A review of the tribes of Deltocephalinae (Hemiptera: Auchenorrhyncha: Cicadellidae)

The classification of the largest subfamily of leafhoppers, Deltocephalinae, including 38 tribes, 923 genera, and 6683 valid species, is reviewed and revised. An updated phylogeny of the subfamily based on molecular (28S, Histone H3) and morphological data and an expanded taxon sample (37 taxa not included in previous analyses) is presented. Based on the results of these analyses and on the morphological examination of many representatives of the subfamily, the classification of the tribes and subtribes of Deltocephalinae is revised. Complete morphological descriptions, illustrations, lists of the included genera, and notes on their distribution, ecology, and important vector species are provided for the 38 recognized tribes and 18 subtribes. A dichotomous key to the tribes is provided. All names in the taxonomic treatments are hyperlinked to online resources for individual taxa which are supported by a comprehensive database for Deltocephalinae compiled using the taxonomic database software package 3I. The online functionality includes an interactive key to tribes and subtribes and advanced database searching options. Each taxon (subspecies through subfamily) has a unique taxon webpage providing nomenclatural information, lists of included taxa, an automated description (if available), images (if available), distributional information, bibliographic references and links to outside resources. Some observations and trends regarding the history of taxonomic descriptions in Deltocephalinae are reported. Four new tribes are described: Bahitini tribe nov. (25 genera), Bonsapeiini tribe nov. (21 genera), Phlepsiini tribe nov. (4 genera), and Vartini tribe nov. (7 genera). The circumscription and morphological characterization of Scaphoideini Oman, 1943 (61 genera) is substantially revised. Eleven new species are described: Acostemma stilleri sp. nov., Arrugada linnavuorii sp. nov., Drabescus zhangi sp. nov., Parabolopona webbi sp. nov., Goniagnathus emeljanovi sp. nov., Hecalus hamiltoni sp. nov., Scaphoideus omani sp. nov., Dwightla delongi sp. nov., Abimwa knighti sp. nov., Gannia viraktamathi sp. nov., and Doratulina dmitrievi sp. nov. Some family-group level taxonomic changes are made: Platymetopiini Haupt, 1929, Anoterostemmini Haupt, 1929, and Allygidiina Dmitriev, 2006 are synonymized with Athysanini Van Duzee, 1892, syn. nov.; Procepitini Dmitriev, 2002 is synonymized with Cicadulini Van Duzee, 1892, syn. nov.; Listrophorini Boulard, 1971 is synonymized with Chiasmini Distant, 1908, syn. nov.; Adamini Linnavuori & Al-Ne’amy, 1983, Dwightlini McKamey, 2003, and Ianeirini Linnavuori, 1978 are synonymized with Selenocephalini Fieber, 1872 syn.nov., and all three are now recognized as valid subtribes in their parent tribe. New placements of many genera to tribe and subtribe are made, and these are described in individual taxon treatments

Paralaevicephalus longistylus Dai, Zhang et Hu

Paralaevicephalus longistylus Dai, Zhang et Hu Figs. 13–16 Paralaevicephalus longistylus Dai, Zhang et Hu, 2005: 407 –408, Figs. 14–18. Distribution. China (Yunnan).Published as part of Xing, Jichun, Dai, Renhuai & Li, Zizhong, 2009, A taxonomic study of the genus Paralaevicephalus Ishihara (Hemiptera: Cicadellidae: Deltocephalinae), with description of four new species from China, pp. 53-61 in Zootaxa 1979 on page 57, DOI: 10.5281/zenodo.18524

Paralaevicephalus nigrifemoratus Matsumura

Paralaevicephalus nigrifemoratus (Matsumura) Figs. 1–4 Deltocephalus nigrifemoratus Matsumura, 1902: 399, Fig. 26. Paralaevicephalus nigrifemoratus Ishihara, 1953: 47, Fig. 4: 4, 17: 5. Metcalf 1967: 1307. Nast 1972: 418. (Note: Fig. 1–4 and 13–16 after Dai, Zhang et Hu, 2005) Lee 1979: 427, Figs. 358, 359. Cai et al. 2001 a: 99. Cai et al. 2001 b: 207. Cai and Shen 2002: 276. Dai et al. 2005: 408. Distribution. China (Jiangxi, Henan, Fujian, Guizhou, Anhui, Sichuan, Taiwan); Japan; Korea.Published as part of Xing, Jichun, Dai, Renhuai & Li, Zizhong, 2009, A taxonomic study of the genus Paralaevicephalus Ishihara (Hemiptera: Cicadellidae: Deltocephalinae), with description of four new species from China, pp. 53-61 in Zootaxa 1979 on pages 54-56, DOI: 10.5281/zenodo.18524

