Review of Neuroleon Navás of West Africa with Descriptions of Four New Species (Neuroptera, Myrmeleontidae)

Zootaxa 3519: 32–52 (2012) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA Copyright © 2012 · Magnolia Press Article ISSN 1175-5334 (online edition) urn:lsid:zoobank.org:pub:E7E9CC12-4ECF-4A13-8E81-F56803F9B0B6 Review of Neuroleon Navás of West Africa with descriptions of four new species (Neuroptera, Myrmeleontidae)

BRUNO MICHEL1 & MASSOUROUDINI AKOUDJIN² 1CIRAD, UMR CBGP (INRA/IRD/CIRAD/ Montpellier SupAgro), Campus International de Baillarguet, CS 30016, F-34988, Montfer- rier-sur-Lez, France. E-mail: [email protected] 2CIRDES, BP 454, Bobo-Dioulasso, Burkina Faso. E-mail: [email protected]

Abstract

The Neuroleon species of sub-Saharan West Africa are reviewed. Eight species are recorded: N. drosimus Navás, N. lapidarius nov. sp., N. modestus (Navás), N. nubilatus (Navás), N. rapax nov. sp., N. raptor nov. sp., N. pardalice (Banks), and N. ruber nov. sp. A ninth species, N. nigericus Navás, of which no specimens could be examined, is tentatively maintained in the genus Neuroleon pending examination of the type. Three new synonymies are established. Seven species are removed from the genus Neuroleon. N. pardalice is recorded from the region for the first time. The species are illustrated and keyed.

Key words: Antlion, Afrotropical Region, Myrmeleontinae, Nemoleontini

Introduction

This contribution to taxonomic knowledge of the antlions of West Africa deals with the genus Neuroleon Navás, 1909, which was described including the following species: N. arenarius (Navás, 1904) (a replacement name for Myrmeleon variegatus Rambur, 1842, nec Klug, 1839), N. ochreatus (Navás, 1904), N sticticus (Navás, 1903) (= N. egenus (Navás, 1914a), a preoccupied name), and N. distichus (Navás, 1903), without designating a type species for the new genus. In the same publication, Navás maintained Myrmeleon nemausiense Borkhausen, 1791 in the genus Myrmeleon with the emendation nemausiensis. Then, Navás (1912) designated the latter species as type species of his genus Nelees, and later he designated N. arenarius as type species of Neuroleon (Navás 1914b). Neuroleon is now the largest genus of the tribe Nemoleontini, with 123 valid species listed by Stange (2004), of which 86 were described by Navás. Among them, 69 are recorded from the Afrotropical Region and 43 from the Palearctic (mainly West Palearctic) Region. Only 14 species are known from the Oriental Region. No species are known from the western hemisphere or Australia. No revisions have been conducted to date, so the systematics of the genus remains unresolved. Stange (2004) listed 12 genera as synonyms of Neuroleon. If synonymies are to be justified, and the placement of some genera or species is to be clarified, further investigation is required to provide a satisfactory knowledge of the systematics of the genus. In this review, four previously described valid species and four new species are recognized from West Africa. A ninth species, Neuroleon nigericus Navás, 1935, is tentatively retained in the genus Neuroleon, pending examination of the type specimen. Three new synonymies are established. Seven species are removed from the genus Neuroleon.

Material and Methods

The region covered by this survey includes the West African countries south of the Sahara, from Senegal to Cameroon, including the Cape Verde Islands and excluding the desert region (Fig. 1).

32 Accepted by A. Contreras-Ramos: 20 Sept. 2012; published: 17 Oct. 2012 FIGURE 1. Area covered by the study.

Material from the following collections was examined (acronyms based on Stange 2004): Natural History Museum, London, England (BMNH) Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Belgium (ISNB) Museo Civico di Storia Naturale Giacomo Doria, Genova, Italy (MCSN) Museo Nacional de Ciencias Naturales, Madrid, Spain (MNMS) Museum für Naturkunde Humboldt-University zu Berlin, Berlin, Germany (ZMHB) Muséum National d’Histoire Naturelle, Paris, France (MNHN) Muséum Royal de l’Afrique Centrale, Tervuren, Belgium (MRAC) Naturhistorisches Museum Wien, Vienna, Austria (NHMW) Nationaal Natuurhistorische Museum Naturalis, Leiden, The Netherlands (RMNH) Leibniz-Zentrum für Agrarlandschaftsforschung, Müncheberg, Germany (LZAM) Zoologisk Museum Københavns Universitet, Copenhagen, Denmark (ZMUC) Zoölogisch Museum , Universiteit van Amsterdam, Amsterdam, The Netherlands (ZMAN) Centre International de Recherche Agronomique pour le Développement, Montpellier, France (CIRAD)

The following institutions provided specimens or photographs: Insect Collection of the IITA Center, Cotonou, Benin (IITA) Naturhistorisches Museum, Brussels, Switzerland (NHMB) Hope Entomological Collections, University Museum, Oxford, England (OXUM)

To examine the terminalia and genitalia, the abdomen was removed from the body, treated with a cold 5% KOH solution and rinsed in distilled water. It was then transferred to glycerine in a microvial for further examination and preservation. For photographs, the abdomen was placed in glycerine on a microscope slide cavity. Habitus photographs were taken using a digital camera Canon® EOS 450D equipped with a Canon® macro lens EF- S 60 mm and fixed on a stand. The lighting system was described by Aberlenc (1980). Photographs of morphological characters were taken using a Leica® M205C stereo microscope equipped with a JAI® AT-200GE digital camera. Photographs of terminalia and genitalia were taken using a Leica® DMIL inverted microscope equipped with a JVC® KY-F75U digital camera. The photographs were processed with Archimed® and retouched with Adobe Photoshop® software.

NEUROLEON OF WEST AFRICA Zootaxa 3519 © 2012 Magnolia Press · 33 Unless otherwise indicated, all specimens were collected by the first author. When no repository is indicated, additional material examined is deposited in the CIRAD-CBGP collection. Abbreviations are as follows: A1, A2, A3—anal veins 1, 2, 3; BC—basal crossvein; CF—cubital fork; CuA—anterior cubitus; CuP—posterior cubitus; M—median; MP2—posterior branch of posterior median; PS—presectorial crossvein; R—radius; RS—radial sector; SC—subcosta (Figs 2, 3).

