Review of the "red" Empoascini leafhoppers in the genera Baguoidea Mahmood, 1967, Dayus Mahmood, 1967 and Homa Distant, 1908 (Hemiptera, Cicadellidae, Typhlocybinae) from Asia and the West Pacific

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Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / https://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2022 ISSN (imprimé / print) : 1280-9551/ ISSN (électronique / electronic) : 1638-9387 549 ZOOSYSTEMA • 2022 • 44 (22) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com urn:lsid:zoobank.org:pub:27DA11CC-1A56-4E2A-BF0E-77F630407E7D Webb M. D. & Xu Y. 2022. — Review of the “red” Empoascini leafhoppers in the genera Baguoidea Mahmood, 1967, Dayus Mahmood, 1967 and Homa Distant, 1908 (Hemiptera, Cicadellidae, Typhlocybinae) from Asia and the West Pacific. Zoosystema 44 (22): 549-563. https://doi.org/10.5252/zoosystema2022v44a22. http://zoosystema.com/44/22 ABSTRACT The “red” Empoascini leafhoppers (Cicadellidae, Typhlocybinae), from Asia and the West Pacific, of the genera Baguoidea Mahmood, 1967, Dayus Mahmood, 1967 and Homa Distant, 1908 are reviewed. The following species and subspecies are treated: Baguoidea rufa (Melichar, 1903) from Sri Lanka and Myanmar, placed as a senior synonym of both Baguoidea rubra Mahmood, 1967 n.syn., from the Philippines and B. yunanensis Qin& Zhang, 2010 n.syn., from China; Dayus euryphaessus (Kirkaldy, 1907) from Australia(?) and Fiji; D. euryphaessus ssp. rubrocincta (Linnavuori, 1960a) from Fiji; D.formosus Dworakowska& Viraktamath, 1978 from China (Hong Kong, new record; Taiwan) and India; D.upoluanus (Osborn, 1934) from Samoa; Homa insignis Distant, 1908 n.stat. (a species revalidated from synonymy with H.haematoptila (Kirkaldy, 1906)) from Sri Lanka, Myanmar (new record), China (new record) and Thailand (new record); H.haematoptila (Kirkaldy, 1906) from Australia (not Sri Lanka, Philippines, Thailand and China as recorded by Xu et al. 2022); H.katoi Dworakowska, 1984 from Malaysia including Sabah (new record); H.rubrodorsata Kato, 1933 from Taiwan; H.sinensis Qin& Zhang, 2011 from China. Checklists for all the species of Homa and Dayus are given, as well as a key for Homa species. Images are given for the first time for the types of Baguoidea rubra, Dayus upoluanus, D. euryphaessus, D.euryphaessus ssp. rubrocincta and Homa haematoptila and genitalia figures provided for the first time for Dayus upoluanus together with revised genitalia figures for Homa insignis, the same as given for H.haematoptila by Xu et al. (2022), in error. Michael D. WEBB Department of Life Sciences (Insects), The Natural History Museum, London, SW7 5BD (United Kingdom) [email protected] (corresponding author) Ye XU Institute of Entomology, Jiangxi Agricultural University, Nanchang 330045 (China) [email protected] Submitted on 3 February 2022 | Accepted on 22 July 2022 | Published 7 December 2022 Review of the “red” Empoascini leafhoppers in the genera Baguoidea Mahmood, 1967, Dayus Mahmood, 1967 and Homa Distant, 1908 (Hemiptera, Cicadellidae, Typhlocybinae) from Asia and the West Pacific KEY WORDS Auchenorrhyncha, Oriental region, eighth abdominal sternite apodemes, Kiwi fruit, new records. 550 ZOOSYSTEMA • 2022 • 44 (22) Webb M. D.