European Journal of Taxonomy 102: 1-37

http://dx.doi.org/10.5852/ejt.2014.102

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2014 • Constant J.

Research article

urn:lsid:zoobank.org:pub:2902587E-8F52-4E91-AA14-36339E54D05E

Revision of the Malagasy lanternfly genus Belbina Stal, 1863, with two

new species (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: [email protected]

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

Abstract. The Malagasy genus Belbina Stal, 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 Stal, 1866 is proposed as a junior

synonym of Belbina. The following new combinations are proposed: Belbina bergrothi (Schmidt, 1911)

comb. nov. and B. nympha (Stal, 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 Stal, 1863 and Cornelia atomaria (Brancsik, 1893) as

a junior synonym of Belbina nympha (Stal, 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 Stal, 1863,

B. laetitiae sp. nov., B. lambertoni Lallemand, 1922 and B. pionneaui Lallemand, 1922.

Keywords. Madagascar, lanternbug, Fulgoroidea, Homoptera, planthopper

Constant J. 2014. Revision of the Malagasy lanternfly genus Belbina Stal, 1863, with two new species (Hemiptera:

Fulgoromorpha: Fulgoridae). European Journal of Taxonomy 102: 1-37. http://dx.doi.org/10.5852.ejt.2Q14.102

Introduction

In the process of identifying material of Fulgoridae from Madagascar, three species attributed to the

genera Belbina Stal, 1863 or Cornelia Stal, 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 Stal (1863a) to accommodate two species: Belbina falleni Stal,

1863 (Madagascar) and Enchophora servillei Spinola, 1839 (patria incognita , now recognized to be

Madagascar). It was redescribed by Stal (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 Stal, 1863 as the type species,

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European Journal of Taxonomy 102: 1-37 (2014)

a view followed by Nast (1977) who clarified this issue according to the dates of publication of Stal’s

papers.

Stal (1866) later described Cornelia for C. nympha Stal, 1866, from Madagascar.

Gerstaecker (1873) described the genus Pyrgoteles for Enchophora sicca Walker, 1851 and considered

the genus close to Belbina Stal, 1863, and to the Oriental genera Aphaena Guerin-Meneville, 1834 and

Prolepta Walker, 1851.

Karsch (1890) synonymized Pyrgoteles under Belbina. Nevertheless, in 1893 the same author described

a new species in the genus Pyrgoteles , P. cristatus Karsch, 1893, thus de facto revalidating the genus

Pyrgoteles.

Melichar (1908) added one species from Africa (Tanzania) to Cornelia'. C. usambarae.

Schmidt (1911) described another species in Cornelia , C. bergrothi from Madagascar and placed

Cornelia in the Aphaenini of the Fulgorinae (note that the concept of the Fulgorinae at that time equals

present Fulgoridae and Aphaenini equals present Aphaeninae).

In 1918, Schumacher reviewed Druentia Stal, 1866 and considered Pyrgoteles Gerstaecker, 1873 and

Belbina Stal, 1863 as junior synonyms of Druentia, the latter synonymy being invalid since Belbina has

priority over Druentia Lallemand (1922) restored Belbina by adding two new species from Madagascar,

B. lambertoni (with a variety, minuta ) and B. pionneaui.

In his catalogue of the Fulgoridae, Metcalf (1947) listed 3 species in Cornelia and 4 in Belbina (and one

variety). He included the species sicca Walker, 1951 in Druentia Stal, 1866 although he also strangely

considered it the type species of Belbina Stal, 1863, even placing the two genera in different subfamilies:

Druentia in the Amyclinae, Amy cl ini and Belbina in the Aphaeninae, Enchophorini.

In 1950, Lallemand added 2 species in Belbina : B. recurva and B.foliacea from Madagascar.

In his revision of the Afrotropical Fulgoridae, Lallemand (1959) considered Druentia as strictly

African and Cornelia and Belbina as restricted to Madagascar. He placed the 3 genera in the subfamily

Enchophorinae Haupt, 1829, which appears to be exclusively Neotropical, with Enchophora Spinola,

1839 as the type genus (type species, E. recurva (Olivier, 1891), described from Surinam; see also

phylogenetic trees based on molecular data in Urban & Cry an 2009). He considered Enchophorinae a

subfamily instead of following Metcalf (1947), who considered it a tribe within the Aphaeninae Blanchard,

1847. He gave a key to separate the genera based on characters such as the width of the head and the

shape of the tegmina and cephalic process. He transferred Belbina foliacea and Enchophora atomaria

Brancsik, 1893 to Cornelia , described Belbina bloetei and B. vicina (the 4 species from Madagascar),

and transferred Cornelia usambarae Melichar, 1908 (from Tanzania) to Druentia. He synonymized

Belbina lambertoni var. minuta Lallemand, 1922 under B. lambertoni Lallemand, 1922. Finally, he

erroneously designated a specimen in his collection as lectotype for Belbina servillei (Spinola, 1839), a

specimen that is not from the type series and not even the right species.

Constant (2004b) gave a key to the genera of Fulgoridae found in Madagascar (mainly based on

Lallemand 1959) and a checklist of the Fulgoridae of Madagascar, including 7 species in Belbina and 4

species in Cornelia , and stated that Cornelia and Druentia could be junior synonyms of Belbina. He also

mentioned the invalid designation by Lallemand (1959) of a lectotype for Belbina servillei.

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CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

This paper aims to clarify the taxonomy of the genus and the specific concept of several taxa based on

illustrated morphological characters, with the description of two new species.

The current concepts of all the taxa mentioned are documented in FTOW (Bourgoin 2014).

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 difficulty to correctly inflate 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:

BMNH

CAS

FSAG

HNHM

INHS

MHNL

MNHN

MRAC

NCSU

NHRS

NMPC

NMW

RBINS

RMNH

SDEI

ZIN

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

California Academy of Sciences, San Francisco, USA (Norman D. Penny)

Faculte des Sciences agronomiques de Gembloux, Gembloux, Belgium (Eric Haubruge,

lea n nin e Bortels)

Hungarian Natural History Museum, Budapest, Hungary (Andras Orosz)

Illinois Natural History Survey, Champaign, USA (Chris Dietrich)

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

Museum National d’Histoire Naturelle, Paris, France (Thierry Bourgoin, Adeline

Soulier-Perkins)

Musee royal de l’Afrique centrale, Tervuren, Belgium (Marc De Meyer, Ugo Dall’Asta)

North Carolina State University, Raleigh, USA (Bob Blinn)

Naturhistoriska Riksmuseet, Stockholm, Sweden (Gunvi Lindberg)

National Museum, Praha, Czech Republic (Petr Kment)

Naturhistorisches Museum Wien, Vienna, Austria (Herbert Zettel)

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

Nationaal Natuurhistorisch Museum (“Naturalis”), Leiden, Netherlands (Jan van Tol,

Yvonne van Nierop)

Senckenberg Deutsches Entomologisches Institut, Mtincheberg, Germany (Stephan

Blank)

Russian Academy of Sciences, Zoological Institute, St. Petersburg, Russia (Vladimir

Gnezdilov)

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European Journal of Taxonomy 102 : 1-37 ( 2014 )

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

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 Gerstaeclcer, 1895 (the latter mentioned as ‘undet. Fulgoridae’ in Urban & Cryan 2009:

fig. 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 ).

Belbina Stal, 1863

Belbina Stal, 1863a: 232. Type species: Belbina falleni Stal, 1863 by subsequent designation by

Lallemand (1959).

Cornelia Stal, 1866: 133 (key), 142. syn. nov. Type species: Cornelia nympha Stal, 1866, by monotypy.

Belbina - Stal 1863b: 580 (redescription). — Stal 1866: 134 (key), 143 (description, key to species). —

Karsch 1890: 61 (listed; erroneous synonymy with Pyrgoteles Gertsaecker, 1873). —Jacobi 1917: 520

(listed). —Tallemand 1922: 62 (listed, description of new species). —Metcalf 1947: 123 (catalogued).

— Tallemand 1950: 84 (description of new species). —Tallemand 1959: 85 (key), 88 (description, key

to species). —Nast 1977: 30 (nomenclature). — Constant 2004b: 31 (key, list of species).

Cornelia- Karsch 1890: 61 (listed). — Schmidt 1911: 241 (listed, description of new species). —Jacobi

1917: 520 (listed). —Metcalf 1947: 122 (catalogued). —Tallemand 1959: 85 (key), 86 (description,

key to species). — Constant 2004b: 31 (key, list of species).

Druentia- Schumacher 1918: 111 pars (erroneous synonymy).

Remarks

Belbina and Cornelia are synonymized because it is not possible to find 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 Gerstaeclcer, 1895 and [Druentia

+ Rhicnophloea] is sister to a larger clade of Old World taxa which contains, among others, a group

4

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

formed by [Samsama Distant, 1906 + Belbina + Metaphaena Schmidt, 1905 + Anecphora + Penthicodes]

(Urban, Bartlett & Cryan unpubl. data).

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 defined 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.