FIGURES 2–3. Wing venation of Nemoleontini. 2—Base of forewing. 3—Base of hind wing. (A1, A2, A3: anal 1, 2, 3. BC: basal crossvein. C: costa. CuA: anterior cubitus. CuF: cubitus fork. CuP: posterior cubitus. M: median. MA: anterior median. MP: posterior median. MP1: anterior branch of MP. MP2: posterior branch of MP. PS: presectorial crossvein. R: radius. RS: radial sector. SC: subcosta).

Taxonomy

A list of 12 synonyms of the genus Neuroleon is given by Stange (2004). If some of them are certainly valid, the correct status of others is pending upon the revision of the whole genus. In this paper the following synonymies are confirmed referring to the West African species:

Neuroleon Navás, 1909 = Nelees Navás, 1912 = Klapalekus Navás, 1912 = Oligoleon Esben-Petersen, 1913

Redescription of the genus Neuroleon. The genus Neuroleon exhibits synapomorphies that characterize the tribe Nemoleontini. In the forewing CuP originates at the extremity of BC and A2 runs close to A1 for short distance, and then bends at sharp angle toward A3. In the hind wing CuA does not reach MP2, but may or may not be connected to it by crossveins. There is only one PS (Figs 2, 3). Among Nemoleontini, the genus Neuroleon can be characterized by the following features. In the forewing the branches of CF diverge. Tibial spurs are absent or at most as long as tarsomeres 1–3 in the foreleg, and at most as long as tarsomeres 1–2 in the middle and hind legs. They are generally slender and slightly bent. Legs slender. For the West African species the ratio “length of posterior tibia/length of first tarsomere” is between 4 and 5.3. The same ratio varies from 5 to 6.2 in the West Palearctic species. The apical margin of the ventral surface of at least tarsomeres 2–4 with a row of stout black setae on each side (Fig. 52). Generally the ventral surface bears other stout setae. Tarsal claws without tooth, capable of closing against distal tarsomere in two species. Ectoproct of male rounded without postventral lobe. Parameres hook-shaped. Gonosaccus with at least 4 long setae (Figs 10, 11). In the female anterior gonapophyses are absent (Fig. 12). The genus Neuroleon can be separated from the genera Nemoleon Navás and Distoleon Banks as follows. In Nemoleon the apical margin of tarsomeres 2–4 bears only one stout seta and the distal tarsomere has two longitudinal rows of stout black setae, the other setae being very thin and slender. Ectoprocts of males have variably developed postventral lobes. In the West African species of Nemoleon the ratio “length of posterior tibia/ length of first tarsomere” is between 2.5 and 3.5. The gonosaccus has two long black setae on each side.

34 · Zootaxa 3519 © 2012 Magnolia Press MICHEL & AKOUDJIN FIGURES 4–12. Neuroleon drosimus Navás. 4—Habitus of male. 5—Thorax and right wings of female. 6—Face. 7—Vertex and pronotum. 8—Hind leg of female. 9—Hind leg of male. 10—Right view of male ectoproct and genitalia. 11. Ventral view of male subgenital plate and genitalia. 12—Female terminalia. (af: apical field. cm: cubital mark. e: ectoprocts. g: gonarcus. gp: gonapophyseal plate. gs: gonosaccus. lg: lateral gonapophyse. p: paramere. pg: posterior gonapophyse. rm: rhegmal mark. sp: subgenital plate).

NEUROLEON OF WEST AFRICA Zootaxa 3519 © 2012 Magnolia Press · 35 In Distoleon the tibial spurs are as long as tarsomeres 1–4 in the forelegs and as long as tarsomeres 1–3 in the middle and hind legs. They are generally stout and strongly arched. In males, the abdominal intersegmental membrane, at least between tergites VI and VII, bears a tubercle with generally two or three long bristles. These setae are present in all the African species of Distoleon examined and are mentioned for Australian species by New (1985). They are absent in the Palearctic and African species of Neuroleon examined.

Neuroleon drosimus Navás, 1912 (Figs. 4–14)

Neuroleon drosimus Navás, 1912: 70. Neuroleon waterloti Navás, 1921: 298, syn. nov.