& Xu Y. RÉSUMÉ Révision des cicadelles Empoascini “rouges” des genres Baguoidea Mahmood, 1967, Dayus Mahmood, 1967 et Homa Distant, 1908 (Hemiptera, Cicadellidae, Typhlocybinae) de l’Asie et du Pacifique ouest. Les cicadelles Empoascini “rouges” de l’Asie et du Pacifique ouest appartenant aux genres Baguoidea Mahmood, 1967, Dayus Mahmood, 1967 and Homa Distant, 1908 sont révisées. Les espèces et sous-espèces suivantes sont traitées: Baguoidea rufa (Melichar, 1903) de Sri Lanka et Myanmar, placée comme synonyme senior de Baguoidea rubra Mahmood, 1967 n.syn., des Philippines et de B. yunanensis Qin& Zhang, 2010 n.syn., de Chine ; Dayus euryphaessus (Kirkaldy, 1907) d’Australie(?) et des îles Fidji ; D. euryphaessus ssp. rubrocincta (Linnavuori, 1960a) des îles Fidji ; D. formosus Dworakowska& Viraktamath, 1978 de Chine (Hong Kong, nouvelle signalisation; de Taïwan) et de l’Inde ; D.upoluanus (Osborn, 1934) de Samoa ; Homa insignis Distant, 1908 n. stat. (une espèce ici sortie de sa synonymie avec H.haematoptila (Kirkaldy, 1906) et revalidée) du Sri Lanka, de Myanmar (nouvelle signalisation), de Chine (nouvelle signalisation) et de Thaïlande (nouvelle signalisation) ; H.haematoptila d’Australie (non du Sri Lanka, des Philippines, de Thaïlande ou de Chine comme l’ont rapporté Xu etal. [2022]) ; H.katoi Dworakowska, 1984 de Malaisie, y compris Sabah (nouvelle signalisation) ; H.rubrodorsata Kato, 1933 de Taiwan ; H.sinensis Qin& Zhang, 2011 de Chine. Le liste des espèces de Homa et Dayus est fournie ainsi qu’une clé d’identification revisée pour les espèces de Homa. Des photos sont proposées pour la première fois pour les types de Baguoidea rubra, Dayus upoluanus, D. euryphaessus, D.euryphaessus ssp. Rubrocincta et Homa haematoptila ; les genitalia sont illustrés pour la première fois pour Dayus upoluanus ; et des figures révisées des genitalia de Homa insignis sont fournies, identiques à celles données par erreur pour H.haematoptila par Xu etal. (2022). INTRODUCTION Species of the leafhopper genera Baguoidea Mahmood, 1967 and Homa Distant, 1908 and some species of Dayus Mahmood, 1967 are mainly red in colour. This coloration makes them conspicuous elements of the Asian and Pacific fauna and readily noticeable in collections. All three genera belong to the “Usharia group” of Empoascini Distant, 1908. This group, first mentioned by Qin etal. (2011), also comprised the Asian genera, Goifa Dworakowska, 1977, Ifugoa Dworakowska& Pawar, 1974, Treufalka Qin& Zhang, 2008 and Usharia Dworakowska, 1977; later Radicafurcus Qin& Zhang, 2010 was also included (Xu etal. 2021b: fig.3). The group was defined by Xu etal. (2017: 468) by the unbranched MP+CuA vein in the hind wing (Fig.3D); a solidly attached or fused aedeagus and connective (Fig.3J) and in Dayus, Homa and Ifugoa the connective arms are also highly modified (Figs4E; 5K). All genera also have all apical cells of the forewing arising from the m cell (Fig.3C) with the 3rd apical cell petiolate in Baguoidea and Dayus (Fig.3C) and the basal group macrosetae of the subgenital plate present, which are truncate apically in Baguoidea (Fig.3L) and some Homa (Fig.5F). In addition, all three genera have the ventral abdominal basal apodemes reduced, being replaced in Baguoidea and Dayus with long divergent dorsal basal apodemes (Fig.3M), and in Dayus an unusual ventral apodeme of the 8 th sternite is also present (Fig.4F), see also Remarks under Baguoidea and Dayus. Previous figures of the basal abdominal apodemes in these genera have either incorrectly stated them as ventral or have not indicated their position. Qin etal. (2014) provided a key including all three genera and Xu et al. (2021b) a phylogeny of Empoascini. The current work arose from identifying red marked typhlocybine specimens from the West Pacific in the Natural History Museum, London collection, specimens which proved to be mainly new species of Dayus and Homa. All species of Homa have distinctive colour pattern (Fig.1J) and the single species of Baguoidea and some species of Dayus are mainly red in colour; these genera and species are reviewed here and a key provided for their separation. A key to all Empoascini, including the “Usharia group” from China is given by Qin etal. (2014). The above three genera were included