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The family Fulgoridae is known for their distinct morphological structures and striking colouration. Despite so, comprehensive documentation of insect-plant interaction from this charismatic family is greatly scarce. Presented here are records of plant association across four species of Fulgoridae from Malaysian Borneo. The current study was based on voucher specimens and field samplings from selected localities in Sarawak and Sabah, Malaysian Borneo. A total of 11 species of plants belonging to 11 genera and nine families were recorded. Three fulgorid species namely Penthicodes quadrimaculata, Pyrops intricatus and Py. sultanus shares the same host plant being the mata kucing fruit tree (Dimocarpus longan ssp. malesianus). The most speciose insect-plant association belongs to Pe. farinosa and Py. sultanus with six species documented. This is the first record of host plants reported for Py. intricatus, Pe. farinosa and Pe. quadrimaculata in Malaysian Borneo. Keywords: Fulgoridae, host plant, insect-plant association, Malaysian Borneo, Sabah, Sarawak

Identification key to the species of Pyrops of Vietnam, Laos, Cambodia and Southern Continental China 1. Cephalic process strongly inflated apically (Figs 9–11)................................................................... 2 – Cephalic process not strongly inflated apically (Figs 12–19)........................................................... 3 2. Abdomen black ventrally (Fig. 9B); tegmina pale yellow-white on disc and with 3 black spots in costal area before nodal line (Fig. 9A); cephalic process yellow (Fig. 9).................................................................................................................................................. Pyrops atroalbus (Distant, 1918) – Abdomen red ventrally (Figs 10B, 11B); tegmina largely black on disc (Fig. 10A), or in the pale forms, bluish white on disc and without black spots in costal area (Fig. 11A, F); cephalic process red-brown to black (Figs 10–11)................................................. Pyrops clavatus (Westwood, 1839) 3. Head entirely green, hind wings bright yellow basally or entirely yellow-orange (Figs 12–13)..... 4 – Head not entirely green, hind wings yellow, white or blue basally (Figs 14–19)............................ 5 4. Hind wings bright yellow with apical third black (Fig. 12A); ground colour of distal half of tegmina black (Fig. 12A); anterior and median legs largely black (Fig. 12A–C)............................................................................................................................................ Pyrops viridirostris (Westwood, 1848) – Hind wings entirely bright yellow-orange (Fig. 13A); ground colour of distal half of tegmina green (Fig. 13A); anterior and median legs green (Fig. 13A–C)............ Pyrops itoi (Satô & Nagai, 1994) 5. Cephalic process black or brown-black dorsally, yellowish ventrally (Figs 4–5, 14–15); pro- and mesonotum with median black or dark brown markings (Figs 4–5, 14–15).................................... 6 – Cephalic process yellow-red to red-brown dorsally, paler, yellow to dark red ventrally (Figs 16–19); pronotum unicolourous, mesonotum sometimes with small brown-black markings on sides (Figs 16– 19)..................................................................................................................................................... 8 6. Apex of cephalic process coloured like the rest of the process (Figs 4–5, 15)................................. 7 – Apex of cephalic process yellow to orange-red, strongly contrasting (Figs 6–7, 14).................................................................................................................................... Pyrops lathburii (Kirby, 1818) 7. Tegmina narrower: LTg/BTg = 2.9 (Figs 4A, 5A); spots on basal half of tegmina not fused into bands (Figs 4A, 5A); cephalic process narrower (LPr/BPrH> 9) (Figs 4–5); hind wings bright yellow (Fig. 4A) or white basally (Fig. 5A)...................................... Pyrops condorinus (Lallemand, 1960) – Tegmina broader: LTg/BTg = 2.53 (Fig. 15A); spots on basal half of tegmina fused into transverse bands (Fig. 15A); cephalic process broader (LPr/BPrH Pyrops spinolae (Westwood, 1842) 8. Hind wings white with sutural margin bordered with black up to the anal lobe (Fig. 1A); cephalic process very narrow (LPr/BPrH = 19) (Fig. 1)............................................ Pyrops buomvoi sp. nov. – Hind wings white, bright yellow or blue with only apical third black (Figs 16A, 17A, 18A, 19A); cephalic process broader (LPr/BPrH Pyrops candelaria (Linné, 1758) – Hind wings white or blue basally (Figs 17A, 18A, 19A); cephalic process dark red-brown and dark red ventrally or dark red, orange ventrally and at apex (Figs 17–19); yellow spots on membrane of tegmina (Figs 18–19) or not (Fig. 17)............................................................................................ 10 10. Tegmina without yellow spots on membrane (Fig. 17A); cephalic process dark red, orange ventrally and at apex (Fig. 17); hind wings pale blue to white basally (Fig. 17A); basal half of tegmina whitish with white-margined green spots in transverse rows, sometimes fused in bands (Fig. 17A)............................................................................................... Pyrops ducalis (Stål, 1863) – Tegmina with yellow spots on membrane (Figs 18A, 19A); cephalic process dark red-brown, dark red ventrally (Figs 18–19); hind wings white or blue basally; basal half of tegmina dark green or dark blue-green with yellow spots (Figs 18A, 19A)................................................................................11 11. Hind wings blue basally (sometimes faded to nearly white in collections specimens) (Fig. 18A); cephalic process very elongate and strongly curved dorsad (Fig. 18); ground colour of tegmina dark blue-green (Fig. 18A)....................................................................... Pyrops coelestinus (Stål, 1863) – Hind wings white basally (Fig. 19A); cephalic process very elongate and curved dorsad (Fig. 19); ground colour of tegmina green (Fig. 19A).................................... Pyrops vitalisius (Distant, 1918), Published as part of Constant, Jérôme & Pham, Hong Thai, 2022, The Lanternfly genus Pyrops in Vietnam: A new species from Central Vietnam, taxonomic changes, checklist, identification key (Hemiptera: Fulgoromorpha: Fulgoridae), pp. 123-154 in European Journal of Taxonomy 813 (1) on pages 138-139, DOI: 10.5852/ejt.2022.813.1741, http://zenodo.org/record/6468100, {"references":["Distant W. L. 1918. The Homoptera of Indo-China. Annals and Magazine of Natural History (Ser. 9) 1: 196 - 200. https: // doi. org / 10.1080 / 00222931808562301","Westwood J. O. 1839. On the family Fulgoridae, with a monograph of the genus Fulgora of Linnaeus. The Transactions of the Linnean Society of London. Second series. Zoology 18: 133 - 153. https: // doi. org / 10.1111 / j. 1095 - 8339.1838. tb 00167. x","Kirby W. F. 1818. A century of insects, including several new genera described from his cabinet. The Transactions of the Linnean Society of London. Second series. Zoology 12 (2): 375 - 453. https: // doi. org / 10.1111 / j. 1095 - 8339.1817. tb 00239. x","Lallemand V. 1960. De quibusdam Fulgoris. Entomologische Mitteilungen aus dem Zoologischen Staatsinstitut und Zoologischen Museum Hamburg 24: 1 - 7."]}

Pyrops buomvoi sp. nov. urn:lsid:zoobank.org:act: 2AFA915E-398C-4F10-A3B7-E68FA9DC2053 Figs 1–3 Diagnosis The species can be separated from all other species of Pyrops by the following combination of characters: (1) posterior wings milky white with apex and area along sutural margin black brown, with brown area paler and narrower towards basosutural angle (Fig. 1B); (2) cephalic process elongate and very slender, moderately curved dorsad (Fig. 1C–G); (3) head yellow with dorsum and sides of cephalic process reddish (Fig. 1); (4) tegmina with 3 irregular yellow bands on basal half, the two more distal ones formed from rows of elongate markings (Fig. 1B). Differential diagnosis The other species of Pyrops showing white posterior wings with dark brown or black apical portion are P. atroalbus (Distant, 1918), P. condorinus (Lallemand, 1960), P. connectens (Atkinson, 1885), P. dohrni (Schmidt, 1905), P. horsfieldii (Westwood, 1839), P. lathburii (Kirby, 1818), P. vitalisius (Distant, 1918) and P. watanabei (Matsumura, 1913) but none of them shows a black band along the posterior margin of the hind wing extending to the basosutural angle and none of them possess a very slender cephalic process. All other species of Pyrops showing a very slender cephalic process, i.e., P. hamdjahi Nagai & Porion, 2002, P. hashimotoi Nagai & Porion, 2002 and P. valerian Nagai & Porion, 2002 have brightly coloured base of posterior wings, red, blue and orange, respectively. Etymology The species epithet buomvoi is a Vietnamese name for the species of the genus Pyrops, with ʻ[con] buom ʼ meaning ʻbutterflyʼ and ʻ[con] voi ʼ meaning ʻelephantʼ. The name literally means ʻbutterfly-elephantʼ and is used as a name in apposition. Material examined Holotype VIETNAM • ♂ (dissected, portion of cephalic process missing); Annam, Cana; [11°22′16″ N, 108°51′11″ E]; 18–22 Aug. 1932; alt. 0–600 m; Pinus merhusii belt; M. Poilane leg.; “Annam-Cana, Prov. Phanrang, VIII-18-22-1932 ”, “ Pinus merkusii belt – alt. 0-600M”, “ M Poilane Coll”; VNMN. Paratype (Fig. 1) VIETNAM • ♀; same collection data as holotype; I.G. 34472; RBINS. Measurements and ratios TL: ♂ (n = 1): 29.6 mm, ♀ (n = 1): 30.9 mm; TL+process: ♀ (n = 1): 41.3 mm; LTg/BTg = 2.79; BF/ BPrH = 5.4; LPr/LF = 3.43; LPr/BPrH = 19. Description HEAD. Yellow with clypeus slightly darker; dorsum and sides of cephalic process reddish; back-side of head and markings around eyes, extending to ocelli, dark brown (Fig. 1C, E–G). Cephalic process elongate and very slender, strongly tapering basally, slightly less than 1.5 times as long as frons and clypeus combined in perpendicular view of frons (Fig. 1G), moderately curved anterodorsad and rather uniform in breadth in lateral view (Fig. 1D); distinct broadening visible in ventral and dorsal views at apical ¾ of process (Fig. 1C, G), at same level, ventral margin slightly sinuate in lateral view (Fig. 1E). Two longitudinal carinae on frons extending on sides of cephalic process up to apex (Fig. 1E–G). Median, ventral carina on apical ⅔ of cephalic process (Fig. 1F–G). Vertex with weak carina extending from middle of disc to base of cephalic process; straight carina along posterior margin (Fig. 1C). Frons subquadrate (Fig. 1F). Clypeus elongate, triangular, smooth with median carina on anteclypeus (Fig. 1F–G). Labium brown-black, elongate and slender, surpassing posterior trochanter (Fig. 1A). Antennae brown, short, with pedicel bulbous (Fig. 1E–G). THORAX. Pronotum yellow with two small, impressed brown points on disc on each side of obsolete median carina, disc slightly wrinkled (Fig. 1C); pair of weak parallel carinae along lateral margin of pronotum and on dorsal portion of lateral lobe of pronotum (Fig. 1E–F); elongate lateral marking along anterior margin of lateral lobe of pronotum, not reaching ventral margin, brown behind eye, turning to black-brown ventrad, behind ocelli; subtriangular black-brown marking in middle of ventral margin of lateral lobe of pronotum (Fig. 1E–G). Mesonotum yellow with brown markings along anterior margin, small black-brown point near base of scutellum along posterior margin and elongate, oblique, blackbrown marking extending laterad from intersection between peridiscal carina and posterior margin; median and peridiscal carinae weak, median one stopped before base of scutellum; disc slightly wrinkled between peridiscal carinae; base of scutellum impressed; lateral fields smooth (Fig. 1C). Tegulae yellow (Fig. 1C–F). TEGMINA (Fig. 1B). Brown with dense reticulum of pale yellow veins and cross-veins; corium with subbasal transverse yellow band followed by two transverse rows of slightly transverse yellow markings (markings can merge together), the more basal one irregular with the marking on disc along clavus, displaced distad (hence the two rows together more or less forming an X-shaped pattern); yellow markings more intensely margined with brown membrane with about 12 irregular round yellow spots (including 2 in costal cell), more distal ones smaller. Tegmina elongate, broadening from base towards apex, rather narrow, with costal margin broadly rounded in distal half, apical margin oblique and apical angles rounded. HIND WINGS (Fig. 1B). Milky white with apex and sutural margin largely brown-black; brown-black area getting paler and narrower from apex towards base along sutural margin and stopping at basosutural angle; veins in brown-black area progressively turning from black-brown in distal portion, to whitish at basosutural angle; veins in milky white area concolourous. Hind wings strongly broader than tegmina. LEGS (Fig. 1A–B). All coxae, trochanters and femora orangish; pro- and mesotibiae and all tarsi dark brown; metatibiae orangish basally progressively turning to brown on distal ⅓. Pro- and mesofemora broader than corresponding tibiae. Metatibiae with 5–6 lateral spines. ABDOMEN (Fig. 1A–B). Terga yellowish; sterna black-brown, narrowly yellow along posterior margin; genital segments black-brown and yellowish. MALE TERMINALIA (Fig. 2). Pygofer higher than long, with posterior margin broadly rounded, sinuate on ventral ¼ in lateral view (Fig. 2A). Gonostyli (Fig. 2 A–B, D–F) elongate, 1.64 times as long as high in lateral view, not surpassing apex of anal tube; dorsal margin broadly rounded, with a small elongate lobe in middle; in lateral view, apical margin rounded and ventral margin straight; fused ventrally on basal ⅕; lateral hooks of gonostyli short, moderately curved and pointing anteroventrally. Aedeagus membranous with pair of elongate ventral endosomal processes widening on distal half (Fig. 2 G–J); connective rodshaped (Fig. 2I); tectiductus well-developed, subtriangular with anterior margin concave in dorsal view, strongly concave ventrally (Fig. 2G–J). Anal tube (Fig. 2 A–C, F) elongate and dorsoventrally flattened, 1.4 times as long as broad in dorsal view (about as broad as long in mid-length), broadest at ⅔ of total length (Fig. 2C); lateral margins broadly rounded (Fig. 2C) and apical margin deeply, roundly notched in dorsal view (Fig. 2C); anal column elongate and narrow, surpassing anal tube posteriorly (Fig. 2A, C–D). Distribution Vietnam, Ninh Thuan Province (Fig. 3). Biology The species was collected at an altitude between 0–600 m above sea level, in a zone of Pinus merkusii Jungh. & de Vriese (Pinaceae Spreng. ex Rudolphi). However it seems very unlikely that this tree could be a host plant of the species as no species of Pyrops has ever been recorded feeding from conifers (Bourgoin 2021)., Published as part of Constant, Jérôme & Pham, Hong Thai, 2022, The Lanternfly genus Pyrops in Vietnam: A new species from Central Vietnam, taxonomic changes, checklist, identification key (Hemiptera: Fulgoromorpha: Fulgoridae), pp. 123-154 in European Journal of Taxonomy 813 (1) on pages 126-130, DOI: 10.5852/ejt.2022.813.1741, http://zenodo.org/record/6468100, {"references":["Distant W. L. 1918. The Homoptera of Indo-China. Annals and Magazine of Natural History (Ser. 9) 1: 196 - 200. https: // doi. org / 10.1080 / 00222931808562301","Lallemand V. 1960. De quibusdam Fulgoris. Entomologische Mitteilungen aus dem Zoologischen Staatsinstitut und Zoologischen Museum Hamburg 24: 1 - 7.","Atkinson E. T. 1885. Notes on Indian Rhynchota. No. 4. Journal and Proceedings of the Asiatic Society of Bengal. Calcutta 54: 127 - 158.","Westwood J. O. 1839. On the family Fulgoridae, with a monograph of the genus Fulgora of Linnaeus. The Transactions of the Linnean Society of London. Second series. Zoology 18: 133 - 153. https: // doi. org / 10.1111 / j. 1095 - 8339.1838. tb 00167. x","Kirby W. F. 1818. A century of insects, including several new genera described from his cabinet. The Transactions of the Linnean Society of London. Second series. Zoology 12 (2): 375 - 453. https: // doi. org / 10.1111 / j. 1095 - 8339.1817. tb 00239. x","Bourgoin T. 2021. FLOW (Fulgoromorpha Lists on The Web): a world knowledge base dedicated to Fulgoromorpha. Version 8, [3 Feb. 2022]. Available from http: // hemiptera-databases. org / flow / [accessed 8 Mar. 2022]."]}

Pyrops condorinus (Lallemand, 1960) Figs 4–5 Fulgora spinolae f. condorina Lallemand, 1960: 7 (described) [type in FSAG]. Fulgora spinolae condorina – Lallemand 1963: 76 (keyed). Pyrops condorina – Nagai & Porion 1996: 24 (catalogued), 29 (taxonomic note), pl. 12 fig. 170 (type illustrated). Pyrops spinolae condorinus – Liang 1998: 45 (taxonomic note). Pyrops condorinus – Constant et al. 2016: 10 (recorded from Cambodia, taxonomy, host plant), fig. 3E (live specimen). Material examined Holotype VIETNAM • ♂ of Fulgora spinolae condorina Lallemand, 1960 (dissected) (Fig. 4); Cochinchine, P. Condore; 14 Aug. 1924; R. Vitalis de Salvaza leg.; “Cochinchine, P. Condore, le 14.VIII.1924, R. Vitalis de Salvaza ”, “Type”, “Holotype ♂ Fulgora spinolae f. condorina Lallemand, 1960, Jérôme Constant det.”; FSAG. Note: “P. Condore” stands for “Poulo Condore”, a former name of Con Son Island in Con Dao Archipelago off Southern Vietnam. Additional material VIETNAM • 1 ♂ (dissected) (Fig. 5); Kiên Giang Province, Phu Quoc National Park; 10°19′30″ N, 103°57′00″ E; 14 Apr. 2013; H.T. Pham leg.; VNMN. Remark Intraspecific variation of the colour of the disc of the hind wings exists in this species, with specimens showing either bright yellow hind wings (Fig. 4) or milky white hind wings (Fig. 5). Conspecificity of both forms was confirmed by the study of the male genitalia., Published as part of Constant, Jérôme & Pham, Hong Thai, 2022, The Lanternfly genus Pyrops in Vietnam: A new species from Central Vietnam, taxonomic changes, checklist, identification key (Hemiptera: Fulgoromorpha: Fulgoridae), pp. 123-154 in European Journal of Taxonomy 813 (1) on page 131, DOI: 10.5852/ejt.2022.813.1741, http://zenodo.org/record/6468100, {"references":["Lallemand V. 1960. De quibusdam Fulgoris. Entomologische Mitteilungen aus dem Zoologischen Staatsinstitut und Zoologischen Museum Hamburg 24: 1 - 7.","Lallemand V. 1963. Revision des Fulgoridae (Homoptera). Deuxieme partie. Faunes asiatique et australienne. Memoires de l'Institut royal des Sciences naturelles de Belgique (2 e serie) 75: 1 - 99.","Nagai S. & Porion T. 1996. Fulgoridae 2: Catalogue illustre des Faunes asiatique et australienne. Sciences Nat, Compiegne.","Liang A. P. 1998. Nomenclatorial notes on the Oriental lantern fly genus Pyrops Spinola (Hemiptera: Fulgoroidea: Fulgoridae). Acta Zootaxonomica Sinica 23 (1): 41 - 47.","Constant J., Phauk S. & Bourgoin T. 2016. Updating lanternflies biodiversity knowledge in Cambodia (Hemiptera: Fulgoromorpha: Fulgoridae) by optimizing field work surveys with citizen science involvement through Facebook networking and data access in FLOW website. Belgian Journal of Entomology 37: 1 - 16."]}