Identification 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 hind¬

winged specimens also have ones with orange hind wings.

The red colour of the abdomen can also fade to orange or yellow in collection specimens, especially if

they have been preserved in ethanol.

1. Hind wings red or orange on disc (Figs 1A, 2A) .2

- Hind wings with disc bluish, greenish or brown (Figs 3A, 6A).10

2. Ground colour of tegmina orange or red (Figs 1A, 7A, 10A) .3

- Ground colour of tegmina brown to pale rosy brown (Figs 2A, 4A) .5

3. Mesonotum with 2 black patches on disc (Fig. 1C); small-sized (less than 22 mm long) .

. Belbina bergrothi (Schmidt, 1911)

- Mesonotum without black patches (Figs 7C, 10C); larger (more than 24 mm long) .4

4. Tegmina orange to red with minute black spots; 3 large black patches on costal cell and one at

the base of clavus (Fig. 7A). Belbina lambertoni Lallemand, 1922

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European Journal of Taxonomy 102: 1-37 (2014)

- Tegmina dark red with membrane pale brown; numerous, irregular, blackish brown spots (Fig.

10A). Belbina pionneauiLaWQmmd, 1922

5. Head with a foliaceous process above the eye, higher than diameter of eye in side view;

median carina of pronotum strongly sinuate and elevated posteriorly in side view (Fig. 5D) .

. Belbina foliacea Lallemand, 1959

- Head without large foliaceous process above the eye, process reduced to a lamina not

higher than half of diameter of eye; median carina of pronotum not strongly sinuate and

elevated posteriorly in side view (Figs 4D, 11D) .6

6. Small-sized, less than 22 mm long; cephalic process directed anterodorsad (Fig. 9D);

abdomen with 2 dorsal rows of black spots; hind wings with 4-6 little black spots on red

zone (Fig. 9A) . Belbina nympha (Stal, 1866)

- Larger, more than 26 mm long; cephalic process angulously directed dorsad or posteriorad

(Fig. 2D), or, if not, dorsum of abdomen red without rows of black spots; hind wings with

2 small black spots or larger black spots (Figs 4A, 8A) .7

7. Cephalic process narrow, elongate, obliquely directed anterodorsad (Fig. 8D); blackish brown

spots with pale center on tegmina (Fig. 8A). Belbina madagascariensis (Westwood, 1851)

- Cephalic process broad, strongly curved dorsad or posteriorad (Fig. 2D); dark spots on tegmina

without pale center (Figs 2A, 4A).8

8. Cephalic process projecting dorsad, apex not surpassing level of anterior margin of eye in

lateral view (Fig. 2D); abdomen without rows of black spots dorsally; no large black patch on

basal half of claval joint; no paler transverse patch at half length of tegmen (Figs 2A, 4A) ...9

- Cephalic process more strongly curved and projecting posterodorsad, reaching level of

posterior margin of eye in lateral view (Fig. 1 ID); abdomen with 2 rows of blackish spots

dorsally; one large black elongate patch on basal half of claval joint; one transverse paler patch

at about half of length of tegmen (Fig. 11 A) . Belbina recurva Lallemand, 1950

9. Ground colour of tegmina and head pale rosy brown; a rectangular paler patch on clavus (Fig.

2A) . Belbina bloetei Lallemand, 1959

- Ground colour of tegmina and head brown; no evident paler patch on clavus (Fig. 4A) .

. Belbina falleni Stal, 1863

10. Head, pro- and mesonotum red (Fig. 12C-E); ground colour of tegmina bluish green (Fig. 12A)

. Belbina servillei (Spinola, 1839)

- Head, pro- and mesonotum brown (Figs 3C-D; 6C-D); ground colour of tegmina bluish green

or brownish (Figs 3A, 6A) .11

11. Ground colour of tegmina bluish green; hind wings nearly totally infuscate with a suturo-

basal, not well limited zone, and spots, paler, milky (Fig. 6A); larger (24 mm long in males,

27 mm long in females) . Belbina laetitiae sp. nov.

- Ground colour of tegmina brownish with a dark reddish band after half of length; hind wings

with paler, bluish green disc very distinctly separated from blackish apex and sutural margin

(Fig. 3A); smaller (19 mm long in males). Belbina bourgoini sp. nov.

6

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

Figs 1-3. — 1. Belbina bergrothi. 2. B. bloetei. 3. B. bourgoini sp. nov. — A. Habitus, dorsal view.

B. Habitus, lateral view. C. Vertex, pro- and mesonotum, dorsal view. D. Head and prothorax, lateral

view. E. Frons, normal view. — Scale bar: 10 mm (C-E not to scale).

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European Journal of Taxonomy 102: 1-37 (2014)

Figs 4-6. — 4. Belkina falleni. 5. B. foliacea. 6. B. laetitiae sp. nov. — A. Habitus, dorsal view.

B. Habitus, lateral view. C. Vertex, pro- and mesonotum, dorsal view. D. Head and prothorax, lateral

view. E. Frons, normal view. — Scale bar: 10 mm (C-E not to scale).

8

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

Figs 7-9. — 7. Belbina lambertoni. 8. B. madagascariensis. 9. B. nympha. — A. Habitus, dorsal view.

B. Habitus, lateral view. C. Vertex, pro- and mesonotum, dorsal view. D. Head and prothorax, lateral

view. E. Frons, normal view. — Scale bar: 10 mm (C-E not to scale).

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European Journal of Taxonomy 102: 1-37 (2014)

Figs 10-12. — 10. Belbina pionneaui. 11. B. recurva. 12. B. servillei. — A. Habitus, dorsal view.

B. Habitus, lateral view. C. Vertex, pro- and mesonotum, dorsal view. D. Head and prothorax, lateral

view. E. Frons, normal view. — Scale bar: 10 mm (C-E not to scale).

10

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

Belbina bergrothi (Schmidt, 1911) comb. nov.

Figs 1A-E, 13-14, 36, 46

Cornelia bergrothi Schmidt, 1911: 242 (type in SDEI).

Cornelia bergrothi - Jacobi 1917: 526 (listed). —Metcalf 1947: 122 (catalogued). —Lallemand 1959:

88, fig. 34 (key, description, lateral view of head). — Constant 2004b: 31 (listed).

Figs 13-18. Male genitalia. — 13-14. Belbina bergrothi. 13. Lateral view. 14. Dorsal view. — 15-16.

B. bloetei. 15. Lateral view. 16. Dorsal view. —17-18. B. bourgoini sp. nov. 17. Lateral view. 18. Dorsal

view. — An = anal tube, Py = pygofer, G = gonostylus. Photographs by Y. Laurent & I. Bachy.

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European Journal of Taxonomy 102: 1-37 (2014)

Diagnostic characters

(1) disc of hind wings orange or red (Fig. 1A); (2) 2 black patches on mesonotum (Fig. 1C); (3) size less

than 22 mm long; (4) cephalic process directed dorsad, narrow in anterior view (Fig. 1D-E).

LT: $ (n = 3) 17.8 mm (17.0-19.0); ? (n = 2) 20.5 mm (20.0-21.0).

Figs 19-23. Male genitalia. — 19-20. Belbina falleni. 19. Lateral view. 20. Dorsal view. — 21.

B.foliacea , gonostyli, normal view. — 22-23. B. laetitiae sp. nov. 23. Lateral view. 24. Dorsal view. —

An = anal tube, Py = pygofer, G = gonostylus. Photographs by Y. Laurent & I. Bachy.

12

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

Material examined

Holotype

MADAGASCAR: $ (on photographs), [Madagasc. Amber Geb.] [Holotypus] [ Cornelia Bergrothi

Schmidt $ Edm. Sc hm idt determ. 1911] [Coll. Breddin] [Dtsch. Entomol. Institut Berlin], Montagne

d’Ambre (Ambohitra), 12°30’ S, 49°10’ E (SDEI).

Figs 24-29. Male genitalia. — 24-25. Belbina lambertoni. 24. Lateral view. 25. Dorsal view. — 26-

27. B. madagascariensis . 26. Lateral view. 27. Dorsal view. — 28-29. B. nympha. 28. Lateral view.

29. Dorsal view. — An = anal tube, Py = pygofer, G = gonostylus. Photographs by Y. Laurent & I. Bachy.

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European Journal of Taxonomy 102: 1-37 (2014)

Additional material

MADAGASCAR: 1 3, 1 ?, E Madagascar, Massif du Marojejy (res. nat. integr. 12), 1300 m, 2-8 Dec.

1972, A. Peyrieras, 14°26’ S, 49°44’ E (MNHN); 1 3, same data, 4 Dec. 1972 (RBINS); 1 ?, same data,

12 Dec. 1972 (RBINS); 1 $, Fianarantsoa province, Parc National Ranomafana, Belle Vue et Talatakely,

Figs 30-35. Male genitalia. — 30-31. Belbinapionneaui. 30. Lateral view. 31. Dorsal view. — 32-33.

B. recurva. 32. Lateral view. 33. Dorsal view. — 34-35. B. servillei. 34. Lateral view. 35. Dorsal view.

— An = anal tube, Py = pygofer, G = gonostylus. Photographs by Y. Laurent & I. Bachy.