Diagnosis. Medium-sized species. General coloration grey and black. Wing membrane hyaline with 4–5 oblique stripes along posterior margin of forewing after the cubital mark (Figs 4, 5). Redescription. Head. Face yellowish. Interantennal mark black extending under and below scapes. A median narrow vertical black stripe on face (Fig. 6). Antennae entirely dark brown. Thorax. Pronotum pattern as in figure 7. Foreleg. Coxa dark dorsally, yellow ventrally with 2–3 long white setae. Femur yellow ventrally often with basal and sub- apical yellow area. Sensory seta long. Tibia yellow, dark dorsally at base, in middle and apically. Tibial spurs slightly shorter than tarsomeres 1–3. Tarsomeres yellow, dark apically. Middle leg. Femur yellow with median dorsal dark ring. Sensory seta long. Tibia yellow. An incomplete dark ring at apical 1/3, dark apically. Tibial spurs as long as tarsomeres 1–2. Tarsomeres as in foreleg. Hind leg (Figs 8, 9). Yellow. Tibia dark apically with longitudinal dark stripe. Tibial spurs as long as tarsomeres 1–2. In female only 4–5 and about 10 stout setae on femur and tibia respectively (Fig. 8). Femur and tibia of male with about 15 and 30 long slender erect setae respectively (Fig. 9). Wings (Figs 4, 5). Forewing ♂ 19 (type of N. waterloti) –23 mm, ♀ 20–23 mm. Hind wing ♂ 19 (type of N. waterloti)–23 mm, ♀ 20–23 mm. Forewing. Generally no crossvein in apical field but some specimens with one to two. Longitudinal veins white with black markings at junctions with crossveins. Crossveins mainly white. Four to five oblique stripes from extremity of A1 along posterior margin after the cubital mark which is generally more marked. Rhegmal mark small. Gradates black. Pterostigma white. Hind wing. Narrower than forewing. Black markings in apical ¼ along posterior margin, clearly more intense in female than in male (Figs 4, 5). Abdomen. ♂ 19–20 mm, ♀ 15–16 mm. Dark grey to black. Distal margin of last segments and ectoprocts yellow. Male. Parameres long, lyre shaped with a stout seta inserted on inner margin. Gonosaccus with numerous long setae. Gonarcus V-shaped basally. Subgenital plate elongate (Figs 10, 11). Female. Gonapophyseal plate straight, arched at extremity (Fig. 12). Type material examined. Neuroleon drosimus Navás, 1912. Type ♂, Badeggi [09°03’N 06°08’E] NIGERIA (Coll. BMNH). Neuroleon waterloti Navás, 1921. Type ♂, BENIN (Coll. MNHN) Additional material examined (51♂♂, 44♀♀). MALI Sikasso [11°19’N 05°40’W] 1♂ 13.I.1994; 1♀ 18.I.1996, attracted to light 04:00 a.m.; 1♂ 03.II.1995, in morning on a wall, attracted to light; 2♂♂ 15.II.1996, in morning on a wall, attracted to light; 1♂ 18.III.1996, light trap, 21:00; 1♀ 08.IV.1996, CMDT factory light, 21:30; 1♀ 17.IV.1996, Wayerma District, light trap 22:30; 1♀ 25.IV.1997, Wayerma District, in morning, attracted to light; 1♂ 16.V.1995, CMDT factory, attracted to light 22:00; 1♀ 18.V.1995, CMDT factory, attracted to light 21:00; 2♀♀ 24.V.1995, CMDT factory, attracted to light 21:30; 1♂ 1♀ 25.V.1995; 1♂ 28.V.1995; 1♂ 3♀♀ 04.VI.1995, attracted to light; 1♀ 23.VI.1995, attracted to light; 1♂ 10.VIII.1994; 1♀ 10.VIII.1996, in the morning, attracted to light; 1♂ 17.VIII.1996, in the morning, attracted to light; 1♂ 24.VIII.1996, in the morning, attracted to light; 1♀ 26.VIII.1996, in the morning, attracted to light; 1♀ 04.IX.1996, CMDT factory, in the morning, attracted to light; 1♂ 09.IX.1994; 1♂ 12.IX.1994; 2♂♂ 14.IX.1994; 1♂ 17.IX.1994; 1♀ 03.X.1995, in the morning, attracted to light; 1♂ 08.X.1997; 1♀ 29.X.??; 1♀ 04.X.1994, side of the airstrip; 2♂♂ 1♀ 05.X.1994, side of the airstrip; 1♀ 02.XI.1994; 1♂ 1♀ 11.XI.1996, side of the airstrip, 17:00; 1♂ 30.XII.1999, Wassoulou Hotel, attracted to light 20:30. About 11 km North Sikasso [11°26’32’’N 05°39’11’’W] 1♂ 03.V.1997; 1♂ 05.V.1997; 1♀ 10.V.1997; 1♂ 10.V.1997; 1♂ 02.X.1995; 1♂ 13.XI.1995; Pégnasso [11°28’N 05°38’W] 1♂ 16.V.1998; 1♂ 20.VIII.1997. Natiékoura [11°20’N 05°51’W] 1♀ 30.I.1996; 1♀ 04.III.1998; 1♀ 06.V.1997; 1♂ 1♀ 08.V.1997; 1♀ 17.V.1998; 1♂ 14.VI.1995; 1♂ 15.VI.1995; 1♀ 22.VIII.1998. Farako

36 · Zootaxa 3519 © 2012 Magnolia Press MICHEL & AKOUDJIN [11°12’27’’N 05°27’00’’W] 3♀♀ 04.II.1997, 16:30, on the ground after bushfire; 1♂ 27.II.1998; 1♂ 18.III.1998; 1♀ 04.IV.1997, light trap; 1♂ 1♀ 08.IV.1996; 3♂♂ 1♀ 29.VIII.1996; 2♂♂ 08.IX.1994; 1♂ 20.IX.1994; 1♀ 03.X.1996. Fana [12°46’N 06°57’W] 1♂ 13.V.1997, CMDT factory, in the morning, attracted to light; 3♀♀ 11.IX.1996, 17:30. Koutiala [12°24’N 05°28’W] 1♀ 06.I.1998, 21:00, attracted to light. Kadiolo [10°32’N 05°44’W] 1♂ 06.IV.1996. BURKINA FASO Bassora [11°43’N 04°05’W] 1♀ 21.V.2003, J. Bouyer leg. Bobo-Dioulasso [11°10’N 04°18’W] 1♂ 19.V.2010, ONEA; 2♂♂ 21.V.2010, ONEA; 1♀ 30.V.2010; 2♂♂ 06.X.2009, M. Akoudjin leg. 1♂ 10.III.2005, Forest of the Kou, gallery forest, J. Bouyer leg. Koumbia [11°14’N 03°41’W] 1♂ 28.VI.2010, Forest of the Mou, M. Akoudjin leg. Farako-Bâ [11°06’ 04°20’W] 2♀♀ 18.X.1994, at light between 18:00–23:00, J.C. Streito leg. IVORY COAST Korhogo [09°26’N 05°38’W] 1♂ 1♀ 10.VI.1995. BENIN Cotonou Calavi IITA station [06°25’N 02°19’W] 1♂ 31.I.1997, light trap; 1♀ 01.XII.1997, light trap, uncultivated land, G. Goergen leg. Ecology. N. drosimus is a species associated with grass savannas and also found in patches of uncultivated land between crop fields. In southern Mali adults were collected throughout the year except in July during the rainy season. Distribution: Widely distributed in the Sudano-Sahelian Region. Recorded from Senegal as Neuroleon extraneus Navás by Auber (1956). Comments. Genitalia of a male of N. lepidus (Kolbe, 1897) from Mozambique were dissected (Fig. 13) and the type specimens of Macronemurus lepidus Kolbe, 1897 ♀ labeled Creagris lepidus TANZANIA (ZMHB), Neuroleon extraneus Navás, 1912 ♀ ZIMBABWE (BMNH) and Campestretus extraneus Navás, 1933 cotype (abdomen missing) MOZAMBIQUE (MNHN) were examined. The morphological characters and the wings markings of all are similar to N. drosimus. But in the specimen dissected, which is the same size of the West African specimens, the genitalia are distinctly stouter with the parameres distinctly more separated and divergent (Fig. 14). Considering this difference N. drosimus and N. lepidus are maintained as separate and valid species. The discontinuity of their area of distribution suggests that both species could be vicariant species. Neuroleon guernei Navás, 1914d from India, redescribed by Ghosh (1984) with the spelling guernii, whose type ♀ is in MNHN is similar to N. drosimus. As in N. lepidus, these are perhaps vicariant species. Examination of more material of N. guernei, including males, should establish the status of both species.

FIGURES 13–14. Neuroleon lepidus (Kolbe). 13—Male from Mozambique. 14—Genitalia of male (right) compared to N. drosimus (left).

