Revision of the Malagasy lanternfly genus Belbina Stål , 1863 , with two new species ( Hemiptera : Fulgoromorpha : Fulgoridae )

The Malagasy genus Belbina Stål, 1863 (Hemiptera: Fulgoridae) is revised, transferred from the Enchophorinae Haupt, 1829 to the Aphaeninae Blanchard, 1847, and two new species, B. bourgoini sp. nov. and B. laetitiae sp. nov., are described. The genus Cornelia Stål, 1866 is proposed as a junior synonym of Belbina. The following new combinations are proposed: Belbina bergrothi (Schmidt, 1911) comb. nov. and B. nympha (Stål, 1866) comb. nov. The combination Belbina foliacea Lallemand, 1950 is restored. Aphana madagascariensis Westwood, 1851 is redescribed, transferred to Belbina and the new combination B. madagascariensis (Westwood, 1851) is proposed. Belbina vicina Lallemand, 1959 is proposed as a junior synonym of B. falleni Stål, 1863 and Cornelia atomaria (Brancsik, 1893) as a junior synonym of Belbina nympha (Stål, 1866). Neotypes are designated for B. madagascariensis (Westwood, 1851) comb. nov. and B. servillei (Spinola, 1839). The genus now comprises 12 species from Madagascar. A list of diagnostic characters, an identification key, illustrations of the male genitalia and distribution maps are provided. The falleni+ species group is defined based on characters of the male genitalia and contains the following 5 species: B. bloetei Lallemand, 1959, B. falleni Stål, 1863, B. laetitiae sp. nov., B. lambertoni Lallemand, 1922 and B. pionneaui Lallemand, 1922.


Introduction
In the process of identifying material of Fulgoridae from Madagascar, three species attributed to the genera Belbina Stål, 1863 or Cornelia Stål, 1866 were found that were not listed in the works of Lallemand (1959) or Constant (2004b).Two appear to be undescribed, while the third one represents the enigmatic Aphana madagascariensis Westwood, 1851.The genus Belbina was described by Stål (1863a) to accommodate two species: Belbina falleni Stål, 1863 (Madagascar) and Enchophora servillei Spinola, 1839 (patria incognita, now recognized to be Madagascar).It was redescribed by Stål (1863b) later the same year, leading to misinterpretation by Metcalf (1947) who erroneously designated Enchophora sicca Walker, 1851 (South Africa) as the type species of Belbina.Lallemand (1959) validly designated Belbina falleni Stål, 1863 as the type species,

Materials and methods
The 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.It was then boiled in water for a few minutes for cleaning and to facilitate removal of pieces of tegument from the last abdominal segments.It was then dried and placed under the specimen in a gelatin capsule after examination.Endosomal characters were not used due to the diffi culty to correctly infl ate the membranous endosome and because it is not indispensible to separate the species in the genus Belbina.Observations were done with a Leica MZ8 stereo microscope.Pictures of specimens were taken with a Canon EOS 300 D camera with a Sigma DG Macro lens and optimized with Photoshop CS3.Pictures of genitalia were taken with a Leica DFC290 camera mounted on a Leica Z6 APO microscope, using LAS V4 software for image capture.Each series of pictures was processed with Combine ZP and optimized with Adobe Photoshop CS5.
The disc designates the central area of the tegmen, roughly covering the zone between the subcostal vein, claval joint and nodal line.
For the transcription of the labels of the types, the exact wording on each label is provided within square brackets.In the results section, species are treated in alphabetical order.
Acronyms used for the collections (name of the curator in parentheses) are as follows:

Results
Class Hexapoda Blainville, 1816 Order Hemiptera Linnaeus, 1758 Suborder Auchenorrhyncha Duméril, 1806Infraorder Fulgoromorpha Evans, 1946Superfamily Fulgoroidea Latreille, 1807 Family Fulgoridae Latreille, 1807 Subfamily Aphaeninae Blanchard, 1847 Morphological characters (shape of pro-and mesonotum, carinae of head, tegmina and wing venation) and especially recent molecular data (Urban & Cryan 2009) lead to the conclusion that Belbina should be transferred to Aphaeninae.The genus is close to the African genera Anecphora Karsch, 1890 and Rhicnophloea Gerstaecker, 1895 (the latter mentioned as 'undet.Fulgoridae' in Urban & Cryan 2009: fi g. 3) and to the Oriental genera Penthicodes Blanchard, 1845 and Kalidasa Kirkaldy, 1900.The colour pattern in Belbina is very similar to that found in Penthicodes, species of Belbina looking somewhat like "Penthicodes with erected cephalic process" (see also Constant 2010 for illustrations of species of Penthicodes).

Remarks
Belbina and Cornelia are synonymized because it is not possible to fi nd any characters allowing consistent separation of these taxa.When all species of both genera are taken into account, a gradient can be observed from the larger species (e.g., falleni and recurva) to the smaller ones (e.g., nympha and bergrothi) in all characters supposed to separate the genera, i.e., (1) the width of the vertex (supposedly narrower than eye in Cornelia and broader than eye in Belbina), (2) vertex prolonged directly on cephalic process (Cornelia) or not (Belbina) and (3) hind margin of tegmina cut transversely (Cornelia) or rounded (Belbina).
The close relationship and possible synonymy of Belbina and Druentia mentioned by Constant (2004) is not supported by the results of recent molecular studies.According to a preliminary analysis (Julie Urban pers. comm., Nov. 2013), Druentia is sister to Rhicnophloea Gerstaecker, 1895 and [Druentia + Rhicnophloea] is sister to a larger clade of Old World taxa which contains, among others, a group formed by [Samsama Distant, 1906+ Belbina + Metaphaena Schmidt, 1905 Five of the twelve currently known species of Belbina seem very close and share very similar male genitalia with the following common characters: (1) pygofer, anal tube and gonostyli red or dark red; (2) gonostyli elongate with ventral margin concave in lateral view and roundly pointed apically; (3) gonostyli with strong basodorsal digitiform process directed dorsolaterally or dorsoposteriorly; (4) anal tube broadening more or less regularly from base to apex.This group is defi ned as the falleni+ group and contains B. bloetei, B. falleni, B. laetitiae sp. nov., B. lambertoni and B. pionneaui.

Diagnostic characters
The genus can be recognized by the following combination of characters: (1) head much narrower than pronotum; (2) frons longer than broad, with longitudinal carinae; (3) cephalic process present, about as long as frons and projecting anterodorsally to posterodorsally; (4) lateral carinae of cephalic process (i.e., prolongation of the lateral carinae of frons and of vertex) fused in a single longitudinal carina before apex of process; (5) pronotum with a strong tectiform median carina and a deeply impressed point on each side of it.
Belbina is very similar to Druentia but can be separated by character (4): in Druentia the lateral carinae of the vertex reach the apex of the cephalic process without fusing together.

Sexual dimorphism
Males 15 to 30% smaller in size than females.

Distribution
Only recorded from Madagascar.

Identifi cation key to the species of Belbina
The disc of the hind wings can be either orange or red in some species (observed in B. bergrothi, B. madagascariensis and B. nympha).It is possible that other species presently known only from red hindwinged specimens also have ones with orange hind wings.

Male genitalia
Black with gonostyli showing large, basoventral ochraceous patch (Fig. 13); pygofer higher than long and with posterior margin sinuate in lateral view (Fig. 13); anal tube elongate, 1.6 times longer than broad at apex and with lateral margins sinuate in dorsal view (Fig. 14); gonostyli elongate, 2.4 times longer than high, not surpassing apex of anal tube and acutely rounded at apex in lateral view (Fig. 13); dorsal margin of gonostyli projecting laterally from base to midlength and with strong tooth directed ventrally ; ventral margin and apical half of dorsal margin broadly rounded in lateral view (Fig. 13).

