Tituria Stål

Jones, Joshua R. & Deitz, Lewis L., 2009, Phylogeny and systematics of the leafhopper subfamily Ledrinae (Hemiptera: Cicadellidae) 2186, Zootaxa 2186 (1), pp. 1-120 : 56-58

Genus Tituria Stål

(Pl. 4A–B, 6F, 11B, 15D, 16A, B, L, M, 17I, 19J)

Tituria Stål, 1865: 158 [as subgenus Petalocephala (Tituria) , elevated to genus by Stål (1866)].

Type species. Petalocephala (Tituria) nigromarginata Stål, 1865: 158 , by subsequent designation of Metcalf (1962b).

Synonymy. Epiclines Amyot and Serville, 1843: 577 (type species: Membracis planata Fabricius, 1794: 11 , by monotypy), NEW SYNONYM. Epiclinata Metcalf 1952: 228 , replacement name for Epiclines Amyot and Serville 1843: 577 , preoccupied [by Epiclines Chevrolat, 1838 ] (type species: Membracis planata Fabricius, 1794: 11 ), NEW SYNONYM.

Description. Linnavuori (1972): “Resembling Petalocephala but body shorter and much robuster [sic]. Crown shorter, +/– triangular or pentagonal in outline. Pronotum very broad, lateral margins strongly diverging caudad, the triangular wing-shaped humeral lobes extending laterad much beyond elytra, body therefore strongly constricted at base of elytra. Anterior margins of pronotum nearly straight.”

Species. [39]: acutangulata Distant ; angulata (Matsumura) ; antica (Walker) ; assamensis Distant ; borneensis Distant ; chersonesia Distant ; chinensis Distant ; clypeata Cai ; colorata Jacobi ; costalis Jacobi ; crinita Cai ; cuneata Distant ; flavimacula Cai & Shen ; flavomarginata (Kuoh & Cai) ; forficula Linnavuori ; fusca Cai & Li ; fuscipennis Kato ; hebes (Walker) ; hyalina Cai & Kuoh ; innotata Cai & Li; javanensis Distant ; laboulbenii (Signoret) ; laticoronata Cai ; lokandu Linnavuori ; maculata Kuoh ; nigricarinata Kuoh ; nigrina Cai & Kuoh ; nigrivena Cai ; nigromarginata (Stål) ; obliqua (Walker) ; obtusa Kato ; plagiata Kuoh ; planata (Fabricius) ; pyramidata Cai ; recta Cai & Mo ; sativa Cai & Shen ; timorensis Distant ; virescens Kuoh ; viridula Linnavuori.

Range. Central African Republic (Boukoko; La Maboke); China (Guizhou: Guiyang City, Sinan Co.; Hainan: Liamui; Henan: Mt. Funiu ; Shandong: Kungyushan , Taishan ) ; Congo (Mayombe; Tshuapa: Ikela); India (Anamalai Hills; Calcutta); Gabon (Bas–Ogooué); Indonesia (Java: Mt. Djampang , Soekaboemi; Sumatra: Benkoelen) ; Japan (Okinawa) ; Laos; Malaysia (Borneo: Lebang hara; Island of Penang; Tebong ) ; Singapore; South Africa (Durban; Natal: Umkomaas); Taiwan (Chipon; Eboshiyama; Ilan; Hualien Fuynan; Kaohsiung; Kueishan; Kuraru; Nantou Hsien: Howang, Jenai Yuaanfeng, Fenghuangku, Meifeng, Shanlinchi; Yuchih Lienhuachi; Pingtung Hsien: Chilushui, Kenting, Nanjenshan, Ouluanpi, Pishshieechao, Shantimen, Tsaopu; Roshan; Taipei: Yangmingshan; Taitung: Chihpen, Peinan Lichia, Taien Tupan, Yenping) ; Thailand (Chiang; Krabi; Mewah Valley; Metah Valley ) ; Vietnam (Chapa: Tonkin; Hoa Binh) .

Host plants. Unknown. The very flat, diamond-shaped nymphs have been seen by the first author on dark green leaves of woody shrubs.

