New Data on African Cheloninae (Hymenoptera, Braconidae) Show a Strong Biogeographic Signal for Taxa with Spined Propodea

New data on African Cheloninae (Hymenoptera, Braconidae) show a strong biogeographic signal for taxa with spined propodea

YVES BRAET1,4, PASCAL ROUSSE2 & MICHAEL SHARKEY3 1 Unité d'Entomologie fonctionnelle et évolutive, Gembloux Agro-Bio Tech, Université de Liège, B-1030 Gembloux, Belgique.; Dépar- tement d’entomologie, IRSNB, Rue Vautier 29, 1000 Bruxelles, Belgique. E-mail: [email protected] 2 UMR 53 PCBMT "Peuplements Végétaux et Bioagresseurs en Milieu Tropical", Cirad Réunion, Pôle de Protection des Plantes, 7 chemin de l'IRAT, 97410 St Pierre, France. E-mail: [email protected] 3 Michael Sharkey, Department of Entomology, University of Kentucky, Lexington KY 40546-0091, USA. E-mail: [email protected] 4 Corresponding author

Abstract

Eight species of Cheloninae (Hymenoptera, Braconidae) are newly described: Ascogaster kibalensis Braet sp. nov., Chelonus (Chelonus) mayi Braet & Rousse sp. nov., Chelonus (Microchelonus) madagasakarensis Braet & Rousse sp. nov., Chelonus (Microchelonus) matilei Braet & Rousse sp. nov. Chelonus (Microchelonus) merdicus Rousse & Braet sp. nov., Odontospha- eropyx gracilis Braet sp. nov., O. leucocoxus Braet sp. nov., and Phanerotomella erena Braet sp. nov. They are all from the Malagasy subregion and several of them share the presence of spine-like apophyses on the propodeum. This character is cate- gorized for future phylogenetic use. The genus Pachychelonus Brues, 1924 is recognized as junior synonym of the genus Odon- tosphaeropyx Cameron, 1910. Three new combinations are proposed Odontosphaeropyx maximus (Zettel, 2002) comb. nov., O. fulviventris (Brues, 1924) comb. nov. and O. flavifasciatus (Zettel, 1990a) comb. nov. for Pachychelonus maximus Zettel, 2002, P. fulviventris Brues, 1924 and P. flavifasciatus Zettel, 1990a, respectively. Updated keys of species Phanerotomella and Odon- tosphaeropyx are provided. New distribution data about Cheloninae from Africa are also given.

Key words: Description; new combinations; keys; spine-like; apophysis; Madagascar; Indian Ocean.

Résumé

Huit nouvelles espèces de Cheloninae (Hymenoptera, Braconidae) sont ici décrites: Ascogaster kibalensis Braet sp. nov., Che- lonus (Chelonus) mayi Braet & Rousse sp. nov., Chelonus (Microchelonus) madagasakarensis Braet & Rousse sp. nov., Chelo- nus (Microchelonus) matilei Braet & Rousse sp. nov., Chelonus (Microchelonus) merdicus Rousse & Braet sp. nov., Odontosphaeropyx gracilis Braet sp. nov., O. leucocoxus Braet sp. nov., et Phanerotomella erena Braet sp. nov. Elles sont toutes présentes dans la sous-région malgache et plusieurs d’entre-elles partagent la présence de longues apophyses ressemblants à des épines sur le propodeum. Ce caractère est défini ici pour un futur usage en analyses cladistiques. Le genre Pachychelonus Brues, 1924 est reconu comme synonyme junior du genre Odontosphaeropyx Cameron, 1910. Trois nouvelles combinaisons sont pro- posées: Odontosphaeropyx maximus (Zettel, 2002) comb. nov., O. fulviventris (Brues, 1924) comb. nov. et O. flavifasciatus (Zettel, 1990a) comb. nov. pour, respectivement, Pachychelonus maximus Zettel, 2002, P. fulviventris Brues, 1924 et P. flavifasciatus Zettel, 1990a. Des clés d’identification des espèces de Phanerotomella et de Odontosphaeropyx sont fournies et mises à jour. De nouvelles données de répartition sur les Cheloninae africains sont également apportées.

Introduction

The cosmopolitan subfamily Cheloninae Förster 1862 sensu stricto contains more than 1500 valid taxa (Yu et al. 2005). However, this number is constantly increasing. Cheloninae are characterized by the presence of a complete postpectal carina in front of the mid coxae, combined with the fusion of the three first metasomal tergites into a carapace, more or less covering the following tergites (van Achterberg 1993). The fore wing has three submarginal cells, though the first cell is sometimes fused with the first discal cell (Walker & Huddleston 1987).