Neuroleon lapidarius Michel & Akoudjin, nov. sp. (Figs. 15–21)

Diagnosis. Medium sized species (Fig. 15). General coloration reddish brown. Abdomen with a large yellowish dot on each side of tergites III to VIII (Fig. 18). Longitudinal veins with black striations more dense and regular on subcosta (Fig. 16).

NEUROLEON OF WEST AFRICA Zootaxa 3519 © 2012 Magnolia Press · 37 FIGURES 15–21. Neuroleon lapidarius, sp. nov. 15—Habitus of male. 16. Fore body and left wings of female. 17—Vertex and thorax. 18—Abdomen. 19—Right view of male ectoproct and genitalia. 20—Ventral view of male genitalia. 21—Female terminalia.

Description. Head. Labrum and clypeus yellowish. Face yellow to reddish. Two arched markings under scapes separated or united medially. Vertex yellowish with rows of brown markings variably fused. Scape

38 · Zootaxa 3519 © 2012 Magnolia Press MICHEL & AKOUDJIN generally with dark marking ventrally. Last joint of labial palp dilated, pointed at apex. Thorax. Pronotum pattern as in figure 17. Legs. Fore coxae with one to three stout white setae. Femora yellowish, darker dorsally. Tibiae brown basally and apically with brown ring at basal 1/3. Tarsomeres brown apically (Fig. 17). Femora and tibiae with stout black or white bristles. Sensory seta short. Spurs of fore and middle legs as long as tarsomeres 1–2. Spurs of hind legs as long as tarsomere 1. Wings (Figs 15, 16). Fore wing ♂ 18–20 mm, ♀ 20–23 mm. Hind wing ♂ 17–18 mm, ♀ 19–22 mm. Forewing. Wide, pointed apically, with three to six crossveins in apical field. Membrane hyaline. Longitudinal veins with white and brown dotted lines, more regular on SC. Some crossveins between R and RS bordered with brown. Rhegmal and cubital marks small. Pterostigma small, more marked in female. Hind wing. Moderately falcate. Longitudinal veins light yellow, dark at junctions with some crossveins (Fig. 16). Abdomen. As long as forewing in male (18–19 mm), shorter than forewing in female (15–16 mm). Tergites reddish brown with two lateral yellowish dots (Fig. 18). Male. Ectoprocts ovoid (Fig. 19). Parameres slender, without setae. Seven to eight long gonosetae (Figs 19, 20). Female. Gonapophyseal plate sinuate (Fig. 21). Type material examined. Holotype ♂ MALI Natiékoura (about 20 km from Sikasso [11°20’N–05°51’W] road to Bougouni) 02.II.1998, between 16:00–17:00 (Coll. CIRAD-CBGP). Paratypes (4 ♂♂, 5 ♀♀) MALI Natiékoura 1♂ 22.II.1997; 1♀ 24.II.1997, between 16:00–17:00; 1♀ 25.II.1997, about 17:00; 1♂ 27.II.1997, 17:00; 1♂ 1♀ 04.III.1998, about 17:00; 1♀ 18.III.1996; 1♂ 25.III.1997. Sikasso about 11 km North of the city [11°26’33’’N–05°39’33’’W], 1♀ 20.III.1998, 17:30, grass savanna along the track to Longorola on the shaded side of a rock (All in coll. CIRAD-CBGP). Etymology. Referring to the behavior of resting on stones (Adjective derived from the Latin name lapis, lapidus = stone) Ecology. Occurs in grass and tree savannas. Adults emerge during the dry season, in February–March. Males and females rest on the shaded side of lateritic stone. Their color similar to that of the stones is probably a cryptic protection against predators. It is interesting to note that this homochromy is shared by some other species of Myrmeleontidae and Ascalaphidae living in the same environment during the dry season and resting also on lateritic stones. Distribution. Known only from two localities of southern Mali which are 25 km apart.

Neuroleon modestus (Navás, 1912) (Figs. 22–33)

Nelees modestus Navás, 1912: 68. Neuroleon modestus Navás, 1912: Krivokhatsky 1996 (new combination) Neuroleon sociorum Hölzel & Ohm, 1983: 243, syn. nov.

Diagnosis. Medium sized species (Fig 22). Abdomen with three yellow markings from tergite III, one median basal and two lateral (Fig. 27). Wing markings variable (Figs 22, 23, 24). Redescription. Head. Face yellow. Two transverse black markings above and below base of antennae. Anterior margin of the latter marking trident-shaped (Fig. 25). Thorax. Pronotum pattern as in figure 26. Foreleg. Coxa yellow with two parallel stripes and an apical dot on posterior surface, more or less distinct. About 12–15 long white setae. Femur with sensory seta long but shorter than femur. Dorsal surface darker, covered by short setae. Posterior surface with five to six stout white setae at extremity. Tibia yellow with dark marking on the dorsal surface. Tibial spurs slender, longer than tarsomeres 1–2. Tarsomeres dark apically. Middle leg. Coxa yellow. Femur with dark dots on the dorsal surface. Tibia and tarsomeres as in foreleg. Hind leg (Fig. 28). Yellow. Femur with about 10 stout setae with black dots at their base. Tibia dark at extremity with about 10 erect black setae. A narrow dark stripe on dorsal surface in the basal 1/3 or 1/2. Tibial spurs as long as T1. Tarsomeres as in other legs. Wings. Forewing ♂ 17–18 mm, ♀ 18–20 mm. Hind wing ♂ 16–17 mm, ♀ 17–19 mm. Membrane hyaline, veins white. Longitudinal veins dark at the junction with some crossveins. Forewing. Two to four crossveins in apical field. Black markings very variable (Figs 22, 23, 24). Generally with at least a basal dot on the sixth crossvein between M and CuA, gradates and base of pterostigma dark (Fig. 22). Other variable black markings on some crossveins between R and RS until extremity of hypostigmatic cell, two between M and CuA, and cubital mark small. Hind wing. Markings at most reduced to some crossveins between R and RS. Abdomen. ♂ 12–13 mm, ♀ 11–13 mm. Brown. Tergite III to VI with three yellow markings, one dorsal at anterior margin and two lateral.

NEUROLEON OF WEST AFRICA Zootaxa 3519 © 2012 Magnolia Press · 39 FIGURES 22–31. Neuroleon modestus (Navás). 22—Male with basal mark on forewing. 23—Female without wing markings. 24—Female with wing markings. 25—Face. 26—Vertex and pronotum. 27—Abdomen. 28—Right hind leg. 29—Right view of male ectoproct and genitalia. 30—Ventral view of male subgenital plate and genitalia. 31—Left view of female terminalia.

