Antianthe

Wallace, Matthew S., 2011, Morphology-based phylogenetic analysis of the treehopper tribe Smiliini (Hemiptera: Membracidae: Smiliinae), with reinstatement of the tribe Telamonini, Zootaxa 3047, pp. 1-42 : 25-32

Antianthe , Hemicardiacus , and Tropidarnis, Smiliinae , incertae sedis

The tribal status of Antianthe , Hemicardiacus , and Tropidarnis is uncertain based on the variable placement of Antianthe and Tropidarnis in the phylogenies of Cast (2002) and this work, and are herein placed Smiliinae, incertae sedis. In Cast’s (2002) molecular phylogeny of the Smiliinae, Antianthe is nested within a clade of Polyglyptini genera (resulting in a paraphyletic Smiliini s.l.), a group sister to the Smiliini s.l. in his analysis, and Tropidarnis is nested in the Telamonini between the basal Telamonanthe and the remaining telamonine genera. Here ( Fig. 1 ), Antianthe and Hemicardiacus form the basal monophyletic group of clade 3, sister to Tropidarnis + Telamonini, while Tropidarnis is the sister group to the Telamonini s. Wallace. These genera were not placed within Telamonini s. Wallace here because their inclusion would have resulted in a weaker concept of the tribe, defined by homoplastic synapomorphies. Nevertheless, these taxa share certain features with the Telamonini, such as an abrupt rise on the dorsal margin of the head near the eyes, enlarged humeral angles, and the forewing veins crowded anteriorly. Despite their doubtful tribal placement, in this analysis, both Antianthe and Tropidarnis form lineages with distinctive distributions, and morphological and biological traits. The only specimen examined of Hemicardiacus was the undissected male holotype, which resulted in missing data for more than 30 characters. The collection of additional specimens of this genus should help determine its tribal placement and relation to Antianthe . Future works with increased taxon sampling should help elucidate the relationships of these genera to other Smiliinae.

Antianthe and Hemicardiacus ( Antianthe group) are distinguished from the Smiliini s. Wallace, Telamonini s. Wallace, and Tropidarnis by the following features: frontoclypeal inferior margin without a ventral projection ( Fig. 12 B, 1); pronotum highly elevated and foliaceous ( Fig. 12 A); and pronotal humeral angles enlarged, long, and often narrowed with the anterior margin contacting the lateral margin of eye ( Fig. 12 A, 1). The placement of Antianthe near various Polyglyptini genera in Cast’s (2002) phylogeny of the Smiliinae deserves further scrutiny considering the differences between these taxa in the characteristics of the female second valvulae (the second valvulae of Antianthe is very similar to the second valvulae of genera in Smiliini s. Wallace and Telamonini s. Wallace), the absence of pronotal longitudinal rugae in Antianthe (present in most Polyglyptini), and the enlarged humeral angles in Antianthe (absent in Polyglyptini), among other features.

Both Antianthe and Hemicardiacus have a primarily Neotropical distribution, apparently localized in high altitudes ( Table 2 ) ( Plummer 1945; McKamey 1998; Cast 2002). Antianthe is the only genus, when compared with the Smiliini s. Wallace and Telamonini s. Wallace, known from the plant families Lauraceae , Rubiaceae , and Solanaceae ( Ballou 1935) , while Hemicardiacus is reported from oak ( Quercus ) ( Plummer 1945). Further, Antianthe is the only taxon from these groups to show subsocial (maternal egg guarding, ant-attended nymphal aggregations) and multivoltine behaviors (Wood 1984; Cast 2002). Most Smiliini and Telamonini are solitary as adults and nymphs, univoltine, and monophagous (Wood 1984; Cast 2002), but Atymna castaneae , Cyrtolobus vau , Ophiderma pubescens , Telamona ampelopsidis , T. unicolor , and Thelia bimaculata are listed as being attended by ants ( Funkhouser 1917).