in the first major work on Oriental Typhlocybinae Kirschbaum, 1868 by Mahmood (1967), see ‘Discussion’. Other externally similar, predominantly reddish, Asian Empoascini include the following: Alebroides rubicundus Ishihara, 1953 (see Dworakowska 1997: 311, figs726-731), A.rubrus Dworakowska, 1994b: 98, Schizandrasca rubrifrons (Matsumura, 1931) (Dworakowska 1982: 53, figs259, 260), and males of Alebrasca actinidiae Hayashi& Okado, 1994 (on Kiwi fruit), Rubiparvus bistigma Xu, Dietrich& Qin, 2016 (see Xu etal. 2016: 585, figs1-4), Nikkotettix galloisi Matsumura, 1931 (see Dworakowska 1982, figs268-275) and N.taibaiensis Qin& Zhang, 2003. MATERIAL AND METHODS “West Pacific” in the text refers to the region between and including Australia and the Philippines. Except where indicated, distribution records ending with (?) relate to records considered dubious by the authors. MOTS CLÉS Auchenorrhyncha, région orientale, apodèmes du huitième sternite abdominal, kiwi, signalisations nouvelles. 551 Revue of the “red” Empoascini leafhopper ZOOSYSTEMA • 2022 • 44 (22) AbbreviAtions The specimens studied or referred to are deposited in the collections abbreviated in the text as follows: EIHU Entomological Institute of Hokkaido University, Sapporo; INHS Illinois Natural History Survey, Champaign, NHM The Natural History Museum, London; NWAFU Entomological Museum, Northwest A&F University, Yangling, Shaanxi; QSBG Queen Sirikit Botanical Garden, Chiang Mai; USNM United States National Museum, Washington. TAXONOMY CheCklist of the “red” West PACifiC generA And sPeCies of the “Usharia grouP” of emPoAsCini Baguoidea rufa (Melichar, 1903); Sri Lanka, Myanmar, Philippines, China. Baguoidea rubra Mahmood, 1967 n. syn. Baguoidea yunanensis Qin& Zhang, 2010 n. syn. Dayus euryphaessus (Kirkaldy, 1907); Australia(?), Fiji Dayus euryphaessus ssp. rubrocincta (Linnavuori, 1960a), Fiji Dayus formosus Dworakowska& Viraktamath, 1978, India, China (Hainan, Hong Kong, new record and Taiwan) Dayus upoluanus (Osborn, 1934), Samoa Homa haematoptila (Kirkaldy, 1906); Australia (not Sri Lanka, Philippines, Thailand and China as recorded by Xu et al. [2022]). Homa insignis Distant, 1908, n. stat., Sri Lanka, Myanmar (new record), China (new record), Thailand (new record) Homa katoi Dworakowska, 1984; Malaysia, including Sabah (new record). Homa rubrodorsata Kato, 1933; Taiwan. Homa sinensis Qin& Zhang, 2011; China. Family CiCAdellidAe Latreille, 1825 Genus Baguoidea Mahmood, 1967 Baguoidea Mahmood, 1967: 40.— Qin etal. 2010: 55; 2014: 1498 (key). tyPe sPeCies.— Baguoidea rubra Mahmood, 1967 by original designation. distribution.— Mainland Asia (Sri Lanka, Myanmar and China), Philippines and Japan(?), see Remarks of B.rufa below. remArks This monotypic genus is tentatively regarded as distinct from Dayus. It differs only in having the basal group setae of the subgenital plate blunt-tipped (also found in some Homa species) and distal macrosetae more numerous (Fig.3L) and in lacking an unusual ventral apodeme of the 8th abdominal sternite, found in Dayus. In other respects, it is similar to Dayus in its overall red colour, having the3rd apical cell in the forewing petiolate (Fig.3C), male pygofer with processes (Fig.3G), form of the subgenital plate and aedeagus and well developed male basal dorsal abdominal apodemes (Fig.3M). The genus was redescribed by Yu& Yang (2013) as there were some errors in Mahmood’s original description and figures, which also had implications on subsequent studies (see Remarks under B.rufa below). Yu& Yang (2013) also noted that Dworakowska (1973) had stated (and figured) that the facial laterofrontal sutures were absent in Baguoidea but were present in their specimens, and are also present in the specimens studied here. This apparent mistake by Dworakowska may have been due to the sutures being obscure against the red facial colour. Baguoidea rufa (Melichar, 1903) (Figs1A-F; 3) Empoasca rufa Melichar, 1903: 212, plate vi, fig.2a, b.