Review of the effusus group of the Lanternfly genus Pyrops Spinola, 1839, with one new species and notes on trophobiosis (Hemiptera: Fulgoromorpha: Fulgoridae)

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.


Introduction
The Lanternfly genus Pyrops spinola, 1839 contains colourful, spectacular and popular species which are immediately recognized as Lanternflies by all entomologists because of their typical cephalic process. It is widely distributed in the Indomalayan region, from sri Lanka to the Himalayas (north India, south China), eastwards to taiwan and Vietnam, and southwards to sulawesi and neighbouring islands through Indonesia and the Philippines. It currently covers 58 species and 16 subspecies (Bourgoin 2015), but despite the publication of revisions and catalogues (e.g., Lallemand 1963;nagai & Porion 1996) its taxonomy is far from comprehensively resolved and still contains many errors and inconsistencies (Constant, unpublished). the genus was divided into 5 species groups by Baker (1925) [at the time under the genus name Fulgora Linnaeus, 1767], a view followed by Lallemand (1963) [also under Fulgora] and by nagai & Porion, 1996, both of the latter with minor changes.
the issue concerning the inconsistent use of the genus names Fulgora Linnaeus, 1767, Pyrops spinola, 1839 and Laternaria Linnaeus, 1764, despite the wise treatment proposed by Kirkaldy (1902), was finally fixed in a similar way by a decision of the International Commission on Zoological Nomenclature (1955), who designated Cicada laternaria Linnaeus, 1758 as the type-species of the genus Fulgora, which is restricted to the Neotropics (the "peanut-headed lanternflies"). It also suppressed the genus name Laternaria because its type species was also Cicada laternaria. Consequently, the genus name Pyrops, with Cicada candelaria Linnaeus, 1758 as type-species, validly grouped the species of south-East asia related to candelaria. However Lallemand (1960aLallemand ( ,b, 1963, Chou et al. (1985) and satô & nagai (1994) continued erroneously using Fulgora instead of Pyrops. nagai & Porion (1996) correctly used Pyrops for all species of sE asia, but left species epithets feminine, an error corrected later by Liang (1998), who erroneously stated that he proposed new combinations (most probably because he was not aware of nagai & Porion's (1996) work) and that the decision of the International Commission on Zoological Nomenclature (1955) contained a formal designation of the genus name Pyrops for the species of the oriental region. sadly enough, some recent papers still contain and spread incorrect use of the name Laternaria, e.g., Leong et al. (2009), who even state that the use of Pyrops is incorrect because of the priority of Laternaria (sic!), or Kemal & Koçak (2012).
the effusus group in the genus Pyrops spinola, 1839 was erected by Baker (1925), at that time as the effusa group in Fulgora. He defined the effusus group for medium to small sized species sharing characters such as the shape of the cephalic process, which is "very short but not very stout", apically more or less swollen or vertically expanded, in most cases strongly bent or curved upwards and at least partly bright blue-green. the tegmina are greenish basally with various yellowish dots or markings, and castaneous apically, the hind wings blue basally and broadly black apically. He included 3 species in the group: P. effusus (Distant, 1891), P. viridicastaneus (Baker, 1925) and P. transversolineatus (Baker, 1925), the latter two species described in the same paper. He actually erroneously identified