Pyrops lathburii (Kirby, 1818) Figs 6–8, 14 Fulgora lathburii Kirby, 1818: 450 (described, compared with Pyrops rogersi (Distant, 1906)) [type in BMNH]. Fulgora astarte Distant, 1914: 409 (described) [type in BMNH]. syn. nov. Fulgora lathburii – Guérin-Méneville 1829: pl. 58 fig. 2 (illustrated). — Latreille 1837: pl. 68 fig. 2 (illustrated). — Spinola 1839: 234 (as “ Fulgora lathburyi ”, treated as a dark form of Pyrops candelaria Linné, 1758 (error!)). — Westwood 1839: 139 (described). — Comte 1840: pl. 68 (listed, illustrated). — Guérin-Méneville 1844: 357 (notes), pl. 58 fig. 2 (illustrated). — White 1844: 425 (recorded from Hong-Kong). — Burmeister 1845: unnumbered page, under genus Fulgora, subgenus Pyrops (as a variety of Fulgora (Pyrops) candelaria (error!)). — Guérin-Méneville 1845: xcvi (as “ Fulgora lothburii ”, close to Pyrops cyanirostris Guérin-Méneville, 1845). — Stål 1870: 741 (as “ Fulgora lathburi ”, compared with Pyrops philippinus Stål, 1870). — Butler 1874: 97 (listed with species with orange posterior hind wings). — Gadeau de Kerville 1881: 43 (listed as “ Fulgora lalhburi ” in “Série I”, species with orange wings, in a census of light-emitting insects). — Atkinson 1885: 129 (described). — Distant 1906: 186 (keyed, nomenclature, described, as “ Fulgora lathburi ”). — Schmidt 1911: 161 (as “ Fulgora lathburi ”, compared with Pyrops peguensis Schmidt, 1911). — Schmidt 1919: 126 (as “ Fulgora lathburi ”, listed from Canton (= Guangzhou)). — Baker 1925: 349 (keyed inside the candelaria species group), 351 (notes), pl. 3 fig. 2 (dorsal habitus), pl. 4 fig. 6 (lateral habitus). — Lallemand 1963: 86 (transferred to the pyrorhynchus species group, keyed, described), pl. xi figs 1–3 (male terminalia). Flata lathburii – Germar 1830: 46 (described). Hotinus lathburii – Amyot & Audinet-Serville 1843: 491 (transferred to Hotinus Amyot & Audinet- Serville). — Walker 1851: 266 (listed, recorded from Silhet (= Sylhet, Bangladesh)). — Dohrn 1859: 57 (listed in catalogue as “ Hotinus lathburi ”). — Girard 1885: 859 (as “ H. lathburi ”, described). Pyrops lathburii – Schaum 1850: 64 (transferred to Pyrops Spinola, 1839). — Nagai & Porion 1996: 25 (catalogued, recorded from Assam and Thailand), figs 194, 196, 198 (dorsal habitus). — Liang 1998: 43 (listed). — Pham 2011: 318 (recorded from Vietnam). — Constant 2015: 9 (note on intraspecific colour variation) — Constant et al. 2016: 15 (in list of species to look for in Cambodia). — Constant & Pham 2017: 18 (host plant). — Wang et al. 2018: 297 (keyed), 301 (nomenclature, measurements, description of male genitalia, diagnostic characters; records in China), figs 18–29 (colour variation, details of head, male terminalia). — Constant 2021: 14 (compared with Pyrops philippinus (Stål, 1870)). — Jiaranaisakul & Constant 2021: 18 (in list of species to look for in Khao Krachom Mountain, Thailand). Laternaria lathburii – Metcalf 1947: 197 (transferred to Laternaria Linné, 1964, catalogued). Fulgora astarte – Distant 1918: 198 (listed from Indochina), 199 (allied to Pyrops vitalisius Distant, 1918). — Lallemand 1963: 85 (placed in the pyrorhynchus species group, keyed, described). Laternaria astarte – Metcalf 1947: 186 (transferred to Laternaria Linné, 1964, catalogued). Pyrops astarte – Nagai & Porion 1996: 25 (transferred to Pyrops, catalogued, recorded from Vietnam and Thailand), figs 195, 197, 199 (dorsal habitus). — Pham & Ta 2004: 58 (in key to Fulgoridae of Vietnam). — Liang 1998: 42 (listed). — Pham 2011: 318 (record from Vietnam). — Constant 2021: 14 (compared with Pyrops philippinus (Stål, 1870)). Note: Wang et al. (2018) erroneously mentioned that the species was attributed to the pyrorhynchus group in Constant & Pham (2017). Material examined Lectotype of Fulgora lathburii Kirby, 1818 (Fig. 6), here designated to provide a stable reference for the taxonomy of the group CHINA • ♀; [China]; [D. Lathbury leg.]; “63 / 41”, “Lathburii K in L. Tr.”, “Kirby.”; BMNH. Note: The specimen was collected in China by D. Lathbury according to Kirby (1818). The code “63 / 41” refers to an entry in the Accession Register in BMNH starting on page 803 of the register and bearing a comment “These insects are the types of Mr Kirby’s Century of Insects [...]” (V. Lemaître pers. com., Jul. 2021). Holotype of Fulgora astarte Distant, 1914 (Fig. 7) VIETNAM • ♀; Indochina, [Lào Cai]; R. Vitalis leg.; “Indo-China (R. Vitalis)”, “ Fulgora astarte Type Dist.”, “Type H.T.”; BMNH. Note: The specimen was collected in Laokay (= Lào Cai) in North Vietnam according to Distant (1914). Additional material LAOS • 1 ♀; Vientiane; [17°58′ N, 102°36′ E]; I.G. 23.993; RBINS. THAILAND • 1 ♀; Phrae Province; May 2004; ex coll. Neef de Sainval; I.G. 30.731; RBINS • 1 ♂, 1 ♀; Chiang Rai Province; Wiang Pa Pao; [19°20′54″ N, 99°30′24″ E]; 10–15 Apr. 2008; ex coll. A. Chaminade; I.G. 31.467; RBINS. VIETNAM • 3 ♂♂, 4 ♀♀; Cao Bang Province, Phia Den; [22°34′26″ N, 105°52′29″ E]; 8 Aug. 2010; J. Constant and P. Limbourg leg.; day time; I.G. 31.668; RBINS • 5 ♂♂, 8 ♀♀; Tay Yen Tu Nature Reserve; 21°11′10″ N, 106°43′25″ E; 7–11 Jul. 2013; J. Constant and J. Bresseel leg.; I.G. 32.454; RBINS • 3 ♂♂; Bac Giang Province, Tay Yen Tu NR, Tram Dong Thong; 3 Jul. 2010; H.T. Pham leg.; VNMN • 1 ♀; [Hai Phong Province], Cat Ba National Park; 20°48′00″ N, 107°00′20″ E; 12–16 Jul. 2013; J. Constant and J. Bresseel leg.; day time; I.G. 32.454; RBINS • 5 ♀♀; [Vinh Phuc Province], Me Linh Biodiversity Station; [21°23′04″ N, 105°42′44″ E]; 20–24 Aug. 2010; J. Constant and P. Limbourg leg.; I.G. 31.668; RBINS • 1 ♀; [Vinh Phuc Province], Me Linh Biodiversity Station; [21°23′04″ N, 105°42′44″ E]; 29 Aug. 2010; J. Constant and P. Limbourg leg.; I.G. 31.668; RBINS • 2 ♂♂, 3 ♀♀; Vinh Phuc Province, Dao Tru, Lap Thach; 17 May 2001; V.T. Hoang leg.; VNMN • 3 ♀♀; Vinh Phuc Province, Me Linh; 24 Apr. 2001; V.T. Hoang leg.; VNMN • 2 ♂♂, 17 ♀♀; Vinh Phuc Province, Me Linh; 26 May 2001; V.T. Hoang leg.; VNMN • 1 ♂, 4 ♀♀; Vinh Phuc Province, Me Linh; 23 Apr. 2007; H.T. Pham leg.; VNMN • 1 ♂, 1 ♀; Vinh Phuc Province, Me Linh; 30–31 May 2007; H.T. Pham leg.; VNMN • 7 ♀♀; Vinh Phuc Province, Me Linh; 26 Jul. 2008; H.T. Pham leg.; VNMN • 2 ♀♀; Vinh Phuc Province, Me Linh; 11 Jun. 2009; H.T. Pham leg.; VNMN • 1 ♂, 10 ♀♀; Vinh Phuc Province, Me Linh; 21 May 2010; H.T. Pham leg.; VNMN • 13 ♀♀; Vinh Phuc Province, Me Linh; 22 May 2010; H.T. Pham leg.; VNMN • 3 ♂♂, 27 ♀♀; Vinh Phuc Province, Me Linh; 4–5 Jun. 2010; H.T. Pham leg.; VNMN • 9 ♀♀; Vinh Phuc Province, Me Linh; 10 Aug. 2011; V.T. Hoang leg.; VNMN • 1 ♀; Vinh Phuc Province, Me Linh; 1 Jun. 2012; V.T. Hoang leg.; VNMN • 8 ♀♀; Vinh Phuc Province, Me Linh; 9–10 Jun. 2012; V.T. Hoang leg.; VNMN • 4 ♀♀; Vinh Phuc Province, Me Linh; 12 Aug. 2011; V.T. Hoang leg.; VNMN • 1 ♀; Vinh Phuc Province, Tam Dao N.P.; May 1998; V.T. Hoang leg.; VNMN • 1 ♀; Vinh Phuc Province, Tam Dao N.P.; Apr. 1999; V.T. Hoang leg.; VNMN • 1 ♀; Vinh Phuc Province, Tam Dao N.P.; May 2003; H.T. Pham leg.; VNMN • 1 ♀; Hanoi, Hoan Kiem; 20 Aug. 2003; V.T. Hoang leg.; VNMN • 2 ♀♀; Thua Thien-Hue Province, Phong Dien, near VNMN station; 16°35′12″ N, 107°20′31″ E; 8–9 Apr. 2017; J. Constant and J. Bresseel leg.; I.G. 33.447; RBINS • 2 ♂♂, 1 ♀; Quang Ninh Province, Dong Son-Ky Thuong nature reserve; 21°11′00.4″ N, 107°07′23.8″ E; 575 m a.s.l.; 26 Jul. 2020; H.T. Pham leg.; VNMN. Remark Intraspecific colour variation of the disc of the hind wings exists in this species, with specimens showing bright yellow hind wings and others, milky white. The colour of the tegmina also varies to a great extent, from very dark forms, nearly black with yellow spots ringed with white, to very pale forms, nearly white with yellow spots or even nearly white with cloudy darker, brownish markings in place of the yellow spots. Conspecificity of the different forms was confirmed by the study of the male genitalia and corroborated by the fact that the different forms are sympatric, as illustrated by a series from Chiang Mai collected in June 1992 in Nagai & Porion (1996: figs 194–199). The comparison of the illustrations of the types specimens (Figs 6A, 7A) with the line illustration of Guérin-Méneville (1829) (Fig. 8) which matches both type specimens is also very convincing. Distribution Northeastern India, southern China, northern Thailand, Laos, North and Central Vietnam. The species is recorded here from Laos for the first time. Checklist of the Vietnamese species of Pyrops Spinola, 1839 Pyrops atroalbus (Distant, 1918) Pyrops buomvoi sp. nov. Pyrops candelaria (Linné, 1758) Pyrops clavatus (Westwood, 1839) Pyrops coelestinus (Stål, 1863) Pyrops condorinus (Lallemand, 1960) Pyrops ducalis (Stål, 1863) Pyrops itoi (Satô & Nagai, 1994) Pyrops lathburii (Kirby, 1818) Pyrops spinolae (Westwood, 1842) Pyrops viridirostris (Westwood, 1848) Pyrops vitalisius (Distant, 1918) Identification key to the species of Pyrops of Vietnam, Laos, Cambodia and Southern Continental China 1. Cephalic process strongly inflated apically (Figs 9–11)................................................................... 2 – Cephalic process not strongly inflated apically (Figs 12–19)........................................................... 3 2. Abdomen black ventrally (Fig. 9B); tegmina pale yellow-white on disc and with 3 black spots in costal area before nodal line (Fig. 9A); cephalic process yellow (Fig. 9).................................................................................................................................................. Pyrops atroalbus (Distant, 1918) – Abdomen red ventrally (Figs 10B, 11B); tegmina largely black on disc (Fig. 10A), or in the pale forms, bluish white on disc and without black spots in costal area (Fig. 11A, F); cephalic process red-brown to black (Figs 10–11)................................................. Pyrops clavatus (Westwood, 1839) 3. Head entirely green, hind wings bright yellow basally or entirely yellow-orange (Figs 12–13)..... 4 – Head not entirely green, hind wings yellow, white or blue basally (Figs 14–19)............................ 5 4. Hind wings bright yellow with apical third black (Fig. 12A); ground colour of distal half of tegmina black (Fig. 12A); anterior and median legs largely black (Fig. 12A–C)............................................................................................................................................ Pyrops viridirostris (Westwood, 1848) – Hind wings entirely bright yellow-orange (Fig. 13A); ground colour of distal half of tegmina green (Fig. 13A); anterior and median legs green (Fig. 13A–C)............ Pyrops itoi (Satô & Nagai, 1994) 5. Cephalic process black or brown-black dorsally, yellowish ventrally (Figs 4–5, 14–15); pro- and mesonotum with median black or dark brown markings (Figs 4–5, 14–15).................................... 6 – Cephalic process yellow-red to red-brown dorsally, paler, yellow to dark red ventrally (Figs 16–19); pronotum unicolourous, mesonotum sometimes with small brown-black markings on sides (Figs 16– 19)..................................................................................................................................................... 8 6. Apex of cephalic process coloured like the rest of the process (Figs 4–5, 15)................................. 7 – Apex of cephalic process yellow to orange-red, strongly contrasting (Figs 6–7, 14).................................................................................................................................... Pyrops lathburii (Kirby, 1818) 7. Tegmina narrower: LTg/BTg = 2.9 (Figs 4A, 5A); spots on basal half of tegmina not fused into bands (Figs 4A, 5A); cephalic process narrower (LPr/BPrH> 9) (Figs 4–5); hind wings bright yellow (Fig. 4A) or white basally (Fig. 5A)...................................... Pyrops condorinus (Lallemand, 1960) – Tegmina broader: LTg/BTg = 2.53 (Fig. 15A); spots on basal half of tegmina fused into transverse bands (Fig. 15A); cephalic process broader (LPr/BPrH Pyrops spinolae (Westwood, 1842) 8. Hind wings white with sutural margin bordered with black up to the anal lobe (Fig. 1A); cephalic process very narrow (LPr/BPrH = 19) (Fig. 1)............................................ Pyrops buomvoi sp. nov. – Hind wings white, bright yellow or blue with only apical third black (Figs 16A, 17A, 18A, 19A); cephalic process broader (LPr/BPrH Pyrops candelaria (Linné, 1758) – Hind wings white or blue basally (Figs 17A, 18A, 19A); cephalic process dark red-brown and dark red ventrally or dark red, orange ventrally and at apex (Figs 17–19); yellow spots on membrane of tegmina (Figs 18–19) or not (Fig. 17)............................................................................................ 10 10. Tegmina without yellow spots on membrane (Fig. 17A); cephalic process dark red, orange ventrally and at apex (Fig. 17); hind wings pale blue to white basally (Fig. 17A); basal half of tegmina whitish with white-margined green spots in transverse rows, sometimes fused in bands (Fig. 17A)............................................................................................... Pyrops ducalis (Stål, 1863) – Tegmina with yellow spots on membrane (Figs 18A, 19A); cephalic process dark red-brown, dark red ventrally (Figs 18–19); hind wings white or blue basally; basal half of tegmina dark green or dark blue-green with yellow spots (Figs 18A, 19A)................................................................................11 11. Hind wings blue basally (sometimes faded to nearly white in collections specimens) (Fig. 18A); cephalic process very elongate and strongly curved dorsad (Fig. 18); ground colour of tegmina dark blue-green (Fig. 18A)....................................................................... Pyrops coelestinus (Stål, 1863) – Hind wings white basally (Fig. 19A); cephalic process very elongate and curved dorsad (Fig. 19); ground colour of tegmina green (Fig. 19A).................................... Pyrops vitalisius (Distant, 1918) Identification key to the species of Pyrops of Vietnam, Laos, Cambodia and Southern Continental China 1. Cephalic process strongly inflated apically (Figs 9–11)................................................................... 2 – Cephalic process not strongly inflated apically (Figs 12–19)........................................................... 3 2. Abdomen black ventrally (Fig. 9B); tegmina pale yellow-white on disc and with 3 black spots in costal area before nodal line (Fig. 9A); cephalic process yellow (Fig. 9).................................................................................................................................................. Pyrops atroalbus (Distant, 1918) – Abdomen red ventrally (Figs 10B, 11B); tegmina largely black on disc (Fig. 10A), or in the pale forms, bluish white on disc and without black spots in costal area (Fig. 11A, F); cephalic process red-brown to black (Figs 10–11)................................................. Pyrops clavatus (Westwood, 1839) 3. Head entirely green, hind wings bright yellow basally or entirely yellow-orange (Figs 12–13)..... 4 – Head not entirely green, hind wings yellow, white or blue basally (Figs 14–19)............................ 5 4. Hind wings bright yellow with apical third black (Fig. 12A); ground colour of distal half of tegmina black (Fig. 12A); anterior and median legs largely black (Fig. 12A–C)............................................................................................................................................ Pyrops viridirostris (Westwood, 1848) – Hind wings entirely bright yellow-orange (Fig. 13A); ground colour of distal half of tegmina green (Fig. 13A); anterior and median legs green (Fig. 13A–C)............ Pyrops itoi (Satô & Nagai, 1994) 5. Cephalic process black or brown-black dorsally, yellowish ventrally (Figs 4–5, 14–15); pro- and mesonotum with median black or dark brown markings (Figs 4–5, 14–15).................................... 6 – Cephalic process yellow-red to red-brown dorsally, paler, yellow to dark red ventrally (Figs 16–19); pronotum unicolourous, mesonotum sometimes with small brown-black markings on sides (Figs 16– 19).........................................................................................................................., Published as part of Constant, Jérôme & Pham, Hong Thai, 2022, The Lanternfly genus Pyrops in Vietnam: A new species from Central Vietnam, taxonomic changes, checklist, identification key (Hemiptera: Fulgoromorpha: Fulgoridae), pp. 123-154 in European Journal of Taxonomy 813 (1) on pages 131-139, DOI: 10.5852/ejt.2022.813.1741, http://zenodo.org/record/6468100, {"references":["Kirby W. F. 1818. A century of insects, including several new genera described from his cabinet. The Transactions of the Linnean Society of London. Second series. Zoology 12 (2): 375 - 453. https: // doi. org / 10.1111 / j. 1095 - 8339.1817. tb 00239. x","Distant W. L. 1906. Rhynchota Vol. 3. Heteroptera-Homoptera. In: C. T. Bingham (ed.) The Fauna of British India, including Ceylon and Burma. Taylor and Francis, London. https: // doi. org / 10.5962 / bhl. title. 48423","Distant W. L. 1914. Some additions to the genera and species in the Homopterous family Fulgoridae. Annals and Magazine of Natural History. London. (Ser. 8) 13: 409 - 424. https: // doi. org / 10.1080 / 00222931408693503","Guerin-Meneville F. E. 1829. Homoptera. In: Cuvier G. L. C. F. D. 1829 Iconographie du regne animal: 58 - 59. Chez J. B. Bailliere, Paris.","Latreille P. A. 1837. The Crustacea, Arachnides & Insecta. In: Cuvier G. L. C. F. D. 1837. The Animal Kingdom, Arranged According to its Organization, Serving as a Foundation for the Natural History of Animals: and an Introduction to Comparative Anatomy. Volume 4. G. Henderson, 1834 - 1837, London. https: // www. biodiversitylibrary. org / item / 89417","Spinola M. 1839. Essai sur les Fulgorelles, sous-tribu de la tribu des Cicadaires, ordre des Rhyngotes. Annales de la Societe Entomologique de France 8: 133 - 337. Available from http: // biodiversitylibrary. org / page / 33710046 [accessed 17 Mar. 2017].","Westwood J. O. 1839. On the family Fulgoridae, with a monograph of the genus Fulgora of Linnaeus. The Transactions of the Linnean Society of London. Second series. Zoology 18: 133 - 153. https: // doi. org / 10.1111 / j. 1095 - 8339.1838. tb 00167. x","Comte J. A. 1840. Regne animal dispose en tableaux methodiques. Fortin Masson et Cie, Paris.","Guerin-Meneville F. E. 1844. Insectes. In: Cuvier G. L. C. F. D. 1844 Iconographie du regne animal: 355 - 370. Chez J. B. Bailliere, Paris.","White A. 1844. Descriptions of some new species of Coleoptera and Homoptera from China. Annals and Magazine of Natural History 14: 422 - 426. https: // doi. org / 10.1080 / 037454809495213","Burmeister H. C. C. 1845. Rhynchota. No. 8. Fam. Fulgorina Gen. Fulgora. In: Burmeister H. C. C. 1838 - 1846 Genera quaedam insectorum. Iconibus illustravit et descripsit Vol. 1. Sumtibus A. Burmeister, Berlin (Germany). https: // doi. org / 10.5962 / bhl. title. 8144","Guerin-Meneville F. E. 1845. (Fulgora cyanirostris. Guer.). Bulletin de la Societe entomologique de France. (Ser. 2) 3: xcvi.","Stal C. 1870. Hemiptera insularum Philippinarum. Bidrag till Philippinska oarnes Hemipter-fauna. Ofversigt af Kongliga Svenska Vetenskaps-Akademiens Forhandlingar. Stockholm 27: 607 - 776. https: // doi. org / 10.5962 / bhl. title. 61898","Butler A. G. 1874. List of the species of Fulgora, with description of new forms in the collections of the British Museum. Proceedings of the Zoological Society of London: 97 - 102. https: // doi. org / 10.1111 / j. 1096 - 3642.1874. tb 02457. x","Gadeau de Kerville H. 1881. Insectes Phosphorescents. Leon Deshays, Rouen.","Atkinson E. T. 1885. Notes on Indian Rhynchota. No. 4. Journal and Proceedings of the Asiatic Society of Bengal. Calcutta 54: 127 - 158.","Schmidt E. 1911. Neue Fulgoriden. Zoologischer Anzeiger 38: 161 - 171.","Schmidt E. 1919. Beitrag zur Kenntnis der Zikadenfauna von Canton (China). Archiv fur Naturgeschichte 85 (7): 121 - 128. Available from https: // www. biodiversitylibrary. org / item / 47756 (accessed 8 Mar. 2022).","Baker C. F. 1925. Remarks on certain Indo-Malayan Fulgora, with special reference to Philippines species. Philippines Journal of Science 28: 343 - 364.","Lallemand V. 1963. Revision des Fulgoridae (Homoptera). Deuxieme partie. Faunes asiatique et australienne. Memoires de l'Institut royal des Sciences naturelles de Belgique (2 e serie) 75: 1 - 99.","Lallemand V. 1960. De quibusdam Fulgoris. Entomologische Mitteilungen aus dem Zoologischen Staatsinstitut und Zoologischen Museum Hamburg 24: 1 - 7.","Germar E. F. 1830. Species Cicadarium enumeratae et sub genera distributae. Thon's Entomologisches Archiv 2 (2): 1 - 57.","Amyot C. & Audinet-Serville J. 1843. Deuxieme partie. Homopteres. Homoptera Latr. Histoire naturelle des Insectes. Hemipteres. Librairie encyclopedique de Roret, Paris. https: // doi. org / 10.5962 / bhl. title. 8471","Walker F. 1851. List of the Specimens of Homopterous Insects in the Collection of the British Museum. Vol. 2: 261 - 636. Trustees of the British Museum (Natural History), London.","Dohrn F. A. 1859. Homoptera. Catalogus Hemipterorum. Herausgegeben von dem entomologischen Vereine zu Stettin 1859: 56 - 93.","Girard M. 1885. Traite elementaire d'Entomologie. Vol. 3. J. - B. Bailliere, Paris.","Schaum H. R. 1850. Fulgorellae. Erster Section A - G. In: Ersch I. S. & Gruber I. G. (eds) Allgemeine Encyklopadie der Wissenschaften und Kunste in alnhaberischen folge von Genannten Schriftstellern bearbeitet und herausgegeben 51: 58 - 73. Brockhaus Verlag, Mannheim.","Nagai S. & Porion T. 1996. Fulgoridae 2: Catalogue illustre des Faunes asiatique et australienne. Sciences Nat, Compiegne.","Liang A. P. 1998. Nomenclatorial notes on the Oriental lantern fly genus Pyrops Spinola (Hemiptera: Fulgoroidea: Fulgoridae). Acta Zootaxonomica Sinica 23 (1): 41 - 47.","Pham H. T. 2011. A checklist of the family Fulgoridae (Homoptera: Auchenorrhyncha: Fulgoroidea) from Vietnam. In: Proceedings of the 3 rd National Scientific Conference on Ecology and Biological Resources Hanoi, 22 October 2009: 317 - 321. [In Vietnamese.]","Constant J. 2015. Review of the effusus group of the Lanternfly genus Pyrops Spinola, 1839, with one new species and notes on trophobiosis (Hemiptera: Fulgoromorpha: Fulgoridae). European Journal of Taxonomy 128: 1 - 23. https: // doi. org / 10.5852 / ejt. 2015.128","Constant J., Phauk S. & Bourgoin T. 2016. Updating lanternflies biodiversity knowledge in Cambodia (Hemiptera: Fulgoromorpha: Fulgoridae) by optimizing field work surveys with citizen science involvement through Facebook networking and data access in FLOW website. Belgian Journal of Entomology 37: 1 - 16.","Constant J. & Pham H. T. 2017. Review of the clavatus group of the lanternfly genus Pyrops (Hemiptera: Fulgoromorpha: Fulgoridae). European Journal of Taxonomy, 305: 1 - 26. https: // doi. org / 10.5852 / ejt. 2017.305","Wang W. Q., Xu S. - L. & Qin D. - Z. 2018. The lanternfly genus Pyrops Spinola (Hemiptera: Fulgoridae) from China with description of a new species. Entomotaxonomia 40 (4): 296 - 309. https: // doi. org / 10.11680 / entomotax. 2018031","Jiaranaisakul K. & Constant J. 2021. The lanternflies (Hemiptera: Fulgoromorpha, Fulgoridae) of Khao Krachom Mountain, Thailand. Far Eastern Entomologist 435: 7 - 19. https: // doi. org / 10.25221 / fee. 435.2","Metcalf Z. P. 1947. General Catalogue of the Homoptera. Fascicle IV Fulgoroidea. Part 9 Fulgoridae. North Carolina State College, Raleigh (USA).","Distant W. L. 1918. The Homoptera of Indo-China. Annals and Magazine of Natural History (Ser. 9) 1: 196 - 200. https: // doi. org / 10.1080 / 00222931808562301","Pham H. T. & Ta H. T. 2004. Key to the species of the family Fulgoridae (Homoptera: Auchenorrhyncha: Fulgoroidea) in Vietnam. Journal of Biology 26 (3 A): 57 - 60. [In Vietnamese.]"]}

