Athis, HUBNER, [1819]

Moraes, Simeão De Souza & Duarte, Marcelo, 2014, Phylogeny of Neotropical Castniinae (Lepidoptera: Cossoidea: Castniidae): testing the hypothesis of the mimics as a monophyletic group and implications for the arrangement of the genera, Zoological Journal of the Linnean Society 170 (2), pp. 362-399 : 390

ATHIS HÜBNER, [1819] ( FIGS 20–22)

Type species

Papilio palatinus Cramer, 1777

Included species

Athis palatinus (Cramer, 1777) , Athis inca ( Walker, 1854) , Athis rutila (R. Felder, 1874) , Athis clitarcha ( Westwood, 1877) , Athis amalthaea ( Druce, 1890) , Athis ahala ( Druce, 1896) , Athis hechtiae ( Dyar, 1910) , Athis delecta ( Schaus, 1911) , Athis bogota ( Strand, 1912) , Athis thysanete ( Dyar, 1912) , Athis fuscorubra ( Houlbert, 1917) , Athis flavimaculata ( Miller, 1972) , Athis axaqua González & Fernández, 1992 , Athis analibiae ( Espinoza & González, 2005) (Imara) comb. nov., Athis pinchoni Pierre & Pierre-Baltus 2003 , and Athis pirrelloi Vinciguerra, 2011 .

Athis superba ( Strand, 1912) incertae sedis.

Comments

This is the most speciose genus in the family. The monophyly of the genus is highly questionable, making this a priority group for future revisionary studies. Although Athis appeared as a monophyletic group (in part) in one of the analyses ( Fig. 38 ), this condition was corroborated only by homoplastic characters (70: 1, 72: 0).

In describing Imara analibiae ( Fig. 22), Espinoza & González (2005) suggested that this species might be more closely related to Athis than to the nominative genus. Indeed, the species included in Athis share with Imara several morphological traits, especially those of the male genitalia, including the shape and length of the uncus, gnathos, and aedeagus, and some elements of the wing pattern.

Jorge González (pers. comm.) suggested that Athis may represent at least four lineages: (i) a Central American group, including A. inca , A. clitarcha , and A. flavimaculata ; (ii) a Caribbean group, including A. pinchoni , A. palatinus , A. axaqua , and A. fuscorubra ; (iii) a south-east American group, including A. therapon (here transferred to the genus Imara ) and A. amalthaea ; and (iv) an Amazonian group, including A. ahala , A. rutila , A. hechtiae , A. bogota , and A. thysanete . Although this hypothesis could not be tested because of a lack of material, it seems reasonable that Athis can be divided into more than one group. Indeed, the Caribbean group, represented in this analysis by A. inca and A. flavimaculata , is supported by the rounded sacculus (48: 4) and triangular vinculum (60: 0).

For A. superba , we sampled only females. The general morphology of the female genitalia, such as the extremely long ductus bursae, shape of the lamella antevaginalis and postvaginalis, and the eighth tergum, is quite different from other Athis species. In our analysis, A. superba was more closely related to Yagra than to the nominative genus. Primarily because we did not include males of this species in our analysis, we prefer to designate Athis superba as incertae sedis and retain this species in the nominative genus.

The recently described A. pirrelloi somewhat resembles A. palatinus ( Fig. 21); however, no morphological attributes other than colour were provided in the original description, an oversight that has been continually repeated for members of the Castniini .

Figures

Figure 38. Cladogram obtained from analysis using weighting (k = 20), not including Tascina metallica Pagenstecher, 1890. Filled circles represent synapomorphies; open circles represent homoplasies. Asterisks next to taxa names represent species of uncertain position.