Accepted by J.T. Jennings: 22 May 2012; published: 11 Jul. 2012 1 The taxonomy of Cheloninae has been extensively reviewed by Zettel (1990a) who recognized four tribes and 14 genera. No major changes have occurred since, except for the description of a new genus, i.e., Siniphaneroto- mella He, Chen et van Achterberg, 1994 and four new subgenera, i.e., Chelonus (Areselonus) Braet, 1999; C. (Bac- ulonus) Braet et van Achterberg, 2001; C. (Carinichelonus) Tobias, 2000 and C. (Scabrichelonus) He, Chen et van Achterberg, 1997. Recent phylogenetic analyses show that the subfamily is part of the microgastroid lineage (Achterberg, 1984; Quicke & van Achterberg, 1990; Shi et al. 2005; Murphy et al. 2008). Some genera (e.g. Che- lonus Panzer, 1806 and Ascogaster Wesmael, 1835) are widespread and diverse in temperate and tropical areas. Conversely, others are known from only few specimens, and/or are confined to limited areas, e.g., Fischeriella Zettel, 1990a; Pachychelonus Brues, 1924 and Pseudophanerotoma Zettel, 1990a. All chelonine wasps are solitary egg-larval koinobiont endoparasitoids of Lepidoptera that are concealed in their larval stage. Their hosts belong to various families such as Noctuidae, Geometridae, Tortricidae, Pyralidae, and Gelechiidae (van Achterberg 1990, Shaw & Huddleston 1991). The present study covers several genera from Africa, including Madagascar. Presently, three tribes, six genera and one subgenus are reported in this region with 120 known species (Yu et al. 2005). The discriminating characters between the two main tribes are listed in Table 1. The Chelonini Förster, 1862 parasitize more than 40 families of Lepidoptera. The genera Ascogaster Wesmael, 1835 and Chelonus Panzer, 1806 occur in Madagascar and mainland Africa. Members of Phanerotomini Baker, 1926 are mostly parasitoids of Pyralidae. Host specialization on Pyralidae, as noted by van Achterberg (1990), may be an apomorphic character of this tribe because parasitization of Pyralidae occurs only exceptionally amongst species of Chelonus (Jones 1985). Two genera are widespread in Africa, i.e., Phanerotoma Wesmael, 1838 and Phanerotomella Szépligeti, 1900. The tribe Odontosphaeropygini Zettel, 1990a was erected for two Ethiopian genera (Odontosphaeropyx Cameron, 1909 and Pachychelonus Brues, 1924). This small tribe is poorly known because no biological data have been recorded and very few specimens are housed in museum collections. It is easily recognized by the presence of a large medio-apical tooth on the clypeus, two transverse sutures on the metasoma (sometimes weakly impressed) and vein 1-R1 of the fore wing short (Zettel 1990a). The two genera were separated by the presence of pectinate tarsal claws and complete occipital carina in Pachychelonus in comparison with claws basally smooth and occipital carina missing dorsally in Odontospaheropyx.

TABLE 1. Diagnostic characters for the two main tribes of African Cheloninae (after van Achterberg 1990, Zettel 1990a, Yu et al. 2005) Characters Chelonini Phanerotomini Occipital carina Separated from hypostomal carina. Joining hypostomal carina. Eye Setose Glabrous Lateral carina of mesoscutum Lamelliform, protruding next axillae Weak, not lamelliform nor protruding Epicnemial carina Not reaching beyond ventral 1/3 of pronotal Reaching half height of pronotal sides, excep- sides tionally reduced Transverse sutures of meta- Usually absent Nearly always two sutures present soma Color of metasoma Usually black Usually mostly yellowish though entirely black in numerous Phanerotomella spp. Host families 40 lepidopteran families, mainly Coleophori- Mainly Pyralidae, also Olethreutidae, Carposini- dae, Gelechidae, Noctuidae, Pyralidae, Tor- dae, Gelechiidae, Oecophoridae, and Coleo- tricidae and Yponomeutidae. phoridae.

Though the taxonomy of Ethiopian Cheloninae has been partially treated by several authors (e.g. de Saeger 1942, 1948; Granger 1949; Sigwalt 1977, 1978; Zettel 1989, 1990a, 1990b), the genera Chelonus, Ascogaster and Phanerotomella are in need of a comprehensive revision and further phylogenetic analyses, including all regional fauna, to clarify some important taxonomic points as stated by Sigwalt (1978) and Zettel (1991, 1992a, 1992b). For example, the genus Microchelonus Szépligeti, 1908 was separated from the genus Chelonus several times (Zettel 1990a, Papp 1995) or include inside as subgenus (van Achterberg & Polaszek 1996). The argument for this separa- tion is based on the presence of an apical aperture on the metasoma in males and the number of flagellomeres fixed at 16 in females. However the two characters are not completely correlated, therefore we follow van Achterberg and Polaszek (1996), and Microchelonus is thus considered as a subgenus to avoid potential paraphyly of Chelonus.

2 · Zootaxa 3385 © 2012 Magnolia Press BRAET ET AL. One of the major impediments for a comprehensive systematic study is the lack of diversified characters, especially for Chelonus. An obvious, but unexploited character, is the presence of two to four apophyses on the transverse propodeal carina. The shape of these apophyses ranges from simple blunt angles to long and fine spines. We describe here several species of Cheloninae exhibiting spined propodea. These species cover the full range of variation for this character and may therefore serve as examples for future taxonomic and phylogenetic studies. Updated keys are provided together with new distributional data for African Cheloninae.

Material and methods

The following abbreviations are used:

CAS: California Academy of Sciences, San Francisco, USA (Robert Zuparko). CIRAD-CBGP: Cirad, UMR CBGP, Montpellier, France (Gérard Delvare). CIRAD-PVBMT: Cirad, UMR PVBMT, Saint Pierre, Réunion, France (Jean-Philippe Deguine). IDR: Insectarium de La Réunion, Le Port, Réunion, France (Jacques Rochat). IFENM: Internationales Forschungsinstitut für Entomologie, Naturhistorisches Museum, Vienna, Austria (Michael Madl). MNHN: Muséum National d’Histoire Naturelle, Paris, France (Claire Villemant). MNHU : Museum für Naturkunde der Humboldt-Universitat zu Berlin, Berlin, Germany (Frank Koch). MRAC: Musée Royal d’Afrique Centrale, Tervuren, Belgium (Eliane De Conninck). PR: Personal collection (Pascal Rousse). YB: Personal collection (Yves Braet).

The material examined comes from the authors’ personal collections, recent collections in Madagascar (CAS), Reunion island (IDR, Cirad-CBGP and Cirad-PVBMT), and from the collections of MRAC, MNHN, MNHU, PR, YB and IFENM. The terminology used in this paper follows van Achterberg (1993).