40 · Zootaxa 3519 © 2012 Magnolia Press MICHEL & AKOUDJIN Tergite VII with only two large lateral yellow markings (Fig. 27). Sternites yellow. Male. Ectoprocts yellow, largely bordered with brown. Parameres short without seta. Gonosaccus with five to six long black setae. Subgenital plate large, rounded at apex (Figs 29, 30). Gonarcus U-shaped. Female. Gonapophyseal plate narrow, strongly arched (Fig. 31). Type material examined. Nelees modestus Navás, 1912. Type ♀, near Porto Novo [06°29’N 02°36’E] BENIN (Coll. MNHN). Neuroleon sociorum Hölzel & Ohm, 1983. Paratype ♂, Bel Air [14°42’N 17°25’W] SENEGAL (Coll. MNHN). Additional material examined (3♂♂, 9♀♀). MALI Sikasso 1♀ 23.I.1996 attracted to light 22:00; 1♀ 15.II.1996 in morning, attracted to light; 1♂ 10.III.1997 Wayerma District, light trap 21:00; 1♀ 24.V.1995 CMDT factory, attracted to light 21:30. Farako 1♂ 13.II.1997 forest, light trap 20:00. Banamba [13°55’N 07°45’W] 3♀♀ 10–12.IV.1992 at light trap, degraded savannah with trees, fields and mango orchards, ½ km from dry stream, J. Beerlink leg. (ZMAN). BURKINA FASO Folonzo (= Folenzo) [09°57’N 04°40’W] 1♀ 30.V.2010, M. Akoudjin leg. BENIN Cotonou Calavi station IITA 1♀ 24.II.1998 fallow, light trap; 1♀ 11.XI.1997 fallow, light trap, G. Goergen leg. IVORY COAST Bouaké [07°41’N 05°01’W] 1♂ ?.V.1982. Ecology. Occurs in grass savannas and in grassy vegetation in cultivated area. In southern Mali adults fly during the dry season from January to May. Distribution. Recorded from northern Guinean and Sudanian Regions including Cape Verde Islands (Hölzel & Ohm 1983, 1991) and from Arabian Peninsula (Hölzel 1988, 1998). Comments. A male of Neuroleon guttatus (Navás, 1914c) from Namibia (ISNB) (Figs 32) and females from Mozambique were examined. The morphological characters and the wing markings are similar to N. modestus although more dark and conspicuous. As in N. lepidus, the genitalia of the specimen from Namibia are distinctly larger with longer parameres (Fig. 33). Considering this difference, N. modestus and N. guttatus are retained as separate and valid species. The discontinuity of their area of distribution suggests that both species could be vicariant species

FIGURES 32–33. Neuroleon guttatus (Navás). 32—Male from Namibia. 33—Genitalia of male (left) compared to N. modestus (right).

Neuroleon nigericus Navás, 1935

Neuroleon nigericus Navás, 1935: 44.

This species is known only by the type specimen from Aderbissinat, between Zinder and Agades in Niger. It is retained as valid species of Neuroleon pending examination of the type borrowed from the MNHN and not yet returned.

NEUROLEON OF WEST AFRICA Zootaxa 3519 © 2012 Magnolia Press · 41 FIGURES 34–40. Neuroleon nubilatus (Navás). 34—Habitus of Male. 35—Face. 36—Vertex and pronotum. 37—Abdomen. 38—Right view of male ectoproct and genitalia. 39—Ventral view of male genitalia. 40—Right view of female terminalia.

Klapalekus nubilatus Navás, 1912: 46. Neuroleon nubilatus (Navás): Stange 2004 (new combination). Oligoleon longipennis Esben-Petersen, 1931: 202, syn. nov. Neuroleon longipennis (Esben-Petersen): Hölzel 1972 (new combination).

Diagnosis. Medium sized species (Fig. 34). Tibial spurs of fore and middle legs shorter than T1. Tibial spurs of hind legs minute or absent. Redescription. Head. Face yellow with black marking surrounding base of antennae (Fig. 35). Thorax. Pronotum pattern as in figure 36. Foreleg. Coxa yellow with row of about 10 long white setae. Femur dark dorsally covered by flat-lying white setae, and four to five black and five to six erect white setae. Tibia with incomplete dark ring at the apical 1/3 and dark distally. Five to six erect black or white setae. Tibial spurs about ½ of T1. Tarsomeres dark apically. Middle leg. As in foreleg. Hind leg. Yellow. Tibial spurs minute (about 0.05 µ) or absent. Wings. Forewing ♂ 15–18 mm, ♀ 12–21 mm. Hind wing ♂ 15–18 mm, ♀ 12–20 mm. Membrane hyaline. Veins yellowish. Longitudinal veins brown at the junctions with crossveins. Forewing. No crossveins in apical field. Cubital mark conspicuous. Gradates brown. Pterostigma brown basally. Hind wing. Longitudinal veins suffused apically along the posterior margin (Fig. 34). In some males shaded area less visible. Abdomen. ♂ 10–13 mm, ♀ 11–13 mm. Brown with large lateral yellow stripes not always visible on dried specimen (Fig. 37). Male. Parameres long with a seta on inner margin. Gonosaccus with six to seven long setae. Extremity of sub-genital plate flat (Figs 38, 39). Gonarcus U-shaped. Female. Gonapophyseal plate stout and straight (Fig. 40). Type material examined. Klapalekus nubilatus Navás, 1912. Type ♀ Zungeru [09°48’N 06°09’E] NIGERIA (Coll. BMNH). Oligoleon longipennis Esben-Petersen, 1931. Type ♀ Port Sudan [19°37’N 37°12’E] SUDAN (Coll. NHMW) Additional material examined (10♂♂, 13♀♀). MALI Sikasso 1♀ 15.II.1996, Wayerma District, light trap. Natiékoura 1♀ 24.II.1997, between 16:00–17:00; 3♂♂ 04.III.1998 ; 3♂♂ 18.III.1996. Koulikoro [13°48’N 07°26’W] 1♀ 04.XI.1995. Banamba 1♀ 08.XII.1991 at light, degraded savannah with trees, fields and mango orchards, ½ km from dry stream, J. Beerlink leg. (ZMAN). BURKINA FASO Bobo-Dioulasso 1♂ 10.III.2005, Forest of the Kou, gallery forest J. Bouyer leg.; 1♀ 26.III.2010; 1♀ 29.III.2010; 1♂ 09.IV.2010; 1♀ 19.V.2010, ONEA; 1♂ 06.X.2009; 1♀ 29.X.2009, all M. Akoudjin leg. W National Parc [11°52’N 02°07’E] 1♂ 5♀♀ 01.IV.2003, J. Bouyer leg. Ecology. In Mali and Burkina Faso adults fly at the end of the rainy season and during the dry season. Distribution. Probably widely distributed within the Sudano-Sahelian region.