14

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

Figs 36-42. Belbina in natura. — 36. B. bergrothi. Vohimana, 4 Jan 2006, photograph by N. Cliquennois.

— 37-38. B.falleni. 37. Montagne d’Ambre, Joffreville, 2009, photograph by R.A. Butler. 38. Montagne

d’Ambre, 27 Oct. 2009, photograph by N. Cliquennois. — 39. B. lambertoni , S, Berenty, 15 Nov.

2008, photograph by J. Cryan. — 40-42. B. recurva. 40. Marojejy, Cascade Humbert, 22 Nov. 2005,

photograph by D. C. Lees. 41. Anajanaharibe Sud, 2 May 2010, photograph by B. Rowe. 42. Masoala

Peninsula, 30 May 2009, photograph by D. Fenolio (©2011 Dante Fenolio /www. anotheca.com , with

permission).

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European Journal of Taxonomy 102: 1-37 (2014)

21°15.99’ S 47°25.2r E, 1020 m, 21-28 Jan. 2002, Malaise trap, secondary forest, R. Harin’Hala

(CAS); 1 S, Moramanga region, Andaside, PL hotel, 18°56.820’ S, 48°25.155’ E, 22 Nov. 2005, 930 m,

Bourgoin, Ouvrard, Attie, Soulier-Perkins (MNHN); 1 $, no data (FSAG); 1 $, Antananarivo province,

46 km NE of Ankazobe, Ambohitantely, 18°11.88’ S, 47°16.89’ E, Malaise trap in sclerophyl forest, 700

m, 16-27 Jan. 2005, Irvin & Harin’Hala (CAS); 1 route d’Anosibe, 11 Nov. 1961 (MHNL).

Examined on photograph

MADAGASCAR: 1 specimen, Vohimana, 4 Jan. 2006, 13:35, Nicolas Cliquennois (Fig. 36), 18°55’ S,

48°30’E.

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 (Figs 13-14); 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.

Distribution

See Fig. 46.

Belbina bloetei Lallemand, 1959

Figs 2A-E, 15-16, 46

Belbina blotei Lallemand, 1959: 92, fig. 40a-b (lateral view of head, wing venation) (type in RMNH).

Belbina blotei - Constant 2004b: 31 (listed).

Note

The species epithet refers to the Dr H.C. Blote (RMNH) and is emended following the article 32.5.2.1

of ICZN (1999).

Diagnostic characters

(1) disc of hind wings red (Fig. 2A); (2) size more than 26 mm; (3) cephalic process projecting dorsad,

broad (Fig. 2D-E); (4) ground colour of tegmina and head pale rosy brown; (5) rectangular pale patch

on clavus (Fig. 2A, C).

LT: S (n = 1) 27.0 mm ; $ (n = 1) 29.5 mm.

16

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

Material examined

Holotype

MADAGASCAR: § (examined on photographs), [Madagascar, coll. Lambert.] [environs de Tananarive]

[Type] [Belbina blotei Lall., V. Lallemand det., 195] [RMNH.INS.780532], Tananarive (Antananarivo),

18°55’ S, 47°3r E (RMNH). -Note: “Lambert” = Lamberton.

Paratypes

MADAGASCAR: 1 $ (examined on photographs), [Madagascar, rec. Lamberton] [environs de

Tananarive] [Belbina blotei Lallemand, V. Lallemand det., 195] [Paratype] [RMNH.INS.780531]

(RMNH); 1 S, 1 ?, [environs de Tananarive] (FSAG). Note: Lallemand (1959) stated that all types are

deposited in RMNH but he actually retained 2 paratypes in his collection (now in FSAG).

Additional material

MADAGASCAR: 1 S, Mahajanga Prov., Foret de Tsimembo, 8.7 km 336° NNW of Soatana, 19°L17”

S, 44°26’26” E, 20 m, 21-25 Nov. 2001, at light in tropical dry forest, Fisher, Griswold et al. (CAS);

2 near Tananarive, Lamberton (RBINS); 2 §§, Morondava a Mahabo, Last 1854-91, 20°20’ S,

44°28’30” E (MNHN); 1 S, Madagascar (NMPC); 1 no data (NCSU).

1 5, erroneously labelled Soekaboemi, Java (INHS).

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 A, 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) from B.falleni , B. laetitiae sp. nov., B. lambertoni and B. pionneaui by

the more strongly concave ventral margin of the gonostyli in lateral view; (2) from B. falleni by the less

excavate dorsal margin of the gonostyli in dorsal view; (3) from B. laetitiae sp. nov. and B. lambertoni

by the more acute apex of the gonostyli in lateral view; (4) from B. pionneaui by the more rounded apex

of the basodorsal digitiform process of the gonostyli.

Distribution

See Fig. 46.

Belbina bourgoini sp. nov.

Figs 3A-E, 17-18, 47

urn:lsid:zoobank.org:act:266C852F-6F78-48E8-AC72-3A286E350836

Diagnostic characters

(1) disc of hind wings light white-bluish (Fig. 3A); (2) small-sized (less than 22 mm long); (3) tegmina

brown with dark red-brown band (Fig. 3A); (4) head, pro- and mesonotum brown (Fig. 3C-E).

17

European Journal of Taxonomy 102: 1-37 (2014)

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.

Material examined

Holotype

MADAGASCAR: S, [Marojezy XII-1972] [Museum Paris, Madagascar Est, mission C.N.R.S., RC.P

n° 225], 14°26’ S, 49°44’ E (MNHN).

Paratypes

MADAGASCAR: 1 S, [Madagascar Est, Massif du Marojejy (res. nat. integr. 12) 1300m, 2/8-XII-

1972, A. Peyrieras] [Museum Paris] (RBINS); 1 S, [Coll. P. Bleuzen, Lakato, Madagascar, 13.1.1962],

19°IE S, 48°26’ E (MHNL).

Description

LT: S (n = 2) 19.2 mm (19.0-19.5).

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.

Thorax. Yellow-brown varied with irregular paler spots (Fig. 3C-D); pronotum carinate anteriorly;

sinuate carinae at anterior margin of disc, reaching anterior margin before median carina (Fig. 3C);

median carina strong, sinuate in lateral view; hind margin strongly elevated above level of mesonotum

medially (Fig. 3D); impressed point on disc on each side of carina; obsolete tubercles at each side of disc

(Fig. 3C); sides of prothorax longitudinally carinate; lateral lobes of pronotum rounded posteriorly (Fig.

3D-E); mesonotum with median carina stopped at scutellum posteriorly; strongly sinuate carina on each

side of disc, Y-shaped anteriorly; scutellum transversely wrinkled (Fig. 3C).

Tegmina (Fig. 3A). Pale yellow-brown with irregular brown-black markings; ground colour red-brown

on middle; clavus paler; white marking along sutural margin at level of nodal line; costal margin slightly

rounded; apical angles rounded; sutural and apical margins sinuate, the latter oblique; ratio FTg/BTg =

2 . 6 .

Hindwings (Fig. 3A). Black-brown with large, light white-buish discal patch of basal half, reaching

costal margin and extending posteriorly along vein Cu without reaching sutural margin; black-brown

elongate marking basally along vein M; big black spot along vein PCu followed by 1-2 smaller spots;

apex rounded, sutural margin sinuate.

Pegs (Fig. 3A). Elongate and slender; femora brown with 3—4 narrow, sinuate and often incomplete pale

yellow rings; pro- and mesotibiae brown with 2 pale yellow rings; metatibiae brown with pale yellow

markings at spines; pro- and mesotarsi black-brown; first metatarsomere pale yellow-brown on basal 2 /3;

rest of metatarsi brown; metatibiae with 6-7 lateral and 7 apical spines; first metatarsomere with 9

apicoventral spines.

18

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

Abdomen (Fig. 3A). Brown, darker ventrally.

Male genitalia

Dark brown (Figs 17-18); 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 Vs (Fig. 17); dorsal margin with basal slender digitiform process

projecting laterodorsally, pointed tooth laterally at middle of process projecting antero ventrally, apex of

digitiform process narrowing apically (Fig. 17); apical % 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.

Distribution

See Fig. 47.

Belbina falleni Stal, 1863

Figs 4A-E, 19-20, 37-38, 47

Belbina falleni Stal, 1863a: 233 (type inNMW).

Belbina vicina Xallemand, 1959: 90, fig. 38 (key, description and dorsal view of head and thorax) (type

in FSAG). syn. nov.

Belbina falleni - Stal 1866: 144 (key, description). —Jacobi 1917: 527 (listed). —Metcalf 1947: 123

(catalogued). — Xallemand 1959: 93, fig. 41 (key, description and lateral view of head). — Constant

2004b: 31 (listed).

Belbina vicina - Constant 2004b: 31 (listed).

Diagnostic characters

(1) disc of hind wings red with black markings (Fig. 4A); (2) ground colour of tegmina pale brown (Fig.

