Petalocephala Stål, 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 : 51-53

Genus Petalocephala Stål

(Pl. 3A–C, 10G, 11A, 13D, L, 16H, 19A–B)

Petalocephala Stål, 1854: 251 .

Type species. P. bohemani Stål, 1854: 251 , by original designation.

Synonymy. Pachyledra Schumacher 1912: 248 (type species P. kamerunensis Schumacher 1912: 249 ).

Description. Stål 1854 (as quoted by Distant 1908): “Body very oblong or a little elongate, depressed; head clypeated, foliaceously produced anteriorly; vertex somewhat flattened; face beneath eyes strongly and abruptly, thence gradually, narrowed, margins very slightly defined; front small, narrow, flattish; eyes small; ocelli situate towards base of vertex, farther removed from the eyes than from each other; pronotum transversely sexangular, not or only slightly anteriorly narrowed, the lateral margins acute, anterior lateral much longer than posterior lateral, anterior margin slightly rounded; scutellum triangular subequilateral; tegmina subcoriaceous, pellucid, densely punctate, tectiform, anteriorly conjointly convex, clavus very broad before the middle, corium obliquely rounded at apex, veins somewhat irregularly anastamosed towards apex; legs somewhat short, anterior coxa free, posterior tibiae remotely dentate.”

Species. [89]: adelungi (Melichar) ; alata Evans ; arcuata Cai & Kuoh ; armata Evans ; bainbriggei Distant ; aluchestanica Dlabola; bazarakana Dlabola ; bicolor Distant ; bipunctata Melichar ; bohemani Stål ; castanea Kato ; cephalotes Distant ; chlorocephala (Walker) ; chlorophana Kuoh ; confusa Distant ; conica (Walker) ; conspersa Kuoh ; conspicua Distant ; convexifrons Schumacher; cultellifera (Walker) ; declivis Walker ; duodiana Kuoh ; engelhardti Kusnezov ; enigmoides Evans ; eurglobata Cai & He ; fasciifrons Melichar; formosana (Matsumura) ; fuscomarginata Cai & Kuoh ; fusiformis (Walker) ; glauca (Melichar) ; gonzalezi Lindberg ; grandiosa Dlabola ; granulosa Distant ; hearsayi Distant ; horishana (Matsumura) ; horne i Distant; insignis Distant ; ixion Linnavuori ; kamerunensis (Schumacher) ; kempi Singh-Pruthi ; koshunensis Schumacher ; latifrons (Walker) ; limbata Evans ; manchurica Kato ; nigrella Evans ; nigrilinea (Walker) ; obtusa Kuoh ; ochracea Cai & Kuoh ; perakensis Distant ; perductalis (Kirby) ; philippina Stål ; pilka Evans ; planata Evans ; porrigens Walker ; potanini Melichar ; pullata Evans ; pulsata Evans ; punctatissima Stål ; quadrimaculata (Matsumura) ; raniceps Jacobi ; remota (Melichar) ; rubromarginata Kato ; rubromarginella Kuoh ; rufa Cen & Cai ; rufomarginata Kuoh ; sanguineomarginata Kuoh ; scutellaris Linnavuori ; signata Distant ; skoba Evans ; spicata Evans ; stellata Evans ; subacta Walker ; subaquila Distant ; tabulata Distant ; taihorensis Schumacher ; taikosana Kato ; tenuifrons (Walker) ; trispicula Evans ; turgida Linnavuori ; umbrosa Distant ; unicolor Cen & Cai ; uniformis Distant ; viridis Cai & He ; viridula Kuoh ; vittata (Matsumura) ; wahlbergi Stål ; walkeri (Melichar) .