specimens of P. whiteheadi (Distant, 1889) under the name effusus and redescribed P. effusus as a new species, P. viridicastaneus. the placement of P. transversolineatus in the group is strange as the shape of the head does not match the characters given for the group and the author even mentioned similarities with the genera Saiva Distant, 1906 andProlepta Walker, 1851 or with two species he placed in the candelaria group: Pyrops maquilinganus (Baker, 1925) and P. delesserti (Guérin-Méneville, 1840). Lallemand (1963) renamed Baker's (1925) effusus group "quatrième groupe: type whiteheadi", most probably because he synonymized effusus under whiteheadi. He restricted the characters of the group to those of the cephalic process: less than 12 mm long, rounded apically, forming a little ball and more or less club-shaped. He included 4 species in the group: P. maquilinganus (Baker, 1925), P. dimotus (Lallemand, 1960(Lallemand, ), P. sapphirinus (schmidt, 1908 and P. whiteheadi. He transferred P. transversolineatus to the genus Saiva Distant, 1906 and synonymized P. effusus and P. viridicastaneus with P. whiteheadi. He also mentioned that P. dimotus shows intermediate characters between Saiva and Pyrops, and that it might only be a variety of P. sapphirinus. Finally, he stated that whiteheadi was a variable species and that in the series of specimens he had identified as whiteheadi from eastern Borneo, one perfectly matched the description of P. effusus while another matched that of viridicastaneus. strangely enough, that statement seems to have been given to justify the synonymy of effusus and viridicastaneus with whiteheadi. actually, Lallemand's collection contains 2 specimens of P. effusus erroneously identified by him as P. whiteheadi. accordingly, the male genitalia illustrated in Lallemand's (1963) work (pl. X, figs 11-13) and erroneously attributed to P. whiteheadi, pertain to P. effusus.
satô & nagai (1994) described and illustrated Fulgora whiteheadi gunjii from Borneo, perpetuating the erroneous use of Fulgora instead of Pyrops, and considered viridicastaneus Baker, 1925 as a subspecies of P. whiteheadi. Nagai & Porion (1996) followed the classification proposed by Lallemand (1963), but correctly used Pyrops as a genus name. In the "groupe IV", as defined by Lallemand, they included 3 species and one subspecies: P. cyanirostris (Guérin-Méneville, 1845), P. maquilinganus, P. whiteheadi and P. whiteheadi gunjii. they synonymized dimotus and sapphirinus with cyanirostris, and followed Lallemand in considering effusus and viridicastaneus as junior synonyms of whiteheadi. they illustrated the types of P. dimotus, P. effusus, P. whiteheadi and P. whiteheadi gunjii, as well as specimens of P. cyanirostris, P. maquilinganus and one specimen of P. effusus from Lallemand's collection erroneously cited under the name P. whiteheadi.
Later, in 2004, nagai & Porion reconsidered the nomenclature of P. whiteheadi and its synonyms and illustrated specimens of P. whiteheadi and P. effusus. they reinstated P. effusus as a valid species with P. viridicastaneus as a junior synonym. they erroneously mentioned 1960 instead of 1963 as the date of publication of Lallemand's revision. the present paper aims at clarifying the nomenclature and proposing an accurately illustrated identification key for the species of the group, and at describing a new species from eastern Borneo. The current knowledge of the distribution of the species of the effusus group is illustrated by a map.
The updated nomenclature of all cited taxa will be documented in FLOW (Bourgoin 2015).

