Hespenedra Kramer, 1966

Jones, Joshua R. & Deitz, Lewis L., 2009, Phylogeny and systematics of the leafhopper subfamily Ledrinae (Hemiptera: Cicadellidae) 2186, Zootaxa 2186 (1), pp. 1-120 : 30-31

Genus Hespenedra Kramer

(Pl. 4H–I, 9C, 12A, G, 13C, K, 18G)

Hespenedra Kramer, 1966: 492 , figs. 62–69.

Type species. Thlasia chilensis Spinola 1852: 277 , by original designation.

Synonymy. None.

Description. Adapted from Kramer (1966): “Moderately large (8.75– 11 mm.) robust leafhoppers; head gross and well-produced beyond eyes, coronal suture and lateral margins carinate, ocelli on crown anterior to eyes, closer to midline than to lateral margins; in lateral view with crown flat, face oblique and nearly flat, area from eye to apex foliaceous, clypeus and clypellus clearly visible; clypellus base bearing paired sublateral carinae converging medially into a single longitudinal carina on apical extension in males, medially depressed longitudinally; in facial view the clypellus, lora, and clypeus slightly tumid, genae broad, lateral frontal sutures terminating at antennal bases, with a variably developed carina on midline from between antennae to apex of head, antennae moderately short, anterior and mesad of eyes, distant from lateral margins of head; pronotum short, widest posteriorly, anterior margin straight, posterior margin shallowly indented, carinate laterally, longitudinally depressed medially, females with a slight longitudinal carina in the depression extending anteriorly to the anterior margin; scutellum large, broader than long; forewings moderately long and broad, punctate-rugulose, with extra crossveinlets apically, appendix absent, claval area strongly pigmented with creamy white pigmentation in males; legs not particularly stout, macrosetae on metathoracic tibiae greatly reduced in number but clearly visible, not hidden by hairs; formula of macrosetae at apex of metathoracic femur 2–1–0. Male genitalia: genital capsule partly withdrawn into abdomen concealing base of valve, anal tube not large, connective modified Y-shape, style long with one apical lobe, aedeagus short and stout with paired ventral processes. Female genitalia: seventh sternum longer than preceding segment and with ovipositors not extending much beyond pygofer.”

Species. [1]: chilensis (Spinola) .

Range. Chile (Arauco Province: Contulme, Palo Botado; Concepcion Province: Hualpen; Isla Chiloe: Dalcahue; Llanquihue Province: Lago Chapo; Valdivia Province: Chesque).

Host plants. Unknown.

Material examined. H. chilensis : 3 males, 1 female, Chile, USNM, JRJ _Led1_060–062, 064 , 1 male, 1 female, Chile, AMNH, JRJ _Led1_063, 065 .

Remarks. Hespenedra shares many similarities with the Petalocephala genus group, particularly the shape of the face, the carination of the frontoclypeus, the shape of the body in lateral and dorsal views, and the straight anterior margin of the pronotum. Males of Hespenedra have the clavus punctate and pigmented very much like some species in Thlasia and a number of unidentified species of Ledrini from China. Females have longer crowns than males, similar to many ledrine genera. These similarities are not well understood, and may be merely convergent features, or retained plesiomorphies.

Despite these similarities, Hespenedra exhibits many character states that are unique among Ledrinae . For example, the plantar surface of metathoracic tarsomere I bears an elongate ovoid pad of short stout white setae not seen in any other group in this study. Because Hespenedra is currently monotypic, however, it is unclear whether or not this character is simply diagnostic for H. chilensis , or if it is also phylogenetically significant for the genus or tribe. Also, the connective is longer and more sclerotized than that of most other ledrines, and the aedeagus and styles are shorter and more robust. These differences taken in context of the topology presented herein seem to indicate its ancestral and removed position within the Ledrinae . The long and narrow median stem of the connective is somewhat like some members of the Petalocephala genus group, and the central area of the proximal end that is not sclerotized and appears hollowed, is similar to that of Proranus (Xerophloeini) , Afrorubria (Afrorubrini) , and Beniledra (Ledrini) . The aedeagus is also short in Thlasia and Hangklipia , and the styles of Hespenedra are apically flexed downward like most Ledrini .

The narrowed and elongate base on which the apical macrosetae of the metathoracic femur are mounted (Pl. 13C) strongly resembles that in Rubria ( Rubrini —Pl. 13E), which may be Hespenedra ’s closest extant relative. ( Figs. 1–3 )

The statement in Kramer’s description that the connective is a “modified Y-shape” may falsely imply a relationship with other leafhopper subfamilies defined by having the connective Y-shaped.

Figures

FIGURE 1. Phylogram of relationships generated from cladistic analysis of entire dataset: {= [Ledrinae sensu Oman et al. (1990) dataset] + [dataset of non-ledrine taxa included to provide resolution to ingroup taxa anticipated to ultimately place outside of Ledrinae (marked in italic bold with an asterisk*)]}. Analysis in PAUP* was rooted to Bathysmatophorus shabliovskii, and resulted in a single most parsimonious tree with a cost of 1719 steps. Bremer support values are given above the branch to which they correspond. Squares, circles and triangles indicate tribal placements of included genera in Oman et al. (1990). The boundary of Ledrinae, as it is here recognized, is marked with a black star.

FIGURE 2. Phylogram of relationships generated from cladistic analysis of pruned dataset, including only species herein recognized to belong to the subfamily Ledrinae. Analysis in PAUP* was again rooted to B. shabliovskii; Aphrodes was included as putative sister group to the Ledrinae. This second analysis resulted in a single most parsimonious tree with a cost of 1185 steps. Bremer support values are given above the branch to which they correspond; the black star marks the phylogenetic boundary of Ledrinae. Ingroup resolution is topologically identical to that in Fig. 1, but branch lengths and Bremer supports vary.

FIGURE 3. Phylogram of summarized relationships among Ledrinae genera generated from a reduced dataset including single exemplars of each available genus. If available, the type species was chosen (marked with an asterisk*). Additional exemplars were included in cases where polyphyly among species was demonstrated in previous analyses, and where a male and a female of different species in the genus were needed to provided all of the pertinent characters (i. e., genitalia). Analysis (in PAUP*, rooted to B. shabliovskii, with Aphrodes included as putative sister group to the Ledrinae) resulted in five equally parsimonious trees (cost 933). The selected topology differs slightly from Fig. 2 in the placement of E. planata as sister to T. antica, in Titiella and Hangklipia not being placed as sister taxa, and in Jukaruka being placed with Platyledra instead of Ledropsella. Numbers at internodes correspond to a list of apomorphies generated from this analysis (see Table 1). Higher taxa within Ledrinae recognized herein, including new tribes (with their distributions), and informal genus and species groups, are indicated by symbols. The inclusion of Dusuna within the Petalocephala genus group is at present uncertain, as indicated by the question marks above the open triangles.