4A); (3) sides of vertex laminate above eye but not foliaceous (Fig. 4C-D); (4) large-sized (more than 26

mm long); (5) cephalic process broad, strongly curved dorsad (Fig. 4D); (6) clavus with colour pattern

similar to the rest of the tegmen (Fig. 4A).

LT: $ (n = 5) 26.6 mm (26.0-27.0); ? (n = 7) 29.1 mm (28.5-30.2).

Material examined

Type material

MADAGASCAR: Holotype of Belbina falleni , $, [Madagascar, Coll. Signoret] [Falleni det. Signoret]

[Type] (NMW).

MADAGASCAR: Holotype of Belbina vicina Xallemand, 1959, S, [Manjakandriana, Madagascar]

[Type.] [Belbina vicina Tall., V. Xallemand det., 1957], 18°55’ S, 47°48’ E (FSAG).

19

European Journal of Taxonomy 102: 1-37 (2014)

Additional material

MADAGASCAR: 1 §, Manjakandriana (FSAG); 2 33 , 1 $, near Tananarive, Lamberton, Tananarive

(Antananarivo), 18°55’ S, 47°31’ E (RBINS); 2 33, Vohemar (Iharaoa), 13°2E S, 50°00’ E (RBINS,

NCSU); 1 5, NMadagascar (BMNH); 1 §, Montagne d’Ambre, Feb. 1930, Sicard, Montagne d’Ambre

(Ambohitra), 12°30’ S, 49° 10’ E (MNHN); 1 §, idem, Dec. 1930 (MNHN); 1 $, no locality, Apr.

1930, Sicard (MNHN); 1 §, Madagascar, coll. De Bergevin (MNHN); 4 §§, Madagascar (2: MNHN,

2: NMW); 1 3 , N Madagascar, 37 km S of de Vohemar, Analalava forest, 25 m, 10-18 Dec. 1968, R

Viette & P. Griveaud, 13°36’ S, 49°59’ E (MNHN); 1 3 , Diego-Suarez, coll. De Bergevin, Diego Suarez

(Ampanolahamirafy), 12°16’ S, 49°17’ E (MNHN); 1 §, Plateau de l’Ankarafantsika, W Madagascar,

coll. De Bergevin 16°16’ S, 46°35’ E (MNHN); 1 ?, N Madagascar (BMNH); 1 $, no data (NCSU).

Examined on photographs

MADAGASCAR: 1 specimen, Montagne d’Ambre, 27 Oct. 2009, 15:05, Nicolas Cliquennois (Fig. 38);

1 specimen, Montagne d’Ambre, Joffreville (Ambohitra), 2009, Rhett A. Butler (Fig. 37).

Male genitalia

Dark red (Figs 19-20); 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 A, 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).

Remarks

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 identified in his collection as

B.falleni ; both specimens were from the same locality (Manjakandriana). Male genitalia are similar and

intraspecific 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.

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.

Distribution

See Fig. 47.

20

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

Belbina foliacea Xallemand, 1950 comb. rev.

Figs 5A-E, 21, 47

Belbina foliacea Xallemand, 1950: 84 (type in FSAG).

Cornelia foliacea - Xallemand 1959: 87, fig. 32a-c (key, description, lateral view of head and male

genitalia). — Constant 2004b: 31 (listed).

Diagnostic characters

(1) disc of hind wings red (Fig. 5A); (2) carina above eye strongly expanded dorsad into foliaceous

process (Fig. 5C-E); (3) cephalic process strongly curved dorsad (Fig. 5D); (4) tegmina variegated

brown-olivaceous; (5) veins of tegmina elevated (Fig. 5A).

XT: f (n = 1) 24.0 mm; $ (n = 1) 29.9 min

Material examined

Holotype

MADAGASCAR: 3, [Maromandia, Madagascar, Xamberton] [Type.] [Cornelia foliacea Tall., V.

Xallemand det., 1954], 18°56’ S, 49°03’ X (FSAG).

Paratype

MADAGASCAR: 1 §, no label, probably same data as holotype.

Male genitalia

The male genitalia were dissected and drawn by Henry Synave for Xallemand’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.

Belbina foliacea can be separated from B. bergrothi , B. madagascariensis, B. nympha and B. recurva by

the strong angle above the medio-dorsal tooth and the broadly rounded apex of the gonostyli.

Distribution

See Fig. 47.

Belbina laetitiae sp. nov.

Figs 6A-E, 22-23, 43M5, 48

um:lsid:zoobank.org:act:87E89C54-19CC-48C8-A2B0-E6CE95E8410A

Belbina servillei - Xallemand 1959: 88, fig. 36a-c (key, description, lateral view of head and pronotum,

male genitalia) [non Belbina servillei (Spinola, 1839)].

21

European Journal of Taxonomy 102: 1-37 (2014)

Diagnostic characters

(1) disc of hind wings milky, base and anal lobe suffused with red (Fig. 6A); (2) head, pro- andmesonotum

brown (Fig. 6C-E); (3) ground colour of tegmina bluish green (Fig. 6A); (4) large-sized (more than 24

mm long); (5) cephalic process broad, projecting dorsad to posterodorsad (Figs 6D-E).

LT: 8 (n = 8) 24.3 mm (23.2-25.4); ? (n = 11) 26.8 mm (25.1-28.4).

Etymology

Dedicated to Mrs Laetitia Despontin, the mother of my c hil dren Emilie and Guillaume.

Material examined

Holotype

MADAGASCAR: 8, [Madagascar, LaMandraka, xii.2000] [Coll. I.R.Sc.N.B., DonT. Neef de Sainval,

I.G. 30.731], 18°55’ S, 47°56’ E (RBINS).

Paratypes

MADAGASCAR: 1 8, same data as holotype; 1 8, [Coll. I.R.Sc.N.B. Madagascar Est, district

Sambava, Marojejy, Ambinanitelo, 500m, XII.58, Raharizonina], 15°2E S, 49°35’ E (RBINS); 1 §,

[Coll. I.R.Sc.N.B. Madagascar Est, district Sambava, Marojejy, Ambodifiakarana, 1600m, XII;58,

Raharizonina], 17°12’ S, 48°1E E (RBINS); 1 $, [Coll. I.R.Sc.N.B. Madagascar Sud-Est, S. kavondro,

225m, Foret Isaka, Fort Dauphin, 24.11.58, R Griveaud], Fort-Dauphin (Tolaoaro), 25°02’ S, 47°00’ E

(RBINS); 1 5, [CASENT 3004835] [Tananarive Madagascar, Oct., 1949], Tananarive (Antananarivo),

18°55’ S, 47°3E E (CAS); 1 8, [Madagascar, Antongil B., Mocquerys] [Madagascar] [1204] [Coll.

Hungarian Nat. Hist. Museum, Budapest], 15°45’ S, 49°50’ E (HMNH); 4 5 ?, [Madagascar, Antongil

B., Mocquerys] [Coll. Hungarian Nat. Hist. Museum,Budapest] (HMNH); 1 j, [212,69] [Museum Paris,

Madagascar, 151 et suiv.-69] [Museum Paris, MNHN (EH), 3948] (MNHN); 1 §, [Goudot, Madagascar,

1832] [Museum Paris, MNHN (EH), 3947] (MNHN); 1 8, [Madagascar, Fenerive, E. Perrot] [Museum

Paris, Oberthur 101-96] [Museum Paris, MNHN (EH), 3950] Fenerive (Fenoarivo Atsinanana), 17°22’

S, 49°25’ E (MNHN); 1 $, [Museum Paris, Madagascar, cote Ouest, Grandidier 1808-71] [Plateau

de FAnkarafantsika, Madagascar (Ouest)] [Museum Paris, MNHN (EH), 3949], 16°16’ S, 46°35’ E

(MNHN); 1 8 , [Madagascar Est, district de Maroantsetra, station forestiere de Farankaraina, route

de Navana, km 16.5, vallee d’Antoroka 100m, 8/18-1-1964, P Viette] [Museum Paris, MNHN (EH),

3951], 15°26’ S, 49°44’ E (MNHN); 1 8, 1 ?, [Mus. Roy. Afr. Centr., Madagascar Est: Ambodivoangy,

1.1960, J. Vadon] [Belbina servillei Spin., H. Synave det. 1963], 15°17 , 50” S, 49 0 36 , 47 ,, E, (MRAC);

1 8, [Coll. Mus. Tervuren, N.E. Madagascar: Ambodivoangy, VIII. 1961, J. Vadon] [Belbina servillei

Spinola, H. Synave det. 1963] (MRAC); 1 $, [Coll. Mus. Tervuren, N.E. Madagascar: Ambodivoangy,

VIII. 1961, J. Vadon] (MRAC); 1 §, [Madagascar: Fampanambo, III. 1961, J. Vadon] [Belbina servillei

Spin., H. Synave det. 1963], 15°22’ S, 49°38’ E (MRAC); 1 8 , [Madagascar Est, massif du Marojejy

(res. nat. integr. 12) 600 m, 9/12-XII-1972, A. Peyrieras] [Museum Paris, MNHN (EH), 3952], 14°26’

S, 49°44’ E, (MNHN); 1 J, [Madagascar Est, massif du Marojejy (res. nat. integr. 12)] [Ambatomitatao,

400 m, A. Peyrieras, 1.1973] [Museum Paris, MNHN (EH), 3953], 13°47 , 30» S, 49°5E E (MNHN); 1

5, [Madag., (on the reverse) 91 158] [Belbina nov. sp. trop abime pour etre decrit, det. V. Lallemand

19] (BMNH); 1 8, [Madagascar, collection Le Moult] [Belbina servillei Spin., V. Lallemand det.]