— Distant 1908: 402.— Metcalf 1968: 351 (see Remarks below). Baguoidea rubra Mahmood, 1967: 42, plate 9, fig.1. n. syn. Baguoidea rufa – Dworakowska 1973: 49, figs1-12, 15; 1994a: 5. Baguoidea yunnanensis Qin& Zhang in Qin etal., 2010: 55, figs1527.— Qin etal. 2014: 1495, figs12, 46, 65, 94. n. syn. distribution.— Mainland Asia (Sri Lanka, Myanmar and China), Philippines and Japan(?) see final comments in Remarks below. mAteriAl exAmined.— Sri Lanka • 1 ♀ ; Peradeniya; IV.1906; Distant Coll; NHM. Myanmar • 1 ♂ ; Myitta, Doherty; coll. Distant; NHM; parasitized • 1 ♂ ; Myitta, Doherty; coll. Distant; NHMUK 013588830. Philippines • 1 ♂ ; Ifugao Prov., Luzon, Banaue; 20.VII.1980; NHM. remArks B. rufa was described from a single specimen from Sri Lanka with the following data (translated from the German): “Peradeniya. This nice Cicadine (1 ♂) was captured by DrUzel on 2 May 1903 in the Botanical Garden on the shrub Dichopsis laevifolia Benth. [=Palaquium laevifolium (Thwaites) Engl. (Sapotaceae)]”. As the recorded host plant in Sri Lanka is an endemic (critically endangered) species and as B. rufa is known from outside Sri Lanka it clearly feeds on other hosts. The specimens recorded from Myanmar by Distant (1908) are probably the same as examined here (Fig.1E, F). Baguoidea rubra was described from the holotype male (Fig.1C, D) and five paratypes (Fig.1A, B) from the Philippines with data: “Baguio, Benguet, Baker” (USNM). The new synonymy of B. rufa and B. rubra is based on the type figures of the former given by Dworakowska (1973), the original description of the latter and images of its holotype sent by J. Zahniser (USNM) and the specimens studied. The differences between the two species, noted by Dworakowska (1973: 49), are either errors in the original description, i.e., Mahmood’s incorrect statement of forewings “mottled with red patches”, which are not present in the holotype images seen (see above) or an acceptable range of species variation, i.e., position of distal aedeagal processes; while the long pygofer processes figured by Mahmood for B.rubra is also probably an error. A specimen from Myanmar examined differs slightly in the male genitalia from the Sri Lanka type of B. rufa (figured by Dworakowska 1973) and the examined Philippine specimen 552 ZOOSYSTEMA • 2022 • 44 (22) Webb M. D.& Xu Y. in having the pygofer process slightly more sinuate apically and in having the lateral fine setae adjacent to the macrosetal row shorter. The same setae are shown longer and greater in number in Qin etal.’s (2010) fig.26 of the junior synonym B.yunnanensis (see reproduced figure here, Fig.3L). The latter species was described from a single specimen from China and distinguished from B.rufa (and B.rubra, the other junior synonym of B.rufa) by the forewing colour (which according to all specimens seen is erroneous) and differences in pygofer and subgenital plate setae and spines at the apex of the aedeagal processes, all differences which fall within the accepted range of species variation. Genitalia figures drawn by Dworakowska (1973) were presumably taken from the holotype, as the only specimen examined, and as shown by Dworakowska’s figs7-9 the base of the aedeagus was damaged when dissected. However, the correct aedeagal base is shown in Fig.3I (lateral view) and Fig.3J (dorsal view) which matches the specimens examined here and which is remarkably similar to that of some Dayus species (see Fig.4D). It should also be noted, that the subgenital plate basal group setae are dorsal (Fig.3L) rather than ventral