Remarks
Male genitalia without basodorsal process on gonostyli.
Belbina bergrothi can be separated (1) from B. recurva by not having the margins of the anal tube produced into a semi-circular lateral plate; (2) from B. madagascariensis and B. nympha by having the gonostyli acutely rounded apically and more elongate; (3) from B. foliacea by not having the gonostyli angulose above mediodorsal tooth.

Male genitalia
Dark red (Figs 15-16); pygofer higher than long and with posterior margin concave in middle in lateral view (Fig. 15); anal tube slightly elongate, 1.16 times longer than broad at apex and with lateral margins bisinuate in dorsal view (Fig. 16); margin of anal opening pointed posteriorly (Fig. 15) and hiding lateral margin apically in dorsal view (Fig. 16); gonostyli elongate, 1.18 times longer than high (dorsal process included), strongly surpassing apex of anal tube and acutely rounded at apex in lateral view (Fig. 15); ventral margin straight on basal 1/5, then strongly excavate (Fig. 15); dorsal margin with basal strong digitiform process projecting laterodorsally, strong hook laterally at middle of process projecting posteroventrally and posterior margin of process excavate between apex and hook (Fig. 15); apex of digitiform process rounded and broad in dorsal view (Fig. 16).

Remarks
Belbina bloetei is a member of the falleni+ group, showing strong basodorsal digitiform process on gonostyli.It can be separated (1)

Etymology
The species is dedicated to Prof. Thierry Bourgoin (MNHN), in acknowledgement of his permanent help and support from the beginning of the work of the author.HEAD.Yellow-brown sometimes suffused with red (Fig. 3C-E); vertex with hind margin and sides strongly carinate (Fig. 3C); sides of vertex bisinuate in lateral view (Fig. 3D); frons with 2 carinae extending on sides of cephalic process (Fig. 3E); cephalic process about 1.5 times as long as diameter of eye, projecting dorsad to posterodorsad (Fig. 3D); apical half carinate anteriorly and strongly emarginate laterally (Fig. 3E); posterior side of process with 2 longitudinal, sinuate carinae; lateral oblique carina between vertex and frons before eye (Fig. 3D); postclypeus with sides strongly carinate and obsolete median carina; anteclypeus with strong median carina (Fig. 3E); labium very elongate, surpassing hind coxae but not reaching apex of abdomen.

Male genitalia
Dark brown ; pygofer higher than long and with posterior margin nearly straight, slightly sinuate dorsally in lateral view (Fig. 17); anal tube elongate, 1.6 times longer than broad at apex and with lateral margins bisinuate in dorsal view (Fig. 18), slightly curved ventrally and with apex broadly rounded in lateral view (Fig. 17); gonostyli elongate, 1.6 times longer than high (dorsal process included), surpassing apex of anal tube and broadly rounded at apex in lateral view (Fig. 17); ventral margin straight, with strong angle at basal ⅛ (Fig. 17); dorsal margin with basal slender digitiform process projecting laterodorsally, pointed tooth laterally at middle of process projecting anteroventrally, apex of digitiform process narrowing apically (Fig. 17); apical ⅔ of dorsal margin sinuate (Fig. 17).

Remarks
Belbina bourgoini sp.nov.can be separated ( 1) from all species of the B. falleni+ group by the brown colour of the genitalia, the more slender basodorsal digitiform process on the gonostyli and by having the gonostyli rounded at the apex, with the dorsal margin sinuate after the process; (2) from B. bergrothi, B. foliacea, B. madagascariensis and B. nympha by the basodorsal digitiform process on the gonostyli; (3) from B. servillei by having the digitiform basodorsal process of the gonostyli slender and narrow, not laminate.