(FSAG); 1 8, [Manjakandriana, Madagascar], 18°55’ S, 47°48’ E (FSAG); 1 8, [Institut Scientifique

Madagascar] [Madagascar Sud, det. Fort Dauphin, Antanimora, 300m, xii-59, Raharizonina] [Belbina

servillei Sign., V. Lallemand det., 1965], 24° 14’ S, 47° 15’ E (FSAG); 1 8 , [Rogez, Madagascar, Mus.

Praha] [Enchophora sp.] (NMPC); 1 8, [Madagascar, Collection Le Moult] [Ne pas etaler, defectueux]

(NCSU); 2 88, [Madagascar, Marsantsetra], Marsantsetra is a misspelling of Maroantsetra (NCSU).

22

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

Examined on photograph

MADAGASCAR: 1 specimen (Figs 43M5), lie Sainte Marie, 25 Nov. 2011, Paul Bertner, 16°50’ S,

49°55’E.

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.

Thorax. Brown with mesonotum and meso- and metapleura red (Fig. 6C-E); pronotum with strong

median carina sinuate in lateral view (Fig. 6C-D); hind margin strongly elevated above level of

mesonotum medially; sides of prothorax longitudinally carinate; lateral lobes of prothorax rounded

posteriorly (Fig. 6D); mesonotum (Fig. 6C) with median carina stopped posteriorly before scutellum;

strongly sinuate carina on each side of disc, sometimes divided into external, sinuate carina and internal,

posterior, angulous carina; scutellum transversely wrinkled.

Tegmina (Fig. 6A). Green-brown to pale olivaceous-brown with irregular, dense, black-brown to black

markings; apical A brown-black with large, unprecise, darker markings and some minute, whitish spots;

small white patch along sutural margin at nodal line with black spot in middle; costal and apical margins

slightly rounded, the latter oblique; apical angles rounded; sutural margin sinuate.

Hind wings (Fig. 6A). Smoky brown-black, darker on apical A; anal lobe and base suffused with red;

milky patch on basal A 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 A

and on disc; apex rounded, sutural margin emarginate at 2 A.

Figs 43-45. Belbina laetitiae sp. nov. in natura , lie Sainte Marie, 25 Nov. 2011, photographs by Paul

Bertner.

23

European Journal of Taxonomy 102: 1-37 (2014)

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 first metatarsomere.

Abdomen (Fig. 6A). Bright red with 2 longitudinal rows of 4 black spots dorsally.

Male genitalia

Red (Figs 22-23); 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 Vs, 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.

Distribution

See Fig. 48.

Belbina lambertoni Lallemand, 1922

Figs 7A-E, 24-25, 39, 49

Belbina lambertoni Lallemand, 1922: 62 (type in FSAG).

Belbina lambertoni var. minuta Lallemand, 1922: 62 (type in FSAG).

Belbina lambertoni - Lallemand 1959: 91, fig. 39a-d (lateral view of head, male genitalia). — Constant

2004b: 31 (listed).

Belbina lambertoni var. minuta - Lallemand 1959: 91 (synonymised with B. lambertoni Lallemand,

1922).

Diagnostic characters

(1) disc of hind wings red (Fig. 7A); (2) tegmina orange to red with 3 large black patches on costal cell

and one at base of clavus (Fig. 7A); (3) cephalic process strongly curved dorsad (Fig. 7D); (4) large¬

sized (more than 24 mm long); (5) head orange (Fig. 7C-E).

LT: $ (n = 12) 27.8 mm (24.6-30.1); $ (n = 17) 34.5 mm (32.5-38.2).

24

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

Material examined

Holotype

MADAGASCAR: §, [Manjakandriana, Madagascar] [Type] [Belbina lambertoni Tall, V. Lallemand

det., 195] (FSAG).

Paratypes

MADAGASCAR: 5 33, 2 [Manjakandriana, Madagascar] [Paratype] (FSAG; 1 §: MRAC); 1

5, [Manjakandriana, Madagascar] [Type] [Belbina lambertoni Tall, V. Xallemand det., 1959] [NHRS-

HEMI000000104] (NHRS); 1 $, [Manjakandriana, Madagascar] [Paratype] [Musee du Congo] [Belbina

lambertoni Tall, V. Lallemand det., 195], 18°55’ S, 47 0 48’ E (MRAC).

Note: Lallemand (1922, 1959) stated that the type and paratypes are in his collection. It seems, however,

that he subsequently gave one paratype to NHRS and two to MRAC.

Additional material

MADAGASCAR: 2 S3, 2 $$, near Tananarive, Lamberton, Tananarive (Antananarivo), 18°55’ S,

47°3T E (FSAG); 1 3, 5 ??, Vohemar, coll. Le Moult, Vohemar (Iharaoa), 13°2ES, 50°00’E (FSAG,

4 NCSU); 2 S3 , Mahajanga Prov., Parc National Tsingy de Bemaraha, 2.5 km 62° ENE of

Belcopaka, Ankidrodroa River, 19°7 , 56” S, 44°48’53” E, 100 m, 11-15 Nov. 2001, tropical dry forest

on Tsingy, Fisher, Griswold et al. (CAS); 1 3, 10 $5, Tananarive (5 $$: NCSU; 1 3, 4 §§: RBINS;

1 ?: MNHN); 5 33, 3 ??, Morondava a Mahabo, Last 1854-91, 20°20’ S, 44°28 , 30” E (MNHN);

1 5, Antalaha region, Dec. 1935, Vadon, 14°53’ S, 50°17’ E (MNHN); 3 $?, Madagascar, coll. De

Bergevin (MNHN); 1 §, Vohemar, coll, de Bergevin (MNHN); 1 3, Ambatofitorano, Jul. 1978,20°49’ S,

47°1E E (RBINS); 2 $$, Ankazoabo, SW Madagascar, K.U. Tyduna, 22°17’ S, 44°3E E (ZIN); 1 3,

Madagascar (NMW); 1 §, Maroantsetra, coll. Le Moult, 15°26’ S, 49°44’ E (NMW); 1 §, Madagascar,

20 Oct. 1905, Dr. Kiderlen (NMW); 1 3, 2 ? ?, no data (NCSU); 2 33, 4 ? ?, Antsalova, Dec. 1992,

18°24’ S, 44°22’ E (MHNL); 2 33, Madagascar (MHNL); 2 $$, idem (INHS); 1 $, Miandrivasa

(Miandrivazo), Mar. 1961, 19°33’20” S, 45°27’03” E (MHNL); 1 3, 3 ??, no label (NCSU).

Examined on photograph

MADAGASCAR: 1 3, Berenty Reserve, riverine forest, 25°00 , 20” S, 46’18’10” E, 15 Nov. 2008,

Jason Cry an (Fig. 39).

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 Vs, 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 (Figs 24-25).

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)

25

European Journal of Taxonomy 102: 1-37 (2014)

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.

Distribution

See Fig. 49.

Belbina madagascariensis (Westwood, 1851) comb. nov.

Figs 8A-E, 26-27, 46

Aphana madagascariensis Westwood, 1851: 208 (neotype in MNHN).

Aphana madagascariensis -Walker 1852: 1134 (listed). — Stal 1866: 263 (listed). —Lallemand 1959:

85 (description, comments). —Constant 2004b: 31 (listed, comments).

Aphaenina madagascariensis - Metcalf 1947: 152 (catalogued).

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).

Material examined

Neotype

MADAGASCAR: 5, neotype of Aphana madagascariensis Westwood, 1851, here designated,

[Goudot Madagascar 1832] [Museum Paris] [Pterobaptes. Amyot -vari-egata Spin. A. Fr. VIII.

225.-Senegal] [Neotype $ Aphana madagascariensis Westwood, 1851 J. Constant des. 2013] [Belbina

madagascariensis (Westwood, 1851) Det. Jerome 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.

Additional material

MADAGASCAR: 1 $, Madagascar] (MNHN); 1 ?, Tenina, J. Vadon, 15°45’ S, 49°40’ E (RBINS);

1 S, 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

26

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

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.

Thorax. Yellow-brown with slightly paler markings (Fig. 8C-E); pronotum with strong median carina

(Fig. 8C); carina very slightly sinuate in lateral view (Fig. 8D) and with strongly impressed point on

each side (Fig. 8C); hind margin elevated above level of mesonotum medially (Fig. 8D); peridiscal

carina anteriorly (Fig. 8C); sides of prothorax with oblique carina; lateral lobe of pronotum rounded

posteriorly (Fig. 8D-E); mesonotum with median carina stopped at scutellum; curved peridiscal carina;

short oblique carina at base of scutellum; scutellum transversely wrinkled (Fig. 8C).

