Review of the Maluku Islands species of the lanternfly genus Birdantis Stål, 1863, with a new species and identification

. A new species of the genus Birdantis Stål, 1863 (Hemiptera: Fulgoridae), B. bhaskarai sp. nov. from Larat Island (Tanimbar), is described. Birdantis collaris (Walker, 1870) stat. rev. and B. trilineata (Schmidt, 1926) stat. rev. are reinstated as valid species, respectively from status of subspecies and as junior synonym of B. delibuta Stål, 1863. These four species, as well as the other one previously described from the Maluku Islands, B. decens Stål, 1863, are illustrated from their type specimens. An identification key, a distribution map, illustrations of habitus and details of male genitalia are provided. The synonymy between Myrilla Distant, 1888 and Birdantis is formally reinstated and all species formerly placed in the subgenus Birdantis ( Myrilla ) are transferred to Birdantis sensu stricto. Birdantis is transferred to the subfamily Aphaeninae Blanchard, 1847 and now contains eighteen species distributed in Maluku (five species), New Guinea and neighbouring islands (ten species) and Australia (three species).


Introduction
The family Fulgoridae counts 143 genera and 762 species, distributed mainly in the tropical and subtropical regions of the world, with some taxa extending to temperate regions.The family contains some of the most spectacular insects, such as the New World 'peanut bug' (Fulgora spp.) surrounded by several legends or the species of the genus Pyrops Spinola, 1839 in the Oriental Region, which are recognized at first glance by all entomologists (O'Brien 2002).Trophobiotic interactions with other insects (Formicidae, Blattodea, Lepidoptera), geckos (Reptilia: Squamata: Gekkonidae) and even snails (Mollusca: Gasteropoda) have been reported in recent years for several species both from the Old and New World (Naskrecki & Nishida 2007;Constant 2015).
The fauna of Fulgoridae in the Maluku Islands is very poor, with only two genera and six species, compared to the fauna of Sulawesi and adjacent islands to the west (5 genera and 30 species) or that of New Guinea (9 genera and 37 species) to the east.Both genera recorded from the Maluku Islands, Birdantis Stål, 1863 andHariola Stål, 1863, are also present in New Guinea, but all Malukan species are endemic in the archipelago (Bourgoin 2018).The Maluku Islands lie in Wallacea, the region marking the transition between the Asian and Australian biogeographical blocks, are geologically relatively young (1-15 million years old) and were never attached to larger landmasses.As they lie between the Weber's and Lydekker's lines, their fauna shows greater affinities with Australasia than with Asia as well as a high degree of endemism.The islands are small in size and mountainous, and are covered in rain forest except the Tanimbar Islands and other southeastern islands, which are arid and sparsely vegetated (Monk et al. 1997).
The genus Birdantis comprises fifteen species distributed in New Guinea, the Maluku Islands and Australia (Cape York Peninsula) (Bourgoin 2018).The description of the first two species of Birdantis from the Maluku Archipelago dates from more than 150 years ago, when Stål (1863) described the genus to accommodate B. decens Stål, 1863 andB. delibuta Stål, 1863.Two additional Malukan species were described, one in the genus Polydictya, P. collaris Walker, 1870, and one in the genus Myrilla, M. trilineata Schmidt, 1926.The latter species were considered as a synonym (trilineata) and a variety (collaris) of B. delibuta, respectively, leaving two valid species of Birdantis for the Maluku Islands: B. decens and B. delibuta (Lallemand 1963;Nagai & Porion 1996).
The study of recent material of Fulgoridae in the collections of the Royal Belgian Institute of Natural Sciences revealed one new species of Birdantis from Larat Island near Tanimbar and led to a critical review of the type material of all species described from the Maluku Islands.
The present paper aims to propose a new subfamily placement of Birdantis, to update the status of the genus, to review the status of all species known from the Maluku Archipelago, to describe the new species from Larat Island and to propose an illustrated identification key to the species of the treated area.