Male genitalia
Dark red ; pygofer higher than long and with posterior margin sinuate in lateral view (Fig. 19); anal tube slightly elongate, 1.27 times longer than broad at apex and with lateral margins bisinuate in dorsal view on apical half (Fig. 20); posterior margin notched in lateral view (Fig. 19), with hind margin of anal opening projecting posteriorly (Fig. 19) and hiding lateral margin apically in dorsal view (Fig. 20); gonostyli elongate, 1.36 times longer than high (dorsal process included), strongly surpassing anal tube and acutely rounded at apex in lateral view (Fig. 19); ventral margin straight on basal 1/5, then slightly sinuate (Fig. 19); dorsal margin with basal strong digitiform process projecting laterodorsally, strong hook laterally at middle of process projecting posteroventrally and posterior margin of process excavate between apex and hook (Fig. 19); apex of digitiform process rounded and broad in lateral view (Fig. 19), slightly compressed laterally in dorsal view (Fig. 20).Lallemand (1959) erroneously mentioned that the type of B. vicina is a female.However it is obvious that the type is a male because he stated that "genitalia have similar shape as those of B. pionneaui and B. lambertoni".Lallemand (1959) described B. vicina based on a specimen of B. falleni presenting the cephalic process more strongly curved posteriorad than in another specimen he had identifi ed in his collection as B. falleni; both specimens were from the same locality (Manjakandriana).Male genitalia are similar and intraspecifi c variability in the direction of the cephalic processs has been observed from series in several species of Belbina.For those reasons, B. vicina is here synonymized with B. falleni.

Remarks
Belbina falleni is a member of the falleni+ group, showing a strong basodorsal digitiform process on the gonostyli.It can be separated (1) from B. bloetei by the less concave ventral margin of the gonostyli in lateral view; (2) from B. laetitiae sp.nov.and B. lambertoni by the laterally more strongly compressed digitiform process and the more acutely rounded apex of the gonostyli in lateral view; (3) from B. pioneaui by the more rounded apex of the basodorsal digitiform process of the gonostyli and the more strongly notched ventroapical margin of the anal tube under the anal opening in lateral view.

Male genitalia
The male genitalia were dissected and drawn by Henry Synave for Lallemand's revision (1959).Only one gonostylus and the phallic complex were illustrated.The organs were preserved in a small glass vial under the specimen.The anal tube is missing and the pygofer is severely damaged.As it has been treated with potassium hydroxide, it is not possible to know the original colour.
Gonostyli elongate, 1.7 times longer than high and broadly rounded at apex in lateral view; ventral margin nearly straight; dorsal margin strongly sinuate basally and with a strong angle at basal third, then straight; strong lateral tooth directed posteroventrally along dorsal margin under angle (Fig. 21).

Remarks
Male genitalia without a baso-dorsal process on the gonostyli.

Description
HEAD.Brown, sometimes slightly suffused with red (Fig. 6C-E); vertex with hind margin strongly carinate and sides laminate, bisinuate in lateral view and elevated above eye (Fig. 6C-D); frons longitudinally wrinkled with 2 carinae extending on sides of cephalic process (Fig. 6E); cephalic process slightly more than twice as long as diameter of eye, projecting posterodorsad to dorsad (Fig. 6D); apical half of process lanceolate in anterodorsal view and with median carina anteriorly (Fig. 6C, E); posterior side of process with 2 carinae fused before apex; strongly sinuate carina under side of vertex, not reaching eye (Fig. 6D); postclypeus with sides strongly carinate and median carina; anteclypeus with strong median carina (Fig. 6E); labium very elongate, surpassing hind coxae but not reaching apex of abdomen.
HIND WINGS (Fig. 6A).Smoky brown-black, darker on apical ⅓; anal lobe and base suffused with red; milky patch on basal ⅔ from costal margin to vein M or Cu; veins darker than ground colour; 4-6 dark brown-black ocelli on clavus with round, waxy spot in middle; irregular white waxy spots on apical ⅓ and on disc; apex rounded, sutural margin emarginate at ⅔. LEGS (Fig. 6A).Elongate and slender; dark brown-black, with obsolete pale yellow-brown rings often only marked by small spots: 3-4 rings on pro-and mesofemora, 2 on mesotibiae and on metafemora; spines of metafemora with pale yellow-brown spots near base; metatibiae with 5 lateral and 7 apical spines; 12 spines apicoventrally on fi rst metatarsomere.ABDOMEN (Fig. 6A).Bright red with 2 longitudinal rows of 4 black spots dorsally.