Tegmina (Fig. 8A). Brown with irregular black-brown, often confluent, small spots with pale yellow-

brown 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.

Hind wings (Fig. 8A). Broad; disc largely bright red or orange; apex and sutural margin brown-black;

2-4 small, brown-black spots on disc; sutural margin bisinuate.

Legs (Fig. 8A). Elongate and slender; pro- and mesofemora brown with 3 narrow, sinuate and often

incomplete pale yellow rings; pro- and mesotibiae brown with 2 pale yellow rings; metafemora brown

with imprecise yellow ring; metatibiae and metatarsi yellow-brown, tibiae darker basally; metatibiae

with 5 (sometimes 4) lateral and 7 apical spines; 8-9 spines apicoventrally on first metatarsomere.

Abdomen (Fig. 8A). Red dorsally, brown-black ventrally.

Male genitalia

Very finely granulose, dark brown, paler along sides of anal tube (Figs 26-27); 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 2 A (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.

Belbina madagascarensis can be separated (1) from B. bergrothi by the less elongate gonostyli, without

large pale marking ventrally and more broadly rounded apex, and 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. nympha by having the margins of the anal tube not rounded laterally and

the dorsal margin of the gonostyli nearly straight on the apical half; (4) from B. recurva by having the

anal tube not produced into a semi-circular lateral plate.

Distribution

Fig. 46.

27

European Journal of Taxonomy 102: 1-37 (2014)

Belbina nympha (Stal, 1866) comb. nov.

Figs 9A-E, 28-29, 50

Cornelia nympha Stal, 1866: 142 (type in NHRS).

Enchophora atomaria Brancsik, 1893: 253, pi. 11: figs 7, 7a (type in HNHM). syn. nov.

Cornelia nympha - Schmidt 1911: 242 (listed). — Jacobi 1917: 526 (listed). — Metcalf 1947: 122

(catalogued). —Lallemand 1959: 87, fig. 33 (key, description, lateral view of head). — Constant 2004b:

31, fig. 4 (listed, habitus).

Enchophora atomaria - Metcalf 1947: 114 (catalogued).

Cornelia atomaria - Lallemand 1959: 88, fig. 35 (key, description, lateral view of head). — Constant

2004b: 31 (listed).

Diagnostic characters

(1) disc of hind wings red or orange (Fig. 9A); (2) ground colour of tegmina brown (Fig. 9A); (3) small¬

sized (less than 22 mm long); (4) cephalic process directed anterodorsad (Fig. 9D); (5) abdomen with 2

rows of black spots (Fig. 9A).

LT: $ (n = 3) 18.9 mm (18.0-19.8); ? (n = 3) 20.6 mm (19.6-21.9).

Material examined

Type material

MADAGASCAR: Holotype of Cornelia nympha , §, [Madag.] [Stal] [Cornelia Stal] [Typus] [NHRS-

HEMI 000000106] (NHRS).

MADAGASCAR: Holotype of Enchophora atomaria , f, [Madagascar, Nossi-Be] [atomaria Brancs.,

Coll. Brancsik] [Holotypus, Enchophora atomaria n.sp., Brancsik, 1893], 13°20’ S, 48°15’ E (HNHM).

Additional material

MADAGASCAR: 1 f, Antongil Bay, Mocquerys, 15°45’ S, 49°50’ E (HNHM); 1 $, Diego-Suarez

Prov., forest area 7 km N of Joffreville, 12°20’ S, 49°15’ E, 360 m, 22-26 Jan. 2001, Malaise trap,

Irwin, Schlinger & Harin’Hala (CAS); 1 f, Nosy-Be (BMNH); 1 §, Diego-Suarez, coll. De Bergevin

(MNHN); 1 f, no data (probably Montagne d’Ambre), Feb., coll. Sicard (MNHN); 1$, Maroantsetra,

15°26’ S, 49°44’ E (FSAG); If, Nosy Komba, flanc. May 1956, A.R., Institut Scientifique Madagascar,

Nosy Komba (Nosy Koba), 12° 12’S, 49° 16’E (FSAG); l$,Nosibe (FSAG); 1 $, Madagascar (MNPC).

Male genitalia

Very finely 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.

28

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

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.

Distribution

See Fig. 50.

Belbina pionneaui Lallemand, 1922

Figs 10A-E, 30-31,51

Belbina pionneaui Lallemand, 1922: 63 (type in FSAG).

Belbina pionneaui -Metcalf 1947: 123 (catalogued). —Lallemand 1959: 93, fig. 42 (key, description,

lateral view of head). — Constant 2004b: 31 (listed).

Diagnostic characters

(1) disc of hind wings red (Fig. 10A); (2) tegmina dark red with membrane pale brown, numerous

black-brown spots (Fig. 10A); (3) cephalic process broad, strongly curved dorsad (Fig. 10D-E); (4)

large-sized (more than 25 mm long).

LT: 8 (n = 3) 25.6 mm (25.1-26.0); ? (n = 13) 30.6 mm (25.8-32.9).

Material examined

Holotype

MADAGASCAR: §, [Type] [Manjakandriana, Madagascar] [Belbinapionneaui Lallem.] (FSAG).

Paratypes

MADAGASCAR: 2 88, 6 $$, [Manjakandriana, Madagascar] [Para-type] (FSAG); 1 §, [Manjak¬

andriana, Madagascar] [Para-type] [Belbina pionneaui Lall, V. Lallemand det., 1959] (NHRS); 1 §,

[Manjakandriana, Madagascar] [Para-type] [Belbina pionneaui Lallemand, V. Lallemand det., 1956],

18°55 , S, 47 0 48’E (NHRS).

Note: Lallemand (1922, 1959) stated that the type and paratypes are in his collection. It seems, however,

that he subsequently gave one of the paratypes to NHRS.

Additional material

MADAGASCAR: 1 §,Madagascar (RBINS); 2 5 Tananarive, Lamberton, Tananarive (Antananarivo),

18°55’ S, 47°3L E (RBINS); 1 8, 1 ?, [“Ambatol...” (unreadable)], Lamberton (RBINS); 1 ?,

Maroantsetra, Lamberton, 15°26’ S, 49°44’ E (RBINS); 1 8, 4 $$, Tananarive (NCSU, 1 $*: MNHN);

2 88, 1 $, no data (NCSU). - The specimen marked with * bears a label [Belbina pionneaui , cotype,

Lallemand]. This specimen is not recognized as a paratype, as Lallemand (1959) stated that all type

specimens were from Manjakandriana.

1 8 mislabeled “Soelcaboemi, Java” has also been examined (INHS).

Examined on photographs

MADAGASCAR: 4 specimens, Tananarive, Lamberton (RMNH); 2 specimens, Maroantsetra,

Lamberton (RMNH).

29

European Journal of Taxonomy 102: 1-37 (2014)

Male genitalia

Dark red (Figs 30-31); pygofer higher than long and with posterior margin straight, slightly rounded

on dorsal % 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 Vs, then sinuate (Fig. 30); dorsal margin with basal digitifonn 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

(Figs 30-31).

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.

Distribution

See Fig. 51.

Belbina recurva Lallemand, 1950

Figs 11A-E, 32-33, 40^12, 52

Belbina recurva Lallemand, 1950: 84 (type in FSAG).

Belbina recurva - Lallemand 1959: 90, fig. 37a-c (key, description, lateral view of head, male genitalia).

— Constant 2004b: 31, fig. 3 (listed, habitus).

Diagnostic characters

(1) disc of hind wings red (Fig. 11 A); (2) ground colour of tegmina variegated brown (Fig. 11A);

(3) veins of tegmina elevated (Fig. 11 A); (4) large-sized (more than 26 mm long); (5) cephalic process

very strongly curved and projecting posterodorsad (Fig. 11D-E); (6) abdomen with 2 rows of black

spots dorsally (Fig. 11 A).

LT: $ (n = 4) 27.3 mm (25.6-28.4); ? (n = 2) 29.8 mm (29.8-29.9).

Material examined

Holotype

MADAGASCAR: $, [Maroantsetra, Madagascar, Lamberton] [Type] [Belbina recurva Lallem., V.

Lallemand det.:], 15°26’ S, 49°44’ E (FSAG).

Paratype

MADAGASCAR: 1 $, [Maroantsetra, Madagascar, Lamberton] [Paratype] (FSAG).

