http://dx.doi.org/10.5852/eit.2015.128

www. europeani ournaloftaxonomv. eu

2015 • Constant C.

This work is licensed under a Creative Commons Attribution 3.0 License.

Research article

urn:lsid:zoobank.org:pub:FBAF2DED-5FD6-4F22-8A3A-E94733A92D73

Review of the effusus group of the Lanternfly genus Pyrops Spinola,

1839, with one new species and notes on trophobiosis

(Hemiptera: Fulgoromorpha: Fulgoridae)

Jerome CONSTANT

Royal Belgian Institute of Natural Sciences, O.D. Phylogeny and Taxonomy, Entomology, Vautier

street 29, B-1000 Brussels, Belgium. E-mail: i [email protected]

urn:lsid:zoobank.org:author:6E6072Al-9415-4C8D-8E60-2504444DB290

Abstract. The species group effusus of the genus Pyrops Spinola, 1 839 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 (Sato & 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 (Guerin-Meneville, 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 (Sato & 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.

Keywords. Indonesia, Malaysia, Lanternfly, Fulgoroidea, citizen science

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.

http://dx.doi.Org/l 0.5852/eit.2015,128

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).

European Journal of Taxonomy 128: 1-23 (2015)

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 (1960a,b, 1963), Chou et al. (1985) and Sato &

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 & Ko£ak (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 and Prolepta Walker, 1851 or with two species he placed in the Candelaria

group: Pyrops maquilinganus (Baker, 1925) and P. delesserti (Guerin-Meneville, 1840).

Lallemand (1963) renamed Baker’s (1925) effusus group “quatrieme groupe: type whiteheadF , 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), 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. Linally, 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 (pi. X,

figs 11-13) and erroneously attributed to P. whiteheadi, pertain to P. effusus.

CONSTANT C., Fulgoridae: Pyrops effusus species group (Hemiptera)

Sato & 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

ofP. whiteheadi.

Nagai & Porion (1996) followed the classification proposed by Tallemand (1963), but correctly used

Pyrops as a genus name. In the “groupe IV”, as defined by Tallemand, they included 3 species and one

subspecies: P. cyanirostris (Guerin-Mene ville, 1845), P. maquilinganus, P. whiteheadi and P. whiteheadi

gunjii. They synonymized dimotus and sapphirinus with cyanirostris, and followed Tallemand 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:

breadth of the frons

BTg =

breadth of the tegmen

BPrH =

breadth of the cephalic process at half length

length of the frons

LTg =

length of the tegmen

LPr

length of the cephalic process

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.

European Journal of Taxonomy 128 : 1-23 ( 2015 )

Acronyms

BMNH =

EUM =

FSAG =

MHNL =

RBINS =

RMNH =

USNM =

used for the collections (name of the curator in parentheses):

Natural History Museum, London, United Kingdom (M. Webb, M. Barclay)

Ehime University Museum, Matsuyama, Japan (H. Yoshitomi)

University of Liege, Gembloux Agro-Bio Technologies, Gembloux, Belgium (J. Bortels)

Museum d’Histoire Naturelle de Lyon, France (C. Audibert)

Royal Belgian Institute of Natural Sciences, Brussels, Belgium (W. Dekoninck)

Nationaal Natuurhistorisch Museum (‘Naturalis”), Leiden, Netherlands (J. van Tol, Y. van

Nierop)

National Museum of Natural History, Washington D.C., U.S.A. (S. McKamey)

Results

Class Hexapoda Blainville, 1816

Order Hemiptera Linnaeus, 1758

Suborder Auchenorrhyncha Dumeril, 1806

Infraorder Fulgoromorpha Evans, 1946

Superfamily Fulgoroidea Latreille, 1807

Family Fulgoridae Latreille, 1807

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.

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 (Guerin-

Meneville, 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.

CONSTANT C., Fulgoridae: Pyrops effusus species group (Hemiptera)

All 4 species included here in the group are restricted to Borneo and adjacent islands. Noteworthy, P.

cyanirostris is recorded from Java (Guerin-Meneville 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.

Fig. 1. Pyrops intricatus $, Kalimantan, Mt Bawan (RBINS). A. Habitus, dorsal view. B. Habitus,

ventral view. C. Habitus, lateral view. D. Head and thorax, lateral view. E. Frons, normal view. Scale

bar = 30 mm (D-E not to scale).

European Journal of Taxonomy 128 : 1-23 ( 2015 )

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.