Pyrops connectens (Atkinson, 1885) Fig. 2 A-B MATERIAL EXAMINED (from photograph). Khao Krachom Mountain: dry evergreen forest, 8.IV 2013, 4 ex., photo S. Ruchisansakul. REMARKS. This species was described from Myanmar (Distant, 1906) and here recorded from Thailand for the first time., Published as part of Jiaranaisakul, K. & Constant, J., 2021, The Lanternflies of Khao Krachom Mountain, Ratchaburi Province, Thailand (Hemiptera: Fulgoromorpha: Fulgoridae), pp. 7-19 in Far Eastern Entomologist 435 on page 13, DOI: 10.25221/fee.435.2, http://zenodo.org/record/7166124

Two new species of Fulgoridae collected in Philippines are described: Egregia palawanica n. sp. from Palawan and Pyrops kozlovi n. sp.from Mindanao., Deux nouvelles espèces de Fulgoridae collectées aux Philippines sont décrites : Egregia palawanica n. sp. de Palawan, et Pyrops kozlovi n. sp. de Mindanao.

Pyrops atroalbus (Distant, 1918) comb. nov. Figs 1, 5–6, 9A Fulgora atroalba Distant, 1918: 199 (type in BMNH). Laternaria atroalba – Metcalf 1947: 187 [transferred to Laternaria; catalogued]. Fulgora watanabei apicalis – Lallemand 1963: 90 [considered as a junior synonym of Fulgora watanabei atroalba (Distant, 1918) (erroneous)]. Fulgora watanabei atroalba – Lallemand 1963: 90 [transferred back to Fulgora and considered as a subspecies of P. watanabei (erroneous)]. Fulgora watanabei var. formosana – Lallemand 1963: 90 [considered as a junior synonym of Fulgora watanabei atroalba (Distant, 1918) (erroneous)]. Pyrops watanabei apicalis – Nagai & Porion 1996: 26 [transferred to Pyrops but still considered as a junior synonym of Fulgora watanabei atroalba (erroneous)]. Pyrops watanabei atroalba – Nagai & Porion 1996: 26, pl. 18, fig. 223 [transferred to Pyrops, but still considered as a subspecies of P. watanabei (erroneous), type illustrated]. — Liang 1998: 22 [considered as a subspecies of P. watanabei (erroneous)]. Pyrops watanabei formosana – Nagai & Porion 1996: 26 [transferred to Pyrops but still considered as a junior synonym of Fulgora watanabei atroalba (erroneous)]. Non Fulgora watanabei apicalis Kato, 1928: 221, pl. 9 fig. 1. Non Fulgora watanabei var. formosana Metcalf, 1947: 208. Note Liang (1998) erroneously stated that the name “ Pyrops watanabei atroalbus (Distant, 1918) ” was a new combination he proposed while the combination had already been proposed by Nagai and Porion (1996). Diagnosis The species is immediately recognized by the following combination of characters: (1) Cephalic process yellow and strongly inflated apically (Fig. 1 D–F). (2) Abdomen black ventrally (Fig. 1B). (3) Tegmina strongly contrasted: pale yellow-white with black markings, including 3 black spots in costal area before nodal line (Fig. 1A). Etymology From Latin ater (adj.) ‘black’ and albus (adj.): ‘white’; the species epithet refers to the black and white colouration of the species. Type material Holotype (Fig. 5) VIETNAM: ♂ [Indo-China, Tonkin, R.V. de Salvaza. 1917–98] [1918–1] [Fulgora atroalba Dist. Type] [Type] (BMNH). Additional material VIETNAM: 1 ♂, 1 ♀, Da Krong Nature Reserve, Quang Tri Province, 10 Jul. 2011, 16°37′ N, 106°47′ E, day collecting, Pham & Hoang (♂ in RBINS, ♀ in VNMN); 1 ♀, Bach Ma National Park, 29 Sep. 2014, 16°12′ N, 107°52′ E, Tuan (VNMN). Material examined from photograph THAILAND: 1 ex. (Fig. 9A), Chiang Mai Prov., Maerim District, 18°54′50″ N, 98°56′42″ E, 14 Mar. 2016, Panaka Jirasuttayaporn. Measurements and ratios TL: ♂ (n = 1): 3.4 cm; ♀ (n = 1): 4.2 cm; LPr: ♂: 1.1 cm; ♀: 1.4 cm; LTg/BTg = 2.55; BF/BPrH = 3.1; LPr/LF = 2.95; LPr/BPrH = 8.8. Male genitalia Pygofer higher than long, with posterior margin regularly rounded dorsally in lateral view (Fig. 6A). Anal tube slightly elongate, 1.1 times as long as broad in dorsal view, broader at 4/5 of total length (Fig. 6C); lateral margins very slightly sinuate and apical margin strongly concave in dorsal view (Fig. 6C). Gonostyli (Fig. 6A) elongate, twice as long as high in lateral view; dorsal margin regularly rounded and posterior margin slightly projecting posteriorly in middle in lateral view (Fig. 6A). Remarks There is a discrepancy between the location given on the label of the specimen (“ Tonkin ”) and the one given in the original description (Distant 1918): “ Indochina, Xieng Klouang (R. Vitalis de Salvaza) ”. The latter location is situated in Laos, not far from the places where we have collected specimens in Central Vietnam. Despite our intensive collecting effort in northern Vietnam, we have never found the species in that region. Hence, it seems that the location given by Distant (1918) is more likely to be the correct one. Nagai & Porion (1996: fig. 223) erroneously gave “central Taiwan ” as the type locality. Distribution The species is known from one location in Laos, one in Northern Thailand, and two in Central Vietnam.

Pyrops clavatus (Westwood, 1839) Figs 2���3, 7, 9 B���G, 10���15 Fulgora clavata Westwood, 1839: 139, pl. 12, fig. 1 (types in OUMNH) [described, illustrated, compared with P. pyrorhynchus (Donovan, 1800), the latter mentioned as the junior synonym Fulgora pyrorhina Westwood, 1839)]. Hotinus ponderosus St��l, 1854: 244 (type in NHRS) [described and mentioned as close to P. clavatus; synonymized by Distant (1906)]. Fulgora woodii Ollenbach, 1929: 279, pl. 1, fig. 13 (types in NFIC) [described, illustrated and mentioned as very near to Pyrops clavatus; synonymized by Lallemand (1963)]. Fulgora nigripennis Chou & Wang in Chou et al., 1985: 33, fig. 4 (type in NWAFU) [described in Chinese, illustrated, compared to P. clavatus], 37 (briefly described in Esperanto, host plant) syn. nov. Fulgora clavata mizunumai Sat�� & Nagai, 1994: 312, figs 3, 12 (type in EUM) [described, illustrated, compared to P. clavatus] syn. nov. Pyrops clavata ��� Burmeister 1845: 4 [transferred to Pyrops]. ��� Schaum 1850: 64 [listed] ��� Kirby 1885: 211 [mentioned from India and briefly described]; 1892: 211 [idem]. ��� Nagai & Porion 1996: 26 [catalogued; distribution]; 31 [very close to Fulgora nigripennis Chou & Wang, 1985], pl. 18, figs 227���228 [illustrated]. Hotinus clavatus ��� Adams 1847: 204 [compared with P. sultanus (Adams & White, 1847)]. ��� Walker 1851: 267 [list of specimens in BMNH]; 1858: 41 [clavatus as the type and single member of a group within Hotinus]. ��� St��l 1854: 244 [close to Hotinus ponderosus St��l, 1854]. ��� Costa 1864: 82 [listed from Assam]. Fulgora (Hotina [sic !]) clavata ��� Westwood 1848: 7, pl. 3, fig. 1 [described, illustrated]. Hotinus ponderosus ��� Walker 1858: 315 [listed from Hindustan]. ��� Distant 1906: 191 [junior synonym of P. clavatus (Westwood, 1839)]. Fulgora clavata ��� Butler 1874: 98 [placed in a section of Fulgora with white hind wings; list of specimens in BMNH; mention of ��� ludicrous ��� cephalic process; probable senior synonym of ��� Fulgora ponderosa ���]. ��� Gadeau de Kerville 1881: 43 [listed as light-producing insect (erroneous)]. ��� Atkinson 1885: 130 [catalogued; described; distribution; intraspecific variation]. ��� Schmidt 1905: 354 [catalogued]. ��� Distant 1906: 191; fig. 83 [keyed; described; habitus, side of head and frons illustrated; senior synonym of Hotinus ponderosus St��l, 1854]. ��� Schumacher 1915: 129 [compared with Fulgora chimara Schumacher, 1915]. ��� Distant 1918: 198 [listed from ���Indochina���], 200 [compared with P. atroalbus (Distant, 1918)]. ��� Paiva 1919: 373 [mentioned from Garo Hills, N India; notes on biology]. ��� Baker 1925: 348 [type of the clavata group in Fulgora], 361 [key to the species of the clavata group; described], pl. 4, fig. 1 [lateral aspect of habitus illustrated]. ��� Ollenbach 1929: 280 [compared with Fulgora woodi Ollenbach, 1929]. ��� Lallemand 1963: 71 [type of 5 th group of Fulgora], 88 [keyed; described; catalogued; senior synonym of Hotinus ponderosus St��l, 1854 and Fulgora woodi Ollenbach, 1929]; pl. 10, figs 4���7 [lateral view of head and male genitalia of type illustrated]. ��� Sat�� & Nagai 1994: 312 [compared with Fulgora clavata mizunumai Sat�� & Nagai, 1994]. Fulgora ponderosa ��� Butler 1874: 98 [probable junior synonym of P. clavatus]. ��� Gadeau de Kerville 1881: 43 [listed as light-producing insect (erroneous)]. ��� Atkinson 1885: 131 [catalogued; described; distribution; probable variety of P. clavatus]. Fulgora ponderosus ��� Matsumura 1913: 54 [close to P. watanabei (Matsumura, 1913)]. Laternaria clavata ��� Metcalf 1947: 193 [catalogued; distribution]. ��� Allnatt 2013: 45, fig. 1 [illustrated from Assam]. Laternaria clavata var. ponderosa ��� Metcalf 1947: 193 [catalogued; distribution; considered as a variety of clavatus (erroneous)]. Laternaria woodi ��� Metcalf 1947: 208 [catalogued]. Pyrops clavata mizunumai ��� Nagai & Porion 1996: 26 [catalogued]; pl. 18, figs 222, 224, 226 [illustrated]. Pyrops clavatus ��� Liang 1998: 42 [catalogued; new combination (erroneous)]. Pyrops clavatus mizunumai ��� Liang 1998: 42 [catalogued; new combination (erroneous)]. Diagnosis The species is immediately recognized by the following combination of characters: (1) Cephalic process red-brown to black, often black with apex red-brown, and strongly inflated apically (Figs 2 D���G, 3C, E). (2) Abdomen red ventrally (Figs 2B, 3B). (3) Tegmina largely black on disc in the dark forms (Fig. 1 A���B); in the pale forms, tegmina bluish white on disc without black spots in costal area (Fig. 3 A���B, D). Etymology clavatus (adj., Latin): clavate. The name refers to the shape of the cephalic process. Material examined Type material BANGLADESH: Lectotype, ♂ of Fulgora clavata Westwood, 1839, here designated to provide a reference standard for the species (examined from photographs, Fig. 10) [Silhet] [Fulgora clavata Westw. Trans. Lin. Soc. 18. P. 139 Pl 12 Fig 5.] [Type] [Type Hem: 596 1/2 Fulgora clavata Westwood, Hope Dept. Oxford] (OUMNH). BANGLADESH: Paralectotype, ♀ of Fulgora clavata Westwood, 1839 (examined from photographs, Fig. 11) [Silhet] [Fulgora clavata Westw. Monogr. M. Hope, Sylhet] [W] [Type] [Type Hem: 596 2/2 Fulgora clavata Westwood, Hope Dept. Oxford] (OUMNH). INDIA: Holotype, ♀ of Hotinus ponderosus St��l, 1854 (Fig. 12) [Ind or] [Saund] [ponderosus St��l] [NHRS-HEMI000000198] (NHRS). MYANMAR: Syntype, ♂ of Fulgora woodii Ollenbach, 1929 (examined from photographs, Fig. 13) [Paga-yo, Tavoy, 27.I.20, O.C. Ollenbach] [Type] [Fulgora woodii (Ollen) ♂ n sp., O.C. Ollenbach det.] (NFRI). CHINA: Holotype, ♂ of Fulgora nigripennis Chou & Wang, 1985 (examined from photographs, Fig. 14) [Fulgora nigripennis Chou, Wang & Huang ��� identified by Io Chou, August 1982 ���] [��� Host plant: coffee ���] [HOLOTYPE] (NWAFU). ��� Parts in italics between ��� ��� were translated from Chinese by D. Qin. The specimen was collected in Ruyuan, Guangdong Province in May 1975 by Lizhong Hua (D. Qin pers. comm., Jun. 2016). THAILAND: Holotype, ♂ of Fulgora clavata mizunumai Sat�� & Nagai, 1994 (examined from photographs, Fig. 15) [Doi Pui, Chiang Mai (N Thailand) May 1987] [Holotype Fulgora clavata mizunumai] (EUM). Additional material INDIA: 1 ♀, Darjeeling (Himalaya), 1869, Higgins (RBINS); 1 ♀, Kurseong, no date, R.P. Wery (RBINS). THAILAND: 5 ♂♂, 15 ♀♀, Chiang Mai, Dec. 2007, local collectors (RBINS). VIETNAM: 1 ♀, Ha Giang, Jul. 2009, local collectors (RBINS); 2 ♀♀, Cuc Phuong National Park, 20��19���00��� N, 105��36���30��� E, 19���23 Jul. 2011, day collecting, on tree trunk, leg. J. Constant and J. Bresseel (RBINS); 2 ♀♀, Cuc Phuong N.P., 25 May 2005, leg. H.T. Pham (VNMN); 3 ♂♂, 3 ♀♀, Lao Cai Prov., Sa Pa Mt, 1800 m, Oct. 2014, leg. Than Le Luong (RBINS); 1 ♀, Vinh Phuc Prov., Me Linh Station, 12 Aug. 2011, leg. H.T. Pham (VNMN); 3 ♂♂, 10 ♀♀, Ba Vi N.P., 21��4���4��� N, 105��21���30��� E, 25���29 Jun. 2015, 1000 m, leg. H.T. Pham (VNMN); 1 ♂, 2 ♀♀, Ba Vi N.P., 21��4���4��� N, 105��21���30��� E, 25���29 Jun. 2015, 600 m, leg. J. Constant and J. Bresseel (RBINS); 1 ♀, Nghe An Prov., Pu Mat N.P., 18��59��� N, 104��40��� E, Jul. 2014, local collector (RBINS); 1 ♀, Tuy��n Quang Prov., Cham Chu Nature Reserve, 22��12��� N, 105��6��� E, 8���12 Jul. 2015, leg. J. Constant and J. Bresseel (RBINS); 2 ♂♂, same data, leg. H.T. Pham (VNMN); 1 ♀, Quang Binh Prov., Phong Nha-Ke Bang N.P., U Bo, 17��32���14��� N, 106��9���4��� E, 550 m, leg. H.T. Pham (VNMN); 3 ♂♂, 3 ♀♀, Son La Prov., Copia Nature Reserve, 21��22���12��� N, 103��30���42��� E, 20���23 Jul. 2016, J. Constant and J. Bresseel (RBINS). Material examined from photographs INDIA: 1 ex., Assam, Mangaldai, 13 Aug. 2014, Mirza Galib (Facebook group: InsectIndia); 1 ex., Manipur, Loktak Lake, 9 Oct. 2014, Sougrakpam Neli (Facebook group: InsectIndia); 1 ex., Nagaland, Aug. 2012, Tarun Karmakar (Facebook group: InsectIndia); 1 ex., Upper Assam, Dibrugarh, Rungagora, no date, H. Stevens (Alnatt 2013: fig. 1). VIETNAM: 1 ex. (Fig. 9 B���C), Me Linh Biological Station, 12 Jul. 2015, on a trunk of Dimocarpus longan (Sapindaceae), J. Constant. Additional data from literature INDIA: Garo Hills above Tura, 3000 ft (Paiva 1919). MYANMAR: Karen Hills (Ollenbach 1929 ��� location of the second syntype of Fulgora woodi). Measurements and ratios TL: ♂ (n = 5): 3.9 cm (3.7���4.0); LPr: 1.2 cm (1.1���1.3); TL: ♀ (n = 5): 5.1 cm (4.5���5.5); LPr: 1.35 cm (1.3���1.5); LTg/BTg = 2.47; BF/BPrH = 2.05; LPr/LF = 3.19; LPr/BPrH = 6.13. Male genitalia Pygofer higher than long, with posterior margin angularly bisinuate dorsally in lateral view (Fig. 7A). Anal tube slightly elongate, 1.1 times as long as broad in dorsal view, broader at 4/5 of total length (Fig. 7C); lateral margins very slightly sinuate and apical margin strongly concave in dorsal view (Fig. 7C). Gonostyli (Fig. 7A) elongate, 1.5 times as long as broad in lateral view; dorsal margin strongly rounded above lateral tooth and posterior margin rounded in lateral view (Fig. 7A). Remarks After examination of the photographs of the type specimens of Fulgora nigripennis Chou & Wang, 1985 and Fulgora woodii Ollenbach, 1929, it was not possible to find any difference between those taxa and the types of P. clavatus and the recorded intraspecific variations of the species. Hence, the first is proposed as a junior synonym of Pyrops clavatus and the synonymy of the second under P. clavatus, as proposed by Lallemand (1963) and followed by Nagai & Porion (1996), is confirmed. Ollenbach (1929) stated that the types of the species described in his paper would later be sent to the British Museum (currently BMNH) but he apparently never did so as none of the types of those species can be found in the BMNH collections (M. Webb pers. comm., 21 May 2013). The examination of numerous specimens of P. clavatus, including large series from Chiang Mai (Figs 2���3) and northern Vietnam, proved that the species shows important intraspecific colour variation of the cephalic process and wings: the tegmina vary from nearly completely black to nearly completely bluish white, the hind wings from white with black apex to completely white, usually tinged with blue or violet basally. The ���subspecies��� mizunumai described by Sat�� & Nagai (1994) only represents the paler extreme of the species and cannot be considered as a subspecies, as it occurs sympatrically with the intermediate and darker forms. It is therefore synonymized under P. clavatus. Specimens showing basally blue-tinged and violet-tinged hind wings were found in the same population, on the same tree in Copia, North Vietnam. The variation is not linked to the sex of the specimens as both males and females showed the two variations. Biology Paiva (1919) stated that the species was ��� not uncommon at 3000 ft. Several specimens sometimes found on a single tree to which they return after disturbance ���. Our observations confirm this statement which is valid for most species of Pyrops we have observed so far. We have found P. clavatus feeding on several unidentified species of tree in Ba Vi National Park (north Vietnam); in Cuc Phuong National Park (north Vietnam), some specimens of P. clavatus were found on a big tree together with numerous Pyrops spinolae (Westwood, 1842), while other trees of the same species and others around did not host any Pyrops specimens (Fig. 9 F���G); at Me Linh Biodiversity Station (north Vietnam), P. clavatus was observed on a big Longan tree trunk (Dimocarpus longan Lour., Sapindaceae) together with P. candelaria (Linnaeus, 1758), P. lathburii (Kirby, 1818) and P. viridirostris (Westwood, 1848). In North Vietnam we found the species at altitudes ranging from 150 to 1200 m. The species was also collected on coffee (Coffea sp., Rubiaceae) in southern China. Distribution Known from N India, Myanmar, N Thailand, S China and N Vietnam. Very probably also present in Laos and maybe in Cambodia (see also Constant et al. 2016 for discussion on Fulgoridae from Cambodia)., Published as part of Constant, J��r��me & Pham, Hong-Thai, 2017, Review of the clavatus group of the lanternfly genus Pyrops (Hemiptera: Fulgoromorpha: Fulgoridae), pp. 1-26 in European Journal of Taxonomy 305 on pages 10-18, DOI: 10.5852/ejt.2017.305, http://zenodo.org/record/3827135, {"references":["Westwood J. O. 1839. On the family Fulgoridae, with a monograph of the genus Fulgora of Linnaeus. The Transactions of the Linnean Society of London, Second series, Zoology 18: 133 - 153.","Stal C. 1854. Nya Hemiptera. Ofversigt af Kongliga Svenska Vetenskaps-Akademiens Forhandlingar 11: 231 - 255.","Distant W. L. 1906. The Fauna of British India, including Ceylon and Burma. Vol. 3. C. T. Birgham. Taylor & Francis, London. https: // doi. org / 10.5962 / bhl. title. 48423","Ollenbach O. C. 1929. Part 3. New species of Cicadidae and Fulgoridae from India and Burma (Hemip.). Indian Forest Records 13 (6): 271 - 282.","Lallemand V. 1963. Revision des Fulgoridae (Homoptera). Deuxieme partie. Faunes asiatique et australienne. Memoires de l'Institut royal des Sciences naturelles de Belgique (2 e serie) 75: 1 - 99.","Chou I., Wang S. & Huang J. 1985. Priskribo de novaj specioj de Fulgoroedoj el Cinio (Homopteroj: Fulgoroedoj). Entomotaxonomia 7 (1): 29 - 38.","Sato M. & Nagai S. 1994. Taxonomic notes on the Southeast Asian species of the genus Fulgora (Homoptera: Fulgoridae). Transactions of the Shikoku Entomological Society 20 (3 - 4): 307 - 314.","Burmeister H. C. C. 1845. Rhynchota. No. 8. Fam. Fulgorina Gen. Fulgora. In: Burmeister H. C. C. 1838 - 1846. Genera Quaedam Insectorum. Iconibus Illustravit et Descripsit 1. Sumtibus. A. Burmeister, Berlin.","Schaum H. R. 1850. Fulgorellae. Erster Section A-G. In: Ersch I. S. & Gruber I. G. (eds) Allgemeine Encyklopadie der Wissenschaften und Kunste in alphabetischer Folge von genannten Schriftstellern bearbeitet und herausgegeben. Brockhaus Verlag, Mannheim.","Kirby W. F. 1885. Elementary Text-book of Entomology. W. Swan Sonnenschein & Co., London https: // doi. org / 10.5962 / bhl. title. 29217","Nagai S. & Porion T. 1996. Fulgoridae 2: Catalogue illustre des Faunes asiatique et australienne. Sciences Nat, Compiegne.","Adams R. N. 1847. Description of a new species of Fulgora. The Annals and Magazine of Natural History 20: 204 - 205.","Walker F. 1851. List of the Specimens of Homopterous Insects in the Collection of the British Museum. Vol. 2: 261 - 636. British Museum, London.","Costa A. 1864. Emitteri in Acquisti fatti durante l'anno 1862. Annuario del Museo Zoologico della R. Universita di Napoli. Naples 2: 8 - 93. Available from http: // biodiversitylibrary. org / page / 11823135 [accessed 20 Mar. 2017].","Westwood J. O. 1848. The Cabinet of Oriental Entomology. Vol. 4. W. Smith, London.","Distant W. L. 1918. The Homoptera of Indo-China. Annals and Magazine of Natural History (Ser. 9) 1: 196 - 200.","Matsumura S. 1913. Thousand Insects of Japan Additamenta. Vol. 1. Keiseisha, Tokyo [in Japanese].","Walker F. 1858. Supplement. List of the Specimens of Homopterous Insects in the Collection of the British Museum. British Museum, London.","Butler A. G. 1874. List of the species of Fulgora, with description of new forms in the collections of the British Museum. Proceedings of the Zoological Society of London 42 (1): 97 - 102. https: // doi. org / 10.1111 / j. 1096 - 3642.1874. tb 02457. x","Gadeau de Kerville H. 1881. Insectes Phosphorescents. Leon Deshays, Rouen.","Atkinson E. T. 1885. Notes on Indian Rhynchota. No. 4. Journal and Proceedings of the Asiatic Society of Bengal 54: 127 - 158.","Schmidt E. 1905. Beitrag zur Kenntnis der Fulgoriden. II. Die im Stettiner Museum vorhandenen Arten des Genus Fulgora Linne. Entomologische Zeitung, herausgegeben von dem entomologischen Vereine zu Stettin 66: 342 - 357.","Schumacher F. 1915. Homoptera in H. Sauter's Formosa-Ausbeute. Supplementa Entomologica 4: 108 - 142.","Paiva C. A. 1919. XXIII. Rhynchota from the Garo Hills, Assam. Records of the Indian Museum 16: 349 - 377. Available from http: // biodiversitylibrary. org / page / 11132445 [accessed 23 Feb. 2017].","Baker C. F. 1925. Remarks on certain Indo-Malayan Fulgora, with special reference to Philippine species. Philippine Journal of Science 28: 343 - 364.","Metcalf Z. P. 1947. General Catalogue of the Homoptera. Fascicle IV Fulgoroidea. Part 9 Fulgoridae. North Carolina State College, Raleigh (U. S. A.).","Allnatt G. 2013. Recuration of the Fulgoridae collection at the Manchester Museum. Journal of Natural Science Collections 1: 44 - 47.","Liang A. - P. 1998. Nomenclatorial notes on the Oriental lanternfly genus Pyrops Spinola (Hemiptera: Fulgoroidea: Fulgoridae). Acta Zootaxonomica Sinica 23 (1): 41 - 47.","Constant J., Phauk S. & Bourgoin T. 2016. Updating lanternflies biodiversity knowledge in Cambodia (Hemiptera: Fulgoromorpha: Fulgoridae) by optimizing field work surveys with citizen science involvement through Facebook networking and data access in FLOW website. Belgian Journal of Entomology 37: 1 - 16."]}