Neuroleon pardalice (Banks, 1911) (Figs. 41–45)

Nemoleon pardalice Banks, 1911: 21. Neuroleon pardalice (Banks): Hölzel 1982 (new combination).

Diagnosis. General coloration dark brown (Fig. 41). Characterized by conspicuous brown markings on forewings and the short, stout antennae (Figs 42, 43). Redescription. The description by Banks (1911) is quite detailed and precise. Only complementary data are provided here. Head. Antennae short, stout. Thorax. Pronotum pattern as in figure 43. Foreleg. Coxa with five long white setae. Tibial spurs about 2/3 of T1. Sensory setae short. Middle leg. As in foreleg. Hind leg. Tibial spurs 1/2 of T1.Wings. Forewing ♂ 16–17 mm. Hind wing ♂ 15–16 mm. Two to four crossveins in apical field. Abdomen. ♂ 12 mm. Tergite III–VII with two lateral round yellowish markings, sometimes difficult to see on dried specimens (Fig 43). Male. Ectoprocts dark ventrally. Parameres short, V-shaped. No seta on the parameres. Gonosaccus with six long black setae. Sub-genital plate rounded apically (Figs 44, 45). Female. Unknown. Type material examined. Nemoleon pardalice Banks, 1911. Type ♂ Har[r]ar [09°18’N 04°07’E] ETHIOPIA (Coll. ZMUC).

NEUROLEON OF WEST AFRICA Zootaxa 3519 © 2012 Magnolia Press · 43 Additional material examined (4 ♂♂). BURKINA FASO Bobo-Dioulasso 2♂♂ 10.III.2005 Forêt du Kou, gallery forest, J. Bouyer leg. NIGERIA Gadau [11°49’N 10°10’E], 1♂ ?.II.1933, P.A. Buxton leg. (BMNH). ERITREA Dorfu [15°28’N 38°57’E] 1♂ 1939 F. Vaccaro leg. (MCSN). Ecology. Unknown. Distribution. Known from Ethiopia and Saudi Arabia (Hölzel 1982, 1998). Recorded here for the first time from Burkina Faso, Nigeria and Eritrea.

FIGURES 41–45. Neuroleon pardalice (Banks). 41—Habitus of male. 42—Face. 43—Forebody. 44—Right view of male genitalia and ectoproct. 45—Ventral view of male genitalia.

44 · Zootaxa 3519 © 2012 Magnolia Press MICHEL & AKOUDJIN FIGURES 46–56. Neuroleon rapax, nov. sp. 46—Habitus female of holotype. 47—Habitus of male. 48—Face. 49—Vertex and thorax. 50—Right foreleg. 51—Right hind leg. 52—Ventral view of tibial spurs and tarsomeres 1–3 of foreleg. 53—Base of abdomen. 54—Left view of male genitalia. 55—Ventral view of male genitalia. 56—Right view of female terminalia. (s = tuft of long setae)

Diagnosis. Medium-sized species (Fig. 46). Tarsal claws capable of closing against last tarsomere. Tibial spurs well developed. Description. Head. Face yellowish with median narrow longitudinal black line sometimes reduced to a dot. A dark line along base of labrum interrupted in middle. A black marking between antennae (Fig. 48). Frons black above antennae. Vertex with rows of black markings. Antennae longer than thorax, scape brown basally or with brown dot. Thorax. Pronotum with four brown markings, the anterior triangular, the posterior more or less comma- shaped, and two dots at transverse furrow (Fig. 49). Meso and metanotum with brown markings (Fig 47, 49). Legs (Figs 50, 51). Foreleg yellowish. Fore and middle legs with incomplete brown ring at basal third. Middle and hind legs yellow. Fore and middle femora with well-developed sensory seta, shorter than femur. Tibial spurs slightly longer than tarsomere 1, curved apically (Fig. 52). Basal tarsomere shorter than distal tarsomere in fore and middle legs, almost as long as distal tarsomere in hind leg. Setae of ventral brush of distal tarsomere tapered, not bend apically (Fig. 50). Claws long, slender. Wings. Narrow. Forewing ♂ 20 mm. Hind wing ♂ 19 mm, ♀ 21–24 mm. One to five crossveins in apical field. Membrane hyaline. Forewing with conspicuous cubital and rhegmal marks. Hind wing with a dark marking in apical third along posterior margin (Fig. 46, 47). The markings darker in male (Fig. 47). Abdomen. Brown with a yellowish dot on segments III–VII (Fig. 53). Male. Ectoprocts rounded. Parameres hook-shaped. Gonosaccus with five or six long setae on each side (Figs 54, 55). Gonarcus U-shaped. Female. Ectoprocts with stout setae. Lateral gonapophyses with short stout setae and two brush-like tufts of long slender setae on posterior area (Fig. 56). Type material examined. Holotype ♀ MALI Sikasso 30.X.1996, CMDT factory, collected in the morning under a lamp on a wall (attracted to light during the night) (Coll. CIRAD-CBGP). Paratypes (1 ♂, 3 ♀♀) MALI Sikasso 1♀ no date; 1♀ 28.X.1997, CRRAS in the morning on a wall under a lamp; 1♀ 07.XI.1994. CAMEROON Guiring [10,6° N–14,4° E] 1♂ 28.IX.1998, Experimental Station of IRAD near Maroua, cotton field with small trees and grass, J.-C. Streito leg (All in coll. CIRAD-CBGP). Etymology. From the Latin adjective rapax = predator. Referring to the grasping legs. Ecology. Unknown. Distribution. Known from southern Mali and North Cameroon.