1 Head with cephalic process longer than half of tegmen (Fig. 1A, C); cephalic process slightly curved

upwards, brown with bright yellow apex, without blue or green (Fig. 1D-E); ratio LPr/LF > 4;

LPr/BPrH > 9 . P. intricatus (Walker, 1857) {pyrorhynchus group]

- Head with cephalic process shorter than half of tegmen (Figs 2A, C, 4A, C); cephalic process strongly

curved upwards, at least partly green or blue (F igs 2D-E, 4D-E); ratio LPr / LF < 2.5; LPr/BPrH <1...

... 2

2 Basal half of head yellow-brown, strongly contrasting with green or blue apical half of cephalic

process (Fig. 2A, C) . 3

- Head unicolorous blue or green, at best slightly paler basally (Figs 4A, C, 5A, C). 4

3 Tegmina green on basal half, getting progressively more yellow on membrane, infuscate along

apical margin; spots on tegmina yellow and rather large, visible on membrane (Fig. 2A) .

. P. effusus (Distant, 1891)

- Tegmina with basal half dark blue-green followed by a contrasting large oval yellow patch

and broadly dark brown along apical margin; spots on tegmina pale yellow-white and rather

small, getting indistinct on membrane (Fig. 3A) . P. gunjii (Sato & Nagai, 1994)

4 Tegmina with transverse yellow bands on corium, the basal one complete (Fig. 4A); pro- and meso-

notum concolorous (Fig. 4A); head dark olivaceous blue-green (Fig. 4A, D-E) P synavei sp. nov.

- Tegmina with small round yellow spots (Fig. 5A); pronotum yellow to orange, mesonotum dark

blue-green (Fig. 5A); head bright blue (often faded in collection specimens) (Fig. 5A, D-E)

. P. whiteheadi (Distant, 1889)

Pyrops effusus (Distant, 1891)

Figs 2A-E, 6-7A-C, 10A, C-F

Fulgora effusa Distant, 1891: 518, pi. XX, figs 4,4a (Type in NHM) [described, illustrated and compared

with P. whiteheadi (Distant, 1889)].

Fulgora viridicastanea Baker, 1925: 359, pi. 3, fig. 5, pi. 4 fig. 8 (Type in USNM) [described, illustrated

and compared with P. whiteheadi erroneously mentioned and illustrated as “ Fulgora effusa'f

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 ofP. 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)],

pi. 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], pi. 1, figs 2, 4 [illustrated from 2 localities].

CONSTANT C., Fulgoridae: Pyrops effusus species group (Hemiptera)

Laternaria effusa — Metcalf 1947: 196 [transferred to Laternaria: catalogued]. —Liang 1998: 45 [listed

as junior synonym ofE 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, pi. X, figs 11-13 [keyed, comments illustration of male

genitalia (based on misidentified specimens of P. effusus)].

Fulgora whiteheadi viridicastanea - Sato & Nagai 1994 [considered as a subspecies of P. whiteheadi

(erroneous) and compared with “ Fulgora whiteheadi gunjiF].

itSs!

Fig. 2. Pyrops effusus $, Kalimantan, Batulicin (RBINS). A. Habitus, dorsal view. B. Habitus, ventral

view. C. Habitus, lateral view. D. Head and thorax, lateral view. E. Frons, normal view. Scale bar = 30

mm (D-E not to scale).

European Journal of Taxonomy 128 : 1-23 ( 2015 )

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, pi. 11, fig. 163 [illustration of the type ofP

effusus (Distant, 1891)].

non Fulgora effusa - Baker 1925: 359, pi. 3, fig. 4, pi. 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. 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).

Material examined

Type material

INDONESIA: Holotype of Fulgora effusa (examined on photographs), f \ [Borneo] [effusa Dist.]

[Distant Coll. 1911—383.] (BMNH).

INDONESIA: Holotype of Fulgora viridicastanea (examined on photographs, Fig. 10C-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 S, E-Borneo, Midden, 10 Oct. 1925, H.C. Siebers

(FSAG); 1 <$, idem, 4 Oct. 1925 (FSAG); 1 f, SE Borneo, Atkinson coll. (BMNH); 1 <$, 1 J, Borneo,

1909 (BMNH); 1 $, North Borneo (BMNH).

Examined on photograph

INDONESIA: 1 specimen (Fig. 10A), Kalimantan, Wehea Protected Forest, 25 Sep. 2010, KaraNorby.

Measurements and ratios

TL: c? (n = 4): 35.1 mm (34.3-36.0); $ (n = 2): 40.2 mm (40.0M0.5); TL+process: S (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. 7A-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).

CONSTANT C., Fulgoridae: Pyrops effusus species group (Hemiptera)

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 Taut Island and Kutai, and the weakly contrasted ones

from Toksado (confirmed by the examination of the specimens in MHNL collections by H. Tabrique,

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).