Pyrops watanabei (Matsumura, 1913) Figs 4, 8, 16–22 Fulgora (Hotinus) watanabei Matsumura, 1913: 54 [described; close to “ Fulgora ponderosus ” (Stål, 1854)], pl. 8, fig. 1 [habitus illustrated] (types in HUIC). Fulgora chimara Schumacher, 1915a: 129 [described] (Types in MFNB and SDEI) [synonymized by Kato (1928); reinstated as good species by Metcalf (1947); re-synonymized by Lallemand (1963)]. Fulgora watanabei var. apicalis Kato, 1928: 221 [described; host plant], pl. 9, fig. 1 [habitus illustrated] (type in UMUT). syn. nov. [name preoccupied by Fulgora apicalis Westwood, 1838, replaced by F. watanabei var. formosana by Metcalf (1947)]. Fulgora watanabei var. formosana Metcalf, 1947: 208 [replacement name for F. watanabei var. apicalis]. syn. nov. Fulgora chimara – Schumacher 1915b: 110 [listed from “ Formosa ” (= Taiwan)]. — Kato 1928: 221 [synonymized under Pyrops watanabei (Matsumura, 1913)]. — Metcalf 1947: 193 [catalogued as a good species]. — Lallemand 1963: 89 [synonymized under Pyrops watanabei (Matsumura, 1913)]. Fulgora watanabei – Kato 1928: 221 [listed; senior synonym of Fulgora chimara Schumacher, 1915; host plant]. — Mutsumura 1931: 1267 [described; illustrated]. — Chou et al. 1985: 118 [described; mentioned from Taiwan and China (Guangxi, Guangdong and Hainan)]. Hotinus watanabei – Matsumura 1931: pl. 8, fig. 12 [illustrated]. Pyrops watanabei – Nagai & Porion 1996: 26 [catalogued; mentioned from China (erroneous)]. — Liang & Suwa 1998: 157 [lectotype designation]. — Yen & Yang 2001 [description; biology; distribution; protection status]. Diagnosis The species is immediately recognized by the following combination of characters: (1) cephalic process yellow and inflated apically (Fig. 4 D–E). (2) abdomen red ventrally (Fig. 4B). (3) tegmina mainly white on disc and with 3 black spots in costal area before nodal line (Fig. 4A). Etymology The species was dedicated to its collector, Kenji Watanabe. Type material TAIWAN: Lectotype, ♀ of Fulgora (Hotinus) watanabei Matsumura, 1913 (examined from photographs, Fig. 16) [Hotinus watanabei Mats.] [Hotinus watanabei Mats.] [Hotinus watanabei det Matsumura] [Formosa Matsumura/ underside: Hoppo, 1 st VII ‘07] [Type Matsumura] [Lectotype Fulgora (Hotinus) watanabei Mats. det. A.P. Liang & M. Suwa 1997] (HUIC). TAIWAN: Paralectotype, ♀ of Fulgora (Hotinus) watanabei Matsumura, 1913 (examined from photographs, Fig. 17) [Hoppo] [Paralectotype Fulgora (Hotinus) watanabei Mats. det. A.P. Liang & M. Suwa 1997] (HUIC). TAIWAN: Paralectotype, ♀ of Fulgora (Hotinus) watanabei Matsumura, 1913 (examined from photographs, Fig. 18) [Formosa Matsumura / underside: Hoppo, 27 VII ‘07] [Paralectotype Fulgora (Hotinus) watanabei Mats. det. A.P. Liang & M. Suwa 1997] (HUIC). TAIWAN: Syntype, ♀ of Fulgora chimara Schumacher, 1915 (Fig. 19) [Formosa, Hoozan, VIII.10, H. Sauter S.G.] [Fulgora chimara * Schum., F. Schumacher det.] [Type] (MFNB). TAIWAN: Syntype, ♀ of Fulgora chimara Schumacher, 1915 (examined from photographs, Fig. 20) [Kosempo, Formosa, H. Sauter, VII. X.1911] [Fulgora chimara * Schum., F. Schumacher det.] [Syntypus] (SDEI). TAIWAN: Holotype, ♀ of Fulgora watanabei var. apicalis Kato, 1928 (examined from photographs, Fig. 21) [Horisha, Formosa (VIII.1919) Col. M. Kato.] [Type No. 95, M. Kato coll.] (UMUT). Additional material TAIWAN: 1 ♀, Taipei city, Congguan Nat. Taiwan University Campus, 2007, leg. W.B. Yeh (RBINS); 1 ♂ (dissected), 1 ♀, Taipei, Tanshui, 14 Jun. 2002, J.H. Chen (NMNS). Material examined from photographs TAIWAN: eggs (Fig. 22A), Taipei City, 28 Jul. 2012, S. Chen; eggs (Fig. 22B), idem, 19 Feb. 2016; 1 nymph (Fig. 22C), Nantou County, 14 Feb. 2012, S. Chen; 1 nymph (Fig. 22D): Taipei City, 7 Jun. 2012, on Triadica sebifera, S. Chen; 1 ex. (Fig. 22 E–F), idem, 16 Aug. 2012, on Triadica sebifera, S. Chen; 1 ex. (Fig. 22G), idem, predated by a Sparassidae spider; 5 ex. (Fig. 22H), Taipei City, 6 Aug. 2011, on Triadica sebifera, S. Chen. Measurements and ratios TL: ♂ (n=1): 3.9 cm; LPr: 1.0 cm; TL: ♀ (n=2): 4.3 cm (4.0–4.6); LPr: 1.1 cm (1.0–1.2); LTg/BTg = 2.31; BF/BPrH = 2.53; LPr/LF = 2.82; LPr/BPrH = 2.55. Male genitalia Pygofer higher than long, with posterior margin sinuate in lateral view (Fig. 8A). Anal tube slightly elongate, 1.1 times as long as broad in dorsal view, broader at 4/5 of total length (Fig. 8C); lateral margins slightly sinuate and apical margin strongly concave in dorsal view (Fig. 8C). Gonostyli (Fig. 8A) elongate, 1.77 times as long as high in lateral view; dorsal margin regularly and broadly rounded and posterior margin slightly projecting posteriorly in middle in lateral view (Fig. 8A). Biology The species was recorded on Triadica sebifera (L.) Small (Euphorbiaceae) by Kato (1928), and on the same tree and Sapium discolor Muell. -Arg. (Euphorbiaceae) by Yen & Yang (2015). Those data were confirmed by multiple observations in the field; freshly laid eggs were observed at the end of July, nymphs in February and June (pers. comm. S. Chen, Dec. 2015) Distribution Taiwan. Remarks This species was mentioned from China by Chou et al. (1985) and Nagai & Porion (1996). However, no specimen of P. watanabei could be found either in the collections of NWAFU, where Chou and co-authors worked (pers. comm. D. Qin, Jun. 2016), in the collections of IZCAS (pers. comm. Z.S. Song, Jun. 2016), or in the collections of MHNL, and there are also no photographic records available supporting the presence of the species on the continent. Hence, the species is removed from the list of Chinese Fulgoridae and is regarded as endemic in Taiwan. Like in P. clavatus, this species shows specimens with posterior wings entirely white and others with the apical third black.

Pyrops watanabei (Matsumura, 1913) Figs 4, 8, 16���22 Fulgora (Hotinus) watanabei Matsumura, 1913: 54 [described; close to ��� Fulgora ponderosus ��� (St��l, 1854)], pl. 8, fig. 1 [habitus illustrated] (types in HUIC). Fulgora chimara Schumacher, 1915a: 129 [described] (Types in MFNB and SDEI) [synonymized by Kato (1928); reinstated as good species by Metcalf (1947); re-synonymized by Lallemand (1963)]. Fulgora watanabei var. apicalis Kato, 1928: 221 [described; host plant], pl. 9, fig. 1 [habitus illustrated] (type in UMUT). syn. nov. [name preoccupied by Fulgora apicalis Westwood, 1838, replaced by F. watanabei var. formosana by Metcalf (1947)]. Fulgora watanabei var. formosana Metcalf, 1947: 208 [replacement name for F. watanabei var. apicalis]. syn. nov. Fulgora chimara ��� Schumacher 1915b: 110 [listed from ��� Formosa ��� (= Taiwan)]. ��� Kato 1928: 221 [synonymized under Pyrops watanabei (Matsumura, 1913)]. ��� Metcalf 1947: 193 [catalogued as a good species]. ��� Lallemand 1963: 89 [synonymized under Pyrops watanabei (Matsumura, 1913)]. Fulgora watanabei ��� Kato 1928: 221 [listed; senior synonym of Fulgora chimara Schumacher, 1915; host plant]. ��� Mutsumura 1931: 1267 [described; illustrated]. ��� Chou et al. 1985: 118 [described; mentioned from Taiwan and China (Guangxi, Guangdong and Hainan)]. Hotinus watanabei ��� Matsumura 1931: pl. 8, fig. 12 [illustrated]. Pyrops watanabei ��� Nagai & Porion 1996: 26 [catalogued; mentioned from China (erroneous)]. ��� Liang & Suwa 1998: 157 [lectotype designation]. ��� Yen & Yang 2001 [description; biology; distribution; protection status]. Diagnosis The species is immediately recognized by the following combination of characters: (1) cephalic process yellow and inflated apically (Fig. 4 D���E). (2) abdomen red ventrally (Fig. 4B). (3) tegmina mainly white on disc and with 3 black spots in costal area before nodal line (Fig. 4A). Etymology The species was dedicated to its collector, Kenji Watanabe. Type material TAIWAN: Lectotype, ♀ of Fulgora (Hotinus) watanabei Matsumura, 1913 (examined from photographs, Fig. 16) [Hotinus watanabei Mats.] [Hotinus watanabei Mats.] [Hotinus watanabei det Matsumura] [Formosa Matsumura/ underside: Hoppo, 1 st VII ���07] [Type Matsumura] [Lectotype Fulgora (Hotinus) watanabei Mats. det. A.P. Liang & M. Suwa 1997] (HUIC). TAIWAN: Paralectotype, ♀ of Fulgora (Hotinus) watanabei Matsumura, 1913 (examined from photographs, Fig. 17) [Hoppo] [Paralectotype Fulgora (Hotinus) watanabei Mats. det. A.P. Liang & M. Suwa 1997] (HUIC). TAIWAN: Paralectotype, ♀ of Fulgora (Hotinus) watanabei Matsumura, 1913 (examined from photographs, Fig. 18) [Formosa Matsumura / underside: Hoppo, 27 VII ���07] [Paralectotype Fulgora (Hotinus) watanabei Mats. det. A.P. Liang & M. Suwa 1997] (HUIC). TAIWAN: Syntype, ♀ of Fulgora chimara Schumacher, 1915 (Fig. 19) [Formosa, Hoozan, VIII.10, H. Sauter S.G.] [Fulgora chimara * Schum., F. Schumacher det.] [Type] (MFNB). TAIWAN: Syntype, ♀ of Fulgora chimara Schumacher, 1915 (examined from photographs, Fig. 20) [Kosempo, Formosa, H. Sauter, VII. X.1911] [Fulgora chimara * Schum., F. Schumacher det.] [Syntypus] (SDEI). TAIWAN: Holotype, ♀ of Fulgora watanabei var. apicalis Kato, 1928 (examined from photographs, Fig. 21) [Horisha, Formosa (VIII.1919) Col. M. Kato.] [Type No. 95, M. Kato coll.] (UMUT). Additional material TAIWAN: 1 ♀, Taipei city, Congguan Nat. Taiwan University Campus, 2007, leg. W.B. Yeh (RBINS); 1 ♂ (dissected), 1 ♀, Taipei, Tanshui, 14 Jun. 2002, J.H. Chen (NMNS). Material examined from photographs TAIWAN: eggs (Fig. 22A), Taipei City, 28 Jul. 2012, S. Chen; eggs (Fig. 22B), idem, 19 Feb. 2016; 1 nymph (Fig. 22C), Nantou County, 14 Feb. 2012, S. Chen; 1 nymph (Fig. 22D): Taipei City, 7 Jun. 2012, on Triadica sebifera, S. Chen; 1 ex. (Fig. 22 E���F), idem, 16 Aug. 2012, on Triadica sebifera, S. Chen; 1 ex. (Fig. 22G), idem, predated by a Sparassidae spider; 5 ex. (Fig. 22H), Taipei City, 6 Aug. 2011, on Triadica sebifera, S. Chen. Measurements and ratios TL: ♂ (n=1): 3.9 cm; LPr: 1.0 cm; TL: ♀ (n=2): 4.3 cm (4.0���4.6); LPr: 1.1 cm (1.0���1.2); LTg/BTg = 2.31; BF/BPrH = 2.53; LPr/LF = 2.82; LPr/BPrH = 2.55. Male genitalia Pygofer higher than long, with posterior margin sinuate in lateral view (Fig. 8A). Anal tube slightly elongate, 1.1 times as long as broad in dorsal view, broader at 4/5 of total length (Fig. 8C); lateral margins slightly sinuate and apical margin strongly concave in dorsal view (Fig. 8C). Gonostyli (Fig. 8A) elongate, 1.77 times as long as high in lateral view; dorsal margin regularly and broadly rounded and posterior margin slightly projecting posteriorly in middle in lateral view (Fig. 8A). Biology The species was recorded on Triadica sebifera (L.) Small (Euphorbiaceae) by Kato (1928), and on the same tree and Sapium discolor Muell. -Arg. (Euphorbiaceae) by Yen & Yang (2015). Those data were confirmed by multiple observations in the field; freshly laid eggs were observed at the end of July, nymphs in February and June (pers. comm. S. Chen, Dec. 2015) Distribution Taiwan. Remarks This species was mentioned from China by Chou et al. (1985) and Nagai & Porion (1996). However, no specimen of P. watanabei could be found either in the collections of NWAFU, where Chou and co-authors worked (pers. comm. D. Qin, Jun. 2016), in the collections of IZCAS (pers. comm. Z.S. Song, Jun. 2016), or in the collections of MHNL, and there are also no photographic records available supporting the presence of the species on the continent. Hence, the species is removed from the list of Chinese Fulgoridae and is regarded as endemic in Taiwan. Like in P. clavatus, this species shows specimens with posterior wings entirely white and others with the apical third black., Published as part of Constant, J��r��me & Pham, Hong-Thai, 2017, Review of the clavatus group of the lanternfly genus Pyrops (Hemiptera: Fulgoromorpha: Fulgoridae), pp. 1-26 in European Journal of Taxonomy 305 on pages 18-23, DOI: 10.5852/ejt.2017.305, http://zenodo.org/record/3827135, {"references":["Matsumura S. 1913. Thousand Insects of Japan Additamenta. Vol. 1. Keiseisha, Tokyo [in Japanese].","Stal C. 1854. Nya Hemiptera. Ofversigt af Kongliga Svenska Vetenskaps-Akademiens Forhandlingar 11: 231 - 255.","Schumacher F. 1915. Homoptera in H. Sauter's Formosa-Ausbeute. Supplementa Entomologica 4: 108 - 142.","Kato M. 1928. Notes on some Formosan Homoptera, with descriptions of new genus and species. Kontyu, Tokyo Entomological Society 2: 221 - 228.","Metcalf Z. P. 1947. General Catalogue of the Homoptera. Fascicle IV Fulgoroidea. Part 9 Fulgoridae. North Carolina State College, Raleigh (U. S. A.).","Lallemand V. 1963. Revision des Fulgoridae (Homoptera). Deuxieme partie. Faunes asiatique et australienne. Memoires de l'Institut royal des Sciences naturelles de Belgique (2 e serie) 75: 1 - 99.","Chou I., Wang S. & Huang J. 1985. Priskribo de novaj specioj de Fulgoroedoj el Cinio (Homopteroj: Fulgoroedoj). Entomotaxonomia 7 (1): 29 - 38.","Matsumura S. 1931. 6000 Illustrated Insects of Japan Empire. Tokoshoin, Tokyo [in Japanese].","Nagai S. & Porion T. 1996. Fulgoridae 2: Catalogue illustre des Faunes asiatique et australienne. Sciences Nat, Compiegne.","Liang A. - P. & Suwa M. 1998. Type specimens of Matsumura's species of Fulgoroidea (excluding Delphacidae) in the Hokkaido University collection, Japan (Hemiptera: Fulgoromorpha). Insecta Matsumurana, Series Entomology, New Series 54: 133 - 166.","Yen S. - H. & Yang P. - S. 2001. Illustrated Identification Guide to Insects Protected by the CITES and Wildlife Conservation Law of Taiwan, R. O. C. Council of Agriculture, Executive Yuan, Taiwan, R. O. C. Available from http: // subject. forest. gov. tw / books / book _ 03 e / insect _ 93. htm [accessed 12 Dec. 2015]."]}