Neuroleon raptor Michel & Akoudjin, nov. sp. (Figs. 57–63)

Diagnosis. Tarsal claws capable of closing against the last tarsomere. This species can be differentiated from the preceding by the absence of tibial spurs (Figs 60, 61). Description. Head. Face yellowish with two brown markings under base of antennae, and a median longitudinal black line. Black above antennae. Vertex with rows of brown markings (Figs 58, 59). Thorax. Pronotum yellowish with two triangular brown markings on anterior lobe and two longitudinal brown lines on posterior lobe. Bordered with brown. Meso and metathorax yellowish and brown (Fig. 58). Wings. Forewing ♂ 19 mm. Hind wing ♂ 19 mm. Membrane hyaline. Forewing. Holotype with no crossveins in the left apical field and one in the right. Some crossveins dark. Longitudinal veins dark at junctions with crossveins. Hind wing. Terminal forks of longitudinal veins dark along the hind margin in apical third (Fig 57). Legs. Tarsal claws capable of closing against last tarsomere. Foreleg (Fig. 60). Femur yellow, brown dorsally. Tibia with a brown ring at the basal third and brown apically. Basal tarsomere longer than distal tarsomere. Setae on ventral surface of tarsomeres 1–4 short. Setae of ventral brush of distal tarsomere tapered, not bend apically. Middle leg. Femur with sub-basal brown ring. Tibia with an incomplete brown ring at basal third. Basal tarsomere as long as distal tarsomere. Hind leg (Fig. 61). Femur with an incomplete brown ring at basal third, without stout setae. Sensory femoral seta short. Tibial spurs absent. Claws long, slender (Fig. 60, 61). Abdomen (Figs 58, 59). Brown. Tergites III–VII with a yellow triangle on each side and narrow median basal line on segments III–IV. The lateral yellow markings are jointed dorsally on the following segments. Sternites yellow with a brown dot. Male. Ectoprocts rounded.

46 · Zootaxa 3519 © 2012 Magnolia Press MICHEL & AKOUDJIN Parameres hook-shaped, elongated. Gonosaccus with five to six long setae on each side (Figs 62, 63). Gonarcus U shaped. Female. Unknown. Type material examined. Holotype ♂ BURKINA FASO Koumbia [11°14’N 03°41’W] 03.I.2010, 14:54, East side of the Forest of the Mou, M. Akoudjin leg (Coll. CIRAD-CBGP). Etymology. From the Latin name raptor = ravisher. Referring to the grasping legs. Ecology. Unknown. Distribution. Known only from the type locality in Burkina Faso.

FIGURES 57–63. Neuroleon raptor, nov. sp. 57—Habitus of male holotype. 58—Dorsal view of forebody. 59—Ventral view of forebody. 60—Right foreleg. 61—Right hind leg and tarsal claw folded. 62—Left view of male genitalia. 63—Ventral view of male genitalia.

NEUROLEON OF WEST AFRICA Zootaxa 3519 © 2012 Magnolia Press · 47 FIGURES 64–72. Neuroleon ruber, nov. sp. 64—Habitus of male holotype. 65—Right wings of female. 66—Face. 67—Vertex and pronotum. 68—Lateral view of thorax. 69—Hind leg. 70—Right view of male ectoproct and genitalia. 71—Ventral view of male genitalia. 72—Right view of female terminalia.

Diagnosis. Medium-sized species (Fig 64). Pronotum with two longitudinal dark lines largely separated (Fig. 67). Most of the crossveins suffused with brown. Body of some individuals orange. Description. General coloration brown to orange. Head. Face yellowish with black marking around base of antennae, a black vertical line in middle of face extending to basal dark margin of labrum (Fig. 66). Antennae orange at least in living specimens. Thorax. Pronotum with two conspicuous longitudinal lines largely separated (Fig. 67). Meso and meta pleurites with two stripe-like markings (Fig. 68). Legs. Yellow. In fore and middle legs tibial spurs as long as basal tarsomere. In hind leg tibial spurs half of basal tarsomere (Fig. 69). No sexual dimorphism in hind leg. Forewing. No crossvein in apical field. Most crossveins suffused with brown. Two rows of brown gradates (Fig. 64, 65). Hind wing. In male some small brown dots along posterior margin in distal part. In female this dark area is more visible (Fig. 65). Abdomen. Brown to orange at least in living specimens. Male. Parameres V-shaped and slender without seta. Gonosaccus with eight to nine long black setae. Subgenital plate triangular (Fig. 70, 71). Female. Gonapophyseal plate narrow, strongly arched (Fig. 72). Type material examined. Holotype ♂ MALI, Bamako 50 km South of the city, road to Bougouni 13.IX.1996 (Coll. CIRAD-CBGP). Paratypes (16 ♂♂, 15 ♀♀) MALI Bamako idem holotype 2♂♂ 1♀ 13.IX.1996. Sikasso 1♂ 05.X.1994, side of the airstrip ; 3♀♀ 24.X.1993; 6♂♂ 2♀♀ 25.X.1994; 2♂♂ 2♀♀ 26.X.1993; 1♀ 01.XI.1995, side of the airstrip; 1♀ 07.XI.1994, side of the airstrip. About 11 km North of the city 2♂♂ 03.X.1995: 1♀ 20.X.1995. Pégnasso [11°28’N 05°38’W] 1♂ 03.X.1997; 1♀ 05.XI.1996, about 17:00. Dandéresso [11°29’N 05°28’W] 1♀ 04.X.1996. Kadiolo [10°33’N 05°46’W] 1♀ 22.X.1996. San [13°23’N 04°54’W] 1♀ 07.X.1996, attracted to light, around 22:00. Fana [12°46’N 06°57’W] 1♂ 12.IX.1996, around 17:30. Sébékoro [12°58’N 08°59’W] 24.IX.1996. Zalogosso [12°44’N 05°02’W] 1♂ 17.IX.1996 (All in coll. CIRAD-CBGP). Additional material examined (7 ♂♂, 17 ♀♀). BURKINA FASO Bobo-Dioulasso 3♂♂ 4♀♀ 06.X.2009; 2♂♂ 29.X.2009. Dédougou Bladi [15°27N 03°28’E] 1♀ 09.XI.2009; 6♀♀ 11.XI.2009. Koumbia 1♀ 12.XI.2009, fallow, East of the Forest of the Mou. Farako-Bâ 1♀ 18.X.1994, attracted to light, J.-C. Streito leg.; Kiéré [11°37’N 02°32’W] 3♀♀ 22.X.1994, attracted to light, J.-C. Streito leg. W National Parc 1♂ 04.X.2002, Cliffs of Tambaga [11°41’N 01°52’E] J. Bouyer leg. GUINEA Bareng [11°10’N 12°31’W] 1♀ 25.XI.1995, on weed, G. Delvare leg. CAMEROON Rhumsiki [10.5° N 13.6° E] 1♂ 30.IX.1998, tree savanna J.C. Streito leg. Etymology: From the Latin adjective ruber = red, referring to the reddish coloration of some individuals, particularly males. Ecology. Species living in grass and tree savannas. In southern Mali adults fly at the end of the rainy season, from early September to early November. Distribution. Widely distributed from Guinea to North Cameroon.