Results

Despite the bad state of the holotype, close examination of Odontosphaeropyx ruficeps Cameron, 1910, show us the presence of at least two basal teeth on the tarsal claws (Fig. 76) of the remaining mid leg. Unfortunately, as the head is missing in the holotype we cannot draw any conclusions about the state of occipital carina (complete or not). Its head was probably present when the type was examined by Zettel (1990a) due to the total length measured. In some specimens of Pachychelonus, when observed dorsally, the occipital carina appears to be missing because the position of the head close to the mesosoma, but this is only an artifact of observation. The shape of metasoma and sculpture is very similar to those observed on some Pachychelonus specimens. These lead us to conclude that Pachychelonus Brues, 1924 is a junior synonym of Odontosphaeropyx Cam- eron, 1909. We describe here eight new species of Ascogaster, Chelonus (Chelonus), Chelonus (Microchelonus), Odon- tosphaeropyx, and Phanerotomella. Updated identification keys to Odontosphaeropyx and Phanerotomella species are provided, including two putative new species of Phanerotomella. These latter two species are not formally described here, awaiting the collection of more specimens. Additionally, we report several new localities for 11 chelonine species.

Ascogaster kibalensis Braet, sp. nov. (Figs 1–8)

FEMALE (Holotype). Body length 4.8 mm, fore wing length 4 mm. Head (Figs 6, 8). Head 0.75x as length as height (in lateral view). Antennal socket inserted close to top of eye. Antenna with 42 flagellomeres. Scapus 4x as long as wide, longer than first flagellomere, without apical lobe. First flagellomere 5x as long as wide, longer than second flagellomere. Penultimate flagellomere as long as wide, 0.04x as long as first segment, 0.60x as long as apical segment. Flagellomeres 1–8 with placodes on all sides, following flagellomeres without placodes on half of their dorsal surface; surface without placodes smooth. Eye length 2.50x length of temple (in dorsal view), 1.50x as high as broad, glabrous. Maxillary palpus of normal length (ending between fore and middle coxae). Clypeus flattened in lateral view, with slightly convex lower margin, without apical teeth (apically blunt and smooth medially), weakly punctate. Malar suture absent. Malar space 1x basal width of mandible, 0.30 x eye height. Face flat in lateral view, densely punctate, with short, sparse setae. Temple not swollen in dorsal view, coarsely punctate, with short, sparse setae. Frons transversely costate/striate, concave (sometimes only weakly), with carina between antennal socket and without lateral carina. POL 2.50x ocellar diameter, 2x OOL. Vertex striate, with strong transverse striate- rugulose sculpture, with short, sparse setae. Occipital carina complete, joining hypostomal carina at mandible. Mesosoma (Figs 4–6). Pronotum mostly punctate but longitudinally striate ventro-laterally, dorsally without modifications. Pronotal furrow absent. Mesosoma 1.43x as long as wide (in lateral view), 1.43x as long as wide (in dorsal view). Mesoscutum sharply raised anteriorly, at right angle to pronotum, medio-anteriorly punctate-areolate, median lobe anteriorly without median groove. Mesoscutum with sparse, short setae. Surface of mesoscutum medially, near scutellar sulcus, entirely punctate to areolate. Notaulus absent. Scutellar sulcus 7.50x as wide as long, 1.40x as long as scutellum, smooth. Scutellar sulcus with 6 complete carina. Scutellum rounded laterally, flattened in lateral view, rugose and punctate. Subalar groove areolate-punctate. Mesopleuron areolate, mostly with short, dense setae but with a median glabrous area anterad mid coxa. Precoxal sulcus absent. Surface of mesosternum weakly punctate. Propodeum 0.46x as long as mesosoma (in dorsal view), rugulose-areolate, carinate areas completely absent. Propodeum vertical posteriorly, with short, sparse setae. Propodeal transverse carina higher than sculpture, with four pointed rounded angles/tubercles (laterally more developed than medially). Median angles present and short (as high as large) equally spaced with lateral ones. Lateral tubercles 0.80x their transverse width, straight with rounded apex. Surface of metapleuron punctate, with short, sparse setae. Metapleural flange absent. Wings: Pterostigma 4x as long as wide. Marginal cell of fore wing closed distally. Vein r 0.83x as long as vein 3- SR, 0.20x as long as vein SR1, 0.45x as long as vein 2-SR. Vein 2-SR of fore wing present. Vein 1-SR+M straight. First discal cell of fore wing 2.18x as long as wide (measured perpendicular to vein 1-CU1+2-CU1). Vein r-m of fore wing with wide bulla. Vein m-cu postfurcal. Vein cu-a of fore wing postfurcal. Vein CU1a (relative to cubital vein (2-CU1) of fore wing) not interstitial, arising behind middle of distal vein of subdiscal cell. Vein CU1b postfurcal with m-cu. First subdiscal cell of fore wing closed distally (vein 2cu-a present). Vein 2–1A long. Hind wing vein 1-SC+R present. Subbasal cell of hind wing medium sized. Hind wing vein M+CU as long as vein 1-M. Hind wing vein r absent, m-cu absent. Basal cell of hind wing closed. Hind wing vein cu-a present. Legs: Femora not swollen. Hind coxa large, dorsally striate/rugose, ventrally smooth. Hind femur 3.75x as long as wide. Hind femur with sparse setae, dorsally and ventrally punctate. Hind tibia 4.50x as long as wide, 2.52x as long as hind basitarsus, with dense stout and rather short setae, apically without specialized patch of setae on its inner side. Hind basitarsus 4.17x as long as wide. Length of hind basitarsomere equal to length of remaining tarsomeres. Metasoma (Figs 2–3, 7). Carapace anteriorly areolate, posteriorly rugulose. Ventral opening of carapace 0.60x as long as its full length. Ovipositor sheath 0.05x as long as metasoma. Ovipositor sheath apically straight, apically with short, thin setae. Color. Scapus, pedicel and first flagellomere yellowish. Head blackish, mandible medially yellowish, maxillary and labial palpus light yellowish. Mesosoma entirely blackish. Fore wing lightly infuscate, veins brownish, full wing weakly infuscate. Leg entirely yellowish. Carapace yellowish but darkened/infuscated on its basal 0.10 and apical 0.20 parts. Ovipositor brownish. Hosts records. None. Morphological variation. Length of fore wing 4.1 mm. Antenna with 40-43 flagellomeres. Flagellomeres 1–10 with placodes on all sides, following flagellomeres without placodes on dorsal side and smooth; scape, pedicel and first flagellomere yellowish.