Materials and methods
type specimens of all species have been examined. the male genitalia were dissected as follows: the pygofer was cut from the abdomen of the softened specimen with a needle blade, then boiled for about one hour in a 10% solution of potassium hydroxide (KoH). the phallic complex was dissected with a needle blade and all pieces examined in ethanol, the whole placed in glycerine for preservation. Observations were done with a Leica MZ8 stereo-microscope. Pictures were taken with a Canon EOS 300 D camera with Sigma DG Macro lens and optimized with Adobe Photoshop CS3. Inflation of the phallus was not performed due to the difficulty of obtaining good and replicable results and because it is not indispensable to separate the species in the genus Pyrops.
For the transcription of the labels of the types, the wording on each single label is given in square brackets.
the measurements were mainly taken as in Constant (2004) and the following abbreviations are used: BF = breadth of the frons BTg = breadth of the tegmen BPrH = breadth of the cephalic process at half length LF = length of the frons LTg = length of the tegmen LPr = length of the cephalic process TL = total length (LF, LPr and TL measured to/from ante-ocular carina at the base of the cephalic process) In the results section, species are treated in alphabetical order.

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.

The effusus species group
The characters of this very consistent group mainly follow Baker's (1925: 348) definition: (1) medium sized species; (2) cephalic process very short, clavate, bent in middle, strongly curved upwards, and with at least apical half blue or green; (3) tegmina green or blue basally, with yellow dots or markings; (4) hind wings blue basally and broadly black apically, the black colour extending towards the base along the main veins; (5) legs brown.
as compared to the list of included species given by nagai & Porion (1996), Pyrops cyanirostris (Guérin-Méneville, 1845) and P. maquilinganus (Baker, 1925) are here removed from the effusus group, the first because it does not match characters (3) and (4), the latter because it does not match characters (2), (4) and (5). P. maquilinganus is returned to the candelaria group as originally proposed by Baker (1925); P. cyanirostris cannot be attributed to any of the currently defined species groups in Pyrops.
all 4 species included here in the group are restricted to Borneo and adjacent islands. noteworthy, P. cyanirostris is recorded from Java (Guérin-Méneville 1845) and sumatra (nagai & Porion 1996) and P. maquilinganus from the Philippines (Baker 1925).
the group seems to be close to the pyrorhynchus species group which shares characters (3) and (4), but contains large sized species with an elongate cephalic process. However, the cephalic process in the latter group is also bent in the middle -even if not so strongly curved upwards -and slightly dilated apically like in effusus group. In some way the members of the effusus group could be regarded as dwarfed members of the pyrorhynchus group. noteworthy, of all members of the pyrorhynchus group, the Bornean species P. intricatus (Walker, 1857) ( Fig. 1) shows the greatest affinities in the pattern of the tegmina with the effusus group.
Identification key to the species of the effusus group a Bornean species pertaining to the pyrorhynchus group, but sharing a number of characters with the species of the effusus group, P. intricatus (cf. above), is included in the key and illustrated to avoid misidentifications.

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. 2D-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).

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. . 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). (satô & nagai, 1994) (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. 3D-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). 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".

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.

Diagnosis
In addition to the characters defining the effusus group, the species can be recognized by: (1) head entirely dark olivaceous blue-green ( Fig. 4D-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 . He was the author of numerous publications on auchenorrhyncha, and of all drawings of Lallemand's revisions of the Fulgoridae (1959,1963).
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.

Biology
No host tree has been identified for this species for which records are available from the months of March, July -august and september. (Distant, 1889) Figs 5A-E, 6, 9A-C, 12A-E  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. 5D-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).

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.
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.

Discussion
the present paper illustrates the critical need for taxonomic work in a spectacular and apparently wellknown group of insects. The production of clear, updated and accurately illustrated identification keys, which requires an in-depth revision, including the examination of all types, is necessary to avoid further spreading of erroneous information and wrong identifications which are currently commonplace, e.g., on the internet. the collaboration with amateur naturalists and nature photographers leads again to very interesting results (see also Constant 2014;Constant & alisto 2015): in the present paper, many data on the distribution and biology of the species including all trophobiosis observations come from such "citizen science" sources. Some data also originate from websites and blogs whose authors are unfortunately difficult to track and contact.
twelve species of Pyrops are currently recorded from Borneo (nagai & Porion 1996(nagai & Porion , 2002(nagai & Porion , 2004. However, several of them are known from single or very few specimens and the biology, life-history and phenology of all is practically undocumented. Hopefully in the future, with the help of local biologists and citizen-scientists, it will be possible to fill those gaps. This, together with the availability of more material and the clarification of some taxonomical issues (Constant, unpublished), will allow the publication of a comprehensive key to the species of Pyrops from Borneo.
Messrs Mick Webb and Max Barclay for their help during my visit at BMnH and after. this study has benefitted from funds from the European Union (Synthesys Project -Grant GB-TAF-1642) for a visit to the collection of BMnH in 2011. I thank all the curators listed above for access to the material under their responsibility; my colleagues Dr Patrick Grootaert and Mr Pol Limbourg (RBIns), and Pr. thierry Bourgoin (MnHn) for their permanent help and support.