Male genitalia
Red ; pygofer higher than long and with posterior margin slightly sinuate in lateral view (Fig. 22); anal tube slightly elongate, 1.39 times longer than broad at apex and with lateral margins nearly straight in dorsal view (Fig. 23); posterior margin strongly notched in lateral view (Fig. 22), with hind margin of anal opening projecting posteriorly and pointed in lateral view (Fig. 22), and hiding lateral margin apically in dorsal view (Fig. 23); gonostyli elongate, 1.22 times longer than high (dorsal process included, 1.72 times longer than high without process), surpassing anal tube and acutely rounded at apex in lateral view (Fig. 22); ventral margin straight on basal 1/5, then slightly sinuate (Fig. 22); dorsal margin, with basal strong digitiform process projecting laterodorsally and slightly curved posteriorly, strong hook laterally at middle of process projecting anteroventrally (Fig. 22); apex of digitiform process rounded and broad (Fig. 22-23).

Remarks
Belbina laetitiae sp.nov. is a member of the falleni+ group, showing a strong basodorsal digitiform process on the gonostyli.It can be separated (1) from B. bloetei by the less concave ventral margin and more rounded apex of the gonostyli in lateral view; (2) from B. falleni by the round cross section of the digitiform process and the more rounded apex of the gonostyli in lateral view; (3) from B. lambertoni by having an anal tube that is more strongly curved posteriorly in lateral view and more elongate in dorsal view; (4) from B. pionneaui by the more rounded apex of the basodorsal digitiform process of the gonostyli and the more strongly notched ventroapical margin of the anal tube under the anal opening in lateral view.

Male genitalia
Bright red (Figs 24-25); pygofer higher than long and with posterior margin bisinuate in lateral view (Fig. 24); anal tube slightly elongate, 1.1 times longer than broad at apex and with lateral margins slightly bisinuate, slightly concave on apical half in dorsal view (Fig. 25); posterior margin slightly notched in lateral view (Fig. 24), with hind margin of anal opening pointed posteriorly in lateral view (Fig. 24) and hiding lateral margin apically in dorsal view (Fig. 25); gonostyli elongate, 1.12 times longer than high (dorsal process included), surpassing anal tube and rounded at apex in lateral view (Fig. 24); ventral margin straight on basal 1/ 6, then slightly sinuate (Fig. 24); dorsal margin with basal, strong digitiform process projecting laterodorsally, strong hook laterally at middle of process projecting posteroventrally and posterior margin of process sinuate between apex and hook (Fig. 24); apex of digitiform process rounded and broad in lateral and dorsal view .

Remarks
Belbina lambertoni is a member of the falleni+ group, showing a strong basodorsal digitiform process on the gonostyli.It can be separated (1) from B. bloetei by the less concave ventral margin and more rounded apex of the gonostyli in lateral view; (2) from B. falleni by the round cross section of the digitiform process and the more rounded apex of the gonostyli in lateral view; (3) from B. laetitiae sp.nov.by the anal tube being less curved posteriorly in lateral view and less elongate in dorsal view; (4) from B. pionneaui by the more rounded apex of the basodorsal digitiform process of the gonostyli and the more strongly notched ventroapical margin of the anal tube under the anal opening in lateral view.