Additional material

MADAGASCAR: 1 §, E Madagascar, Sambava district, Marojejy, Ambinanitelo, 500 m, Dec. 1958,

Raharizonina, 15°2L S, 49°35’ E (RBINS); 1 $, Fianarantsoa, Ranomafana National Park, Talatakely,

trail FF, 915-1000 m, 4-20 Nov. 1998, Lee & Ribardo, 21°15’ S, 47°27’ E (CAS); 1 ?, E Madagascar,

30

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

Ambodivoangy, Jan. 1960, J. Vadon, 15° 17’50” S, 49°36 , 47” E (MRAC); Antsianaka forest, 1964,

17°30’ S, 48°30’ E (MRAC); 1 S, 1 ?, Madagascar, E coast, Grandidier, 1309-71 (MNHN); 1 S, E

Madagascar, Mananara district, N Antanambe, Vadon & Peyrieras, 16°26’ S, 49°5E E (MNHN); 1 S,

E Madagascar, massif du Marojejy (res. nat. integr. 12), Ambatomitatao, 400 m, Jan. 1973, A. Peyrieras,

13°47’30” S, 49°51’ E (MNHN); 1 ?, Antongil Bay, Mocquerys, 15°45’ S, 49°50’ E (HNHM); 1 $,

Analamazotra, Perinet, Nov. 1930, Perinet (Andasibe), 18°56’ S, 48°25’ E (ZIN); 1 5, Antsiranana,

Parc National de Marojejy, Manantenina River, 28.0 km 38° NE of Andapa, 8.2 km 333° NNW of

Manantenina, 14°26’12” S, 49 o 46’30” E, 450 m, Malaise trap in rainforest, 12-25 Nov. 2003, B.L.

Fischer et al. (CAS); 2 $ $, Rogez, 18°48’ S, 48°37’ E (MNPC).

Examined on photographs

MADAGASCAR: 1 specimen, Masoala Peninsula, 30 May 2009, Dante Fenolio (Fig. 42), 15°30’48” S,

50°07 , 20” E; 1 specimen, Anajanaharibe Sud, 2 May 2010, Becky Rowe (Fig. 41), 14°46’ S, 49°26’ E;

1 specimen, Marojejy, near Cascade Humbert, 22 Nov. 2005 at 20:41, David C. Lees (Fig. 40), 14°26’

S, 49°42 , E.

Male genitalia

Black-brown with gonostyli slightly paler (Figs 32-33); 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 2 A into semi-circular lamina in dorsal view, apical

A 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 semi¬

circular lateral plate.

Distribution

See Fig. 52.

Belbina servillei (Spinola, 1839)

Figs 12A-E, 34-35, 53

Enchophora servillei Spinola, 1839: 227, pi. 2, fig. 3a-c (neotype in HMNH).

Enchophora servillei -Amyot & Serville 1843: 496 (listed). —White 1846: 331 (comparative note). —

Walker 1851: 271 (catalogued). —Dohrn 1859: 57 (listed).

Phrictus servillei - Schaum 1850: 65 (listed).

Belbina servillei - Stal 1863a: 233 (description). — Stal 1866: 144 (key, description). — Jacobi 1917:

527 (listed). —Metcalf 1947: 123 (catalogued). — Constant 2004b: 31 (listed, comments on type).

non Belbina servillei - Lallemand 1959: 88, fig. 36a-c (key, description, lateral view of head, male

genitalia) - [Misidentification of Belbina laetitiae sp. nov.]

31

European Journal of Taxonomy 102: 1-37 (2014)

Diagnostic characters

(1) disc of hind wings turquoise (Fig. 12A); (2) head, pro- and mesonotum red (Fig. 12C-E); (3) ground

colour of tegmina bluish-green (Fig. 12A); (4) cephalic process strongly curved, projecting dorsad (Fig.

12D).

LT: 3 (n = 1) 20.4 mm; $ (n = 13) 23.3 mm (21.7-24.8).

Material examined

Neotype

MADAGASCAR: 3 , neotype of Belbina servillei (Spinola, 1839), here designated, [Madagascar,

Antongil B., Mocquerys], 15°45’ S, 49°50’ E (HMNH).

A neotype is designated here in order to ensure stability in the nomenclature of the group, following rule

75.3 of the International Code of Zoological Nomenclature (ICZN 1999). The origin of the specimen

described by Spinola (1839) is unknown and Stal (1866) was the first to give a location for the species

(Madagascar). Lallemand (1959) designated a lectotype in his collection (FSAG) that is invalid according

to nomenclatural rules. Furthermore, the only labelled specimen of this species, [Belbina servillei Sign.,

V. Lallemand det., 1963], in Lallemand’s collection is in fact a specimen of Belbina laetitiae sp. nov., as

demonstrated by the genitalia illustrated in Lallemand (1959). In order to avoid further nomenclatural

confusion, a neotype is designated which perfectly matches the illustrations given by Spinola (1839) and

which is deposited in a public collection (HMNH). I chose a male with genitalia in good condition to

facilitate further work on the species.

Additional material

MADAGASCAR: 1 3,2 no data (MNHN); 1 $, near Tamatave, A. Raffray, 1884, Tamatave

(Toamasina), 18°10’ S, 49°23’ E (MNHN); 1 3 §?, Madagascar, coll. De Bergevin (MNHN); 1

Tampina, coll. De Bergevin, 18°30’ S, 49°16’E (MNHN); 3 $ §, E Madagascar, forest, coll. De Bergevin

(MNHN); 1 5, Diego-Suarez, Ch. Alluaud, 1893, Diego Suarez (Ampanolahamirafy), 12° 16’ S, 49° 17’

E (MNHN); 2 §§, Maroantsetra region, Rantabe, Nov. 1935, Vadon, 15°42’ S, 49°39’ E (MNHN,

RBINS); 1 5, Antanambe, Mocquerys, 16°26’ S, 49°5E E (MNHN); 1 3,2 $$, Manjakandriana,

18°55’ S, 47°48’ E (FSAG); 2 33, 1 $, Madagascar (NMW; NHRS; ?: NCSU); 2 ?$, Maroantsetra

(NCSU).

1 3 mislabelled “S. America, Coll. Signoret” has also been examined (NMW).

Male genitalia

Brown (Figs 34-35); pygofer higher than long and with posterior margin concave in middle in lateral

view (Fig. 34); anal tube as long as wide at apex and with lateral margins roundly produced in middle

in dorsal view (Fig. 35); posterior margin of anal tube acutely rounded posteriorly in lateral view (Fig.

34) and hiding lateral margin apically in dorsal view (Fig. 35); gonostyli elongate, 1.33 times longer

than high (including dorsal process, 1.88 times without process), strongly surpassing apex of anal tube

and rounded at apex in lateral view (Fig. 34); ventral margin rounded on basal A, then nearly straight

(Fig. 34); dorsal margin with basal, strong, laminate process projecting dorsally, strong hook laterally

at middle of process projecting ventrally; dorsal half of process slightly excavate and narrowly rounded

dorsally, dorsal margin of gonostyli bisinuate after process (Fig. 34); gonostyli minutely rugulose on

ventral half (Fig. 34).

Remarks

Belbina servillei can be separated (1) from all species of the B. falleni+ group by the brown colour

of the genitalia, the laminate basodorsal digitiform process on the gonostyli and having the gonostyli

32

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

12'S

14"S

16'S ’

10'S

20 'S

22‘S

24 5 - -

V V M

, • . . .

I - ■ • • ’4. -A- 1m

d % ■ vA' i A ':®

46

\J| £0

26 r S ■

42'E 44'E 46 J E 40‘E

• Belbina bergrothi

• Belbina biotei

• Belbina madagascariensis

50" E

l£S

M’S

16‘S

10'S

2D"5

22"S

24'$

26'S

42'E

44'E

40‘E

48" E

50'E

• Belbma bourgoini $p. nov.

• Beibma falleni

• Belbina foliacea

2&'& J ---i-i=-i-1—

42'E 44'E 46'E 40‘E 50‘E

Belbina laetitiae sp, nov.

12'S

14 r S

15"S

IBS

20 ‘S

22' S

24 "S

2S'S

42’E 44'E

46’E 4B‘- 50"E

Belbina lambertoni

Figs 46-49. Distribution maps. — 46. Belbina bergrothi , B. bloetei and B. madagascariensis.

47. B. bourgoini sp. nov., B.faiieni and B. foliacea. 48. B. laetitiae sp. nov. 49. B. lambertoni.

33

European Journal of Taxonomy 102: 1-37 (2014)

IE'S

14‘£ ■

IB'S

10'S

20 'S

22 ‘S

24 5 -

25 r S

m

2000

1500

1000

500

0

42 J E 44 'E 46 J E

Belbina nympha

4B'E 50 L E

Belbina pionneaui

1Z'S

14 r E

16S

IB'S

20 5

22 r S

24'S

2B‘S

42'E 44"E 46’E 43’E 50'E

Beto/na recun/a

12S

14 r S

16'5

IB'S

20'S

22*3

24'S

2B‘S

42'E 44"E 46’E 4£’E 50'E

Belbina servliiei

Figs 50-53. Distribution maps. — 50. Belbina nympha. 51. B. pionneaui. 52. B. recurva. 53. B. servillei.

34

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

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 laminate process on the gonostyli; (3) from B.

bourgoini sp. nov. by the laminate basodorsal process of the gonostyli.

Distribution

See Fig. 53.

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, 1906 and 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 field 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.

