgenera of Eurema , Abaeis Hübner and Pyrisitia Butler , as distinct genera pending further investigation of their monophyly. Lamas’ classification is tentatively followed here, although it should be noted that a recent cladistic analysis of molecular characters provided strong evidence that Eurema is paraphyletic, with the Eurema clade (i.e. Eurema (Terias) + Eurema (Eurema) ) including the genus Leucidia Doubleday , as well as Terioco ­ lias and Pyrisitia (Braby et al., unpublished data). Although Abaeis was not included in that study, it is clear that the status of other taxa currently or formerly included in Eurema will need to be revised. Two possible solutions are either to raise the Old World subgenus Terias to generic status, or to treat Pyrisitia , Teriocolias and Leucidia all as subgenera of Eurema . Historically, Klots (1933: 184) regarded Pyrisitia and Teriocolias as subordinate taxa of Eurema , but stated that “the relationships of Leucidia are rather hard to trace. It may be an offshoot from the ancestral stem of Eurema , to some species of which the genitalia are very similar.” implying that Leucidia may be the sister taxon to Eurema . The analysis of Braby et al. (unpublished data) shows that Leucidia belongs within the Eurema group, being most closely related to New World species of Eurema (Eurema) .

Klots (1933) proposed three subgenera within the predominantly Neotropical genus Phoebis Hübner ( Phoebis , Rhabdodryas Godman & Salvin , Aphrissa Butler ) but treated Prestonia Schaus as a synonym of it. Lamas (2005) did not recognise any subgenera in his comprehensive checklist of the Neotropical Pieridae , treating Rhabdodryas and Aphrissa as well as Prestonia as distinct genera, although emphasised that his taxonomic elevations were partly a matter of convenience until more phylogenetic information becomes available. The status of these four taxa will almost certainly be revised in future. Klots (1933) noted that ‘his’ subgenera were somewhat heterogenous and that Phoebis showed greater intraspecific differences in genitalia and other characters compared to differences observed between other genera. A recent molecular cladistic analysis (Braby et al., unpublished data) showed that Phoebis and Aphrissa comprise sister taxa in a well­supported monophyletic group, although that study did not include Rhabdodryas or Prestonia in their analysis.

Klots (1933) placed three species of Colias Fabricius in the subgenus Zerene Hübner (with type species caesonia Stoll). The male genitalia of Zerene , illustrated by Klots (1933), differ fundamentally from Colias , and several authors (e.g. Berger 1986; Cheong 1990; Ferris 1993; Pollock et al. 1998) have advocated full generic status. The molecular phylogenetic analysis of 10 Holarctic species by Pollock et al. (1998) revealed that Colias and Zerene were sister taxa and formed a well­supported monophyletic group. Petersen (1963), Berger (1986) and Verhulst (2000) revised the higher­level classification of Colias sensu stricto. Berger (1986) divided the genus into eight subgenera, five of which were newly described. The subgenera are distinguished by few morphological characters, each showing rather minor differences, and the validity of some of these taxa has been seriously questioned (Yamauchi et al., unpublished data; G. Lamas, pers. comm. 2004). For instance, Shapiro (1993), Lamas (2005) and Yamauchi et al. (unpublished data) did not recognise subgenus Protocolias Petersen, and Pollock et al. (1998) concluded that Neocolias Berger would be better subsumed under Eriocolias E. Y. Watson. The subgenus Eriocolias, however, was not recognised by either Klots (1933) or Lamas (2005). Six of the subgenera, besides Colias (Colias) , are listed tentatively until further study.

Hemming (1964) introduced Klotsius Hemming as a subgenus of Anteos Hübner (with type species menippe Hübner ), but this was not accepted by Lamas (2005) on nomenclatural grounds. Yamauchi and Yata (2000) revised the systematics of Gandaca Moore , although did not propose changes at the generic or subgeneric level.