Paralaevicephalus Ishihara

<i>Paralaevicephalus</i> Ishihara <p> <i>Paralaevicephalus</i> Ishihara, 1953: 14.</p> <p> <i>Khasiana</i> Rao, 1989: 81, synonymized by Dai, Zhang <i>et</i> Hu, <i>Paralaevicephalus</i> Ishihara, 2005: 404. <b>Type species:</b> <i>Deltocephalus nigrifemoratus</i> Matsumura, 1902.</p> <p> <b>Description.</b> Dirty yellow species, often with light veins on forewings. Head produced in front, slightly wider and longer than pronotum. Vertex triangular, each side near fore margin with oblique streak. Eyes black, fairly long and oblique; ocelli semitransparent, located on the lateral margin of vertex, separated from adjacent eye by approximately their own diameter. Face with dark, often confluent, transverse streaks. Frontoclypeus narrow, longer than width between eyes, anteclypeus slightly narrowed apically. Antennae arising near lower corner of eye. Pronotum with sides short, its anterior margin produced and posterior margin concave. Scutellum triangular, its length slightly shorter than length of pronotum, with transverse suture curved and depressed. Forewings with four apical cells and three subapical cells, 3.8–4.2 times as long as wide, apical cells sometimes with dark streaks, appendix well developed.</p> <p>Male pygophore slightly longer than high, with several macrosetae on dorsal two thirds. Valve large, triangular. Subgenital plate short, with macrosetae irregularly arranged towards apex, with an elongate stout process from inner margin, directed posteriorly. Apical process of style short to long. Connective, loop-shaped with arms fused apically or Y-shaped with arms forked apically, stem present or absent. Aedeagal shaft short to elongate, without processes at apex, gonopore apical on ventral margin.</p> <p> <b>Size.</b> Length (including tegmen): 2.7– 3.5 mm.</p> <p> <b>Diagnosis.</b> This genus is similar to <i>Laevicephalus</i> DeLong in appearance, but can be distinguished from the latter by the unique subgenital plate, which has an elongate stout process arising from the inner margin, directed posteriorly, and by the aedeagal shaft without processes at the apex.</p> <p> <b>Distribution.</b> China, Japan, Korea, India.</p> <p> <b>Discussion.</b> <i>Paralaevicephalus</i> Ishihara is distributed in the Oriental and Palaearctic Regions. The genus belongs to the tribe Paralimini. In this group, the connective usually has the anterior branches fused apically. The form of the connective has long been considered a stable diagnostic trait that separates Paralimnini from other tribes of Deltocephalinae.</p> <p> The four new species described below are the only known Paralimnini having a Y-shaped connective. This is the first report of such extensive variation in connective shape within one genus of Paralimnini. The new species would normally be placed in another tribe based on their having a Y-shaped connective, but they all belong to <i>Paralaevicephalus</i> based on the unique subgenital plate and the form of the aedeagus. The new species described in this paper represent the only known case where a single genus of Deltocephalinae includes some species with the anterior connective arms divergent and others with the arms fused.</p>Published as part of <i>Xing, Jichun, Dai, Renhuai & Li, Zizhong, 2009, A taxonomic study of the genus Paralaevicephalus Ishihara (Hemiptera: Cicadellidae: Deltocephalinae), with description of four new species from China, pp. 53-61 in Zootaxa 1979</i> on pages 53-54, DOI: <a href="http://zenodo.org/record/185247">10.5281/zenodo.185247</a&gt

Paralaevicephalus grossus Xing, Dai & Li, 2009, sp. nov.

Paralaevicephalus grossus sp. nov. Figs. 30–35 Description. Color and external features as in generic description. Male genitalia: Subgenital plate short, outer and apical margins rounded, macrosetae irregularly arranged towards apex; elongate stout process arising basally from inner margin, strongly curved dorsad at apex (Fig. 32). Style short, apex broad, rounded, exceeding apical margin of subgenital plate, based broad (Fig. 32). Connective with arms forked, Y-shaped, stem and arms of approximately equal length. (Fig. 35). Aedeagus with shaft short and slender, curved dorsad (Figs. 33, 34). Measurement. Length (including tegmen): 3, 3.0– 3.2 mm; Ƥ, 3.1–3.3 mm. Host. Weeds. Type Material. Holotype 3, China: Shanxi Prov., Xi’an City, Cuihuashan, 26 August 2008, coll. Jichun Xing (IEGU). Paratypes: 63,4 Ƥ, same data as holotype. Diagnosis. This species is similar to Paralaevicephalus prima (Rao), but can be distinguished from the latter by the following characteristics: aedeagal shaft slender, connective with arms forked, Y-shaped and the end of style is crassi and developed. Etymology. The species name is derived from the Latin words grossus, indicating the end of style is broad and well developed.Published as part of Xing, Jichun, Dai, Renhuai & Li, Zizhong, 2009, A taxonomic study of the genus Paralaevicephalus Ishihara (Hemiptera: Cicadellidae: Deltocephalinae), with description of four new species from China, pp. 53-61 in Zootaxa 1979 on pages 59-60, DOI: 10.5281/zenodo.18524