4 · Zootaxa 3385 © 2012 Magnolia Press BRAET ET AL. FIGURES 1–8. Ascogaster kibalensis Braet sp. nov., Holotype female. 1. full imago; 2. metasoma, dorsal; 3. metasoma, lateral ; 4. medio transversal carina of propodeum ; 5. anterior part of mesosoma, dorsal; 6. head and mesosoma lateral; 7. apex of metasoma; 8. head, face.

CHELONINAE WITH SPINED PROPODEA Zootaxa 3385 © 2012 Magnolia Press · 5 FIGURES 9–14. Chelonus (Chelonus) mayi Braet & Rousse sp. nov., Holotype female. 9. full imago ; 10. apex of metasoma ; 11. anterior part of mesosoma, dorsal ; 12. head and mesosoma lateral; 13. metasoma, lateral ; 14. metasoma, dorsal

Comments. Ascogaster kibalensis sp. nov. does not have spine-like apophyses on the propodeum but we describe it here because its overall similarity with Chelonus (C.) mayi sp. nov. Ascogaster kibalensis sp. nov. is closely related to A. mayamotensis De Saeger, 1948 as indicated by the presence of an obvious transverse ridge on propodeum with four strong and rounded angles. But it differs clearly by the color of the carapace (yellowish rather than blackish). The yellow color also occurs in C. (C.) mayi sp. nov., but the species can be distinguished by the presence of fore wing vein 1-SR+M and the fewer number of flagellomeres in C. (C.) mayi sp. nov. (all characters separating Ascogaster from the Chelonus). Etymology. From the name of the locality. Distribution record. Ethiopian (Uganda). Type material. Holotype: & (MRAC), "Uganda, Kibale, Kanyawara Biol Stat, alt 1509m, leg. S. Katusale & Co, Malaise 3, 00°33'54.4''N 30°21'29.8''E, 3–7.III.2010". Paratypes: & (Coll. YB), "Uganda, Kibale, Kanyawara Biol Stat, alt 1509m, leg. S. Katusale & Co, Malaise 3, 00°33'54.4''N 30°21'29.8''E, 3–7.III.2010"; & (Coll. YB), "Uganda, Kibale, Kanyawara Biol Stat, alt 1509m, leg. S. Katusale & Co, Malaise 3, 00°33'54.4''N 30°21'29.8''E, 16–23.V.2010"; & (MNHN), "Uganda, Kibale, Kanyawara Biol Stat, alt 1509m, leg. S. Katusale & Co, Malaise 3,

6 · Zootaxa 3385 © 2012 Magnolia Press BRAET ET AL. 00°33'54.4''N 30°21'29.8''E, 21–28.III.2010"; % (MNHN), "Uganda, Kibale, Kanyawara Biol Stat, alt 1509m, leg. S. Katusale & Co, Malaise 3, 00°33'54.4''N 30°21'29.8''E, 18–25.IV.2010".

Chelonus (Microchelonus) madagasakarensis Braet et Rousse, sp. nov. (Figs 15–21)