as shown in Dworakowska’s (1973) figs3, 4 and that the abdominal apodemes described by Qin etal. (2010), and shown in their figure 27 (and reproduced here, Fig.3M), are dorsal, and are a feature of the genus (see generic Remarks). Finally, the references for Japan for this species by Esaki (1932, 1950), Esaki& Ito (1954) and Kato (1933b) need to be confirmed due to the similarity of some other red marked Empoascini (see Introduction). The reference of the species from Japan (Matsumura 1934) presumably refers to Dayus takagii Dworakowska, 1971, as this species was described from material in Matsumura’s collection from Japan and also Hong Kong (see Remarks under D.takagii). Genus Dayus Mahmood, 1967 Dayus Mahmood, 1967: 39.— Qin& Zhang 2007: 43.— Yu& Yang 2013: 2.— Qin etal. 2014: 1498 (key); 2021a: 229. t yPe sPeCies .— D. elongatus Mahmood, 1967, by original designation. distribution.— Widespread in Asia and the Pacific. remArks This genus is tentatively regarded as distinct from Baguoidea (see Remarks under that genus). It differs only in having the basal group setae of the subgenital plate not blunt-tipped and in having an unusual ventral apodeme of the 8th abdominal sternite with a pair of short anterior lobes (Fig.4F), and also found here with a series of short setae at the posterior lateral corner of the 8 th sternite. There is considerable variation among species in proportion of the vertex, size and colour, for example, in Dayus formosus the vertex is distinctly longer medially than next to eye but more or less the same in D.trifurcatus Yu& Yang, 2013 while the latter is yellow and not red as in some other congeners and very much bigger (twoparatypes of D. trifurcatus [♂ and ♀] examined, NHM). A key to the Chinese species of the genus was given by Yu& Yang (2013). CheCklist And distribution of DayUs sPeCies D.bifurcatus Yu& Yang, 2013: 3, figs1-9. China (Zhejiang). D.elongatus Mahmood, 1967: 39, plate 8, fig.1. Malaysia (Singapore). D.euryphaessus (Kirkaldy, 1907): 68. Fiji, Australia(?). D.formosus Dworakowska& Viraktamath, 1978: 544, figs3341. India, China (Hainan, Hong Kong new record, Taiwan). D.furcatus Xu, Dietrich& Qin, 2021a: 231, figs10-17. Thailand. D.lamellatus Qin& Zhang, 2007: 48, figs22-31. China (Fujian, Zhejiang). D.lii Qin& Zhang, 2007: 45, figs1-12. China (Fujian). D.membranaceus Qin& Zhang, 2007: 45, figs13-21. China (Fujian, Jiangxi). D.serratus Yu& Yang, 2013: 4, figs24-30. China (Hainan). D.takagii Dworakowska, 1971: 501, figs1-11. China (Sichuan, Hong Kong, Taiwan), Japan. D.trifurcatus Yu& Yang, 2013: 3, figs10-23. China (Chongqing). D.upoluanus (Osborn, 1934): 190, fig.15. Western Samoa. Dayus euryphaessus (Kirkaldy, 1907) (Fig.2A-F) Cicadula euryphaessa Kirkaldy, 1907: 68; 1908: 383. Empoasca euryphaessa – Linnavuori 1960a: 17, Fig.5f, h-j.— Evans 1966: 266 (misidentification(?) see Remarks below).— Wilson 2009: 46. Empoasca euryphaessa rubrocincta Linnavuori, 1960a: 18.— Dworakowska 1971: 501.— Wilson 2009: 46. Dayus euryphaessus – Dworakowska 1971: 501. distribution.— Fiji, Australia(?) see Remarks below. mAteriAl exAmined.— Fiji • 1 ♂ ; Labasa; R. Veitch; VII.1921; NHM • 1 ♀; Lautoka; W. Greenwood; 6.XII.1921; breeding on leaves of Glochidion sp.; NHM • 1 ♀; Loloti; W. Greenwood; 19.XII.1920; NHM. remArks This species was described from an unknown number of specimens (syntypic) from Fiji with data: “Viti Levu, Rewa (Mar.