Diagnostic characters
(1) disc of tegmina red or orange (Fig. 8A); ( 2) cephalic process elongate and narrow, projecting anterodorsad (Fig. 8D-E); (3) ground colour of tegmina brown (Fig. 8A); ( 4) large-sized (more than 26 mm long); ( 5) black-brown spots with white center on tegmina (Fig. 8A).(Westwood, 1851) Det.Jérôme Constant 2013] (MNHN).Note: A neotype is here designated in order to stabilise the nomenclature in the group, following rule 75.3 of the International Code of Zoological Nomenclature (ICZN 1999).The specimen chosen here is probably the one on which Westwood (1851) based his description.It was collected by Jules Prosper Goudot in 1832 and included in the collections of the MNHN in 1834 (Adeline Soulier-Perkins pers.comm.).Westwood (1851) stated that the specimen he examined was from the collections of the "Mus.Jardin des Plantes, Paris", the name of the MNHN at that time.No specimen labelled Aphana madagascariensis was found in the collections of the MNHN, but the specimen chosen here perfectly matches the original description.Lallemand (1959) assumed that the type was lost and stated that, according to the description, the species probably belonged to Belbina or Cornelia.As it is not totally certain that the specimen is the one on which the original description was based, I designate it here as a neotype rather than recognize it as the holotype.NE Madagascar, Fampanambo, Jul. 1959, J. Vadon, 15°22' S, 49°38' E (MRAC).

Redescription
LT: ♂ (n = 1) 26.3 mm; ♀ (n = 1) 28.3 mm.HEAD.Yellow-brown with 2 darker patches on disc of vertex (Fig. 8C-E); yellow-orange patch around insertion of antennae (Fig. 8D); vertex with hind margin slightly carinate and obsolete median carina on disc extending posterad beyond hind margin; sides of vertex strongly carinate, laminate above eye and extending anteriorly to apex of cephalic process; sides of vertex slightly bisinuate in lateral view (Fig. 8C-D); frons with 2 carinae extending ventrally on sides of process (Fig. 8E); cephalic process about 3.2 times as long as diameter of eye, projecting anterodorsad, curved, elongate and narrow; apical half ventrally with median carina getting laminate near apex (Fig. 8D-E); transverse wrinkles dorsally at base of process (Fig. 8C); postclypeus with sides strongly carinate and slight median carina; anteclypeus with median carina (Fig. 8E); labium very long, reaching or surpassing apex of abdomen.
TEGMINA (Fig. 8A).Brown with irregular black-brown, often confl uent, small spots with pale yellowbrown center; center of spots covered with white waxy secretion in fresh specimens; no spots beyond nodal line of cross-veins except at apicosutural angle; apex of clavus with large black-brown marking followed by white patch along sutural margin; costal margin straight, slightly rounded after nodal line; apical margin oblique, straight in middle and with angles rounded; sutural margin sinuate.

Male genitalia
Very fi nely granulose, dark brown, paler along sides of anal tube ; pygofer higher than long and with posterior margin broadly rounded in lateral view (Fig. 26); anal tube slightly elongate, 1.31 times longer than broad at apex and with lateral margins sinuate in dorsal view (Fig. 27); posterior margin obliquely rounded and underside nearly straight in lateral view (Fig. 26); gonostyli elongate, 1.6 times longer than high, not surpassing anal tube and broadly rounded at apex in lateral view (Fig. 26); ventral margin slightly sinuate on basal ⅔ (Fig. 26); dorsal margin obliquely directed dorsally on basal half, then straight after angle, pointed hook-like tooth at half of basal oblique part curved lateroventrally (Fig. 26); gonostyli nearly not visible from above (Fig. 27).

Remarks
Male genitalia without basodorsal process on gonostyli.

Male genitalia
Very fi nely granulose, dark brown, with gonostyli slightly paler dorsally and ventrally (Figs 29-30); pygofer higher than long and with posterior margin broadly sinuate in lateral view (Fig. 29); anal tube elongate, 1.43 times longer than broad at apex and with lateral margins diverging on basal half, then broadly rounded in dorsal view (Fig. 30); curved ventrally and with posterior margin acutely rounded apically in lateral view (Fig. 29); gonostyli elongate, 1.63 times longer than high, not surpassing anal tube and broadly rounded at apex in lateral view (Fig. 29); all margins broadly rounded except dorsal margin obliquely straight on basal half (Fig. 29); lateral hook-shaped tooth at base of dorsal margin curved lateroventrally (Fig. 29); gonostyli nearly not visible from above (Fig. 30).