MALE (Holotype). Body length 4.5 mm, fore wing length 3.2 mm. Head (Figs 18–19). Head 0.71x as length as height (in lateral view). Antennal socket inserted close top of eye. Antenna with 24 flagellomeres (left antenna missing apical segment). Scapus 2.43 as long as wide, longer than first flagellomere, without apical lobe. First flagellomere 5x as long as wide, shorter than second. Penultimate flagellomere 1.50x as long as wide, 0.18x as long as first flagellomere, shorter than apical flagellomere. Apical flagellomere acute. Eye length 1.9x length of temple (in dorsal view), 1.7x as high as broad, setose (white setae). Maxillary palpus of normal length (ending between fore and middle coxa). Clypeus flat in lateral view, with horizontal lower margin, without apical teeth, weakly punctate. Malar suture absent. Malar space equal to basal width of mandible, 0.22x eye height. Face straight in lateral view (with small median bump between toruli), finely rugose, with short, sparse setae. Temple not swollen in dorsal view, striate/rugose, with short, sparse setae. Frons costate/striate, weakly concave, without carina between toruli, without lateral carina. POL 1.75x ocellar diameter, 2.5x OOL. Vertex striate, with strong transverse sculpture, with short, sparse setae. Occipital carina complete, joining hypostomal carina at mandible. Mesosoma (Figs 19–20). Pronotum areolate and punctate, dorsally without modifications. Pronotal furrow absent. Mesosoma 1.34x as long as wide in lateral view, 1.17x as long as wide in dorsal view. Mesoscutum sharply raised anteriorly, at right angle with pronotum, medio-anteriorly coriaceous/acinose (with some weak transverse rugae posteriorly on median lobe), median lobe anteriorly without median groove. Mesoscutum with dense, short setae. Mesoscutum medially, near scutellar sulcus, mostly with large punctures or areola, somewhat longitudinally reticulate. Notaulus present anteriorly, not impressed posteriorly, anteriorly areolate, shortened, ending at middle of mesoscutum; notauli not meeting each other, ending their course in large longitudinally rugulose-reticulate area on medio-posterior part of mesoscutum. Scutellar sulcus transversely long and narrow, 10x as wide as long, 1.67x as long as scutellum, smooth, with 8 complete carina. Scutellum with lateral carinae, weakly raised in lateral view (convex), smooth and punctate. Subalar groove areolate. Mesopleuron areolate above sternaulus, areolate to punctate below sternaulus, with short, sparse setae. Precoxal sulcus absent but surface of mesopleuron at this level slightly depressed and with several large, transverse areola looking (this area could be viewed as a faint and large precoxal sulcus transversely rugose/scrobiculate, at least partly) Surface of mesosternum weakly punctate. Propo- deum 0.2x as long as mesosoma (in dorsal view), completely or partly sculptured, rugose/areolate, carinated areas completely absent. Propodeum vertical posteriorly, with short, sparse setae. Propodeal transverse carina short, without acute angles or apophysis. Surface of metapleuron rugulose, with short, sparse setae. Metapleural flange absent. Wings: Pterostigma 2.3x as long as wide. Marginal cell of fore wing closed distally. Vein r 1.25x as long as vein 3-SR, 0.25x as long as vein SR1, 0.5x as long as vein 2-SR. Vein 2-SR of fore wing present. Vein 1-SR+M absent. Vein r-m of fore wing with wide bulla. Vein m-cu interstitial. Vein cu-a of fore wing postfurcal. Vein CU1a (relative to cubital vein (2-CU1) of fore wing) not interstitial, arising beyond middle of distal vein of subdiscal cell. Vein CU1b postfurcal with m-cu. First subdiscal cell of fore wing closed distally (vein 2cu-a present). Vein 2–1A long. Hind wing vein 1-SC+R present. Subbasal cell of hind wing of medium size. Hind wing vein M+CU 1.67x as long as vein 1-M. Hind wing vein r absent, m-cu absent. Basal cell of hind wing closed. Hind wing vein cu-a present, subbasal cell entirely glabrous. Legs: Fore femur and mid femur not swollen. Hind coxa shortened (less than one third length of metasoma), dorsally and ventrally punctate. Hind femur not swollen, 2.92x longer than wide. Hind femur with sparse setae, dorsally and ventrally punctate. Hind tibia 5x as long as wide, 2.35x longer than hind basitarsus, with dense, stout and rather short setae, apically without specialized patch of setae on its inner side. Hind basitarsus 5.33x as long as wide, 0.80 as long as tarsomeres 2–5. Tarsal claws with basal lobes. Metasoma (Figs 16–17, 21). Metasoma without sutures on carapace, shape of carapace in dorsal view rectangular, 2.33x as long as large. Carapace anteriorly longitudinally rugose-areolate, posteriorly with small areolate rugae. Carapace with large, oval apical opening, length of apical opening 1.44x as long as wide. Carapace ventral opening 0.64x as long as total length of carapace, presence of ventral border. Apical opening mostly visible in lateral view.

CHELONINAE WITH SPINED PROPODEA Zootaxa 3385 © 2012 Magnolia Press · 7 FIGURES 15–21. Chelonus (Microchelonus) madagasakarensis Braet & Rousse sp. nov., Holotype male. 15. full imago, laterally; 16. metasoma, lateral ; 17. metasoma, dorsal; 18. head, face; 19. head and mesosoma lateral; 20. anterior part of mesosoma, dorsal; 21. apex of metasoma.

8 · Zootaxa 3385 © 2012 Magnolia Press BRAET ET AL. Color. Head entirely orange-yellowish, palpi yellow, mandible light brownish-yellow but brownish apically. Mesosoma yellowish orange, metanotum yellowish, propodeum mostly dark-reddish. Scapus, pedicellus yellowish; flagellomeres light brown. Fore wing lightly infuscate, veins at base of fore wing whitish, veins and pterostigma brownish, parastigma yellowish. Membrane of fore wing weakly infuscate in apical half, more infuscate below stigma, presence of large patch without dark setae (and then the membrane looking lighter) below parastigma until vein CU1. Hind wing veins yellowish, membrane hyaline. Fore tarsus, mid coxa, mid and hind trochanters, and mid tibia all yellowish; mid femur mostly, mid tarsus light brown except tarsus 5 darker; hind coxa, hind femur, apical third of hind tibia dark-brownish; hind tarsus strongly infuscate; hind tibia on its basal 2/3, hind tibial spurs whitish. Basal 0.1 part of carapace with semicircular black patch and apical 0.58 blackish. Morphological variations. The paratype is entirely similar to holotype. Hosts records. None. Comments. This species may be distinguished from all Cheloninae from Madagascar and mainland Africa by the rectangular shape of metasoma and the body color. The apical opening of the metasoma suggests it to be a member of the subgenus Microchelonus. But it differs from all other Microchelonus by the number of flagellomeres. Etymology. From the name of the country in local language. Distribution records. Ethiopian (Madagascar). Material examined. HOLOTYPE % (IFENM), Verbatim label data: "Madagascar: Prov. Mahajanga, Umgeb. Katsepy, 17–28.XII.1993, Madl". PARATYPES: % (IFENM), same label as the Holotype. Both specimens are in good condition.