Pyrops gunjii (Sato & Nagai, 1994) stat. nov.

Figs 3A-E, 6, 8A-C, 10B

Fulgora whiteheadi gunjii Sato & Nagai 1994: 308, figs 2, 9 (Type in EUM) [described, illustrated and

compared with P. whiteheadi and P. effusus , the latter mentioned as “A whiteheadi viridicastanea ”].

Pyrops whiteheadi gunjii - Nagai & Porion, 1996: 26, pi. 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. 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. 3 A); (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: Sato & 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 SS, 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 o 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

European Journal of Taxonomy 128 : 1-23 ( 2015 )

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: c? (n = 2): 35.1 mm (34.5-35.6); ? (n = 1): 38.6 mm; TL+process: c? (n = 2): 39.7 mm (39.0^10.3);

$ (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. 8A-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).

30 mm

Fig. 3. Pyrops gunjii $, E Kalimantan, Samarinda, Mt Beratus (RBINS). A. Habitus, dorsal view.

B. Habitus, ventral view. C. Habitus, lateral view. D. Head and thorax, lateral view. E. Frons, normal

view. Scale bar = 30 mm (D-E not to scale).

CONSTANT C., Fulgoridae: Pyrops effusus species group (Hemiptera)

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.

Pyrops synavei sp. nov.

urn:lsid:zoobank.org:act:B502AF69-AEC2-4245-B440-57F75BF8559E

Figs 4A-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. 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 (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. 11A-B): Kalimantan, Long Alango, Lalut Birai Sacred Forest, 2 Sep. 2011, Joost

Woltering; 1 specimen (Fig. 11C-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

European Journal of Taxonomy 128: 1-23 (2015)

Fig. 4. Pyrops synavei sp. nov. §, holotype (RBINS). A. Habitus, dorsal view. B. Habitus, ventral view.

C. Habitus, lateral view. D. Head and thorax, lateral view. E. Frons, normal view. Scale bar = 30 mm

(D-E not to scale).

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.

30 mm

CONSTANT C., Fulgoridae: Pyrops effusus species group (Hemiptera)

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. 4A-B). Dark olivaceous brown. Metatibiae with 6 lateral and 7 apical spines.

Abdomen (Fig. 4A-B). Tergites brown-black; sternites black; genital segments tinged with yellow-

brown.

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 5A-E, 6, 9A-C, 12A-E

Fulgora whiteheadi Distant 1889: 422 (Type inNHM) [described and compared to P. delesserti (Guerin-

Meneville, 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 ]. — Sato & 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], pi. 11, fig. 160 [type

illustrated]. — Liang 1998: 45 [listed]. — Nagai & Porion 2004: 6 [comparison with P. effusus ], pi.

1, fig. 6 [illustration of habitus].

non Fulgora whiteheadi - Lallemand 1963: pi. X, figs 11-13 [illustration of the genitalia of P. effusus

erroneously identified as P. whiteheadi ].

non Pyrops whiteheadi - Nagai & Porion 1996: pi. 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. 5D-E); (2) pronotum yellow-brown,

mesonotum blue-brown (Fig. 5A); (3) tegmina green with small yellow spots also beyond nodal line;

European Journal of Taxonomy 128: 1-23 (2015)

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: S, 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).

Fig. 5. Pyrops whiteheadi Sabah, Danum Valley (RMNH). A. Habitus, dorsal view. B. Habitus,

ventral view. C. Habitus, lateral view. D. Head and thorax, lateral view. E. Frons, normal view. Scale

bar = 30 mm (D-E not to scale).

CONSTANT C., Fulgoridae: Pyrops effusus species group (Hemiptera)

Additional material

MATAYSIA: 1 Sabah, Kinabalu National Park, Poring Hot Spring, 21 Apr. 1999, T. Trilar (RBINS);

1 5, 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 S, Sabah, Danum Valley, 70 km W of Tahad Datu, rainforest along

narrow stream, at light, next to bridge Nature Trail, 50 m, 1 Dec. 1989, M.J. & J.P Duffels (RMNH).

“

^ Pyrops effusus (Distant, 1891)

0 Pyrops gunjii (Sato & Nagai, 1994)

O Pyrops synavei sp. nov.

^ Pyrops whiteheadi (Distant, 1889)

Fig. 6. Pyrops effusus group, distribution map.

European Journal of Taxonomy 128: 1-23 (2015)

Figs 7-9. Pyrops effusus group, S genitalia. 7. Pyrops effusus. 8. Pyrops gunjii. 9. Pyrops whiteheadi. —

A. Lateral view. B. Dorsal view. C. Posteroventral view. An = anal tube, G = gonostylus, Py = pygofer.

Scale bar = 1 mm. Photographs by J. Brecko.