Paralaevicephalus prima Rao

<i>Paralaevicephalus prima</i> (Rao) <p>Figs. 5–8</p> <p> <i>Khasiana prima</i> Rao, 1989: 82, Figs. 105–108.</p> <p> <i>Paralaevicephalus prima</i> (Rao), a new combination, by Dai,Zhang <i>et</i> Hu, 2005: 405.</p> <p> <b>Distribution.</b> China (Guangxi, Hainan, Yunnan); India.</p> <p> <b>Host.</b> Weeds.</p> <p> <b>Material examined.</b> 13,1Ƥ, China, Hainan Prov., Dongfang City, Datian (19°25′N, 108°35′E), 9 July 2007, coll. Jichun Xing (IEGU); 1Ƥ, Hainan Prov., Dongfang City, Datian (19°25′N, 108°35′E), 9 July 2007, coll. Bin Zhang (IEGU); 13, Hainan Prov., Wuzhishan City, Wuzhishan (18°49′N, 109°36′E), 14 July 2007, coll. Hui Zhang (IEGU); 1Ƥ, Hainan Prov., Diaoluoshan City, Diaoluoshan (18°40′N, 109°54′E), 17 July 2007, coll. Jichun Xing (IEGU); 13,1Ƥ, Yunnan Prov.,Puer, 15 July 2007, coll. Jianda Li (IEGU); 13, Yunnan Prov., Mengyang, 15 July 2007, coll. Jianda Li (IEGU); 2Ƥ, Yunnan Prov.,Mengla, 18 July 2007, coll. Jianda Li (IEGU); 1Ƥ, Yunnan Prov.,Mengla, 18 July 2007, coll. Xiaohong Jiang (IEGU).</p>Published as part of <i>Xing, Jichun, Dai, Renhuai & Li, Zizhong, 2009, A taxonomic study of the genus Paralaevicephalus Ishihara (Hemiptera: Cicadellidae: Deltocephalinae), with description of four new species from China, pp. 53-61 in Zootaxa 1979</i> on page 56, DOI: <a href="http://zenodo.org/record/185247">10.5281/zenodo.185247</a&gt

Paralaevicephalus lamellatus Xing, Dai & Li, 2009, sp. nov.

<i>Paralaevicephalus lamellatus</i> sp. nov. <p>Figs. 36–41</p> <p> <b>Description.</b> General appearance as in <i>Paralaevicephalus grossus</i> <b>sp. nov.</b>.</p> <p>Male genitalia: Subgenital plate short, outer and apical margins rounded, macrosetae irregularly arranged towards apex; elongate stout process arising basally from inner margin, strongly curved dorsad at apex (Fig.</p> <p>38). Style short, apical process narrow, rounded, not exceeding apical margin of subgenital plate, base narrow (Fig. 38). Connective with arms forked, without stem (Fig. 41). Aedeagus with shaft short, curved dorsad, apex lamellar. (Figs. 39, 40).</p> <p> <b>Measurement.</b> Length (including tegmen): 3, 2.7–2.9 mm; Ƥ, 2.9–3.1 mm.</p> <p> <b>Host.</b> Weeds.</p> <p> <b>Type Material.</b> Holotype 3, China: Henan Prov., Jiyuan City, Wangwushan, 21 August 2008, coll. Jichun Xing (IEGU). Paratypes: 23,4Ƥ, same data as holotype.</p> <p> <b>Diagnosis.</b> This species is similar to <i>Paralaevicephalus prima</i> (Rao), but the connective is Y-shaped and the apex of the aedeagus is lamellar. It also differs from <i>Paralaevicephalus grossus</i> <b>sp. nov.</b> in lacking the stem of the connective and in is the apically lamellar aedeagus.</p> <p> <b>Etymology.</b> The species name is derived from the Latin words <i>lamellatus</i>, indicating the end of aedeagus is lamellar.</p>Published as part of <i>Xing, Jichun, Dai, Renhuai & Li, Zizhong, 2009, A taxonomic study of the genus Paralaevicephalus Ishihara (Hemiptera: Cicadellidae: Deltocephalinae), with description of four new species from China, pp. 53-61 in Zootaxa 1979</i> on pages 60-61, DOI: <a href="http://zenodo.org/record/185247">10.5281/zenodo.185247</a&gt

Paralaevicephalus bisubulatus Xing & Li 2011, sp.nov.

<i>Paralaevicephalus bisubulatus</i> Xing & Li sp.nov. <p> <i>Paralaevicephalus bisubulatus</i> Xing & Li, 2011 in Li, Dai & Xing, 2011: 165; unavailable</p> <p>Holotype: ♂, China: Guizhou Province, Kuankuoshui Nature Reserve, Baishaogou, 2 September 2010, coll. Jichun Xing.</p> <p>Type repository: Institute of Entomology, Guizhou University, Guiyang, China (GUGC).</p>Published as part of <i>Dmitriev, Dmitry A., Li, Zizhong, Dai, Renhuai & Xing, Jichun, 2024, Validation of taxon names described in " Deltocephalinae from China (Hemiptera: Cicadellidae) ", pp. 269-278 in Zootaxa 5403 (2)</i> on page 274, DOI: 10.11646/zootaxa.5403.2.5, <a href="http://zenodo.org/record/10561826">http://zenodo.org/record/10561826</a&gt