Morphology-based phylogenetic analysis of the treehopper tribe Smiliini (Hemiptera: Membracidae: Smiliinae), with reinstatement of the tribe Telamonini

Abstract

Members of the Smiliini, the nominotypical tribe of the large New World subfamily Smiliinae, are predominately Nearctic in distribution. This tribe included 169 mostly tree-feeding species in 23 genera. A parsimony-based phylogenetic analysis of an original dataset comprising 89 traditional and newly discovered morphological characters for 69 species, including representatives of 22 of the 23 described genera of Smiliini and five other previously recognized tribes of the subfamily, resulted in a single most parsimonious tree with three major clades. The broad recent concept of Smiliini (including Telamonini as a junior synonym) was not recovered as monophyletic by the analysis. Instead, the analysis supported narrower definitions of both Telamonini, here reinstated from synonymy, and Smiliini. A key and diagnoses are given to define these tribes, along with discussions of their phylogeny, biogeography, and host plant associations. The genera Antianthe Fowler, Hemicardiacus Plummer, Smilirhexia McKamey, and Tropidarnis Fowler are placed as Smiliinae, incertae sedis. Based on the phylogeny, several genera from both tribes including Atymna Stål, Cyrtolobus Goding, Heliria Stål, and Telamona Fitch are not monophyletic. Diagnostic characters emphasizing the morphological differences between the Smiliini and Telamonini include the dorsal margin of the head, the shape of the pronotum, the size of the pronotal humeral angles, the presence or absence of pronotal longitudinal rugae, the size of forewing cells, variations in the fusion of veins R and M apically in both the foreand hind wing, and the shape of the apex of the female second valvulae. Mapping geographic distribution onto the phylogeny suggests that the common ancestor of the ingroup (all three clades) occurred in Central America and Mexico, with multiple dispersals to temperate North America. Many Smiliini and Telamonini feed on various species of oak (Quercus) and the close evolutionary association between these insects and their hosts is discussed.

Vanduzea segmentata (Fowler) (Hemiptera: Membracidae): Seasonality and Habits in the Southeastern United States, with Review of Its U.S. Distribution and Host Plants

“…In Costa Rica, V. segmentata has been taken on sunflower (Helianthus) (Ballou 1936), an asteraceous genus that served as a host in Bulloch County, Georgia. Composites and legumes also are used by other amastrine treehoppers (Wallace 2011). The large numbers of V. segmentata on leguminous crops in Hawaii, such as alfalfa (Medicago sativa L.) and soybean (Glycine max (L.) Merr.…”

“…Goding is a small genus of the New World membracid subfamily Smiliinae, tribe Amastrini (Metcalf and Wade 1965, McKamey 1998, Wallace 2011. Five species are found in the Nearctic Region (Deitz and Wallace 2012).…”

The identity of three South American “smiliine” treehoppers (Hemiptera, Membracidae) and related taxonomic changes, including description of a new genus in Thuridini

“…Four species of the related tribe Smiliini (Smiliinae) ( sensu Wallace 2011) were also described from South America: Ophiderma fascipennis Funkhouser from Bolivia, Atymna pilosa Funkhouser from Peru, Cyrtolobus (Atymna) atromarginata Goding from Ecuador, and Thelia planeflava Fairmaire (Telamonini but most recently placed in the genus Ophiderma of Smiliini) from Brazil. Smiliines are principally Nearctic, with numerous species occurring in the mountains of Central America, and throughout their range usually feed on oaks ( Quercus sp.).…”

A striking new treehopper genus Mutilifolia (Hemiptera: Membracidae: Smiliinae: Telamonini), from Costa Rica

“…Since 2004, numerous articles have been published improving the taxonomy of the second largest treehopper subfamily, the Smiliinae. Most of the recent taxonomic work has focused on the tribes Amastrini (Evangelista and Sakakibara 2007b), Ceresini (Andrade 2004a, b;2005;2008), Smiliini (Wallace 2011), and Telamonini (Wallace 2011). Only two Smiliinae genera were described during this period, Aurimastris Evangelista and Sakakibara (2007a) Telamonines are known for their large size (8-10 mm) and striking pronotal projections, sometimes assuming a quadrate, sinuate, tonguelike, or thornlike shape (Ball 1931;Wallace 2011).…”

“…), and they are distinguished from other Smiliinae by these additional diagnostic features: head dorsal margin with abrupt elevation mesad of eyes; pronotal longitudinal rugae distinct (in most); humeral angles enlarged; forewing with vein R 4+5 confluent with vein M 1+2 distad of or at beginning of M fork; hind wing with veins R 4+5 and M 1+2 free; dorsal and ventral margins of second valvulae apex converging proximally, apex acuminate; and male style clasp angular with a large, recurved tooth. Telamonines feed on various trees and some vines, notably many species from the genus Quercus (oak) (Wallace 2011(Wallace , 2014).…”