Material examined. T. antica : 2 males, 2 females, South Africa, SANC, Led 1_184–187; T. forficula : 1 (abdomen missing), Congo, AMNH, JRJ _Led1_192 , 3 females (including type specimen—195), Central African Republic, MNHN, JRJ _ Led 1_193–195; Tituria sp .: 2 males, 2 females, China, Japan, Taiwan, NCSU, JRJ _Led1_188–190, 192 .

Remarks. Metcalf, in his Catalogue (1962b), recognized the monobasic genus Epiclinata with type species Membracis planata Fabricius, 1794 . He had earlier (1952) given the name Epiclinata as a replacement name for the genus Epiclines Amyot & Serville , which was preoccupied by Epiclines Chevrolat, 1838 . It appears, however, that neither Metcalf, nor any author before him, has properly fixed the type species of Epiclines Amyot & Serville according to the rules of the Code of Zoological Nomenclature, as explained below.

In 1843 Amyot & Serville gave Ledra planata (Fabricius) [ Membracis planata Fabricius, 1794 ] as the sole representative (and thus type species by implicit monotypy) for their new genus Epiclines . However, in their description, they gave as an identifying character “prothorax incliné presque verticalement.” This feature matches the later described New World genus Proranus Spinola, 1850 (see Pl. 4J, 6C) but not the Old World species M. planata (Pl. 4B, 6F), a fact pointed out by subsequent authors ( Stål 1865, 1866; Distant 1907, 1908). Amyot and Serville likely relied solely on Fabricius’ original description to identify their specimen, as no illustrations of M. planata appear to have been in existence at that time ( Metcalf 1962b).

It is clear that Amyot & Serville misidentified a specimen, possibly of Proranus , as M. planata , but the exact species they had in hand when they made their identification is not known. In such cases of misidentification, previous versions of the Code required authors to submit a case in question to a committee of the Commission of Zoological Nomenclature for their judgment in assigning a type species. Neither Metcalf, nor any of several other authors who recognized M. planata as the type species of Epiclines (see Metcalf 1962b), applied to the Commission for such resolution.

The most recent version of the Code (Fourth Edition, 1999) provides for authors to subsequently fix the type species of a genus in the case of misidentification. Article 70.3 allows for two options: selecting the species indicated by the name chosen by the original author(s) (70.3.1), or selecting the taxonomic species actually described (70.3.2) The latter option assumes that an author knows which species was actually described by the original authors. In this case, the species is not known; thus, that path is not available. The former article becomes the only option. M. planata is here fixed as the type of Epiclines Amyot & Serville (and thus Epiclinata Metcalf ). This accords with Metcalf’s earlier designation, which was performed without authority, and maintains stability with existing names (i.e., Proranus , Epiclinata ).

In the phylogenetic analyses performed in this paper, M. planata has been demonstrated to belong in a clade with species of Tituria , and it is herein transferred to Tituria . Epiclines becomes the senior synonym for this genus, but as a junior homonym, is not available. Tituria is the next oldest name, and take priority. Epiclinata becomes a junior synonym of Tituria . Epiclinata flavomarginata Kuoh & Cai, 1992 is also here moved to Tituria .

Tituria is the third largest genus of Ledrini , and like Petalocephala and Ledra , has a broad distribution. Many of its recently described Asian species appear to belong in Neotituria (see remarks for Neotituria ), but the phylogenetic limits of both genera have not yet been adequately established. Tituria antica (Walker) , placed by Linnavuori (1972: 225) in its own monobasic species group within Tituria , strongly resembles members of Thlasia in aedeagal structure and was placed near to, but not with, Thlasia in the phylogenetic analysis ( Figs. 1–3 ). Tituria needs to be revised, and with it Thlasia and Neotituria . A taxon-based (nonregional) species–level treatment of all members of the Petalocephala genus group is warranted.

Among the borrowed material of Tituria were some very large specimens, with one unidentified female from Gabon (MNHN collection) measuring 24 millimeters in length and being quite robust. Some unidentified females of Ledra from Sumatra measured 23.5 mm in length, but were not quite as robust. These are among the largest species of Ledrinae and Cicadellidae .

Tituria flavomarginata ( Kuoh & Cai, 1992) , NEW COMBINATION. Synonymy: Epiclinata flavomarginata Kuoh & Cai, 1992: 136 , new species.

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.