Material and methods
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 some minutes in a 10% solution of potassium hydroxide (KOH).The aedeagus was dissected with a needle blade and all pieces examined in ethanol, the whole preparation placed in glycerine for preservation.Observations were done with a Leica MZ8 stereo microscope.Photographs of specimens were taken with a Canon EOS 700 D camera with a Sigma DG Macro lens and of genitalia with a Leica EZ4W stereo microscope with an integrated camera, stacked with CombineZ software and optimized with Adobe Photoshop CS3.The metatibiotarsal formula gives the number of spines on (side of metatibia) apex of metatibia/apex of first metatarsus/apex of second metatarsus.The terminology for the venation follows Bourgoin et al. (2015).Lallemand (1959) synonymized Myrilla under Birdantis and proposed a key to the species of Birdantis.However, Fennah (1977) considered Myrilla as a subgenus of Birdantis, separating them on characters of the vertex, i.e., anterior margin of vertex straight, without deep transverse sulcus behind it in Myrilla; anterior margin of vertex concave, with a deep transverse sulcus behind it in Birdantis s. str.It is therefore currently formally divided into two subgenera: Birdantis (ten species and one subspecies) and Myrilla (two species) (Fennah 1977;Nagai & Porion 1996).The genus is externally close to the Oriental Polydictya Guérin-Méneville, 1844 and Gebenna Stål, 1863, and the Australian Desudaboides Musgrave, 1927 (Lallemand 1963;Nagai & Porion 1996;Constant 2010Constant , 2011)).Nagai & Porion (1996) followed the classification in Fennah (1977), but in my paper on the species of Birdantis from Australia (Constant 2011), I questioned the value of the subgenera as defined by Fennah (1977), because intermediate species exist, and followed the views of Lallemand (1959) but without formally reinstating the synonymy between Birdantis and Myrilla.Accordingly, three species remained without subgeneric affiliation (Bourgoin 2018) within the last formally accepted classification of Fennah (1977).

Remarks
The genus Birdantis is currently placed in the Poiocerina Haupt, 1929of the Poiocerini Haupt, 1929, in the subfamily Poiocerinae Haupt, 1929(Lallemand 1963;Constant 2011;Bourgoin 2018).However, Urban & Cryan (2009) have shown, based on molecular data, that Fulgoridae can be separated into two main clades, one grouping all New World taxa with strong support, the other containing all Old World taxa.Hence, as the type genus of the Poiocerinae is the Neotropical genus Poiocera Laporte, 1832, the suprageneric placement of Birdantis needs to be reconsidered.
The classifications proposed by Metcalf (1947), Lallemand (1963) and Nagai & Porion (1996) were based on a very small number of characters of the head, especially the presence/absence and shape of the cephalic process.The genus Birdantis is here transferred to the subfamily Aphaeninae, following the conclusions of the DNA study by Urban & Cryan (2009), which place the genus close to Desudaba Walker, 1858.However, the latter study did not contain any species of the genera Polydictya, Gebenna or Desudaboides, which are putatively considered closely related to Birdantis based on morphological characters.Hence, an accurate tribal placement requires a complete study of the suprageneric relations between the genera of Aphaeninae with the inclusion of more Old World taxa in the molecular analysis and the integration of morphological data (Urban & Cryan 2009).

Diagnosis
This species can be separated from all other species of Birdantis by the following combination of characters: (1) frons entirely yellow-brown without lines or spots (Fig. 1D) (2) bulge between frons and vertex coloured as vertex (Fig. 1B) (3) abdomen mostly orange dorsally (Fig. 1A) and ventrally (Fig. 1C) (4) tegmina opaque on basal half and infuscate on distal half (Fig. 1A) (5) hind wings largely smoky, but not hyaline or with a large black area (Fig. 1A)

Etymology
This species is dedicated to Mr Edy Bhaskara (Indonesia) in acknowledgment of his generous contribution to the present work.HeAd (Fig. 1B, D, F).Pale yellow-brown with small black marking at posterior angles of vertex, a larger black marking behind eyes, median portion of clypeus darker and more reddish, with a yellowish central line, labium pale yellow-brown turning to brown on two apical segments and antennae brown.Vertex with deep transverse groove and all margins carinate, shorter in middle than on sides.Frons finely wrinkled, with two smooth longitudinal carinae slightly diverging towards dorsum and a slight longitudinal groove on each side between carina and lateral margin; broadest near base, above clypeus; convex in dorsal view and with dorsal margin rounded in perpendicular view; bulge between frons and vertex only visible dorsally.Ocelli under eyes.Antennae with scape short and cylindrical, and pedicel inflated and reniform.Clypeus narrower than frons, reaching apex of procoxae.Labium with penultimate segment surpassing hind coxae (Fig. 1C).