1 1 1 1 1 1 1 1 1 1 2 continued next page 1 1 1 1 1 1 1 1 1 1 2 Ceresa vitulus 2 1 2 1 1 1 1 1 1 2? 1 2 1 1 2 1??? Smiliini sp. 1 2 1 1 1 1 1 2 1 1 2 1 1 1 2 2 2 1??? Smiliini sp. 2 2 1 2 1 1 1 1 2 2 2 2 1 2 2 1 2 1??? Spissistilus festinus 2 1 2 1 1 1 1 1 1 2? 1 2 1 1 2 1??? Telamonini sp. 2 2 1 2 2 1 1 1 1 2 2 1 2 2 1 2 2 2 1 1 continued

2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 3 3 4

1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 Antianthe expansa ??????? 2 1 1? 2 3 1 1&2 1 2 2 2 1 Antianthe foliacea ??????? 2 1 1? 2 3 1 1 1 2 2 2 1 Archasia auriculata ??????? 2 1 1? 2 1 2 2 1 2 1 2 1 Archasia belfragei ??????? 2 1 1? 2 1 2 2 1 2 1 2 1 Archasia pallida ??????? 2 1 2 1 2 1 2 2 1 2 1 2 1 Ashmeadea carinata ??????? 2 1 1? 1? 1 2 1 2 2 2 1 Atymna castaneae ??????? 2 1 1? 1? 1 2 1 2 2 2 1 Atymna distincta ??????? 2 1 1? 1? 1 1 1 2 2 2 1 Atymna helena ??????? 3 1 1? 1? 1 1&2 1 2 2 2 1 Bryantopsis ensigera 1 2??? 1 1 1 1 2 2 1? 1 2 2 2 1 1 1 Carynota mera ??????? 2 1 2 1 2 1 2 2 1 2 1 2 1 Cyrtolobus arcuatus ??????? 2 1 1? 1? 1 2 1 2 2 2 1

continued next page Xantholobus inflatus ??????? 2&3 2 1? 1? 2 2 1 2 2 2 1 Xantholobus muticus ??????? 3 1 1? 1? 1 2 1 2 2 2 1 Xantholobus tumidus ??????? 3 1 1? 1? 1 2 1 2 2 2 1

Ceresa vitulus ??????? 3 1 1? 1? 1 1 1 1 2 2 2 continued next page 2 2 2 2 2 2 2 2 2 3 3 3 3 3 3 3 3 3 3 4

1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 Smiliini sp. 1??????? 2 1 1? 2 2 1 2 1 2 2 2 1 Smiliini sp. 2??????? 3 1 1? 1? 1 2 1 2 2 2 1 Spissistilus festinus ??????? 3 1 1? 1? 1 1 1 1 2 2 2 Telamonini sp. 2? 2 1 2 2 1 1 1 2 1 2 2 1 2 1 2 1 2 1 continued.