Identification key to the Neuroleon species of West Africa (excluding N. nigericus)

1 Tarsal claws capable of closing against distal tarsomere ...... 2 – Tarsal claws not capable of closing against distal tarsomere ...... 3 2 Tibial spurs as long as basal tarsomere ...... N. rapax, nov. sp. – Tibial spurs absent...... N. raptor, nov. sp. 3 Tibial spurs of hind leg minute or absent ...... N. nubilatus (Navás) – Tibial spurs of hind leg at least one half of distal tarsomere...... 4 4 Three yellow markings on the abdominal tergite III...... N. modestus (Navás) – At most two dots on the abdominal tergite III...... 5 5 Abdominal tergite III entirely dark ...... 6 – Abdominal tergite III with two yellow dots ...... 7 6 Legs entirely yellow ...... N. ruber, nov. sp. – Legs yellowish with black markings...... N. drosimus Navás 7 Forewing with conspicuous brown markings ...... N. pardalice (Banks) – Forewing without conspicuous brown markings ...... N. lapidarius, nov. sp.

After examining type specimens and additional material from West Africa (material examined indicated in parentheses below), the following species mentioned as Neuroleon by Stange (2004) are removed from the genus Neuroleon. The correct placement of these species is pending upon the revision of the genera they belong to.

Neuroleon clignyi Navás, 1922: 251. (Type specimen MNHN) Neuroleon ducorpsi Navás, 1914e: 204. (Type specimen MNHN) Neuroleon hyalinus Navás, 1921b: 135. (Type specimen MNHN) Neuroleon infidus (Walker, 1853): 365. (Type specimen BMNH) Neuroleon linarixius (Navás, 1924): 378. (Type specimen MNHN + 9♂♂, 17♀♀) Neuroleon serrandi (Navás, 1921a): 301. (Type specimen MNHN) Neuroleon striolatum (Navás, 1914c): 96. (Type specimen NHMW + 3♂♂, 3♀♀)

Two other species mentioned by Stange (2004) as Neuroleon have already been transferred from this genus: Ganguilus imperator (Navás, 1914c) and Ganguilus pulchellus (Banks, 1911) (Michel & Mansell 2010).

Conclusion

In his catalogue of the antlions of the world Stange (2004) lists 15 valid species of Neuroleon from West Africa from Senegal to Nigeria. The present study of collections and examination of all the type specimens of West African species except one showed that some are synonyms or belong to other genera. In this review eight species of Neuroleon are recorded from sub-Saharan Africa from Senegal to Cameroon and a ninth species is tentatively included pending examination of the type specimen to establish its correct placement. Most of the West African species of Neuroleon are recorded from the Sudano-Sahelian region, the fauna of which is now well known. However, the fauna of the Guinean region remains very poorly known. No species are recorded from Guinea-Bissau, Ghana, Liberia and Sierra Leone and only a few species are recorded from the rest of the sub-region. One can assume that surveys in this area will provide new data on the distribution of the species and probably discovery of new taxa. Regarding the possible vicariant species (N. drosimus and N. lepidus, N. modestus and N. guttatus), it is interesting to note that in both cases the same evolutionary trends were observed. The species of the eastern and southern part of Africa are generally darker with more conspicuous wing markings and, even if the individuals are of same size, the genitalia are distinctly larger with morphological differences. This suggests that the populations of West Africa are clearly separated from the populations of East and southern Africa and that they have evolved independently. In this publication the modification of the last tarsomere with closing claws in N. rapax and N. raptor is considered to represent an evolutionary trend within the genus Neuroleon and not an apomorphy justifying the establishment of a new genus. This decision is based on the fact that both species possess other characters correlating with the genus Neuroleon such as the rounded ectoproct and hook shaped parameres in males. But the length of the tibial spurs, of the different tarsomeres and of the setae on the ventral surface of tarsi are quite different, and the true affinities of both species remain unclear. The closing claws must be interpreted as a convergent evolution found in several genera of antlions (Megistopus Rambur, 1842, Gymnocnemia Schneider, 1845, Visca Navás, 1927, Paraglenurus Van der Weele, 1909, Negrokus Navás, 1930). This feature is described here for the first time in species of sub-Saharan Africa. In all the genera harboring such a character, the morphological modification is quite similar. This modification has probably evolved in response prey capture. But no observation has been made to confirm this hypothesis. This publication is a first contribution to the knowledge of the genus Neuroleon in Africa. Even if it deals with the West African species only, it shows that the systematics of the genus is not yet satisfactory and that Neuroleon is probably not a monophyletic group. A revision of the whole West African antlion fauna has been undertaken using as much as possible gene sequencing techniques. Such a work would probably result in the reinstatement of some genera synonymized with Neuroleon and more generally result in a better understanding of the systematics and the distribution of the African species.

This research received support from the SYNTHESYS project http://www.synthesys.info, which is financed by the European Community Research Infrastructure Action under the FP6 and FP7 “Structuring the European Research Area” Programmes. We are grateful to the following persons of the institutions visited: J. Constant (ISNB, Bruxelles), E. De Coninck (MRAC, Tervuren), R. de Vries (RMNH, Leiden), D. Goodger (BMNH, London), W. Hogenes (ZMAN, Amsterdam), N.P. Kristensen (ZMUC, Copenhagen), M. Ohl (ZMHB, Berlin), M. París (MNMS, Madrid), S. Randolf (NHMW, Vienna). The RIPIECSA project supported by the French Ministry of Foreign Affairs facilitated field studies in Burkina Faso. We thank the following persons for providing access to collections in their care and information on specimens: J. Legrand (MNHN, Paris), R. Poggi (MCSND, Genova), A. Taeger (LZAM, Müncheberg), D. Burckhardt (NHMB, Basel) and Z. Simmons and A. Spooner (OXUM, Oxford). We are grateful to J. Bouyer (CIRAD, Senegal), G. Delvare (CIRAD, Montpellier), G. Goergen (IITA, Cotonou) and J.C. Streito (CBGP, Montpellier) for providing us with specimens they collected and to M. W. Mansell (National Insect Collection, Pretoria) for the language reviewing.

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