Genus Pyrops Spinola, 1839 Pyrops Spinola, 1839: 231. Type species: Pyrops candelaria (Linnaeus, 1758) by subsequent designation by Duponchel (1840: 200). Hotinus Amyot & Serville, 1843: 490 (type species Pyrops candelaria (Linnaeus, 1758) by original designation) synonymized by Blanchard 1845: 425. After comparison with the classification proposed by Lallemand (1963) and Nagai & Porion (1996), the genus Pyrops is here removed from the subfamily Fulgorinae and not attributed to any of the currently defined subfamilies, following the conclusions of the DNA study by Urban & Cryan (2009). The subfamily Fulgorinae is found in the New World, with the Neotropical genus Fulgora Linnaeus, 1767 as type. Diagnostic characters The definition of the genus given by Constant (2015) is followed: head with cephalic process, sometimes very long, narrowing progressively beyond the eyes; apically it can be dilated or even spherical. Vertex about 4 times as broad as an eye. Before eyes, genae truncate, with a transverse carina which sometimes extends to vertex. Two longitudinal carinae on frons, a third median one starting on base of cephalic process. Fronto-clypeal suture usually slightly bisinuate; median carina on clypeus. Pronotum with median carina (sometimes obsolete) and a small but strongly impressed point on each side of it. Mesonotum with median and peridiscal carinae, sometimes obsolete. Tegmina at most 3 times as long as broad, with apical margin more or less rounded and with transverse veinlets on all surfaces. Clavus open and elongate, vein A 1+ A 2 extending far towards apex. Legs slender. The clavatus species group This group was defined by Baker (1925: 348) with the following set of characters: (1) medium sized species; (2) cephalic process short, very stout, strongly clavate, black or olive green above and with red or ochraceous apex; (3) tegmina largely black. It seems worth mentioning that Baker did not examine any specimen of P. atroalbus comb. nov. or P. watanabei. Lallemand (1963: 88) restricted the definition to characters of the cephalic process only: ��� cephalic process rather short, much shorter than body, gradually narrowing, strongly dilated apically into a quite large ball��� (translated from French). Nagai & Porion (1996) followed Lallemand���s (1963) definition. After examination of the types of all species placed in the group by previous authors, the combination of the following characters is given to define the group: (1) medium sized species; (2) cephalic process rather short, progressively narrowing towards apex and strongly swollen apically; (3) apical third of hind wings black or white. The Philippine species P. polillensis is removed from the group based on the broad black area of the hind wing extending all along the sutural margin (see illustrations in Baker 1925), in contradiction with character (3), and not attributed to any of the currently defined species groups of Pyrops. The three species included here in the group are distributed in a zoogeographically consistent zone extending from northern India eastwards to Taiwan through Bangladesh, Myanmar, northern Thailand, Laos and southern China, and southwards to central Vietnam. Identification key to the species of the Pyrops clavatus group 1. Abdomen black ventrally (Fig. 1B); tegmina pale yellow-white on disc and with 3 black spots in costal area before nodal line (Fig. 1A); cephalic process yellow (Fig. 1 D���F) (known from Laos, Thailand and Vietnam) ������������������������������������������������ Pyrops atroalbus (Distant, 1918) comb. nov. ��� Abdomen red ventrally (Fig. 2B) ���������������������������������������������������������������������������2 2. Tegmina largely black on disc (Fig. 2A), or in the pale forms (Fig. 3A, D), bluish white on disc and without black spots in costal area; cephalic process red-brown to black (Figs 2 D���G, 3C, E) (known from N India, Bangladesh, N Myanmar, N Thailand, S China and N Vietnam) ��������� ��������������������������������������������������������������������������� Pyrops clavatus (Westwood, 1839) ��� Tegmina mainly white on disc and with 3 black spots in costal area before nodal line (Fig. 4A, F); cephalic process yellow (Fig. 4 D���E) (known from Taiwan) ��� Pyrops watanabei (Matsumura, 1913), Published as part of Constant, J��r��me & Pham, Hong-Thai, 2017, Review of the clavatus group of the lanternfly genus Pyrops (Hemiptera: Fulgoromorpha: Fulgoridae), pp. 1-26 in European Journal of Taxonomy 305 on pages 3-4, DOI: 10.5852/ejt.2017.305, http://zenodo.org/record/3827135, {"references":["Spinola M. 1839. Essai sur les Fulgorelles, sous-tribu de la tribu des Cicadaires, ordre des Rhyngotes. Annales de la Societe Entomologique de France 8: 133 - 337. Available from http: // biodiversitylibrary. org / page / 33710046 [accessed 17 Mar. 2017]","Duponchel P. A. J. 1840. Essai sur les fulgorelles, par Maximilien Spinola. Revue Zoologique, par la Societe cuvierienne; Association Universelle pour l'Advancement de la Zoologie, de l'Anatomie comparee et de la Palaeontologie; Journal mensuel 2: 199 - 206.","Amyot C. & Serville J. 1843. Deuxieme partie. Homopteres. Homoptera Latr. Histoire Naturelle des Insectes. Hemipteres. Librairie encyclopedique de Roret, Paris. https: // doi. org / 10.5962 / bhl. title. 8471","Blanchard E. 1845. Septieme ordre les Hemipteres. In: Blanchard E. Histoire des insectes, traitant de leurs moeurs et de leurs metamorphoses en general et comprenant une nouvelle classification fondee sur leurs rapports naturels 2: 424 - 427. Available from http: // biodiversitylibrary. org / page / 25155881 [accessed 20 Mar. 2017]","Lallemand V. 1963. Revision des Fulgoridae (Homoptera). Deuxieme partie. Faunes asiatique et australienne. Memoires de l'Institut royal des Sciences naturelles de Belgique (2 e serie) 75: 1 - 99.","Nagai S. & Porion T. 1996. Fulgoridae 2: Catalogue illustre des Faunes asiatique et australienne. Sciences Nat, Compiegne.","Urban J. & Cryan J. 2009. Entomologically famous, evolutionarily unexplored: the first phylogeny of the lanternfly family Fulgoridae (Insecta: Hemiptera: Fulgoroidea). Molecular Phylogenetics and Evolution 50: 471 - 484. https: // doi. org / 10.1016 / j. ympev. 2008.12.004","Constant J. 2015. Review of the effusus group of the lanternfly genus Pyrops Spinola, 1839, with one new species and notes on trophobiosis (Hemiptera: Fulgoromorpha: Fulgoridae). European Journal of Taxonomy 128: 1 - 23. https: // doi. org / 10.5852 / ejt. 2015.128","Baker C. F. 1925. Remarks on certain Indo-Malayan Fulgora, with special reference to Philippine species. Philippine Journal of Science 28: 343 - 364.","Distant W. L. 1918. The Homoptera of Indo-China. Annals and Magazine of Natural History (Ser. 9) 1: 196 - 200.","Westwood J. O. 1839. On the family Fulgoridae, with a monograph of the genus Fulgora of Linnaeus. The Transactions of the Linnean Society of London, Second series, Zoology 18: 133 - 153.","Matsumura S. 1913. Thousand Insects of Japan Additamenta. Vol. 1. Keiseisha, Tokyo [in Japanese]."]}

Pyrops clavatus (Westwood, 1839) Figs 2–3, 7, 9 B–G, 10–15 Fulgora clavata Westwood, 1839: 139, pl. 12, fig. 1 (types in OUMNH) [described, illustrated, compared with P. pyrorhynchus (Donovan, 1800), the latter mentioned as the junior synonym Fulgora pyrorhina Westwood, 1839)]. Hotinus ponderosus Stål, 1854: 244 (type in NHRS) [described and mentioned as close to P. clavatus; synonymized by Distant (1906)]. Fulgora woodii Ollenbach, 1929: 279, pl. 1, fig. 13 (types in NFIC) [described, illustrated and mentioned as very near to Pyrops clavatus; synonymized by Lallemand (1963)]. Fulgora nigripennis Chou & Wang in Chou et al., 1985: 33, fig. 4 (type in NWAFU) [described in Chinese, illustrated, compared to P. clavatus], 37 (briefly described in Esperanto, host plant) syn. nov. Fulgora clavata mizunumai Satô & Nagai, 1994: 312, figs 3, 12 (type in EUM) [described, illustrated, compared to P. clavatus] syn. nov. Pyrops clavata – Burmeister 1845: 4 [transferred to Pyrops]. — Schaum 1850: 64 [listed] — Kirby 1885: 211 [mentioned from India and briefly described]; 1892: 211 [idem]. — Nagai & Porion 1996: 26 [catalogued; distribution]; 31 [very close to Fulgora nigripennis Chou & Wang, 1985], pl. 18, figs 227–228 [illustrated]. Hotinus clavatus – Adams 1847: 204 [compared with P. sultanus (Adams & White, 1847)]. — Walker 1851: 267 [list of specimens in BMNH]; 1858: 41 [clavatus as the type and single member of a group within Hotinus]. — Stål 1854: 244 [close to Hotinus ponderosus Stål, 1854]. — Costa 1864: 82 [listed from Assam]. Fulgora (Hotina [sic !]) clavata – Westwood 1848: 7, pl. 3, fig. 1 [described, illustrated]. Hotinus ponderosus – Walker 1858: 315 [listed from Hindustan]. — Distant 1906: 191 [junior synonym of P. clavatus (Westwood, 1839)]. Fulgora clavata – Butler 1874: 98 [placed in a section of Fulgora with white hind wings; list of specimens in BMNH; mention of “ ludicrous ” cephalic process; probable senior synonym of “ Fulgora ponderosa ”]. — Gadeau de Kerville 1881: 43 [listed as light-producing insect (erroneous)]. — Atkinson 1885: 130 [catalogued; described; distribution; intraspecific variation]. — Schmidt 1905: 354 [catalogued]. — Distant 1906: 191; fig. 83 [keyed; described; habitus, side of head and frons illustrated; senior synonym of Hotinus ponderosus Stål, 1854]. — Schumacher 1915: 129 [compared with Fulgora chimara Schumacher, 1915]. — Distant 1918: 198 [listed from “Indochina”], 200 [compared with P. atroalbus (Distant, 1918)]. — Paiva 1919: 373 [mentioned from Garo Hills, N India; notes on biology]. — Baker 1925: 348 [type of the clavata group in Fulgora], 361 [key to the species of the clavata group; described], pl. 4, fig. 1 [lateral aspect of habitus illustrated]. — Ollenbach 1929: 280 [compared with Fulgora woodi Ollenbach, 1929]. — Lallemand 1963: 71 [type of 5 th group of Fulgora], 88 [keyed; described; catalogued; senior synonym of Hotinus ponderosus Stål, 1854 and Fulgora woodi Ollenbach, 1929]; pl. 10, figs 4–7 [lateral view of head and male genitalia of type illustrated]. — Satô & Nagai 1994: 312 [compared with Fulgora clavata mizunumai Satô & Nagai, 1994]. Fulgora ponderosa – Butler 1874: 98 [probable junior synonym of P. clavatus]. — Gadeau de Kerville 1881: 43 [listed as light-producing insect (erroneous)]. — Atkinson 1885: 131 [catalogued; described; distribution; probable variety of P. clavatus]. Fulgora ponderosus – Matsumura 1913: 54 [close to P. watanabei (Matsumura, 1913)]. Laternaria clavata – Metcalf 1947: 193 [catalogued; distribution]. — Allnatt 2013: 45, fig. 1 [illustrated from Assam]. Laternaria clavata var. ponderosa – Metcalf 1947: 193 [catalogued; distribution; considered as a variety of clavatus (erroneous)]. Laternaria woodi – Metcalf 1947: 208 [catalogued]. Pyrops clavata mizunumai – Nagai & Porion 1996: 26 [catalogued]; pl. 18, figs 222, 224, 226 [illustrated]. Pyrops clavatus – Liang 1998: 42 [catalogued; new combination (erroneous)]. Pyrops clavatus mizunumai – Liang 1998: 42 [catalogued; new combination (erroneous)]. Diagnosis The species is immediately recognized by the following combination of characters: (1) Cephalic process red-brown to black, often black with apex red-brown, and strongly inflated apically (Figs 2 D–G, 3C, E). (2) Abdomen red ventrally (Figs 2B, 3B). (3) Tegmina largely black on disc in the dark forms (Fig. 1 A–B); in the pale forms, tegmina bluish white on disc without black spots in costal area (Fig. 3 A–B, D). Etymology clavatus (adj., Latin): clavate. The name refers to the shape of the cephalic process. Material examined Type material BANGLADESH: Lectotype, ♂ of Fulgora clavata Westwood, 1839, here designated to provide a reference standard for the species (examined from photographs, Fig. 10) [Silhet] [Fulgora clavata Westw. Trans. Lin. Soc. 18. P. 139 Pl 12 Fig 5.] [Type] [Type Hem: 596 1/2 Fulgora clavata Westwood, Hope Dept. Oxford] (OUMNH). BANGLADESH: Paralectotype, ♀ of Fulgora clavata Westwood, 1839 (examined from photographs, Fig. 11) [Silhet] [Fulgora clavata Westw. Monogr. M. Hope, Sylhet] [W] [Type] [Type Hem: 596 2/2 Fulgora clavata Westwood, Hope Dept. Oxford] (OUMNH). INDIA: Holotype, ♀ of Hotinus ponderosus Stål, 1854 (Fig. 12) [Ind or] [Saund] [ponderosus Stål] [NHRS-HEMI000000198] (NHRS). MYANMAR: Syntype, ♂ of Fulgora woodii Ollenbach, 1929 (examined from photographs, Fig. 13) [Paga-yo, Tavoy, 27.I.20, O.C. Ollenbach] [Type] [Fulgora woodii (Ollen) ♂ n sp., O.C. Ollenbach det.] (NFRI). CHINA: Holotype, ♂ of Fulgora nigripennis Chou & Wang, 1985 (examined from photographs, Fig. 14) [Fulgora nigripennis Chou, Wang & Huang “ identified by Io Chou, August 1982 ”] [“ Host plant: coffee ”] [HOLOTYPE] (NWAFU). – Parts in italics between “ ” were translated from Chinese by D. Qin. The specimen was collected in Ruyuan, Guangdong Province in May 1975 by Lizhong Hua (D. Qin pers. comm., Jun. 2016). THAILAND: Holotype, ♂ of Fulgora clavata mizunumai Satô & Nagai, 1994 (examined from photographs, Fig. 15) [Doi Pui, Chiang Mai (N Thailand) May 1987] [Holotype Fulgora clavata mizunumai] (EUM). Additional material INDIA: 1 ♀, Darjeeling (Himalaya), 1869, Higgins (RBINS); 1 ♀, Kurseong, no date, R.P. Wery (RBINS). THAILAND: 5 ♂♂, 15 ♀♀, Chiang Mai, Dec. 2007, local collectors (RBINS). VIETNAM: 1 ♀, Ha Giang, Jul. 2009, local collectors (RBINS); 2 ♀♀, Cuc Phuong National Park, 20°19′00″ N, 105°36′30″ E, 19–23 Jul. 2011, day collecting, on tree trunk, leg. J. Constant and J. Bresseel (RBINS); 2 ♀♀, Cuc Phuong N.P., 25 May 2005, leg. H.T. Pham (VNMN); 3 ♂♂, 3 ♀♀, Lao Cai Prov., Sa Pa Mt, 1800 m, Oct. 2014, leg. Than Le Luong (RBINS); 1 ♀, Vinh Phuc Prov., Me Linh Station, 12 Aug. 2011, leg. H.T. Pham (VNMN); 3 ♂♂, 10 ♀♀, Ba Vi N.P., 21°4′4″ N, 105°21′30″ E, 25–29 Jun. 2015, 1000 m, leg. H.T. Pham (VNMN); 1 ♂, 2 ♀♀, Ba Vi N.P., 21°4′4″ N, 105°21′30″ E, 25–29 Jun. 2015, 600 m, leg. J. Constant and J. Bresseel (RBINS); 1 ♀, Nghe An Prov., Pu Mat N.P., 18°59′ N, 104°40′ E, Jul. 2014, local collector (RBINS); 1 ♀, Tuyên Quang Prov., Cham Chu Nature Reserve, 22°12′ N, 105°6′ E, 8–12 Jul. 2015, leg. J. Constant and J. Bresseel (RBINS); 2 ♂♂, same data, leg. H.T. Pham (VNMN); 1 ♀, Quang Binh Prov., Phong Nha-Ke Bang N.P., U Bo, 17°32′14″ N, 106°9′4″ E, 550 m, leg. H.T. Pham (VNMN); 3 ♂♂, 3 ♀♀, Son La Prov., Copia Nature Reserve, 21°22′12″ N, 103°30′42″ E, 20–23 Jul. 2016, J. Constant and J. Bresseel (RBINS). Material examined from photographs INDIA: 1 ex., Assam, Mangaldai, 13 Aug. 2014, Mirza Galib (Facebook group: InsectIndia); 1 ex., Manipur, Loktak Lake, 9 Oct. 2014, Sougrakpam Neli (Facebook group: InsectIndia); 1 ex., Nagaland, Aug. 2012, Tarun Karmakar (Facebook group: InsectIndia); 1 ex., Upper Assam, Dibrugarh, Rungagora, no date, H. Stevens (Alnatt 2013: fig. 1). VIETNAM: 1 ex. (Fig. 9 B–C), Me Linh Biological Station, 12 Jul. 2015, on a trunk of Dimocarpus longan (Sapindaceae), J. Constant. Additional data from literature INDIA: Garo Hills above Tura, 3000 ft (Paiva 1919). MYANMAR: Karen Hills (Ollenbach 1929 – location of the second syntype of Fulgora woodi). Measurements and ratios TL: ♂ (n = 5): 3.9 cm (3.7–4.0); LPr: 1.2 cm (1.1–1.3); TL: ♀ (n = 5): 5.1 cm (4.5–5.5); LPr: 1.35 cm (1.3–1.5); LTg/BTg = 2.47; BF/BPrH = 2.05; LPr/LF = 3.19; LPr/BPrH = 6.13. Male genitalia Pygofer higher than long, with posterior margin angularly bisinuate dorsally in lateral view (Fig. 7A). Anal tube slightly elongate, 1.1 times as long as broad in dorsal view, broader at 4/5 of total length (Fig. 7C); lateral margins very slightly sinuate and apical margin strongly concave in dorsal view (Fig. 7C). Gonostyli (Fig. 7A) elongate, 1.5 times as long as broad in lateral view; dorsal margin strongly rounded above lateral tooth and posterior margin rounded in lateral view (Fig. 7A). Remarks After examination of the photographs of the type specimens of Fulgora nigripennis Chou & Wang, 1985 and Fulgora woodii Ollenbach, 1929, it was not possible to find any difference between those taxa and the types of P. clavatus and the recorded intraspecific variations of the species. Hence, the first is proposed as a junior synonym of Pyrops clavatus and the synonymy of the second under P. clavatus, as proposed by Lallemand (1963) and followed by Nagai & Porion (1996), is confirmed. Ollenbach (1929) stated that the types of the species described in his paper would later be sent to the British Museum (currently BMNH) but he apparently never did so as none of the types of those species can be found in the BMNH collections (M. Webb pers. comm., 21 May 2013). The examination of numerous specimens of P. clavatus, including large series from Chiang Mai (Figs 2–3) and northern Vietnam, proved that the species shows important intraspecific colour variation of the cephalic process and wings: the tegmina vary from nearly completely black to nearly completely bluish white, the hind wings from white with black apex to completely white, usually tinged with blue or violet basally. The “subspecies” mizunumai described by Satô & Nagai (1994) only represents the paler extreme of the species and cannot be considered as a subspecies, as it occurs sympatrically with the intermediate and darker forms. It is therefore synonymized under P. clavatus. Specimens showing basally blue-tinged and violet-tinged hind wings were found in the same population, on the same tree in Copia, North Vietnam. The variation is not linked to the sex of the specimens as both males and females showed the two variations. Biology Paiva (1919) stated that the species was “ not uncommon at 3000 ft. Several specimens sometimes found on a single tree to which they return after disturbance ”. Our observations confirm this statement which is valid for most species of Pyrops we have observed so far. We have found P. clavatus feeding on several unidentified species of tree in Ba Vi National Park (north Vietnam); in Cuc Phuong National Park (north Vietnam), some specimens of P. clavatus were found on a big tree together with numerous Pyrops spinolae (Westwood, 1842), while other trees of the same species and others around did not host any Pyrops specimens (Fig. 9 F–G); at Me Linh Biodiversity Station (north Vietnam), P. clavatus was observed on a big Longan tree trunk (Dimocarpus longan Lour., Sapindaceae) together with P. candelaria (Linnaeus, 1758), P. lathburii (Kirby, 1818) and P. viridirostris (Westwood, 1848). In North Vietnam we found the species at altitudes ranging from 150 to 1200 m. The species was also collected on coffee (Coffea sp., Rubiaceae) in southern China. Distribution Known from N India, Myanmar, N Thailand, S China and N Vietnam. Very probably also present in Laos and maybe in Cambodia (see also Constant et al. 2016 for discussion on Fulgoridae from Cambodia).