4 4 4 4 4 4 4 4 4 5 5 5 5 5 5 5 5 5 5 6

1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 9 0 Antianthe expansa 3 1 2 2 1 1? 2 1 2 2 1 2 2 2 2 1 2 1 2 Antianthe foliacea 3 1 2 2 1 1? 2 1 2 2 1 2 2 2 2 1 2 1 2 Archasia auriculata 2 1 1 2 1 2 2 2 2 2 2 1 2 2 2 2 1 2 1 2 Archasia belfragei 2 1 1 2 1 2 2 2 2 2 2 1 2 2 2 2 1 2 1 2 Archasia pallida 2 1 1? 1 2 2 2 2 2 2 1 2 2 2 2 1 2 1 1 Ashmeadea carinata 3 1 2 2 1 1? 2 1 2 2 1 1 2 2 1 1 2 1 1 Atymna castaneae 3 1 2 2 1 2 1 2 1 2 2 1 1 2 2 2 1 2 1 1 Atymna distincta 3 1 2? 1 2 1 2 1 2 2 1 1 2 2 2 1 2 1 1 Atymna helena 3 1 2 2 1 2 1 2 1 2 2 1 1 2 2 2 1 2 1 1 Bryantopsis ensigera 3? 2? 1 1? 2 1 2 2 1 1 2 2 1? 1 2 1 Carynota mera 2 1 1 2 1 2 2 2 2 2 2 1 2 2 2 2 1 2 1 2 Cyrtolobus arcuatus 3 1 2? 1 2 1 2 1 2 2 1 1 2 2 2 1 2 1 1 Cyrtolobus dixianus 3 1 2? 1 2 1 2 1 2 2 1 1 2 2 2 1 2 1 1 Cyrtolobus fenestratus 3 1 2 2 1 2 1 2 1 2 2 1 1 2 2 2 1 2 1 1 Cyrtolobus inermis 3 1 2 2 1 2 1 2 1 2 2 1 1 2 2 2 1 2 1 1 Cyrtolobus tuberosus 3 1 2 2 1 2 1 2 1 2 2 1 1 2 2 2 1 2 1 1 Cyrtolobus vanduzii 3 2 2? 1 2 1 2 1 1 2 1 1 2 2 2 1 2 1 1 Cyrtolobus vau 3 1 2 2 1 2 1 2 1 2 2 1 1 2 2 2 1 2 1 1 Glossonotus acuminatus 2 1 1 2 1 2 2 2 2 2 2 1 2 2 2 2 1 2 1 2 Glossonotus univittatus 2 1 1 2 1 2 2 2 2 2 2 1 2 2 2 2 1 2 1 2 Godingia guerreroensis 3 1 2? 1 2 1 2 1 2 2 2 1 2 2 2 1 2 1? Grandolobus grandis 3 1 2 2 1 2 1 2 1 2 2 1 1 2 2 2 1 2 1 1 Heliria cornutula 2 1 1 2 1 2 2 2 2 2 2 1 2 2 2 2 2 2 1 2 Heliria cristata 2 1 1 1&2 1 2 2 2 2 2 2 1 2 2 2 2 2 2 1 2 Heliria mexicana 2 1 1? 2 1? 2 1 2 2 1 1 2 2 2? 2 1 2 Heliria praealta 2 1 1 2 1 2 2 2 2 2 2 1 2 2 2 2 1 2 1 2 Heliria scalaris 2 1 1 2 1 2 2 2 2 2 2 1 2 2 2 2 2 2 1 2 Heliria sinuata 2 1 1? 1 2 2 2 1 2 2 1 2 2 2 2 1 2 1 2 Helonica excelsa 2 1 1 2 1 2 2 2 2 2 2 1 2 2 2 2 1 2 1 1 Hemicardiacus saundersi 2 1 1 2 1 1? 2 1 2 2 1? 2 2 2???? Heranice miltoglypta 3 1 2 2 1 1? 2 1 2 2 1 1 2 2 2 1 1 2 1 Idioderma virescens 3 1 2 2 1 2 2 2 1 2 2 1 1 1 2 2 1 2 1? Ophiderma pubescens 3 1 2 2 1 2 1 2 1 2 2 2 1 2 2 2 1 2 1 2 Ophiderma salamandra 3 1 2 2 1 2 1 2 1 2 2 2 1 2 2 2 1 2 1 1 Palonica nogalana 2 1 1? 1 2 2 2 2 2 2 1 2 2 2 2 1 2 1 1

continued next page Tropidarnis tectigera 3 1 1 2 1 1? 2 1 2 2 1 1&2 1 2 2 1 2 1 1 Xantholobus inflatus 2&3 1 2 2 1 1? 2 1 2 2 2 1 2 2 2 1 2 1 1 Xantholobus muticus 3 1 2 2 1 2 1 2 1 2 2 2 1 2 2 2 1 2 1 1 Xantholobus tumidus 3 1 2 2 1 2 1 2 1 2 2 2 1 2 2 2 1 2 1 1

continued.