-Apr.) Navna (Feb. Muir’s No. 53) on a native tree, also on Saccarum officinarum” (Sugarcane) (BPBM). The data label on an imaged specimen seen (BPBM) labelled both “type” and “Holotype”, is as follows: “Fiji Is 1905 [printed] 53[handwritten]” (Fig.2A, B). An image of four other syntype series specimens (labelled “Paratype”) has been seen (BPBM). In its description Kirkaldy noted the following: “Allied to C.rufa (Melichar) [=Dayus rufus] but the head is longer, pronotum shorter, legs pale, etc. Bright scarlet, vertex pale testaceous with a medio-longitudinal suffused scarlet stripe, which forks at the base of the frons and extends all over the face suffusedly (sometimes the red stripe is obsolete at the fork, the frons then being entirely pale, only the genae and the clypeus, etc., red.)”. 553 Revue of the “red” Empoascini leafhopper ZOOSYSTEMA • 2022 • 44 (22) Fig. 1. — Baguoidea Mahmood, 1967 and Homa Distant, 1908 species: A-F, Baguoidea rufa (Melichar, 1903), A, B, paratype of B. rubra Mahmood, 1967, A, dorsal habitus, B, labels; C, D, holotype of B. rubra Mahmood, C, male genitalia (on mount), D, fore and hind wings on mount; E, dorsal habitus (non-type: Myanmar); F, labels of same; G-I, Homa insignis Distant, 1908 (lectotype), G, dorsal habitus; H, fore and hind wings on mount; I, labels; J-K, Homa katoi Dworakowska, 1984 (paratype), J, dorsal habitus, K, labels; L-M, Homa haematoptila (Kirkaldy, 1906) (holotype), L, dorsal habitus, M, labels. Scale bars: A, L, 1 mm; E, G, J, 2 mm. A B E F G J K L M HI CD 554 ZOOSYSTEMA • 2022 • 44 (22) Webb M. D.& Xu Y. Fig. 2. — Dayus Mahmood, 1967 species: A-F, Dayus euryphaessus (Kirkaldy, 1907), A, B, holotype, dorsal habitus and labels respectively; C-F, types Empoasca euryphaessa rubrocincta Linnavuori, 1960, C, D, allotype, dorsal habitus and labels respectively, E, F, holotype, dorsal habitus and labels respectively; G-N, Dayus upoluanus (Osborn, 1934), G, dorsal habitus of female paratype; H, labels of same; I, male basal dorsal apodemes; J, male pygofer, lateral view; K subgenital plate, ventral view; L, Xth segment, ventral view; M, aedeagus, lateral view; N, aedeagus, connective and style, ventral view. Scale bars: A, C, E, 0,75 mm; G, 2 mm. A B C GH J K L M N I D E F 555 Revue of the “red” Empoascini leafhopper ZOOSYSTEMA • 2022 • 44 (22) Dworakowska (1971: 501) noted her placement of D.euryphaessus in Dayus was based on the original description, presumably its colour, proportions of the head and thorax and venation. In the original description it was also noted that the venation was similar to Baguoidea rufa and this is confirmed by Linnavuori’s (1960a) redescription, which reads “third apical cell of elytra triangular and stalked” and the material examined here. However, the species is only tentatively retained in Dayus as the male genitalia show certain differences to that genus, i.e., the pygofer lacks a caudo-dorsal lobe, the subgenital plate lacks a basal setal group and the aedeagal shaft is very short without processes, all characters also found in the similar D.upoluana, although the aedeagus and connective are fused (Fig.2N) and dorsal abdominal apodemes are present (Fig.2I), as in other congeners. Linnavuori (1960a) identified (and figured) D.euryphaessus from 29 specimens from Fiji, Lami, as part of a collection from Fiji sent to him by BPBM (see Introduction in Linnavuori 1960a) and not the type series, also BPBM (see above). The identification is however considered correct based on the distinctive marking of the species. It is also worth noting that Linnavuori’s figure of the aedeagus is in lateral view, but its attached connective is shown in dorsal view, while his figure of long sternal abdominal apodemes are dorsal, as in other Dayus species. Linnavuori (1960a) also described a new subspecies (Empoasca euryphaessa rubrocincta) from the holotype and allotype (BPBM) (Fig.2C-F) and one paratype in his own collection, from the same locality as the nominate subspecies, Fiji, Lami. It seems strange that Linnavuori (1960a), when referring to his new subspecies, should say “As the nominate form…” as he