Chelonus (Chelonus) mayi Braet et Rousse, sp. nov. (Figs 9–14)

FEMALE (Holotype). Body length 3.25 mm, fore wing length 2.8 mm. Head (Fig. 12). Head 0.77x as length as height (in lateral view). Antennal socket inserted near middle level of eye. Antenna with 14 flagellomeres. Scapus 2.10x as long as wide, apically narrower than its width at mid-length, longer than first flagellomere, without apical lobe. First flagellomere 4x as long as wide, longer than second flagellomere. Penultimate flagellomere twice as long as wide, 0.33x as long as first segment, shorter than apical segment, apical segment rather robust and weakly acute apically. Nine last flagellomeres with placodes on half side only, surface lacking placodes smooth. Eye length 2.50x length of temple (in dorsal view), eye 1.47x as high as broad, setose with dense, long erect setae. Maxillary palpus of normal length, ending between fore and middle coxa. Clypeus flattened in lateral view, with horizontal lower margin without apical teeth, shining with fine punctures. Malar suture absent but malar space with several rugae between eyes and mandible. Malar space 1.33x basal width of mandible, 0.36x eye height. Face straight in lateral view, smooth, shining, finely punctate, with short, sparse setae. Temple not swollen in dorsal view, smooth, shining and very finely sparsely punctate, with short, sparse setae. Frons smooth medially with short transverse striae between antenna and anterior ocellus. Frons flat, without carina between antennal socket or lateral carina. POL 2x ocellar diameter, 4x OOL. Vertex smooth (weakly and sparsely punctate), with faint and poorly defined transverse striate sculpture (transversely close to posterior ocelli), with short, sparse setae. Occipital carina complete, joining hypostomal carina at mandible. Mesosoma (Figs 11–12). Pronotum smooth (latero-ventrally), medio-laterally sparsely striate/costate and dorsally punctate (all shining). Pronotal furrow small. Mesosoma 1.30x as long as wide in lateral view, 1.53x as long as wide in dorsal view. Mesoscutum sharply raised anteriorly, at right angle with pronotum (median lobe weakly protruding anteriorly, in lateral view), medio-anteriorly with large punctures, median lobe without median groove anteriorly. Mesoscutum with short, sparse setae. Surface of mesoscutum areolate medially, near scutellar sulcus. Notaulus entirely absent. Scutellar sulcus 8x as wide as long, 1.70x as long as scutellum, smooth. Scutellar sulcus with six complete carina. Scutellum rounded laterally, flattened in lateral view, shining and very finely and sparsely punctate. Subalar groove areolate. Mesopleuron areolate but rather smooth and punctate ventrally, with short, sparse setae. Precoxal sulcus absent. Surface of mesosternum weakly punctate. Propodeum 0.22x as long as mesosoma (in dorsal view), rugose-areolate, carinated areas absent. No visible transverse carina dividing propodeum. Propodeum vertical posteriorly, with short, sparse setae. Propodeum laterally with two long spines and medially two (rather acute) apophyses. Median tubercle present and short (as long as wide). Distance between median

CHELONINAE WITH SPINED PROPODEA Zootaxa 3385 © 2012 Magnolia Press · 9 FIGURES 22–29. Odontosphaeropyx leucocoxus Braet sp. nov., Holotype male. 22. full imago, laterally ; 23. fore wing ; 24. metasoma, laterally ; 25. metasoma, dorsal ; 26. head, face ; 27. mesosoma, lateral ; 28. head and anterior part of mesosoma, dorsal ; 29. head, lateral.

10 · Zootaxa 3385 © 2012 Magnolia Press BRAET ET AL. tubercles shorter than distance between median and lateral tubercles. Propodeal lateral tubercle twice as long as its transverse width, straight, with rounded apex. Surface of metapleuron areolate, with short, sparse setae. Metapleu- ral flange absent. Wings: Pterostigma 2.7x as long as wide. Marginal cell of fore wing closed distally. Vein r as long as vein 3-SR, 0.15x as long as vein SR1, 0.36x as long as vein 2-SR. Vein 2-SR of fore wing present. Vein 1-SR+M absent. Vein r-m of fore wing, with wide bulla. Vein m-cu antefurcal or interstitial. Vein cu-a of fore wing, postfurcal. Vein CU1a (relative to cubital vein (2-CU1) of fore wing) not interstitial, arising behind middle of distal vein of subdiscal cell. Vein CU1b interstitial or postfurcal with m-cu. First subdiscal cell of fore wing closed distally (vein 2cu-a present). Vein 2–1A long. Hind wing vein 1-SC+R present. Subbasal cell of hind wing distinctly enlarged. Hind wing vein M+CU 1.47x as long as vein 1-M. Hind wing vein r absent, m-cu absent. Basal cell of hind wing closed. Hind wing vein cu-a present. Legs: Femora not swollen. Hind coxa shortened, shining, dorsally and ventrally smooth. Hind femur 3.70x as long as wide. Hind femur with sparse setae, smooth and sparsely punctate. Hind tibia 4.70x as long as wide, 2.30x as long as hind basitarsus, with dense, stout and rather short setae, apically without specialized patch of setae on its inner side. Hind basitarsus 6x as long as wide, 0.75x as long as tarsomeres 2–5. Metasoma (Figs 10, 13–14). Metasoma without visible sutures on carapace. Carapace areolate anteriorly (with two short median carina), smooth posteriorly (shining). Carapace without apical opening. Carapace ventral opening as long as metasoma, presence of ventral border, carapace weakly emarginate ventro-apically. Hypopigium large and exerted. Genitalia and ovipositor: Ovipositor sheath 0.43x as long as metasoma. Ovipositor sheath apically weakly spatulated and with small setae, straight, with unmodified, short and thin setae. Color. Scapus mainly yellowish; apex of scapus, pedicellus brownish, flagellum dark brownish. Head blackish, mandible yellowish (margins only weakly brownish), palpi whitish. Mesosoma entirely blackish. Fore wing hyaline, veins of fore wing brownish, parastigma of fore wing and veins of hind wing yellowish. Legs yellowish; fore tarsus, outer side of mid tibia, mid tarsus, rather infuscate to brownish; hind femur apically testaceous; hind femur mostly whitish on inner side, infuscate elsewhere; hind tarsus entirely infuscate; hind tibial spurs whitish. Carapace entirely yellowish but brownish on its shining apex. Ovipositor entirely yellowish-brown but apically brownish. Hosts records. None. Morphological variation. The paratype has the scapus rather orange and the flagellum rather dark yellowish (especially distal segments). Comments. This species is also very distinctive among all species of African Chelonus. It is the only species which has a yellowish metasoma (except apically) (vs. usually entirely blackish or sometimes with one or two white patch(es) basally) , shining apex of metasoma (usually it is rugose or finely areolate). Other species of Chelo- nus with entirely yellowish carapaces can be found in Europe (e.g., Chelonus erythrogaster Lucas, 1849), Africa (e.g., Chelonus mediterraneus Schmiedeknecht et Fahringer, 1934, C. (M.) ensifer Granger, 1949), Oriental region (van Achterberg, personal communication) and at least one undescribed neotropical species. But none of them have a dark patch apically on carapace. Distribution record. Ethiopian (La Réunion). Etymology. A random combination of letters. Type material. Holotype: & (MNHN): "La Réunion, K, Sainte-Suzanne, forêt Dugain, 780m""Piège Malaise au sol, taillis de Psidium cattleianum"02–08.XII.2001, Attié Marc leg.""Chelonus sp3 &, G. Delvare det.". Paraty- pes: & (MNHN), "St Denis / Le Brulé, sur fougère, alt. 1150m, VIII.2003, coll. IDR".

