Hovanini Sanborn, Marshall & Moulds

Sanborn, Allen F., Marshall, David C., Moulds, Maxwell S., Puissant, Stéphane & Simon, Chris, 2020, Redefinition of the cicada tribe Hemidictyini Distant, 1905, status of the tribe Iruanini Boulard, 1993 rev. stat., and the establishment of Hovanini n. tribe and Sapantangini n. tribe (Hemiptera: Cicadidae), Zootaxa 4747 (1), pp. 133-155 : 149

Hovanini Sanborn, Marshall & Moulds , n. tribe

Type genus Hovana Distant 1905g: 279 .

Diagnosis

Head including eyes narrow, much less than lateral angles of pronotal collar; supra-antennal plates extending to eye; lateral ocelli widely spaced, closer to eyes than to each other; postclypeus apex angular in lateral view. Pronotum with median groove, pronotal collar with paranota weakly developed; mid lateral tooth absent. Opercula small, not covering the tympanal cavity, with convex lateral margin, not S-shaped, not surrounding meracanthus. Meracanthus rudimentary. Foreleg primary spine prostrate. Forewing costal vein equal in width to R+Sc; CuP and 1A fused in part. Hind wing anal lobe narrow with anal vein 3 straight; hind wing cubital cell 1 width at distal end not twice or more the width of cubital cell 2; hind wing RP and M veins fused at base. Male abdominal tergites with sides convex in cross section, tergites 2–4 similar in size, tergites 4–7 reducing in width posteriorly; sternite I hidden; epipleurites deeply reflexed to ventral surface. Timbal covers absent, partially formed ridge on dorsal and ventral posterior timbal cavity. Pygofer distal shoulder undeveloped; upper lobe well-developed; basal lobes small; dorsal beak present. Uncus elongated, not retractable within pygofer, curving downward, with laterally expanding apex; claspers absent. Aedeagus lacking a ventrobasal pocket ( Fig. 7 ).

Distinguishing features

Hovanini n. tribe differs from other tribes in having, in combination, a mesonotum with a triangular scutellum extending over the anterior abdominal tergites, opaque forewings, well-developed pygofer upper lobes, a dorsal beak, and a downward curving non-retractable uncus widening at its apex.

Hovanini n. tribe can be distinguished from the Lacetasini by the male operculum that fully encapsulates the meracanthus, the epipleurites reflexed to the ventral surface, the well-developed pygofer upper lobe, the presence of a dorsal beak, and an extended uncus widening at the apex. Hovanini n. tribe can be distinguished from the Malagasiini Moulds & Marshall, 2018 (in Marshall et al. 2018) by the comparatively wide pronotal collar, the widely separated costal and R+Sc veins in the forewings, the prostrate primary spine of the fore-femur, opercula that do not cover the tympanal cavity, the tapering posterior abdomen, the lack of any development in the pygofer distal shoulder, the presence of pygofer upper lobes, and an extended uncus widening at the apex. The Hovanini n. tribe differs from the Tettigomyiini in the forewings being significantly longer than the body, the lack of an inflated male abdomen, the deeply reflexed epipleurites, the lack of an extended distal shoulder to the pygofer, the well-developed pygofer upper lobes, the presence of a dorsal beak, and the extended uncus widening at the apex. Finally, the Hovanini n. tribe differs from the Ydiellini Boulard, 1973 in the presence of timbals, the lack of thickened forewing veins adjoining the apical and ulnar cells, the lack of large ulnar cells in the forewings, and the lack of a row of teeth on the hindwing costa.

Comments

Malagasiini appears to be the most closely related tribe based on the similarities in characters that distinguish both Malagasiini and Hovanini n. tribe from the other Tettigomyiinae tribes. Both tribes are found in Madagascar but differ in the diagnostic characters as outlined above.

The presence of the reticulated distal forewings in Hovana and Hemidictya alone does not warrant their classification in the same tribe. There are other cicada genera that possess reticulated, many-celled forewings that are not classified in the same tribes. For example, in the Cicadinae , Talainga Distant, 1890a and Paratalainga He, 1984 are in Gaeanini Distant, 1905c , whereas Angamiana Distant, 1890b and Polyneura Westwood, 1840 are in Polyneurini Amyot & Audinet-Serville, 1843 . Note that the genitalia of these two pairs of genera are much more similar to each other than the genitalia of Hovana and Hemidictya are to each other. The differences in the forewing morphology of Hovana and Hemidictya are distinctive and suggest that these genera evolved the leaf-like appearance of their forewings through convergence rather than from a common ancestor. Boulard (2000) recognised this when he described the leaf-like appearance of the forewings of Hemidictya , Hovana , Lacetas , and Cystosoma Westwood, 1842 as parallel evolution from independent ancestors rather than being derived from a similar ancestor as proposed by Ruschel & Campos (2019). Leaf mimicry is common in insects being found in a number of diverse clades and dates back at least to the Permian ( Garrouste et al. 2016). The structure of the genitalia alone suggests that Hovana should be classified in a different tribe and subfamily from Hemidictya , a view supported by the number of significant morphological differences outlined above.

Figures

FIGURE 7. Lateral view of pygofer (A) and dissected aedeagus (B) of Hovana distanti (MNHN). (bl) basal lobe; (db) dorsal beak; (un) uncus; (upl) upper pygofer lobe.