Pyrops atroalbus (Distant, 1918) comb. nov. Figs 1, 5���6, 9A Fulgora atroalba Distant, 1918: 199 (type in BMNH). Laternaria atroalba ��� Metcalf 1947: 187 [transferred to Laternaria; catalogued]. Fulgora watanabei apicalis ��� Lallemand 1963: 90 [considered as a junior synonym of Fulgora watanabei atroalba (Distant, 1918) (erroneous)]. Fulgora watanabei atroalba ��� Lallemand 1963: 90 [transferred back to Fulgora and considered as a subspecies of P. watanabei (erroneous)]. Fulgora watanabei var. formosana ��� Lallemand 1963: 90 [considered as a junior synonym of Fulgora watanabei atroalba (Distant, 1918) (erroneous)]. Pyrops watanabei apicalis ��� Nagai & Porion 1996: 26 [transferred to Pyrops but still considered as a junior synonym of Fulgora watanabei atroalba (erroneous)]. Pyrops watanabei atroalba ��� Nagai & Porion 1996: 26, pl. 18, fig. 223 [transferred to Pyrops, but still considered as a subspecies of P. watanabei (erroneous), type illustrated]. ��� Liang 1998: 22 [considered as a subspecies of P. watanabei (erroneous)]. Pyrops watanabei formosana ��� Nagai & Porion 1996: 26 [transferred to Pyrops but still considered as a junior synonym of Fulgora watanabei atroalba (erroneous)]. Non Fulgora watanabei apicalis Kato, 1928: 221, pl. 9 fig. 1. Non Fulgora watanabei var. formosana Metcalf, 1947: 208. Note Liang (1998) erroneously stated that the name ��� Pyrops watanabei atroalbus (Distant, 1918) ��� was a new combination he proposed while the combination had already been proposed by Nagai and Porion (1996). Diagnosis The species is immediately recognized by the following combination of characters: (1) Cephalic process yellow and strongly inflated apically (Fig. 1 D���F). (2) Abdomen black ventrally (Fig. 1B). (3) Tegmina strongly contrasted: pale yellow-white with black markings, including 3 black spots in costal area before nodal line (Fig. 1A). Etymology From Latin ater (adj.) ���black��� and albus (adj.): ���white���; the species epithet refers to the black and white colouration of the species. Type material Holotype (Fig. 5) VIETNAM: ♂ [Indo-China, Tonkin, R.V. de Salvaza. 1917���98] [1918���1] [Fulgora atroalba Dist. Type] [Type] (BMNH). Additional material VIETNAM: 1 ♂, 1 ♀, Da Krong Nature Reserve, Quang Tri Province, 10 Jul. 2011, 16��37��� N, 106��47��� E, day collecting, Pham & Hoang (♂ in RBINS, ♀ in VNMN); 1 ♀, Bach Ma National Park, 29 Sep. 2014, 16��12��� N, 107��52��� E, Tuan (VNMN). Material examined from photograph THAILAND: 1 ex. (Fig. 9A), Chiang Mai Prov., Maerim District, 18��54���50��� N, 98��56���42��� E, 14 Mar. 2016, Panaka Jirasuttayaporn. Measurements and ratios TL: ♂ (n = 1): 3.4 cm; ♀ (n = 1): 4.2 cm; LPr: ♂: 1.1 cm; ♀: 1.4 cm; LTg/BTg = 2.55; BF/BPrH = 3.1; LPr/LF = 2.95; LPr/BPrH = 8.8. Male genitalia Pygofer higher than long, with posterior margin regularly rounded dorsally in lateral view (Fig. 6A). Anal tube slightly elongate, 1.1 times as long as broad in dorsal view, broader at 4/5 of total length (Fig. 6C); lateral margins very slightly sinuate and apical margin strongly concave in dorsal view (Fig. 6C). Gonostyli (Fig. 6A) elongate, twice as long as high in lateral view; dorsal margin regularly rounded and posterior margin slightly projecting posteriorly in middle in lateral view (Fig. 6A). Remarks There is a discrepancy between the location given on the label of the specimen (��� Tonkin ���) and the one given in the original description (Distant 1918): ��� Indochina, Xieng Klouang (R. Vitalis de Salvaza) ���. The latter location is situated in Laos, not far from the places where we have collected specimens in Central Vietnam. Despite our intensive collecting effort in northern Vietnam, we have never found the species in that region. Hence, it seems that the location given by Distant (1918) is more likely to be the correct one. Nagai & Porion (1996: fig. 223) erroneously gave ���central Taiwan ��� as the type locality. Distribution The species is known from one location in Laos, one in Northern Thailand, and two in Central Vietnam., Published as part of Constant, J��r��me & Pham, Hong-Thai, 2017, Review of the clavatus group of the lanternfly genus Pyrops (Hemiptera: Fulgoromorpha: Fulgoridae), pp. 1-26 in European Journal of Taxonomy 305 on pages 5-10, DOI: 10.5852/ejt.2017.305, http://zenodo.org/record/3827135, {"references":["Distant W. L. 1918. The Homoptera of Indo-China. Annals and Magazine of Natural History (Ser. 9) 1: 196 - 200.","Metcalf Z. P. 1947. General Catalogue of the Homoptera. Fascicle IV Fulgoroidea. Part 9 Fulgoridae. North Carolina State College, Raleigh (U. S. A.).","Lallemand V. 1963. Revision des Fulgoridae (Homoptera). Deuxieme partie. Faunes asiatique et australienne. Memoires de l'Institut royal des Sciences naturelles de Belgique (2 e serie) 75: 1 - 99.","Westwood J. O. 1839. On the family Fulgoridae, with a monograph of the genus Fulgora of Linnaeus. The Transactions of the Linnean Society of London, Second series, Zoology 18: 133 - 153.","Nagai S. & Porion T. 1996. Fulgoridae 2: Catalogue illustre des Faunes asiatique et australienne. Sciences Nat, Compiegne.","Liang A. - P. 1998. Nomenclatorial notes on the Oriental lanternfly genus Pyrops Spinola (Hemiptera: Fulgoroidea: Fulgoridae). Acta Zootaxonomica Sinica 23 (1): 41 - 47.","Matsumura S. 1913. Thousand Insects of Japan Additamenta. Vol. 1. Keiseisha, Tokyo [in Japanese].","Kato M. 1928. Notes on some Formosan Homoptera, with descriptions of new genus and species. Kontyu, Tokyo Entomological Society 2: 221 - 228."]}

The clavatus group of Pyrops Spinola, 1839 is reviewed and redefined. The new combination Pyrops atroalbus (Distant, 1918) comb. nov. is proposed, as atroalbus is reinstated as a full species from status of subspecies of Pyrops watanabei (Matsumura, 1913). Pyrops nigripennis (Chou & Wang, 1985) and Pyrops clavatus mizunumai (Sato & Nagai, 1994) are proposed as junior synonyms of P. clavatus (Westwood, 1839). The Philippine species P. polillensis (Baker, 1925) is removed from the group and not attributed to any of the currently defined species groups. Hence, the clavatus group is restricted to continental Southeast Asia and Taiwan and contains three species: P. atroalbus comb. nov., P. clavatus and P. watanabei . A key to the species of the group and illustrations of the male genitalia are provided. The intraspecific colour variation in the group is discussed and illustrated. The genus Pyrops is removed from the subfamily Fulgorinae and not attributed to any of the currently defined subfamilies of Fulgoridae.

A new Fulgoridae, Pyrops silighinii n. sp., collected in Mindanao (Philippines) is described and illustrated., Un nouveau Fulgoridae, Pyrops silighinii n. sp., collecté à Mindanao (Philippines), est décrit et illustré.

A new Fulgoridae, Pyrops nishiguroi n. sp., collected in Palawan (Philippines) is described and illustrated. Pyrops alboroseus Liang, 1998 stat. nov. is elevated to specific rank. Some new localities are given for Pyrops oculatus., Un nouveau Fulgoridae, Pyrops nishiguroi n. sp., collecté à Palawan (Philippines), est décrit et illustré. Pyrops alboroseus Liang, 1998 stat. nov. est élevé au rang d’espèce. Des localités nouvelles sont données pour Pyrops oculatus.

A new Fulgoridae collected in Mindoro (Philippines) is described and illustrated: Pyrops priscilliae n. sp. Besides, Pyrops agusanensis (Baker, 1925), a very rare species is illustrated with a specimen from Mindanao (Philippines)., Un nouveau Fulgoridae, collecté à Mindoro (Philippines), est décrit et illustré : Pyrops priscilliae n. sp. Une autre espèce, Pyrops agusanensis (Baker, 1925), très rare, est illustrée à partir d’un spécimen de Mindanao (Philippines).

Pyrops effusus (Distant, 1891) Figs 2 A–E, 6–7A–C, 10A, C–F Fulgora effusa Distant, 1891: 518, pl. XX, figs 4, 4a (Type in NHM) [described, illustrated and compared with P. whiteheadi (Distant, 1889)]. Fulgora viridicastanea Baker, 1925: 359, pl. 3, fig. 5, pl. 4 fig. 8 (Type in USNM) [described, illustrated and compared with P. whiteheadi erroneously mentioned and illustrated as “ Fulgora effusa ”]. Synonymized under P. effusus by Nagai & Porion (2004). Fulgora effusa – Schmidt 1905: 349 [comparative note with other species]; 354 [mention of a specimen in the Stettiner Museum]. — Lallemand 1963: 88 [synonymized under whiteheadi (erroneous)]. — Liang 1998: 45 [listed as junior synonym of P. whiteheadi (erroneous)]. Pyrops effusa (sic!) – Schmidt 1926: 47 [transferred to Pyrops; compared with P. peltzeri Schmidt, 1926, the latter synonymized under P. pythicus (Distant, 1891) by Lallemand (1963)]. — Nagai & Porion 1996: 26 [transferred back to Pyrops; catalogued as junior synonym of P. whiteheadi (erroneous)], pl. 11, fig. 163 [type of P. effusus illustrated under the name of P. whiteheadi]. — Nagai & Porion 2004: 6 [removal from synonymy with P. whiteheadi; notes on morphology; senior synonym of P. viridicastaneus and distribution], pl. 1, figs 2, 4 [illustrated from 2 localities]. Laternaria effusa – Metcalf 1947: 196 [transferred to Laternaria; catalogued]. — Liang 1998: 45 [listed as junior synonym of P. whiteheadi (erroneous)]. Laternaria viridicastanea – Metcalf 1947: 207 [transferred to Laternaria; catalogued]. — Liang 1998: 45 [listed as junior synonym of P. whiteheadi (erroneous)]. Fulgora viridicastanea – Lallemand 1963: 88 [synonymized under P. whiteheadi (erroneous) and compared with P. whiteheadi and P. effusus; transferred back to Fulgora (erroneous)]. — Liang 1998: 45 [listed as junior synonym of P. whiteheadi (erroneous)]. Fulgora whiteheadi – Lallemand 1963: 88, pl. X, figs 11–13 [keyed, comments illustration of male genitalia (based on misidentified specimens of P. effusus)]. Fulgora whiteheadi viridicastanea – Satô & Nagai 1994 [considered as a subspecies of P. whiteheadi (erroneous) and compared with “ Fulgora whiteheadi gunjii ”]. Pyrops viridicastanea – Nagai & Porion 1996: 26 [transferred back to Pyrops; catalogued as junior synonym of P. whiteheadi (erroneous)]. — Nagai & Porion 2004: 6 [synonymized under P. effusus, notes]. Pyrops whiteheadi (Distant, 1889) – Nagai & Porion 1996, pl. 11, fig. 163 [illustration of the type of P. effusus (Distant, 1891)]. non Fulgora effusa – Baker 1925: 359, pl. 3, fig. 4, pl. 4, fig. 7 [described, illustrated (misidentification of Pyrops whiteheadi)]. Diagnosis In addition to the characters defining the effusus group, the species can be recognized by: (1) head yellow-brown with apical half of cephalic process blue (often faded to greenish in collection specimens) (Fig. 2 D–E); (2) pro- and mesonotum yellow-brown (Fig. 2A); (3) tegmina green with yellow spots also beyond nodal line; on basal half, spots tranverse and sometimes coalescent (Fig. 2A); (4) tegmina without large pale yellow patch at nodal line and apical margin slightly infuscate (Fig. 2A). Material examined Type material INDONESIA: Holotype of Fulgora effusa (examined on photographs), ♂: [Borneo] [effusa Dist.] [Distant Coll. 1911 —383.] (BMNH). INDONESIA: Holotype of Fulgora viridicastanea (examined on photographs, Fig. 10 C–F), ♀: [Banjermassin, Borneo] [Fulgora viridicastanea Baker] (USNM). Additional material INDONESIA: 2 ♂♂, 2 ♀♀: Kalimantan, Batulicin, Apr. 2008 (RBINS); 2 ♂♂: Laut Island, Jan. 2008 (RBINS); 1 ♀, Laut Island, Jun. 2004 (RBINS); 1 ♂, E-Borneo, Midden, 10 Oct. 1925, H.C. Siebers (FSAG); 1 ♂, idem, 4 Oct. 1925 (FSAG); 1 ♂, SE Borneo, Atkinson coll. (BMNH); 1 ♂, 1 ♀, Borneo, 1909 (BMNH); 1 ♀, North Borneo (BMNH). Examined on photograph INDONESIA: 1 specimen (Fig. 10A), Kalimantan, Wehea Protected Forest, 25 Sep. 2010, Kara Norby. Measurements and ratios TL: ♂ (n = 4): 35.1 mm (34.3–36.0); ♀ (n = 2): 40.2 mm (40.0–40.5); TL+process: ♂ (n = 4): 42.5 mm (41.0–43.2); ♀ (n = 2): 46.7 mm (46.4–47.0); LTg/BTg = 2.33; BF/BPrH = 3.0; LPr/LF = 2.18; LPr/ BPrH = 6.92. Male genitalia Pygofer and anal tube dark brown; gonostyli brown, paler dorsally and dark brown basoventrally. Pygofer higher than long, with posterior margin sinuate in lateral view (Fig. 7A). Anal tube slightly elongate, 1.20 times as long as broad, broader at 2/3 of total length (Fig. 7 A–B); lateral margins slightly sinuate (Fig. 7B) and apical margin strongly notched in dorsal view (Fig. 7B). Gonostyli (Fig. 7A) elongate with posterior margin broadly rounded (Fig. 7A, C). Distribution Known from Borneo, South Kalimantan and southern East Kalimantan, and Laut Island (Fig. 6). Biology No host tree has been identified for this species, for which records are available for the months January, April, June, September and October. Note Nagai & Porion (2004) mentioned the existence of 2 types of specimens showing (1) strongly and (2) weakly contrasted colour of the tegmina and illustrated both (Nagai & Porion 2004: figs 2, 4). They mentioned the strongly contrasted ones from Laut Island and Kutai, and the weakly contrasted ones from Loksado (confirmed by the examination of the specimens in MHNL collections by H. Labrique, pers. comm. Mar. 2015). Since then strongly contrasted specimens were found in several locations on Borneo and weakly contrasted ones on Laut Island, which seems to confirm that they occur together. The occurrence of pale and dark forms in the same species was already known from other species of Pyrops, e.g., P. sultanus (Adams & White, 1847) (see also Nagai & Porion 1996: figs 189, 191), and P. lathburii (Kirby, 1818) (see also Nagai & Porion 1996: figs 194, 196, 198).