Holotype
THOrAx (Fig. 1B, D, F).Pronotum pale yellow-brown, with median carina and impressed point on each side of latter; small back-brown spot on each side of disc, short black-brown line on median carina on anterior half and brown marking behind eye.Mesonotum dark brown, with median and peridiscal carinae well marked, concolorous; wrinkled in the area limited by peridiscal carinae; mesothoracic sternites pale yellow-brown.Metathoracic sternites dark brown.Tegulae yellow-brown.TegMiNA (Fig. 1A, C).Elongate, broader at nodal line and acutely rounded apically.Corium and clavus with numerous cross-veins, brown variegated with darker patches and a blackish line along external side of vein Pc+CP; veins and cross-veins yellow.Membrane smoky, roundly extending inside corium medially and with brown spots on distal part and brown markings on nodal line; veins yellow turning to black-brown after nodal line.
HiNd wiNgS (Fig. 1A, C).Light smoky brown with veins black-brown, darker at basal angle and with anal area with a grey hue and cross-veins whitish; maximal breadth near base; slightly broader than tegmina.
LegS (Fig. 1A, C).Elongate and slender.Pale yellow-brown with apex of pro-and mesofemora black distally; pro-and mesotarsi black; metatibiae brown apically, with six lateral spines and six apical spines, all spines dark brown apically; metatarsi with first tarsomere yellow-brown, brown apically and with nine apical spines ventrally, second and third tarsomeres dark brown, the second one with eight apical spines ventrally.Metatibiotarsal formula: (6) 6/9/8.

Male genitalia
Pygofer higher than long, about 1.4 times as high as maximum length, longer ventrally and abruptly narrowing on dorsal ¼ in lateral view; dorsal margin of pygofer oblique in lateral view (Fig. 2A).Anal tube elongate, about 2.1 times as long as maximum breadth, curved ventrally near base; lateral margins subparallel, only slightly sinuate and with narrowest portion at basal ¼; apical margin strongly emarginate in dorsal view, acutely rounded in lateral view (Fig. 2A-B).Gonostyli subtriangular in lateral view, with posterior margin rounded; anterodorsal margin emarginate after lateral process; lateral process laminate, directed laterally and apically pointed, with apical point directed lateroventrally (Fig. 2A-B).Aedeagus mostly membranous; phallobase with two ventral, elongate processes; aedeagus s. str.strongly reduced but endosoma well developed with six membranous digitiform processes, with posteroventral left process with secondary process directed centrally (Fig. 2C-F).

Diagnosis
This species can be separated from all other species of Birdantis by the following combination of characters: (1) frons yellowish, with five longitudinal black lines not reaching ventral margin of frons, the three central ones merging together dorsally in a transverse line and central one dilated in middle (Fig. 5F) (2) bulge between frons and vertex coloured as vertex (Fig. 5C) (3) abdomen mostly black dorsally (Fig. 5A) and brown ventrally (Fig. 5B) (4) tegmina opaque on basal half and hyaline on distal half (Fig. 5A) (5) hind wings largely hyaline, with basal angle red (Fig. 5A)

Diagnosis
This species can be separated from all other species of Birdantis by the following combination of characters: (1) frons yellow-brown with three dark brown lines limited dorsally to level of middle of eyes, and with the two lateral ones extending on to clypeus (Fig. 6B) (2) bulge between frons and vertex dark brown and vertex yellow-brown (Fig. 6C) (3) tegmina opaque on basal half and infuscate on distal half (Fig. 6A) (4) hind wings largely smoky, with a basal red marking (Fig. 6A) (5) anterior margin of frons in dorsal view rounded but not strongly protruding (Fig. 6C)

Male genitalia
Pygofer higher than long, about 2.0 times as high as maximum length, longer ventrally and abruptly narrowing on dorsal ½ in lateral view; dorsal margin of pygofer horizontal in lateral view (Fig. 9A).Anal tube elongate, about 1.5 times as long as maximum breadth, nearly straight with ventral margin sligtly curved in lateral view; progressively broadening from base towards ¾ of length in dorsal view, then with lateral margins converging; apical margin strongly, roundly emarginate in dorsal view, acutely rounded in lateral view (Fig. 9A-B).Gonostyli slightly elongate in lateral view, with ventral margin slightly concave and posterior margin broadly rounded; anterodorsal margin slightly emarginate after lateral process; lateral process laminate, curved laterally and apically pointed, with apical point directed lateroventrally (Fig. 9A-B).Aedeagus mostly membranous (Fig. 9C-F).

Discussion
Together with the previous work on Australian species of Birdantis (Constant 2011), a large part of the genus has now been reviewed, although three species from Maluku are still only documented from their female type specimens, which were collected about 150 years ago.
The Papuan species were treated by Fennah (1977), but the conclusions of this work need to be assessed and refined by a thorough study of the type material of the species and their male genitalia, especially the taxa described by Schmidt (1906Schmidt ( , 1907Schmidt ( , 1911) ) and Lallemand (1959Lallemand ( , 1963)), the types of which were not studied by Fennah.
The suprageneric position and the relationships with the other genera, notably the Oriental Gebenna and Polydictya and the Australian Desudaba and Desudaboides, will need further study, especially based on molecular data.
The biology of the species of Birdantis remains nearly completely undocumented in terms of development, phenology, host plants, behaviour etc., and the male genitalia still need to be described for several species.