Atlides

Martins, Ananda Regina P., Duarte, Marcelo & Robbins, Robert K., 2019, Phylogenetic Classification of the Atlides Section of the Eumaeini (Lepidoptera, Lycaenidae), Zootaxa 4563 (1), pp. 119-134 : 121-122

The Atlides Section

Six synapomorphies characterize the Atlides Section in the phylogenetic results ( Table 1, Fig. 8), including a cleft anal angle of the hindwing ( Martins et al. 2018). This trait was used historically to characterize the group ( Godman & Salvin 1887 –1901; Robbins 2004a). It occurs in all species of the Atlides Section (illustrated for each genus, Figs. 9–15 ). The cleft anal angle occurs at vein 2A, which usually terminates on the distal side of the indentation. Indentations are unreported in other Eumaeini , except for two species of Panthiades ( Godman & Salvin 1887 – 1901). The depth of the cleft anal angle was treated as three states in the phylogenetic analysis ( Martins et a. 2018). As noted by Godman & Salvin (1887 –1901), a deep indentation occurs in Arcas ( Fig. 9 ), a shallow indentation occurs in Brangas ( Figs. 10 ), and an “intermediate” indentation occurs in all other members of the Atlides Section ( Figs. 11–15 ).

Atlides Section

Male, frons, color: muted green

Male, paraocular area, color: muted green

Female, hindwing, ventral surface, submarginal region, red band: absent

Hindwing, outer margin split at intersection with vein 2A: present

Male, hindwing, number of tails: two tails (M3-Cu1 and Cu1-Cu2)

Valvae, posterior margin, lateral view: rounded

Theritas

Forewing, dorsal surface, scent patch distal of the discal cell between veins M1 and M3, not intermixed with regular wing

scales: present

Male, forewing, ventral surface, discal cell band: present

Male, forewing, origin of vein R2: discal cell apex

Female, forewing and hindwing, dorsal surface, ground color: brown

Male, hindwing, ventral surface, postmedian band in cell Sc+R1-Rs: basally displaced

Arcas

Male, frons, color: emerald green

Male, paraocular area, color: emerald green

Male, forewing, ventral surface, postmedian band: absent

Female, forewing, ventral surface, postmedian band: absent

Male, hindwing, ventral and dorsal surface, cell Cu1-Cu2 submarginal macule: present

Hindwing, anal region, size of split at vein 2A: reaching postmedian region

Male, hindwing, shape of anal lobe structure: triangular

Saccus, length (cm): Long (0.30 - 0.40)

Ductus bursae, shape: in S shape

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Brangas

Male, forewing, costal margin: straight

Male and female, forewing ventral surface at the base, red scaling between vein Sc and the discal cell: present

Male and female, hindwing ventral surface at the base, red scaling between vein Sc+R1 and the discal cell: present

Male, hindwing, ventral surface, cell 2A-3A with a small pencil of hairs: present

Hindwing, anal region, size of split at vein 2A: not reaching submarginal region

Valvae, posterior margin, lateral view: finger-like

Saccus, length (cm): short (0.05 0.07)

Vinculum, dorsal projection, location and shape: lies under and on the inner edge of brush organs

Atlides

Forewing, dorsal surface, scent patch distal of the discal cell between veins M1 and M3, not intermixed with regular wing scales: present

Forewing, dorsal surface, scent pad: present

Male, hindwing, ventral surface, veins covered by black scales: all veins

Male, hindwing, postmedian band: absent

Male, abdomen, ventral surface, ground color: orange

Denivia

Ductus bursae, posterior region, membranous area: present

We do not propose a subtribe name for the Atlides Section. It is one of numerous unresolved clades with long branches (Quental, unpublished dissertation). Genomic sequencing currently in progress may resolve eumaeine subtribe classification, but it is currently premature to propose a more extensive higher classification. For now, we adhere to Eliot’s (1973) classification based on sections.

Robbins (2004a, b) excluded Brangas from the Atlides Section based on morphological differences of the genitalia and male secondary sexual traits. As noted, Brangas possesses a hindwing anal cleft, and the phylogenetic results strongly support the inclusion of Brangas ( B. caranus and B. getus ) in the Atlides Section as the sister of Atlides . The morphological differences that distinguish Brangas from the other Atlides Section genera appear to be autapomorphies.

The phylogeny based on DNA sequences in Quental (unpublished dissertation) was rooted on the lineage leading to Pseudolycaena . If one roots the Quental phylogeny where the tree of Martins et al. (2018) was rooted, the cladograms are the same, albeit with significantly fewer species in Quental. This congruence lends support to the phylogenetic results of Martins et al. (2018). Despite the different placement of the root, which may be due to the different ingroup sampling densities, the genera that we recognize are monophyletic in all analyses.

We here treat the Atlides Section as consisting of seven genera containing 71 described species.

Figures

FIGURES 9–15. Hindwing anal lobe cleft (black arrows). 9. Arcas imperialis, 10. Brangas getus, 11. Atlides cosa (with a forewing double scent pad in the discal cell—red arrow), 12. Theritas drucei, 13. Lucilda danaus, 14. Pseudolycaena damo, 15. Denivia silma.