Genus Pyrops Spinola, 1839 Pyrops Spinola, 1839: 231. Type species: Pyrops candelaria (Linnaeus, 1758) by subsequent designation by Duponchel (1840: 200). Hotinus Amyot & Serville, 1843: 490 [synonymized by Blanchard 1845: 425]. Type species: Pyrops candelaria (Linnaeus, 1758) by original designation. Diagnostic characters The characters defining the genus as given by Lallemand (1963) under Fulgora are here slightly modified, i.e., head with cephalic process, sometimes very long, narrowing progressively beyond the eyes; apically it can be dilated or even spherical. Vertex about 4 times as broad as an eye. Before eyes, genae truncate with a transverse carina which sometimes extends to vertex. Two longitudinal carinae on frons, a third median one starting on base of cephalic process. Fronto-clypeal suture usually slightly bisinuate; median carina on clypeus. Pronotum with median carina (sometimes obsolete) and a small but strongly impressed point on each side of it. Mesonotum with median and peridiscal carinae, sometimes obsolete. Tegmina at most 3 times as long as broad, with apical margin more or less rounded and with transverse veinlets on all surfaces. Clavus open and elongate, vein A1+A2 extending far towards apex. Legs slender., Published as part of Constant, J��r��me, 2015, Review of the effusus group of the Lanternfly genus Pyrops Spinola, 1839, with one new species and notes on trophobiosis (Hemiptera: Fulgoromorpha: Fulgoridae), pp. 1-23 in European Journal of Taxonomy 128 on page 4, DOI: 10.5852/ejt.2015.128, http://zenodo.org/record/3785153, {"references":["Duponchel P. A. J. 1840. Essai sur les fulgorelles, par Maximilien Spinola. Revue Zoologique, par la Societe cuvierienne; Association Universelle pour l'Advancement de la Zoologie, de l'Anatomie comparee et de la Palaeontologie; Journal mensuel 2: 199 - 206.","Blanchard E. 1845. Septieme ordre, les Hemipteres. In: Blanchard E. 1845. Histoire des insectes, traitant de leurs moeurs et de leurs metamorphoses en general et comprenant une nouvelle classification fondee sur leurs rapports naturels 2: 424 - 427.","Lallemand V. 1963. Revision des Fulgoridae (Homoptera). Deuxieme partie. Faunes asiatique et australienne. Memoires de l'Institut royal des Sciences naturelles de Belgique (Series 2) 75: 1 - 99."]}

The species group effusus of the genus Pyrops Spinola, 1839 is reviewed and the nomenclatural history of the genus Pyrops is briefly summarized. One new species from eastern Borneo, Pyrops synavei sp. nov., is described. P. gunjii (Satô & Nagai, 1994) stat. nov. is proposed as a valid species instead of a subspecies of P. whiteheadi (Distant, 1889). P. maquilinganus (Baker, 1925) is removed from the effusus group and placed back into the candelaria group. P. cyanirostris (Guérin-Méneville, 1845) is removed from the group and not attributed to any of the currently defined species groups. An illustrated key to the species of the group with the addition of P. intricatus (Walker, 1857) and a distribution map are given. The effusus group is restricted to Borneo and adjacent Laut Island and presently contains 4 species: P. effusus (Distant, 1891), P. gunjii (Satô & Nagai, 1994) stat.nov., P. synavei sp. nov. and P. whiteheadi (Distant, 1889). Trophobiosis observations with the gecko Gehyra mutilata (Wiegmann, 1835) (Reptilia: Squamata: Gekkonidae) and two species of cockroaches (Insecta: Blattodea), one Dorylaea sp. and an unidentified species of Pseudophyllodromiinae, are reported and illustrated for P. whiteheadi; observation with a cockroach, Dorylaea sp., is reported for P. intricatus.

Pyrops gunjii (Satô & Nagai, 1994) stat. nov. Figs 3 A–E, 6, 8A–C, 10B Fulgora whiteheadi gunjii Satô & Nagai 1994: 308, figs 2, 9 (Type in EUM) [described, illustrated and compared with P. whiteheadi and P. effusus, the latter mentioned as “ F. whiteheadi viridicastanea ”]. Pyrops whiteheadi gunjii – Nagai & Porion, 1996: 26, pl. 17, fig. 210 [transferred to Pyrops; listed, illustrated]. — Liang 1998: 45 [listed, erroneous mention of new combination already proposed by Nagai & Porion (1996)]. Diagnosis In addition to the characters defining the effusus group, the species can be recognized by: (1) head yellow-brown with apical half of cephalic process blue (often faded to green in collection specimens) (Fig. 3 D–E); (2) pro- and mesonotum yellow-brown (Fig. 3A); (3) tegmina dark blue-green on corium, with small yellow-white spots; on basal half, spots slightly tranverse and rarely coalescent, spots barely distinct on membrane (Fig. 3A); (4) tegmina with large pale yellow-brown patch at nodal line and apical margin broadly infuscate (Fig. 3A). Material examined Type material Holotype ♂ (examined on photographs): [Embalut, E. Kalimantan, INDONESIA, Sep. 22. 1988] [Holotype Fulgora whiteheadi gunjii, M. Sato & S. Nagai, 1994] (EUM). Note: Satô & Nagai (1994) stated that the type was collected by S. Okada and suggested that the type location was probably “Ambalut, Samarinda” rather than “Embalut”. Additional material INDONESIA: 2 ♂♂, E Kalimantan, Samarinda, Mt Beratus, Aug. 2006, S. Jakl (RBINS); 1 ♀, no data (RMNH). Examined on photographs INDONESIA: 1 specimen (Fig. 10B), East Kalimantan, 0°32’4.70”S 117°27’46.73”E, 3 Jul. 2011, Martin Hauskrecht; 1 specimen, East Kalimantan, Kutai National Park, 7 Jan. 2009, Martin Dancak; 1 specimen, East Kalimantan, Kutai National Park, 26 Sept. 2009, “Sophie & Dale”; 1 specimen, East Kalimantan, Kutai National Park, 28 Jan. 2010, Darwyn Moffatt-Mallett; 1 specimen, East Kalimantan, Penajam Paser Utara, 17 Mar. 2010, “Emjanda”. Measurements and ratios TL: ♂ (n = 2): 35.1 mm (34.5–35.6); ♀ (n = 1): 38.6 mm; TL+process: ♂ (n = 2): 39.7 mm (39.0–40.3); ♀ (n = 1): 44.0 mm; LTg/BTg = 2.53; BF/BPrH = 3.0; LPr/LF = 1.88; LPr/BPrH = 6.97. Male genitalia Dark brown with gonostyli paler dorsally. Pygofer higher than long, with posterior margin sinuate in lateral view (Fig. 8A). Anal tube slightly elongate, 1.10 times as long as broad, broader at 4/5 of total length (Fig. 8 A–B); lateral margins slightly sinuate (Fig. 8B) and apical margin strongly notched in dorsal view (Fig. 8B). Gonostyli (Fig. 8A) elongate with posterior margin roundly pointed (Fig. 8A, C). Distribution Known from Borneo, south-eastern East Kalimantan (Fig. 6). Biology No host tree has been identified for this species, for which records are available from the months of January, March, July, August and September. Remarks The species epithet refers to the Japanese entomologist Mr Yoshiaki Gunji.

Genus Pyrops Spinola, 1839 Pyrops Spinola, 1839: 231. Type species: Pyrops candelaria (Linnaeus, 1758) by subsequent designation by Duponchel (1840: 200). Hotinus Amyot & Serville, 1843: 490 [synonymized by Blanchard 1845: 425]. Type species: Pyrops candelaria (Linnaeus, 1758) by original designation. Diagnostic characters The characters defining the genus as given by Lallemand (1963) under Fulgora are here slightly modified, i.e., head with cephalic process, sometimes very long, narrowing progressively beyond the eyes; apically it can be dilated or even spherical. Vertex about 4 times as broad as an eye. Before eyes, genae truncate with a transverse carina which sometimes extends to vertex. Two longitudinal carinae on frons, a third median one starting on base of cephalic process. Fronto-clypeal suture usually slightly bisinuate; median carina on clypeus. Pronotum with median carina (sometimes obsolete) and a small but strongly impressed point on each side of it. Mesonotum with median and peridiscal carinae, sometimes obsolete. Tegmina at most 3 times as long as broad, with apical margin more or less rounded and with transverse veinlets on all surfaces. Clavus open and elongate, vein A1+A2 extending far towards apex. Legs slender.

Pyrops synavei sp. nov. urn:lsid:zoobank.org:act: B502AF69-AEC2-4245-B440-57F75BF8559E Figs 4 A–E, 6, 11A–D Diagnosis In addition to the characters defining the effusus group, the species can be recognized by: (1) head entirely dark olivaceous blue-green (Fig. 4 D–E); (2) pro- and mesonotum bluish-brown (Fig. 4A); (3) tegmina green with yellow spots also beyond nodal line; on corium, basal row of spots fused in a transverse band, next rows fused into incomplete tranverse bands (Fig. 4A); (4) tegmina without large pale yellow patch at nodal line and apical margin very slightly infuscate (Fig. 4A). Etymology The species epithet refers to the late Belgian hemipterist Henry Synave (1921–1981). He was the author of numerous publications on Auchenorrhyncha, and of all drawings of Lallemand’s revisions of the Fulgoridae (1959, 1963). Material examined Type material INDONESIA: Holotype, ♀: [N.O. Borneo, S. Peleben, 7.u.8.1935, Baron Plessen] [R.I.Sc.N.B., I.G. 23.860] [H. Synave det., 1968, Laternaria intricata Walk.] (RBINS). Coordinates of S Peleben: 2°47’N 116°35’E. Examined on photographs 1 specimen (Fig. 11 A–B): Kalimantan, Long Alango, Lalut Birai Sacred Forest, 2 Sep. 2011, Joost Woltering; 1 specimen (Fig. 11 C–D): Kalimantan, Lalut Birai Forest, 13 Mar. 2009, Dave Forney. Coordinates of Lalut Birai Forest: 3°12’N 115°30’E. Measurements and ratios TL: ♀ (n = 1): 40.9 mm; TL+process: ♀ (n = 1): 47.7 mm; LTg/BTg = 2.50; BF/BPrH = 2.78; LPr/LF = 2.48; LPr/BPrH = 6.78. Description HEAD. Dark olivaceous blue-green progressively turning more brownish basally (Fig. 4A, D–E). Cephalic process slightly longer than frons and clypeus together in normal view of frons (Fig. 4E), strongly curved anterodorsad and bent in middle dorsally (Fig. 4D); apical half slightly dilated and rounded apically in lateral view (Fig. 4D), with sides parallel in anteroventral view (Fig. 4E). Two longitudinal carinae on frons extending on sides of cephalic process up to apex, undulated. Median, ventral undulated carina on apical half of cephalic process (Fig. 4E). Frons subquadrate (Fig. 4E). Clypeus elongate with smooth median carina. Labium brown-black, slightly surpassing posterior trochanter. Antennae brown. THORAX (Fig. 4A, D). Olivaceous brown, pro- and mesonotum greener in middle. Pronotum irregularly wrinkled with smooth longitudinal carina and 2 strongly impressed points on anterior half of disc (Fig. 4A), dorsolateral and lateral carinae well marked (Fig. 4A, D). Mesonotum with disc wrinkled and sides smooth; 3 smooth longitudinal carinae not reaching anterior margin, median one reaching base of scutellum posteriorly; slight impression along posterior margin at base of scutellum; scutellum slightly elevated apically. Tegulae olivaceous brown. TEGMINA (Fig. 4A). Dark green-brown with veins green turning to brown near apical margin. 3 more or less interrupted transverse yellow bands on corium; more basal one complete; median one sometimes cut at costal vein or/and claval suture; third band in 2–3 parts often not aligned. Row of 3 more or less oval yellow spots on nodal line. Yellow markings on corium and nodal line bordered with dark brown. 8–9 round yellow spots on membrane. Apical margin oblique with angles broadly rounded; costal margin slightly rounded. HIND WINGS (Fig. 4A). Bright blue with apex and sutural margin largely brown-black; brown black extending into blue along main veins; cross-veinlets blue in brown-black area on baso-sutural half. Hind wings slightly broader than tegmina. LEGS (Fig. 4 A–B). Dark olivaceous brown. Metatibiae with 6 lateral and 7 apical spines. ABDOMEN (Fig. 4 A–B). Tergites brown-black; sternites black; genital segments tinged with yellowbrown. Distribution Known from Borneo, North Kalimantan (see map Fig. 6). Biology No host tree has been identified for this species for which records are available from the months of March, July – August and September.

Pyrops whiteheadi (Distant, 1889) Figs 5 A���E, 6, 9A���C, 12A���E Fulgora whiteheadi Distant 1889: 422 (Type in NHM) [described and compared to P. delesserti (Gu��rin- M��neville, 1840)]. Fulgora whiteheadi ��� Distant 1891: 518 [compared to P. effusus]. ��� Lallemand 1963: 88 [considered as senior synonym of P. effusus and P. viridicastaneus (erroneous); described, keyed; interpretation of the species mixing P. effusus and P. whiteheadi]. ��� Sat�� & Nagai 1994: 308 [compared with P. gunjii]. Fulgora effusa ��� Baker 1925: 359 [comparison between P. effusus and P. whiteheadi based on specimens of P. whiteheadi erroneously identified as P. effusus]. Laternaria whiteheadi ��� Metcalf 1947 [transferred to Laternaria; catalogued]. Pyrops whiteheadi ��� Nagai & Porion 1996: 26 [transferred to Pyrops; listed], pl. 11, fig. 160 [type illustrated]. ��� Liang 1998: 45 [listed]. ��� Nagai & Porion 2004: 6 [comparison with P. effusus], pl. 1, fig. 6 [illustration of habitus]. non Fulgora whiteheadi ��� Lallemand 1963: pl. X, figs 11���13 [illustration of the genitalia of P. effusus erroneously identified as P. whiteheadi]. non Pyrops whiteheadi ��� Nagai & Porion 1996: pl. 17, fig. 213 [illustration of a specimen of P. effusus erroneously identified as P. whiteheadi]. Diagnosis In addition to the characters defining the effusus group, the species can be recognized by: (1) head entirely blue (often faded to greenish in collection specimens) (Fig. 5 D���E); (2) pronotum yellow-brown, mesonotum blue-brown (Fig. 5A); (3) tegmina green with small yellow spots also beyond nodal line; spots rounded and not coalescent (Fig. 5A); (4) tegmina without large pale yellow patch at nodal line and with apical margin slightly infuscate (Fig. 5A). Material examined Type material MALAYSIA: Lectotype: ♂, here designated to provide a reference standard for the species (examined on photographs): East Malaysia, [Kina Balu (Whitehead)] [Whiteheadi Dist.] [Distant Coll. 1911��� 383.] [Type] (BMNH). MALAYSIA: Paralectotypes: 2 ♂♂, 2 ♀♀ (examined on photographs): East Malaysia, [Kina Balu (Whitehead)] [Whiteheadi Dist.] [Distant Coll. 1911���383.] (BMNH). Additional material MALAYSIA: 1 ♂, Sabah, Kinabalu National Park, Poring Hot Spring, 21 Apr. 1999, T. Trilar (RBINS); 1 ♀, Sabah, 23 km W of Sandakan, Sepilok tree tower, 5��49���N 118��06���E, 1 Nov. 1987, 0���100 m, J. Huisman & R. de Jong (RMNH); 1 ♂, Sabah, Danum Valley, 70 km W of Lahad Datu, rainforest along narrow stream, at light, next to bridge Nature Trail, 50 m, 1 Dec. 1989, M.J. & J.P. Duffels (RMNH). Note One specimen labelled: Selangor, Banting, 2 Oct. [19]89, Ismail & Ruslan in the collections of BMNH, has been examined on photograph. The location is in West Malaysia and the specimen is assumed to be mislabelled. Examined on photographs EAST MALAYSIA: 1 specimen (Fig. 12 A���B), Sabah, Danum Valley, 4 Jul. 2013, Anton Sorokin; 1 specimen, idem, 13 Jun. 2013; 1 specimen, idem, 3 Oct. 2010, Hok Ping Guek; 1 specimen, idem, 4 Oct. 2013; 1 specimen, idem, 18 Aug. 2008, Thomas Bridle; 2 specimens, idem, 27 Sept. 2009, Tim Moss; 1 specimen, idem, 3 Jun. 2007, Karin Saner; 1 specimen, idem, Aug. 2004, Dirk Mezger; 1 specimen (Fig. 12C), Sabah, Danum Valley, 15 Dec. 2008, Ch���ien C. Lee; 1 specimen (Fig. 12D), Sabah, Lower Kinabatangan River, 9 Aug. 2012, Nicola Messina; 1 specimen (Fig. 12E), Sabah, Sepilok, 6 Jun. 2013, Ch���ien C. Lee; 2 specimens, Keningau, 24 Sep. 2011, Khairin Saili; 2 specimens, Kinabatangan, 14 Feb. 2008, Christophe Maerten; 1 specimen, idem, 21 Jan. 2008, Andrea Ferrari; 1 specimen, idem, 17 Oct. 2009, ���Pietra & Paoli���; 1 specimen, idem, 24 May 2010, Frank Joas; 1 specimen, Kinabatangan, Sukau, 12 Aug. 2011, J.M. Gayman; 1 specimen, Sandakan, 27 Mar. 2013, Gavin Golden; 1 specimen, Sandakan, Sukau, 14 Aug. 2009, Samuel Tan; 1 specimen, Taman Bukit Tawau, 17 Aug. 2012, Subki Abdul Hadi; 1 specimen, Sepilok, 1 Aug. 2010, J.P. Lawrence; 1 specimen, idem, 15 Aug. 2011. Measurements and ratios TL: ♂ (n = 2): 34.8 mm (34.3���35.2); ♀ (n = 1): 38.0 mm; TL+process: ♂ (n = 2): 39.8 mm (38.7���40.9); ♀ (n = 1): 42.2 mm; LTg/BTg = 2.73; BF/BPrH = 3.0; LPr/LF = 2.5; LPr/BPrH = 5.4. Male genitalia Brown with gonostyli paler dorsally and ventrally. Pygofer higher than long, with posterior margin sinuate in lateral view (Fig. 9A). Anal tube slightly elongate, 1.25 times as long as broad, broader at 4/5 of total length (Fig. 9 A���B); lateral margins very slightly sinuate (Fig. 9B) and apical margin strongly notched in dorsal view (Fig. 9B). Gonostyli (Fig. 9A) elongate with posterior margin rounded, slightly narrowing (Fig. 9A, C). Distribution Borneo, known from Sabah (Fig. 6). Biology No host tree has been identified for this species, for which records are available from all months of the years, with a higher number of records (8/26) in August. However, we do not conclude that the species is more abundant in August; as most data are from photographs taken by tourists, it is possible that the higher amount of available data is biased by the fact that August is a holiday period in many countries. Notes on trophobiosis with Pyrops whiteheadi and P. intricatus Trophobiosis with ants commonly occurs in Hemiptera, and ant-attendance involving Fulgoromorpha was reviewed by Bourgoin (1997) who recorded the phenomenon in 5 families of Fulgoromorpha (Cixiidae, Delphacidae, Issidae, Hypochthonellidae and Tettigometridae). He recognized 4 main types of ant-attendance, from opportunistic attendance by ants to long term attendance where ants collect honeydew directly from the anal opening of the planthopper. At the time, no interaction was known to involve Fulgoridae. Trophobiotic interactions between Fulgoridae and other animals attracted by the droplets of honeydew produced by the lanternflies were only recently reported by (1) Roth & Naskrecki (2001), involving Neotropical species of the genera Copidocephala St��l, 1869 and Enchophora Spinola, 1839, with Blattodea; (2) Naskrecki & Nishida (2007), involving Neotropical species of the genera Copidocephala, Enchophora and Phrictus Spinola, 1839, with Blattodea, Lepidoptera and gastropods; (3) Kemal & Ko��ak (2012) between Pyrops candelaria (Linnaeus, 1758) and the gecko Hemidactylus platyurus (Schneider, 1792) (Squamata: Gekkonidae) in Thailand. Interactions between geckos (5 species belonging to 3 different genera) and another family of Fulgoroidea, Flatidae, in Madagascar, were previously reported and analyzed by F��lling et al. (2001). Similar interactions have been observed and documented between Pyrops whiteheadi and two species of Blattodea in Sabah, Danum Valley: one species of the subfamily Pseudophyllodromiinae (C.C. Lee pers. comm., Fig. 12C) and one species of the genus Dorylaea (A. Sorokin pers. comm., Fig. 12 A���B), possibly D. magna (Schelford, 1909) (identifications of the Blattodea by G. Beccaloni, Dec. 2014). Interactions between one species of gecko (Squamata: Gekkonidae), Gehyra mutilata (Wiegmann, 1835) (identification by O. Pauwels, Dec. 2014) and P. whiteheadi was also observed and documented in Sabah, Sepilok (C.C. Lee pers. comm., Fig. 12E). Geckos generally are attracted by sugar-rich food (O. Pauwels pers. comm., 2014). Although it does not belong to the effusus group, I should also mention that interactions between Pyrops intricatus (Walker, 1857) and Blattodea of the genus Dorylaea have been documented in Sarawak, Mulu National Park (C.C. Lee pers. comm., Fig. 12F). The unavailabily of such observations for the three other species of the effusus group can probably be explained by the fact that P. whiteheadi and P. intricatus are present and apparently rather common in several preserved areas (e.g., Danum Valley, Sepilok, Kinabatangan for the first, Mulu for the second) which are extensively visited by naturalists and ecotourists, while the other species are found in more remote regions. All observations reported here come from such sources and it is likely that the same interactions will be documented for the other species in the future., Published as part of Constant, J��r��me, 2015, Review of the effusus group of the Lanternfly genus Pyrops Spinola, 1839, with one new species and notes on trophobiosis (Hemiptera: Fulgoromorpha: Fulgoridae), pp. 1-23 in European Journal of Taxonomy 128 on pages 13-21, DOI: 10.5852/ejt.2015.128, http://zenodo.org/record/3785153, {"references":["Distant W. L. 1889. XLIX. Descriptions of new species of Rhynchota collected on or near the Kina Balu Mountain, North Borneo. Annals and Magazine of Natural History (Series 6) 3: 419 - 422.","Distant W. L. 1891. XXIII. Descriptions of four new species of the genus Fulgora. Transactions of the Entomological Society of London 1891: 517 - 519.","Lallemand V. 1963. Revision des Fulgoridae (Homoptera). Deuxieme partie. Faunes asiatique et australienne. Memoires de l'Institut royal des Sciences naturelles de Belgique (Series 2) 75: 1 - 99.","Sato M. & Nagai S. 1994. Taxonomic notes on the Southeast Asian species of the genus Fulgora (Homoptera: Fulgoridae). Transactions of the Shikoku Entomological Society 20 (3 - 4): 307 - 314.","Baker C. F. 1925. Remarks on certain Indo-Malayan Fulgora, with special reference to Philippines species. Philippines Journal of Science 28: 343 - 364.","Metcalf Z. P. 1947. General Catalogue of the Homoptera. Fascicle IV Fulgoroidea. Part 9 Fulgoridae. North Carolina State College, Raleigh (U. S. A.).","Nagai S. & Porion T. 1996. Fulgoridae 2: Catalogue illustre des faunes asiatique et australienne. Sciences Nat, Compiegne.","Liang A. - P. 1998. Nomenclatorial notes on the Oriental lantern fly genus Pyrops Spinola (Hemiptera: Fulgoroidea: Fulgoridae). Acta Zootaxonomica Sinica 23 (1): 41 - 47.","Nagai S. & Porion T. 2004. Fulgoridae 2, supplement 2: Nouveaux Fulgoridae d'Asie du Sud-Est. Hillside Books, Canterbury.","Bourgoin T. 1997. Habitat and ant-attendance in Hemiptera: a phylogenetic test with emphasis on trophobiosis in Fulgoromorpha. In: Grandcolas P. (ed.) The Origin of Biodiversity in Insects: Phylogenetic Tests of Evolutionary Scenarios. Memoires du Museum National d'Histoire Naturelle 173: 109 - 124. Museum National d'Histoire Naturelle, Paris.","Roth L. & Naskrecki P. 2001. Trophobiosis between a blatellid cockroach (Macrophyllodromia spp.) and fulgorids (Enchophora and Copidocephala spp.) in Costa Rica. Journal of Orthoptera Research 10: 189 - 194.","Naskrecki P. & Nishida K. 2007. Novel trophobiotic interactions in lantern bugs (Insecta: Auchenorrhyncha: Fulgoridae). Journal of Natural History 41 (37 - 40): 2397 - 2402. http: // dx. doi. org / 10.1080 / 00222930701633570","Kemal M. & Kocak A. O. 2012. A report on the mutualism between Hemidactylus (Gekkonidae) and","Folling M., Knogge C. & Bohme W. 2001. Geckos are milking honeydew-producing planthoppers in Madagascar. Journal of Natural History 35 (2): 279 - 284. http: // dx. doi. org / 10.1080 / 00222930150215378"]}