described the red marking of the two subspecies differently, confirmed by the images reproduced here (Fig.2A, C, E). However, his observation that the male genitalia were the same in the two subspecies, and their same type locality, suggests they are the same taxon with unaccountably different markings. In synonymising the two, Dworakowska (1971) noted the type-series of Empoasca euryphaessa rubrocincta had been studied but from personal communication a single specimen was studied. The record of the species from Australia (Kuranda) by Evans (1966: 266), could be incorrect and could be the specimen of Homa haematoptila, from Kuranda, noted below. Dayus formosus Dworakowska& Viraktamath, 1978 Dayusformosus Dworakowska& Viraktamath, 1978: 544, figs3341.— Qin& Zhang 2007: 51. distribution.— India, China (Hainan, Hong Kong new record), Taiwan. mAteriAl exAmined.— India • 2 ♀ ; T. Nadu; Yercaud; I. Dworakowska det.; NHM. China • 1 ♂ ; Hong Kong (new record); NHM. remArk This species was described from the holotype male and onemale, three female paratypes from India. An additional male from Taiwan was also recorded. Dayus takagii Dworakowska, 1971 Empoasca rufa – Matsumura 1934: 5 (nec Melichar 1903, misidentification). Dayustakagii Dworakowska, 1971: 501, figs1-11.— Lee 1979: 565-566, plate 48, fig.264, figs528-530 (Korea).— Dworakowska 1982: 54, figs282-288 (Japan).— Qin& Zhang 2007: 48, figs3240, China. distribution.— Japan, Korea, China, Hong Kong(?) see Remarks below. mAteriAl exAmined.— Japan • 1 ♀ ; Kyushu; Fukuoka; 10.I.1958; NHM. remArks This species was described from the holotype male and several paratypes (male and female) from Kyushu (Japan) and two female paratypes from Hong Kong. Although no depository for the types is indicated in the original description, it states in the acknowledgements to the paper that the type series was borrowed from EIHU, and this is presumed to be Matsumura’s collection. Therefore, Matsumura’s misidentification given in the original description (see above) refers to Matsumura (1934) where Empoasca rufa Melichar is listed from Japan and not (as stated) Matsumura (1931) where it is listed from Formosa. The female paratypes from Hong Kong could equally be D.formosus examined here from a male from Hong Kong and hence the question mark under Distribution.The colour of D. takagii was not originally described except that the species resembled D. euryphaessus but with paler coloration. The female tentatively identified here as this species (based on its locality Japan), is mainly reddish in colour. Dayus upoluanus (Osborn, 1934) (Fig.2G-N) Homa upoluana Osborn, 1934: 190, fig.15. Empoasca (Homa) upoluana – Metcalf 1968: 442. Dayus upoluanus – Dworakowska 1971: 501.— Qin& Zhang 2007: 44. distribution.— Samoa. mAteriAl exAmined.— Paratypes • 2 ♂ , 1 ♀ , 1 specimen missing from mount; “Upolu Samoa”; “Apia, 9-15-23”; “Swezey& Wilder”; “Brit. Mus. 1930-467”; NHMUK 013588829 • 1 ♂ ; “Savaii, Samoa” “Safune, v.2.24”; “rain forest; 2000-4000”; “H. Bryan Jr”; “Brit. Mus. 1930-467”; NHM. remArks This species was described from 16 specimens of both sexes (holotype, allotype and paratypes) with original data: “Apia, 562 ZOOSYSTEMA • 2022 • 44 (22) Webb M. D.& Xu Y. Acknowledgements The authors thank Valérie A.Lemaître (NHM), Chandra Viraktamath (Agricultural University, Bangalore, India), Chris Dietrich and Dmitry Dmitriev (INHS) and Murray Fletcher (formerly of the Orange Agricultural Institute, New South Wales, Australia) for constructive comments on earlier drafts of the manuscript, James Zahniser (USNM) for imaging types of Baguoidea rubra, Dr I. Dworakowska (formerly of the Museum and Institute of Zoology of the Polish Academy of Sciences, Warsaw, Poland and of the Forest Sciences Department of the University of British Columbia, Vancouver, Canada) and Masami Hayashi (formerly of the Department of Biology, Faculty of Education, Saitama University, Urawa 338, Japan) for helpful discussion and Radvile Markeviciute (Nature Research Centre, Lithuania) for preparing the plates. Special mention is made of Jim Boone (BPBM) who prematurely passed away, soon after sending images of BPBM types. The work by the second author was funded by the National Natural Science Foundation of China (32160121). REFERENCES dietriCh C. h. 2005.