6 6 6 6 6 6 6 6 6 7 7 7 7 7 7 7 7 7 continued next page continued next page 6 6 6 6 6 6 6 6 6 7 7 7 7 7 7 7 7 7

1 2 3 4 5 6 7 8 9 0 1 2 3 4 5 6 7 8 Xantholobus inflatus 1 2 2 2 2 1 1 2 1 2 3 1 1 1 2 1 3 1 Xantholobus muticus 2 1 2 2 2 2 1 2 1 2 3 1 1 1 1 1 2 1 Xantholobus tumidus 2 1 1 2 2 1 1 2 1 2 3 1 1 1 2 1 2 1

Ceresa vitulus 1 1 2 1?????? 3 1 2 1 1 1 1 2 Smiliini sp. 1?????????? 3 1 1 1 2 1 2 1 Smiliini sp. 2?????????? 3 1 1 1 2 1 2 1 Spissistilus festinus 1 1 2 2 1 1 1? 1 1 3 1 1 2 1 1 1 2 Telamonini sp. 1 2 1 2 2 1 1 1 2 1 3 1 1 1 1 1 4 1 continued.

continued next page The genus Tropidarnis is characterized by a simple, single cucullate row of setae in row I of the metathoracic tibia ( Fig. 12 D). In all Smiliini s. Wallace, Telamonini s. Wallace, and Antianthe + Hemicardiacus specimens examined in the analysis, this row was irregular or double. Other features that distinguish it from genera in Telamonini s. Wallace include the absence of a median anterior pronotal horn or projection and pronotal longitudinal rugae; vein R4+ 5 in the forewing confluent with vein M basad of M fork; and a rounded, vertically oriented (from a posterior aspect; Fig. 14 B, 1) style clasp with a small recurved tooth. Tropidarnis is found in the Neotropical and Nearctic regions (southwest United States), and both the nymphs and adults have been collected from oak ( Quercus ) ( Table 2 ).

Figures

FIGURE 1. Phylogenetic relationships generated from cladistic analysis of the entire dataset, illustrating the new proposed reclassification of the Smiliini s. l. Analysis in PAUP * resulted in a single most parsimonious tree with 311 steps. Bremer support and synapomorphies for nodes are listed in Table 6.

FIGURE 12. Antianthe group and Tropidarnis I: A, Hemicardiacus saundersi Plummer, head and pronotum from anterior aspect, holotype, # 71 - 363 d 3: 1, pronotal humeral angles long, narrowed, and with anterior margin contacting lateral margin of eye. B, Antianthe expansa (Germar), head from anterior aspect, # 11 - 005 aƤ: 1, frontoclypeus ventral margin appearing carinate, without ventral projection; 2, pronotal lateral supraocular callosities complete. C, Tropidarnis tectigera Fowler, head from anterior aspect, # 72 - 10 fƤ: 1, frontoclypeus ventral margin with rounded projection. D, T. tectigera, row 1 of right metathoracic leg, # 72 - 10 fƤ.

FIGURE 14. Antianthe group and Tropidarnis II: A, Antianthe expansa (Germar), male style from posterior aspect, # 09 - 183 a 3: 1, style clasp from posterior aspect angled laterally. B, Tropidarnis tectigera Fowler, male style from posterior aspect, # 72 - 10 g 3: 1, style clasp from posterior aspect vertical. C, A. expansa, aedeagus from lateral aspect, # 09 - 183 a 3. D, Tropidarnis tectigera, aedeagus from lateral aspect, # 72 - 10 g 3. E, A. expansa, connective from dorsal aspect # 09 - 183 a 3. F, A. expansa, female second valvulae from lateral aspect, # 09 - 183 bƤ: 1, second valvulae with middle and apical large teeth distinctly closer; 2, second valvulae basal teeth. a, aedeagus. c, style clasp. cn, connective.