Acknowledgements

I thank here all the curators listed above for the loan of the material. I also thank Mr Nicolas Cliquennois

(Madagascar), Dr Rhett A. Butler (USA), Dr David C. Tees (France), Dr Dante B. Fenolio (USA), Dr

Jason Cryan (USA), Mr Paul Bertner (Canada) and Mrs Becky Rowe (UK) for the permission to use

their photographs, Dr Patrick Grootaert (RBINS) and Mr Pol Timbourg (RBINS) for their support and

comments, Mrs Yvo nn e van Nierop (RMNH) and Mrs Gunvi Tindberg (NHRS) for sending photographs

of several specimens, and Mr Yves Laurent and Mrs Isabelle Bachy, with the kind permission of Dr

Maurice Leponce (RBINS), for taking and processing the photographs of the genitalia.

This study has also benefited from funds from the European Union for a visit to the collections of the

MNHN (COLPARSYST project) and NHRS (Synthesys project), and from MHNL for a visit to its

collections so kindly arranged by Messrs Cedric Audibert, Joel Clary and Harold Labrique.

Drs Julie Urban, Jason Cryan and Charles Bartlett are warmly thanked for sharing preliminary results of

their molecular studies on Fulgoridae.

References

Amyot C.J.B. & Serville J. 1843. Histoire naturelle des hemipteres. Hemiptera Linn. - Latr. Histoire

Naturelle des Insectes 8: 1-676. http://dx.doi.org/10.5962/bhl.title.8471

Bourgoin T. 2014. FLOW (Fulgoromorpha Lists on The Web): a world knowledge base dedicated to

Fulgoromorpha. V.8, updated [14 Apr. 2014], http://hemiptera-databases.org/flow/

Brancsik K. 1893. Beitrage zur Kenntniss Nossibe ’s und dessen F auna nach Sendungen und Mittheilungen

des Herrn P. Frey. Jahresheft des Naturwissenschaftlichen Vereines des Trencsener Comitates 15-16:

135-159.

Constant J. 2004a. Revision des Eurybrachidae (I). Le genre Amychodes Karsch, 1895 (Homoptera:

Fulgoromorpha: Eurybrachidae). Bulletin de VInstitut royal des Sciences naturelles de Belgique 74:

11-28.

35

European Journal of Taxonomy 102 : 1-37 ( 2014 )

Constant J. 2004b. Fulgoridae of Madagascar: description of a new species representing a new genus,

key to the genera and list of the species (Homoptera: Fulgoromorpha: Fulgoridae). Bulletin de l Tnstitut

royal des Sciences naturelles de Belgique 74: 29-34.

Constant J. 2010. The lanternfly genus Penthicodes : key to the species and review of the “ Ereosoma

group” with two new species and one new subspecies (Hemiptera: Fulgoromorpha: Fulgoridae). Zootaxa

2523: 1-26. http://www.mapress.eom/zootaxa/2010/l/zt02523p026.pdf

Dohrn F. 1859. Catalogus Hemipterorum. Herrcke & Lebeling, Entomologischer Verein zu Stettin.

http: //dx. doi. org/10.5962/bhl. title .8515

Gerstaecker C.E.A. 1873. Baron Carl Claus von der Deckens Reisen in Ost-Afrika in den Jahren

1859-1865. Vol. 3, issue 2. C.F. Winter’sche Verlagshandlung, Leipzig & Heidelberg, http://dx.doi.

org/10.5962/bhl.title.33914

ICZN. 1999. International Code of Zoological Nomenclature. Fourth Edition. The International Trust

for Zoological Nomenclature, London, U.K. http://iczn.org/iczn/index.jsp

Jacobi A. 1917. Die Zilcadenfauna Madagascars und der Comoren. Reise in Ostafrika in den Jahren

1903-1905 mit Mitteln der Hermann und Elise geb. Heckman Wentzel-Stiftung ausgefurt von Professor

Dr. Alfred Voeltzkow. Wissenschaftliche Ergebnisse. Systematische Arbeiten 3 (5): 519-552. http://www.

biodiversitvlibrarv.org/item/131294#page/329/mode/lup

Karsch F. 1890. Afrikanische Fulgoriden. Berliner Entomologische Zeitschrift 35 (1): 57-70. http://

biodiversitylibrary.org/page/25 049326

Karsch F. 1893. Beitrage zur Fauna des Togolandes nach den Sammlungen der Herren Hauptmann

E. Kling und Dr. R. Biittner. Mitteilungen von Forschungsreisen und Gelehrten aus den Deutschen

Schutzgebieten 4 (3): 219-229.

Lallemand V. 1922. Homopteres nouveaux. Bulletin du Museum National d’Histoire Naturelle 28: 62-

68. http://biodiversitvlibrarv.org/page/34159669

Lallemand V. 1950. Contribution a F etude des homopteres de Madagascar. Memoires de l Tnstitut

Scientifique de Madagascar 4A: 83-95.

Lallemand V. 1959. Revision des especes africaines de la famille Fulgoridae (super-famille Fulgoroides

- sous-ordre des homopteres). Publicagoes Culturais da Companhia de Diamantes de Angola 41: 37-

124.

Latreille RA. 1807. Sectio secunda. Familia quarta. Cicadariae. Cicadaires. Genera Crustaceorum et

Insectorum secundum ordinem naturalem in familias disposita, iconibus exemplisqueplurimis explicata

3: 1-258. http://dx.doi.org/10.5962/bhl.title.11558

Melichar L. 1908. Nove rody a druhy Homopter z vychodni Afriky. Casopis Ceske Spolecnosti

Entomologicke, Praha 5: 1-15. http://biodiversitylibrary.org/page/12738193

Metcalf Z.P. 1947. General Catalogue of the Homoptera. Fascicle IV. Fulgoroidea. Part 9. Fulgoridae.

North Carolina State College, Raleigh.

Nast J. 1977. Homopterological notes XII-XX. Annales Zoologici 34 (2): 27-37.

SchaumH. 1850. Fulgorellae. AllgemeineEncyklopadie der Wissenschaften undKunste inalphabetischen

Folge von Genannten Schriftstellern bearbeitet und herausgegeben von I.S. Ersch und I.G. Gruber mit

Kupfern und Charten. Erster Section A-G, 51: 58-73.

Schmidt E. 1911. Beitrag zur Kenntnis der Homopteren. (Neue Gattungen und Arten). Stettiner

Entomologische Zeitung 72: 238-307. http://biodiversitylibrary.org/page/9013241

36

CONSTANT J., Fulgoridae of Madagascar: Belbina (Hemiptera)

Schumacher F. 1918. Die Fulgoriden-Gattung Druentia Stal (. Pyrgoteles Gerst.). Deutsche Entomolo-

gische Zeitschrift 1918 (4): 110-112. http://biodiversitylibrary,org/page/33047045

Spinola M. 1839. Essai sur les Fulgorelles, sous-tribu de la tribu des Cicadaires, ordre des Rhyngotes.

Annalesde la Societe Entomologique de France 8:133-337. http://biodiversitvlibrarv.org/page/33710046

Stal C. 1863a. Beitrag zur Kenntniss der Fulgoriden. Stettiner Entomologische Zeitung 24: 230-251.

http: //biodiversity library, org/page/35950043

Stal C. 1863b. Hemipterorum exoticorum generum et specierum no nn ull arum novarum descriptiones.

Transactions of the Entomological Society of London, Series 3, 1: 571-603. http://biodiversitvlibrarv.

org/page/32121525

Stal C. 1866. Hemiptera Homoptera Tatr. Hemiptera Africana 4: 1-276. http://biodiversitvlibrarv.org/

page/8695280

Urban J. & Cryan J. 2009. Entomologically famous, evolutionarily unexplored: the first phylogeny

of the lanternfly family Fulgoridae (Insecta: Hemiptera: Fulgoroidea). Molecular Phylogenetics and

Evolution 50: 471M84. http://dx.doi.org/10.1016/j.ympev.2008.12,004

Walker F. 1851. List of the Species of Homopterous Insects in the Collection of the British Museum 2:

261-636. http://dx.doi.org/10.5962/bhl.title.9063

Walker F. 1852. Supplement. List of the Species of Homopterous Insects in the Collection of the British

Museum 4: 1119-1188. http://dx.doi.org/10.5962/bhl.title.9063

Westwood J.O. 1851. Description of some new species of exotic homopterous insects. Annals and

Magazine of Natural History, Series 2, 7: 207-210. http://biodiversitvlibrarv.org/page/13787797

White A. 1846. Description of some apparently new species of homopterous insects in the collection of

the British Museum. Annals and Magazine of Natural History 17: 330-333. http://biodiversitvlibrarv.

org/page/22161952

Manuscript received: 7 July 2014

Manuscript accepted: 29 August 2014

Published on: 14 November 2014

Topic editor: Koen Martens

Desk editor: Danny Eibye-Jacobsen

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; National Botanic Garden

of Belgium, Meise, Belgium; Royal Museum for Central Africa, Tervuren, Belgium; National History

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

Natural History Museum of Denmark, Copenhagen, De nm ark

37