Range. Angola; Benin (Parakou); Bhutan; Cameroon; Central African Republic (Banqui; Kapou; Kivu; La Maboke); China (Hainan: Ta Hian; Guizhou: Pingtang Co.; Hong Kong; Kiangsi; Kwangtung); Congo (Dimonika; Elisabethville; Faradje; Flandria; Gorge de la Pelenge; Gwanga; Lulua: Kapanga; Ubangl: Bosobolo); Côte d’Ivoire [ Ivory Coast] (Blekoun); Ghana; Guinea; India (Cherangode; Coimbatore; Kerala; Kodaikanal; Nedungadu; S. Malabar); Indonesia (Borneo: Kuching; Java: Batavia, Soekaboemi, Wijnkoopsbay; Sumatra: Mt. Simasopa, Prapat); Japan (Loochoo Island); Kenya; Liberia (Bomboma; Mt. Coffee; Suakoko); Mozambique (Delagoa Bay [Maputo Bay]); New Guinea (Wisselmeren); Nigeria; Philippines (Luzon: Butuan, Limay, Mt. Banahao, Mt. Makiling; Mindanao: Surigao; Negros: Dumaguete); Rwanda (Nduga); Singapore; Sudan; Taiwan (Heng Chun; Nantou Hsien: Howang, Jenai Chungyang, Meifeng, Mt. Po Jing Cha, Puli, Sungkang, Tungpu, Wushe; Pingtung Hsien: Kenting; Suisha; Taichung: Anmashan, Chingshan; Taitung: Yenping); Thailand (Chiang); Togo; Uganda (Ruwenzore); Vietnam (Hoa Binh); Yemen; Zimbabwe [ Rhodesia].

Host plants. Acoranthera schimperi v. deflersi (Apocynaceae) , Indigofera oblongifolia (Fabaceae) , Theobroma cacao (Malvaceae) .

Material examined. P. bohemani : 1 female, Indonesia (Java), BMNH, JRJ _ Led 1_116 , 1 female, New Guinea, ASCU, JRJ _ Led 1_121 , 1 female, New Guinea, BPBM, JRJ _ Led 1_291; P. conspicua : 1 male, Singapore, USNM, JRJ _ Led 1_187 , 1 male, (location not given), MNHN, JRJ _ Led 1_132 , 4 male, China, India, NCSU, JRJ _ Led 1_134–136, 138; P. raniceps : 1 female, Congo, OSUC, JRJ _ Led 1_130 , 1 male, 1 female, Central African Republic, MNHN, JRJ _ Led 1_126, 131, 1 (abdomen missing), Nigeria, AMNH, JRJ _ Led 1_128 , 3 males, 2 females, Côte d’Ivoire [ Ivory Coast], Liberia, USNM, JRJ _ Led 1_123–125, 127, 129 .

Remarks. Petalocephala is the largest genus (87 described species) in Ledrini and has a distribution almost as great as that of Ledra (except it is not found in Europe). Many of its species are indistinguishable except by dissection and examination of the male genitalia. It is probable that several of its described species are redundant. At present, the taxonomic limits of Petalocephala are not yet well understood (see discussion for Destinia Nast above).

In general body shape and with its quadrate (in cross section) non-foliaceous tibia, Petalocephala strongly resembles Hespenedra and Rubria . It is possible that its torpedo-like shape and reduced features represent an ancestral ground plan in Ledrinae , and that Petalocephala therefore represents a basal ledrine lineage. In some early phylogenetic analyses utilizing fewer characters and taxa (not shown), Petalocephala was, in fact, placed at the base of the Ledrini clade, closer to Hespenedra and Rubria . However, in the final analyses with all of the taxa and characters ( Figs. 1–3 ), it was placed in a position as one of the most derived taxa within the Petalocephala genus group. Other features of Petalocephala actually display similarities with more derived taxa—in characters of the male genitalia, for example, some Petalocephala species are very similar to Ledra . Based on all the evidence assembled here, it seems more likely that the reduced features and shape of Petalocephala , Hespenedra and Rubria represent an ecologically generalized condition on which many ledrine (and non-Ledrinae) taxa have converged.

Petalocephala , which lacks lateral extensions on the pronotum, was placed as sister group to Neotituria , which retains the pronotal extensions, in the phylogenetic analyses ( Figs. 1–3 ). It seems logical to hypothesize that lateral extensions from the pronotum are a derived condition in the Petalocephala genus group; however, this was not demonstrated in the analyses.

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.