Pyrops synavei sp. nov. urn:lsid:zoobank.org:act: B502AF69-AEC2-4245-B440-57F75BF8559E Figs 4 A���E, 6, 11A���D Diagnosis In addition to the characters defining the effusus group, the species can be recognized by: (1) head entirely dark olivaceous blue-green (Fig. 4 D���E); (2) pro- and mesonotum bluish-brown (Fig. 4A); (3) tegmina green with yellow spots also beyond nodal line; on corium, basal row of spots fused in a transverse band, next rows fused into incomplete tranverse bands (Fig. 4A); (4) tegmina without large pale yellow patch at nodal line and apical margin very slightly infuscate (Fig. 4A). Etymology The species epithet refers to the late Belgian hemipterist Henry Synave (1921���1981). He was the author of numerous publications on Auchenorrhyncha, and of all drawings of Lallemand���s revisions of the Fulgoridae (1959, 1963). Material examined Type material INDONESIA: Holotype, ♀: [N.O. Borneo, S. Peleben, 7.u.8.1935, Baron Plessen] [R.I.Sc.N.B., I.G. 23.860] [H. Synave det., 1968, Laternaria intricata Walk.] (RBINS). Coordinates of S Peleben: 2��47���N 116��35���E. Examined on photographs 1 specimen (Fig. 11 A���B): Kalimantan, Long Alango, Lalut Birai Sacred Forest, 2 Sep. 2011, Joost Woltering; 1 specimen (Fig. 11 C���D): Kalimantan, Lalut Birai Forest, 13 Mar. 2009, Dave Forney. Coordinates of Lalut Birai Forest: 3��12���N 115��30���E. Measurements and ratios TL: ♀ (n = 1): 40.9 mm; TL+process: ♀ (n = 1): 47.7 mm; LTg/BTg = 2.50; BF/BPrH = 2.78; LPr/LF = 2.48; LPr/BPrH = 6.78. Description HEAD. Dark olivaceous blue-green progressively turning more brownish basally (Fig. 4A, D���E). Cephalic process slightly longer than frons and clypeus together in normal view of frons (Fig. 4E), strongly curved anterodorsad and bent in middle dorsally (Fig. 4D); apical half slightly dilated and rounded apically in lateral view (Fig. 4D), with sides parallel in anteroventral view (Fig. 4E). Two longitudinal carinae on frons extending on sides of cephalic process up to apex, undulated. Median, ventral undulated carina on apical half of cephalic process (Fig. 4E). Frons subquadrate (Fig. 4E). Clypeus elongate with smooth median carina. Labium brown-black, slightly surpassing posterior trochanter. Antennae brown. THORAX (Fig. 4A, D). Olivaceous brown, pro- and mesonotum greener in middle. Pronotum irregularly wrinkled with smooth longitudinal carina and 2 strongly impressed points on anterior half of disc (Fig. 4A), dorsolateral and lateral carinae well marked (Fig. 4A, D). Mesonotum with disc wrinkled and sides smooth; 3 smooth longitudinal carinae not reaching anterior margin, median one reaching base of scutellum posteriorly; slight impression along posterior margin at base of scutellum; scutellum slightly elevated apically. Tegulae olivaceous brown. TEGMINA (Fig. 4A). Dark green-brown with veins green turning to brown near apical margin. 3 more or less interrupted transverse yellow bands on corium; more basal one complete; median one sometimes cut at costal vein or/and claval suture; third band in 2���3 parts often not aligned. Row of 3 more or less oval yellow spots on nodal line. Yellow markings on corium and nodal line bordered with dark brown. 8���9 round yellow spots on membrane. Apical margin oblique with angles broadly rounded; costal margin slightly rounded. HIND WINGS (Fig. 4A). Bright blue with apex and sutural margin largely brown-black; brown black extending into blue along main veins; cross-veinlets blue in brown-black area on baso-sutural half. Hind wings slightly broader than tegmina. LEGS (Fig. 4 A���B). Dark olivaceous brown. Metatibiae with 6 lateral and 7 apical spines. ABDOMEN (Fig. 4 A���B). Tergites brown-black; sternites black; genital segments tinged with yellowbrown. Distribution Known from Borneo, North Kalimantan (see map Fig. 6). Biology No host tree has been identified for this species for which records are available from the months of March, July ��� August and September., Published as part of Constant, J��r��me, 2015, Review of the effusus group of the Lanternfly genus Pyrops Spinola, 1839, with one new species and notes on trophobiosis (Hemiptera: Fulgoromorpha: Fulgoridae), pp. 1-23 in European Journal of Taxonomy 128 on pages 11-13, DOI: 10.5852/ejt.2015.128, http://zenodo.org/record/3785153

Pyrops effusus (Distant, 1891) Figs 2 A���E, 6���7A���C, 10A, C���F Fulgora effusa Distant, 1891: 518, pl. XX, figs 4, 4a (Type in NHM) [described, illustrated and compared with P. whiteheadi (Distant, 1889)]. Fulgora viridicastanea Baker, 1925: 359, pl. 3, fig. 5, pl. 4 fig. 8 (Type in USNM) [described, illustrated and compared with P. whiteheadi erroneously mentioned and illustrated as ��� Fulgora effusa ���]. Synonymized under P. effusus by Nagai & Porion (2004). Fulgora effusa ��� Schmidt 1905: 349 [comparative note with other species]; 354 [mention of a specimen in the Stettiner Museum]. ��� Lallemand 1963: 88 [synonymized under whiteheadi (erroneous)]. ��� Liang 1998: 45 [listed as junior synonym of P. whiteheadi (erroneous)]. Pyrops effusa (sic!) ��� Schmidt 1926: 47 [transferred to Pyrops; compared with P. peltzeri Schmidt, 1926, the latter synonymized under P. pythicus (Distant, 1891) by Lallemand (1963)]. ��� Nagai & Porion 1996: 26 [transferred back to Pyrops; catalogued as junior synonym of P. whiteheadi (erroneous)], pl. 11, fig. 163 [type of P. effusus illustrated under the name of P. whiteheadi]. ��� Nagai & Porion 2004: 6 [removal from synonymy with P. whiteheadi; notes on morphology; senior synonym of P. viridicastaneus and distribution], pl. 1, figs 2, 4 [illustrated from 2 localities]. Laternaria effusa ��� Metcalf 1947: 196 [transferred to Laternaria; catalogued]. ��� Liang 1998: 45 [listed as junior synonym of P. whiteheadi (erroneous)]. Laternaria viridicastanea ��� Metcalf 1947: 207 [transferred to Laternaria; catalogued]. ��� Liang 1998: 45 [listed as junior synonym of P. whiteheadi (erroneous)]. Fulgora viridicastanea ��� Lallemand 1963: 88 [synonymized under P. whiteheadi (erroneous) and compared with P. whiteheadi and P. effusus; transferred back to Fulgora (erroneous)]. ��� Liang 1998: 45 [listed as junior synonym of P. whiteheadi (erroneous)]. Fulgora whiteheadi ��� Lallemand 1963: 88, pl. X, figs 11���13 [keyed, comments illustration of male genitalia (based on misidentified specimens of P. effusus)]. Fulgora whiteheadi viridicastanea ��� Sat�� & Nagai 1994 [considered as a subspecies of P. whiteheadi (erroneous) and compared with ��� Fulgora whiteheadi gunjii ���]. Pyrops viridicastanea ��� Nagai & Porion 1996: 26 [transferred back to Pyrops; catalogued as junior synonym of P. whiteheadi (erroneous)]. ��� Nagai & Porion 2004: 6 [synonymized under P. effusus, notes]. Pyrops whiteheadi (Distant, 1889) ��� Nagai & Porion 1996, pl. 11, fig. 163 [illustration of the type of P. effusus (Distant, 1891)]. non Fulgora effusa ��� Baker 1925: 359, pl. 3, fig. 4, pl. 4, fig. 7 [described, illustrated (misidentification of Pyrops whiteheadi)]. Diagnosis In addition to the characters defining the effusus group, the species can be recognized by: (1) head yellow-brown with apical half of cephalic process blue (often faded to greenish in collection specimens) (Fig. 2 D���E); (2) pro- and mesonotum yellow-brown (Fig. 2A); (3) tegmina green with yellow spots also beyond nodal line; on basal half, spots tranverse and sometimes coalescent (Fig. 2A); (4) tegmina without large pale yellow patch at nodal line and apical margin slightly infuscate (Fig. 2A). Material examined Type material INDONESIA: Holotype of Fulgora effusa (examined on photographs), ♂: [Borneo] [effusa Dist.] [Distant Coll. 1911 ���383.] (BMNH). INDONESIA: Holotype of Fulgora viridicastanea (examined on photographs, Fig. 10 C���F), ♀: [Banjermassin, Borneo] [Fulgora viridicastanea Baker] (USNM). Additional material INDONESIA: 2 ♂♂, 2 ♀♀: Kalimantan, Batulicin, Apr. 2008 (RBINS); 2 ♂♂: Laut Island, Jan. 2008 (RBINS); 1 ♀, Laut Island, Jun. 2004 (RBINS); 1 ♂, E-Borneo, Midden, 10 Oct. 1925, H.C. Siebers (FSAG); 1 ♂, idem, 4 Oct. 1925 (FSAG); 1 ♂, SE Borneo, Atkinson coll. (BMNH); 1 ♂, 1 ♀, Borneo, 1909 (BMNH); 1 ♀, North Borneo (BMNH). Examined on photograph INDONESIA: 1 specimen (Fig. 10A), Kalimantan, Wehea Protected Forest, 25 Sep. 2010, Kara Norby. Measurements and ratios TL: ♂ (n = 4): 35.1 mm (34.3���36.0); ♀ (n = 2): 40.2 mm (40.0���40.5); TL+process: ♂ (n = 4): 42.5 mm (41.0���43.2); ♀ (n = 2): 46.7 mm (46.4���47.0); LTg/BTg = 2.33; BF/BPrH = 3.0; LPr/LF = 2.18; LPr/ BPrH = 6.92. Male genitalia Pygofer and anal tube dark brown; gonostyli brown, paler dorsally and dark brown basoventrally. Pygofer higher than long, with posterior margin sinuate in lateral view (Fig. 7A). Anal tube slightly elongate, 1.20 times as long as broad, broader at 2/3 of total length (Fig. 7 A���B); lateral margins slightly sinuate (Fig. 7B) and apical margin strongly notched in dorsal view (Fig. 7B). Gonostyli (Fig. 7A) elongate with posterior margin broadly rounded (Fig. 7A, C). Distribution Known from Borneo, South Kalimantan and southern East Kalimantan, and Laut Island (Fig. 6). Biology No host tree has been identified for this species, for which records are available for the months January, April, June, September and October. Note Nagai & Porion (2004) mentioned the existence of 2 types of specimens showing (1) strongly and (2) weakly contrasted colour of the tegmina and illustrated both (Nagai & Porion 2004: figs 2, 4). They mentioned the strongly contrasted ones from Laut Island and Kutai, and the weakly contrasted ones from Loksado (confirmed by the examination of the specimens in MHNL collections by H. Labrique, pers. comm. Mar. 2015). Since then strongly contrasted specimens were found in several locations on Borneo and weakly contrasted ones on Laut Island, which seems to confirm that they occur together. The occurrence of pale and dark forms in the same species was already known from other species of Pyrops, e.g., P. sultanus (Adams & White, 1847) (see also Nagai & Porion 1996: figs 189, 191), and P. lathburii (Kirby, 1818) (see also Nagai & Porion 1996: figs 194, 196, 198)., Published as part of Constant, J��r��me, 2015, Review of the effusus group of the Lanternfly genus Pyrops Spinola, 1839, with one new species and notes on trophobiosis (Hemiptera: Fulgoromorpha: Fulgoridae), pp. 1-23 in European Journal of Taxonomy 128 on pages 6-9, DOI: 10.5852/ejt.2015.128, http://zenodo.org/record/3785153, {"references":["Distant W. L. 1891. XXIII. Descriptions of four new species of the genus Fulgora. Transactions of the Entomological Society of London 1891: 517 - 519.","Distant W. L. 1889. XLIX. Descriptions of new species of Rhynchota collected on or near the Kina Balu Mountain, North Borneo. Annals and Magazine of Natural History (Series 6) 3: 419 - 422.","Baker C. F. 1925. Remarks on certain Indo-Malayan Fulgora, with special reference to Philippines species. Philippines Journal of Science 28: 343 - 364.","Nagai S. & Porion T. 2004. Fulgoridae 2, supplement 2: Nouveaux Fulgoridae d'Asie du Sud-Est. Hillside Books, Canterbury.","Lallemand V. 1963. Revision des Fulgoridae (Homoptera). Deuxieme partie. Faunes asiatique et australienne. Memoires de l'Institut royal des Sciences naturelles de Belgique (Series 2) 75: 1 - 99.","Liang A. - P. 1998. Nomenclatorial notes on the Oriental lantern fly genus Pyrops Spinola (Hemiptera: Fulgoroidea: Fulgoridae). Acta Zootaxonomica Sinica 23 (1): 41 - 47.","Nagai S. & Porion T. 1996. Fulgoridae 2: Catalogue illustre des faunes asiatique et australienne. Sciences Nat, Compiegne.","Metcalf Z. P. 1947. General Catalogue of the Homoptera. Fascicle IV Fulgoroidea. Part 9 Fulgoridae. North Carolina State College, Raleigh (U. S. A.).","Sato M. & Nagai S. 1994. Taxonomic notes on the Southeast Asian species of the genus Fulgora (Homoptera: Fulgoridae). Transactions of the Shikoku Entomological Society 20 (3 - 4): 307 - 314."]}

Pyrops gunjii (Sat�� & Nagai, 1994) stat. nov. Figs 3 A���E, 6, 8A���C, 10B Fulgora whiteheadi gunjii Sat�� & Nagai 1994: 308, figs 2, 9 (Type in EUM) [described, illustrated and compared with P. whiteheadi and P. effusus, the latter mentioned as ��� F. whiteheadi viridicastanea ���]. Pyrops whiteheadi gunjii ��� Nagai & Porion, 1996: 26, pl. 17, fig. 210 [transferred to Pyrops; listed, illustrated]. ��� Liang 1998: 45 [listed, erroneous mention of new combination already proposed by Nagai & Porion (1996)]. Diagnosis In addition to the characters defining the effusus group, the species can be recognized by: (1) head yellow-brown with apical half of cephalic process blue (often faded to green in collection specimens) (Fig. 3 D���E); (2) pro- and mesonotum yellow-brown (Fig. 3A); (3) tegmina dark blue-green on corium, with small yellow-white spots; on basal half, spots slightly tranverse and rarely coalescent, spots barely distinct on membrane (Fig. 3A); (4) tegmina with large pale yellow-brown patch at nodal line and apical margin broadly infuscate (Fig. 3A). Material examined Type material Holotype ♂ (examined on photographs): [Embalut, E. Kalimantan, INDONESIA, Sep. 22. 1988] [Holotype Fulgora whiteheadi gunjii, M. Sato & S. Nagai, 1994] (EUM). Note: Sat�� & Nagai (1994) stated that the type was collected by S. Okada and suggested that the type location was probably ���Ambalut, Samarinda��� rather than ���Embalut���. Additional material INDONESIA: 2 ♂♂, E Kalimantan, Samarinda, Mt Beratus, Aug. 2006, S. Jakl (RBINS); 1 ♀, no data (RMNH). Examined on photographs INDONESIA: 1 specimen (Fig. 10B), East Kalimantan, 0��32���4.70���S 117��27���46.73���E, 3 Jul. 2011, Martin Hauskrecht; 1 specimen, East Kalimantan, Kutai National Park, 7 Jan. 2009, Martin Dancak; 1 specimen, East Kalimantan, Kutai National Park, 26 Sept. 2009, ���Sophie & Dale���; 1 specimen, East Kalimantan, Kutai National Park, 28 Jan. 2010, Darwyn Moffatt-Mallett; 1 specimen, East Kalimantan, Penajam Paser Utara, 17 Mar. 2010, ���Emjanda���. Measurements and ratios TL: ♂ (n = 2): 35.1 mm (34.5���35.6); ♀ (n = 1): 38.6 mm; TL+process: ♂ (n = 2): 39.7 mm (39.0���40.3); ♀ (n = 1): 44.0 mm; LTg/BTg = 2.53; BF/BPrH = 3.0; LPr/LF = 1.88; LPr/BPrH = 6.97. Male genitalia Dark brown with gonostyli paler dorsally. Pygofer higher than long, with posterior margin sinuate in lateral view (Fig. 8A). Anal tube slightly elongate, 1.10 times as long as broad, broader at 4/5 of total length (Fig. 8 A���B); lateral margins slightly sinuate (Fig. 8B) and apical margin strongly notched in dorsal view (Fig. 8B). Gonostyli (Fig. 8A) elongate with posterior margin roundly pointed (Fig. 8A, C). Distribution Known from Borneo, south-eastern East Kalimantan (Fig. 6). Biology No host tree has been identified for this species, for which records are available from the months of January, March, July, August and September. Remarks The species epithet refers to the Japanese entomologist Mr Yoshiaki Gunji., Published as part of Constant, J��r��me, 2015, Review of the effusus group of the Lanternfly genus Pyrops Spinola, 1839, with one new species and notes on trophobiosis (Hemiptera: Fulgoromorpha: Fulgoridae), pp. 1-23 in European Journal of Taxonomy 128 on pages 9-11, DOI: 10.5852/ejt.2015.128, http://zenodo.org/record/3785153, {"references":["Sato M. & Nagai S. 1994. Taxonomic notes on the Southeast Asian species of the genus Fulgora (Homoptera: Fulgoridae). Transactions of the Shikoku Entomological Society 20 (3 - 4): 307 - 314.","Nagai S. & Porion T. 1996. Fulgoridae 2: Catalogue illustre des faunes asiatique et australienne. Sciences Nat, Compiegne.","Liang A. - P. 1998. Nomenclatorial notes on the Oriental lantern fly genus Pyrops Spinola (Hemiptera: Fulgoroidea: Fulgoridae). Acta Zootaxonomica Sinica 23 (1): 41 - 47."]}

Five new Asian Fulgoridae (Hemiptera : Fulgoromorpha) Five new species of Fulgoridae are described and illustrated, based on specimens collected in Borneo, Sulawesi and Luzon., Cinq nouvelles espèces de Fulgoridae sont décrites et illustrées, à partir de spécimens collectés à Bornéo, Sulawesi et Luzon., Chew Kea Foo Steven, Porion Thierry, Audibert Cédric. Cinq nouveaux Fulgoridae asiatiques fil (Hemiptera : Fulgoromorpha). In: Les Cahiers du Musée des Confluences. Etudes Scientifiques, tome 1, 2010. pp. 51-63.