Chelonus (Microchelonus) matilei Braet et Rousse, sp. nov. (Figs 30–36)

FEMALE (Holotype). Body length 2.6 mm, forewing length 2.5 mm. Head (Figs 31, 36). Head 0.76x as length as height (in lateral view). Antennal socket inserted near middle level of eye, not acute. Antenna with 14 flagellomeres. Scapus 2.40x as long as wide, equal to first flagellomere, without apical lobe. First flagellomere 6x as long as wide, equal to second (five first flagellomeres almost equal in length). Penul- timate flagellomere 2x as long as wide, 0.50x as long as first segment, shorter than apical segment. Eye length 3x length of temple (in dorsal view), 1.90x as high as broad, with dense, short setae. Maxillary palpus of normal length

CHELONINAE WITH SPINED PROPODEA Zootaxa 3385 © 2012 Magnolia Press · 11 (ending between fore and middle coxa). Clypeus flattened in lateral view, with slightly convex lower margin, without apical teeth, weakly punctate. Malar suture absent but presence of several fine rugae running from base of eye to base of mandible. Malar space 1.50x basal width of mandible, 0.43x eye height. Face straight in lateral view, punctate (but presence of weak longitudinal striae between base of eye and base of mandible), with dense, short setae. Temple not swollen in dorsal view, smooth (with scattered punctations), with short, sparse setae. Frons smooth, flat, without carina between antennal socket, without lateral carina. POL 1.70x ocellar diameter, 0.83x OOL. Vertex smooth, punctate, with short, sparse setae. Occipital carina complete, joining hypostomal carina at mandible.

FIGURES 30–36. Chelonus (Microchelonus) matilei Braet & Rousse sp. nov., Holotype female. 30. full imago, laterally ; 31. head, face ; 32. propodeum and metasoma, laterally ; 33. apex of metasoma; 34. metasoma, dorsal ; 35. mesosoma with spines of propodeum, dorsal ; 36. head and anterior part of mesosoma, dorsal.