— Keys to the families of Cicadomorpha and subfamilies and tribes of Cicadellidae (Hemiptera: Auchenorrhyncha). Florida Entomologist 88 (4): 502-517. https://doi. org/10.1653/0015-4040(2005)88[502:KTTFOC]2.0.CO;2 dietriCh C. h. & dmitriev d. 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Forewing reddish in basal half to two-thirds (Fig. 1A, E) or with a ‘T’ shaped red mark (Fig. 2E); third apical cell petiolate (Fig. 3C); male pygofer with a ventral process; subgenital plate slightly produced basolaterally but without a distinct lobe (Figs 3L; 4I); aedeagal shaft elongate or very short .................................................... 2 — Forewing marked with red in corium basally, along clavus distally and continued across wing (Fig.1H, J); third apical cell not petiolate (Fig. 5D); male pygofer without a ventral process; subgenital plate with a distinct basolateral lobe (Figs 5F; 6A); aedeagal shaft elongate (Homa Distant, 1908) ..................................................... 5 — Subgenital plate basal group macrosetae truncate apically; lateral macrosetae uniseriate and few in number in basal two thirds, irregularly arranged and numerous in distal third (Fig. 3L); aedeagal shaft elongate (Fig.3I); male 8th abdominal sternite without apodeme .............................................. Baguoidea rufa (Melichar, 1903) — Subgenital plate basal group macrosetae acute apically, lateral macrosetae uniseriate throughout and few in number (Fig.4I); male 8th abdominal sternite with apodeme (Fig.4F) (Dayus Mahmood, 1967) ............... 3 2. Aedeagal shaft very long with a pair of apical subparallel processes (see Dworakowska& Viraktamath, 1978, fig.36); India, China (Hainan, Taiwan and Hong Kong) ................. Dayus formosus Dworakowska & Viraktamath, 1978 — Aedeagal shaft very short without a pair of apical processes ......................................................................... 4 3. Aedeagal shaft evenly curved in lateral view (Fig. 2M); Samoa ..................... Dayus upoluanus (Osborn, 1934) — Aedeagal shaft angularly curved in lateral view (see Linnavuori 1960: fig.5i); Fiji .......................................... ................................................................................................................ Dayus euryphaessus (Kirkaldy, 1907) 4. Subgenital plate with latero-basal lobe narrow, with group of lateral macrosetae near midlength (Fig. 5M); China ....................................................................................................... Homa sinensis Qin & Zhang, 2011 — Subgenital plate with latero-basal lobe broad, without group of lateral macrosetae near midlength (Fig. 6G) .... 6 5. Aedeagal shaft relatively broad in lateral view (Fig. 6A); mainland Asia .............. Homa insignis Distant, 1908 — Aedeagal shaft very narrow in lateral view (see Dworakowska 1978: fig.54); Malaysia .................................... .................................................................................................................... Homa katoi Dworakowska, 1984 563 Revue of the “red” Empoascini leafhopper ZOOSYSTEMA • 2022 • 44 (22) Hungarica 55: 93-215. 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