Remarks
Male genitalia without basodorsal process on gonostyli.
Belbina nympha can be separated (1) from B. bergrothi by the less elongate gonostyli, without large pale marking ventrally and more broadly rounded apex, and the less elongate anal tube; (2) from B. foliacea by having gonostyli more broadly rounded apically and without a strong angle above the mediodorsal tooth; (3) from B. madagascariensis by having the margins of the anal tube rounded laterally and the dorsal margin of the gonostyli broadly rounded on the apical half; (4) from B. recurva by not having the anal tube produced into a semi-circular lateral plate.

Male genitalia
Dark red (Figs 30-31); pygofer higher than long and with posterior margin straight, slightly rounded on dorsal ¼ in lateral view (Fig. 30); anal tube slightly elongate, 1.28 times longer than broad at apex and with lateral margins bisinuate in dorsal view (Fig. 31); posterior margin broadly rounded in lateral view (Fig. 30) with hind margin of anal opening slightly projecting posteriorly (Fig. 30) and hiding lateral margin apically in dorsal view (Fig. 31); gonostyli elongate, 1.35 times longer than high (dorsal process included), strongly surpassing anal tube and rounded at apex in lateral view (Fig. 30); ventral margin straight on basal 1/ 5, then sinuate (Fig. 30); dorsal margin with basal digitiform process slightly curved posteriorly and projecting laterodorsally, strong hook laterally at middle of process projecting ventrally (Figs 30-31); digitiform process narrowing from base to apex and acutely rounded apically .

Remarks
Belbina pionneaui is a member of the falleni+ group, showing a strong basodorsal digitiform process on the gonostyli.It can be separated from all species of the group by having the basodorsal digitiform process of the gonostyli narrowing from the base to the apex and the apical margin of the anal tube not notched under the anal opening in lateral view.

Male genitalia
Black-brown with gonostyli slightly paler ; pygofer higher than long and with posterior margin nearly straight in lateral view (Fig. 32); anal tube slightly transverse, 0.93 times as long as broad and with margins strongly projecting laterally on basal ⅔ into semi-circular lamina in dorsal view, apical ⅓ with lateral margins diverging towards apex (Fig. 33); apical margin acutely rounded in lateral view (Fig. 32); strong hump dorsally near base (Fig. 32); gonostyli elongate, 1.93 times longer than high, reaching apex of anal tube (Fig. 32); ventral margin broadly rounded to apex and dorsal margin strongly sinuate and with ante-apical notch in lateral view (Fig. 32); dorsal margin with strong, hook-shaped tooth at midlength pointing anteroventrally (Fig. 32); dorsal margin projecting laterally, semi-circular in dorsal view, with basal lobe and slightly folded internally near notch (Fig. 33).

Remarks
Male genitalia without basodorsal process on gonostyli.
Belbina recurva can be separated from all other species by having the anal tube produced into a semicircular lateral plate.

Discussion
The genus Belbina contains most of the species of Fulgoridae recorded from Madagascar, 12 out of 17 (Constant 2004b), with the 5 other species belonging to the genera Antsalovasia Constant, 2004, Radamana Distant, 1906and Zanna Kirkaldy, 1902.Belbina, Antsalovasia and Radamana are endemic to Madagascar while Zanna is also found in Afrotropical and Oriental regions.
Although the species of Belbina often have a wide distribution range (see maps on Figs 46-53), they are not regularly collected, probably due to their cryptic colour when sitting on tree trunks (see Figs 36,38,42).More fi eld work should be conducted to document their life-history and to make observations on the host plants and the biology of these insects.Although they are actively searched by collectors, the feeding habits, larval stages and eggs remain unknown for all species.