CONSTANT C., Fulgoridae: Pyrops effusus species group (Hemiptera)

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.

Fig. 10. —A. Pyrops effusus , in natura, Borneo, Kalimantan, Wehea Protected Forest, 25 Sep. 2010, K.

Norby. — B. Pyrops gunjii , in natura, Borneo, East Kalimantan, 0°32’4.70”S 117°27 , 46.73 ,, E, 3 Jul.

2011, M. Hauskrecht. — C-F. Fulgora viridicastanea, holotype (Photographs A. Kennedy). C. Dorsal

view. D. Left lateral view. E. Head, normal view of frons. F. Labels.

European Journal of Taxonomy 128: 1-23 (2015)

Examined on photographs

EAST MALAYSIA: 1 specimen (Fig. 12A-B), Sabah, Danum Valley, 4 Jul. 2013, Anton Sorokin; 1

specimen, idem, 13 Jun. 2013; 1 specimen, idem, 3 Oct. 2010, HokPing Guek; 1 specimen, idem, 4 Oct.

Fig. 11. Pyrops synavei sp. nov. in natura. A-B. Borneo, Kalimantan, Long Alango, Lalut Birai Sacred

Forest, 2 Sep. 2011, J. Woltering. C-D. Borneo, Kalimantan, Lalut Birai Forest, 13 Mar. 2009, D. Forney.

CONSTANT C., Fulgoridae: Pyrops effusus species group (Hemiptera)

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,

Sulcau, 12 Aug. 2011, J.M. Gayman; 1 specimen, Sandakan, 27 Mar. 2013, Gavin Golden; 1 specimen,

Sandakan, Sulcau, 14 Aug. 2009, Samuel Tan; 1 specimen, Taman Bulcit Tawau, 17 Aug. 2012, Sublci

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 (38.7^10.9);

$ (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. 9A-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 Stal, 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 &

Kogalc (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 ofFulgoroidea,

Flatidae, in Madagascar, were previously reported and analyzed by Foiling et al. (2001).

European Journal of Taxonomy 128: 1-23 (2015)

Fig. 12. Pyrops whiteheadi and P. intricatus, in natura, trophobiosis. — A-E. P. whiteheadi. A-B. Tended

by cockroaches (Blattodea, Dorylaea , probably D. magna (Schelford, 1909)), Sabah, Danum Valley, 4

Jul. 2013, A. Sorokin. C. Tended by small cockroach (Blattodea, Pseudophyllodromiinae), Sabah, Danum

Valley, 15 Dec. 2008, C.C. Lee. D. Sabah, Lower Kinabatangan River, 9 Aug. 2012, N. Messina. E. Tended

by an adult gecko, Gehyra mutilata (Wiegmann, 1835) (Squamata: Gekkonidae), Sabah, Sepilok, 6

Jun. 2013, C.C. Lee. — F . P. intricatus tended by cockroach (Blattodea, Dorylaea sp.), Sarawak, Mulu

National Park, 15 Oct. 2009, C.C. Lee.

CONSTANT C., Fulgoridae: Pyrops effusus species group (Hemiptera)

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. Tee

pers. comm.. Fig. 12C) and one species of the genus Dorylaea (A. Sorokin pers. comm., Fig. 12A-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. Tee 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. Tee 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 well-

known 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, 2002, 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.

Acknowledgements

I warmly thank Mrss Kara Norby and Ashley Kennedy, Messrs Joost Woltering, Dave Forney, Martin

Hauskrecht, Nicola Messina, CITien C. Tee ( www.wildborneo.com.my I and Anton Sorokin for sharing

their photographs, observations and/or data, as well as all the persons who published their photographs

and data on the internet (listed under “material examined on photographs”); Mr Jonathan Brecko

for taking the photographs of the male genitalia; Dr. Hiromi Yoshitomi for sending photographs of

the type of P. gunjii ; Mrs Jeannine Bortels for the photographs of the specimens in the Lallemand’s

collection (FSAG); Mr Harold Tabrique for the information on Porion’s collection (MHNT); Dr Olivier

S.G. Pauwels for the identification of the gecko species and Dr George Beccaloni for the Blattodea;

European Journal of Taxonomy 128: 1-23 (2015)

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 2011.1 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.

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Manuscript received: 27 February 2015

Manuscript accepted: 8 April 2015

Published on: 8 July 2015

Topic editor: Koen Martens

Desk editor: Kristiaan Hoedemakers

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; Botanic Garden Meise,

Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London,

United Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History

Museum of Denmark, Copenhagen, Denmark.