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 (Guérin-Méneville, 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]. — Satô & 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 Stål, 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 & Koçak (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 Fölling 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, Jérôme, 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