12 · Zootaxa 3385 © 2012 Magnolia Press BRAET ET AL. Mesosoma (Figs 32, 35, 36). Pronotum punctate, dorsally without modifications. Ventral sides of pronotum laterally smooth. Mesosoma 1.44x as long as wide in lateral view, 1.37x as long as wide in dorsal view. Mesoscu- tum sharply raised anteriorly, at right angle with pronotum, medio-anteriorly areolate, median lobe without median groove anteriorly. Mesoscutum with short, sparse setae. Surface of mesoscutum medially areolate near scutellar sulcus, punctate elsewhere. Notaulus entirely absent. Mesonotal suture present and straight. Scutellar sulcus 7.50x as wide as long, 1.50x as long as scutellum, smooth. Scutellar sulcus with six complete carina. Scutellum laterally rounded, flattened in lateral view, areolate (laterally) but smooth medially. Subalar groove areolate. Mesopleuron areolate, with short, sparse setae. Precoxal sulcus absent. Surface of mesosternum weakly punctate-areolate. Propo- deum 0.38x as long as mesosoma (in dorsal view), areolate, carinated areas completely absent anteriorly. Propo- deum vertical posteriorly, with short, sparse setae. Propodeal transverse carina distinct (as high as neighboring sculpture) with two long spines laterally and two (sometimes acute) angles/tubercles medially; median tubercles short (as long as wide) and closer to each other than to lateral ones. Lateral spines 4x their transverse width, straight, with rounded apex. Surface of metapleuron areolate with short, sparse setae. Metapleural flange absent. Wings: Pterostigma 3.10x as long as wide. Marginal cell of fore wing closed distally. Vein r 0.60x as long as vein 3- SR, 0.16x as long as vein SR1, 0.36x as long as vein 2-SR. Vein 2-SR of fore wing present. Vein 1-SR+M absent. Vein r-m of fore wing with wide bulla. Vein cu-a of fore wing postfurcal. Vein CU1a (relative to vein 2-CU1 of fore wing) not interstitial, arising behind middle of distal vein of subdiscal cell (vein CU1a present basally as short sclerotized stub). Vein CU1b postfurcal with m-cu. First subdiscal cell of fore wing closed distally (vein 2cu-a present). Vein 2–1A long. Hind wing vein 1-SC+R present. Subbasal cell of hind wing distinctly enlarged. Hind wing vein M+CU 0.75x as long as vein 1-M. Hind wing vein r absent, m-cu absent. Basal cell of hind wing closed. Hind wing vein cu-a present. Legs: Femora not swollen. Hind coxa shortened, dorsally and ventrally smooth. Hind femur 3.50x as long as wide. Hind femur with sparse setae, dorsally and ventrally smooth (somewhat sparsely punctate). Hind tibia 5x as long as wide, 3x as long as hind basitarsus, without very long setae (not longer than tibia width), apically without specialized patch of setae on its inner side. Hind basitarsus 5x as long as wide, 0.65x as long as tarsomeres 2–5. Metasoma (Figs 32–34). Metasoma without visible sutures on carapace. Carapace longitudinally rugose-areolate anteriorly, areolate dorso-posteriorly, with transverse weak rugae apically in rear view. Carapace without apical opening. Carapace with short setae, its ventral opening 0.9x as wide as its full length. Ovipositor sheath 0.13x as long as metasoma. Ovipositor sheath apically weakly spatulate and with straight, short, thin setae. Color. Scapus and pedicel yellowish, all flagellomeres brownish. Head blackish, mandible (excepted apex) and palpi yellowish, mandible apically brownish. Mesosoma blackish. Pronotum ventrally, tegula and apex of lateral spines on propodeum yellowish. Fore wing hyaline, veins and stigma yellowish. Coxae, femora, tibiae yellowish but hind tibia infuscate; fore tarsus yellowish, mid and hind tarsi strongly infuscate; hind tibial spurs whitish. Carapace mainly blackish apically, with long medio anterior rufous patch and presence on lighter small patch on the middle of carapace. Ovipositor sheath rufous. Hosts records. None. Morphological variation. Carapace with apical opening (on male) large and oval, opening 3x as long as wide. Rugae on vertex deeply impressed, face heavily punctate, hind coxa pale brownish latero-ventrally. The white patch on metasoma 0.60-0.80x as long as carapace length. Carapace baso-laterally often whitish. Comments. Although more than 500 species (24 in Ethiopian region) have been described, the actual affinities of Chelonus (Microchelonus) spp. cannot be properly determined without a comprehensive revision. Chelonus (M.) matilei sp. nov. and C. (M.) merdicus sp. nov. together form a new species-group characterized by the presence of small lateral spine-like apophyses and a large dorsal white patch on carapace. We have seen at least two other undescribed taxa from Reunion differing in the shape of the apical opening of carapace, the apical sculpture of carapace, the sculpture on vertex, and the color of scapus and hind coxa. They superficially resemble Chelonus (Microchelo- nus) curvimaculatus Cameron, 1906 (Figs 45–47) but the undescribed species from the Reunion have no propodeal spines. Etymology. From the name of the collector, L. Matile. Distribution record. Ethiopian (Mauritius Island). Type material. Holotype: & (MNHN): "Muséum Paris, Ile Maurice, Forêt de Macchabé, 19.III.1981, L. Mat- ile". Paratypes: 2 %% (MNHN), "Muséum Paris, Ile Maurice, Forêt de Macchabé, 19.III.1981, L. Matile"; & (Cirad-PVBMT), Reunion "St Pierre / Bassin Martin, Station expérimentale de l’Armeflhor, parcelle AB, alt.

CHELONINAE WITH SPINED PROPODEA Zootaxa 3385 © 2012 Magnolia Press · 13 290m, XII.2010, tente Malaise"; % (Cirad-PVBMT), Reunion "St Pierre / Montvert les Hauts, verger de manguiers AB Valatchy (partie non fauchée), alt. 600m, III. 2011, tente Malaise"; % (Cirad-PVBMT ) Reunion "St Paul / Tan Rouge, exploitation AB Glénac, alt. 800m, II.2011, tente Malaise". Additional material: % (IDR), Reunion "St Denis / La Providence, sur fougère aigle, alt. 600m, II.2006, battage".

Chelonus (Microchelonus) merdicus Rousse et Braet sp. nov. (Figs 37–42)

FEMALE (Holotype). Body length 3.5 mm, fore wing length 3.0 mm. Head (Figs 38, 40, 42). Head 0.65x as length as height (in lateral view). Antennal socket acute and inserted near middle level of eye. Antenna with 14 flagellomeres. Scapus 2.10x longer than wide, equal to first flagellomere, without apical lobe. First flagellomere 5x as long as wide, equal to second (first 3 segments almost equal in length). Penultimate flagellomere 3x as long as wide, 0.40x as long as first segment, 0.60x as long as apical segment. Basal flagellomeres thin, following flagellomeres enlarged. Eye length 2.20x length of temple (in dorsal view), 1.57x as high as broad, with dense, short setae. Maxillary palpus of normal length (ending between fore and middle coxa). Clypeus flattened in lateral view, with slightly convex lower margin, without apical teeth, weakly punctate. Malar suture absent. Malar space 1.5x basal width of mandible, 0.4x eye height. Face straight in lateral view, smooth, punctate, with short, dense setae. Temple not swollen in dorsal view, smooth, with some punctation, with short, sparse setae. Frons smooth, flat, with carina between antennal socket, without lateral carina. POL 1.67x ocellar diameter, 0.83x OOL. Vertex smooth and finely punctate, presence of weak transverse striae near stemmaticum, with short, sparse setae. Occipital carina meeting hypostomal carina at base of mandible. Mesosoma (Figs 40–42). Pronotum punctate, dorsally without modifications. Pronotal furrow absent, ventral sides of pronotum laterally smooth. Mesosoma 1.28x as long as wide in lateral view, 1.45x as long as wide in dorsal view. Mesoscutum sharply raised anteriorly, at right angle with pronotum, medio-anteriorly areolate, median lobe without median groove anteriorly,of row of punctures in place of groove. Mesoscutum with short, sparse setae. Surface of mesoscutum medially, near scutellar sulcus, mostly punctate or areolate. Notaulus entirely absent,but row of punctures looking like notaulus. Scutellar sulcus 4.30x as wide as long, 1.30x as long as scutellum, smooth. Scutellar sulcus with four complete carina. Scutellum rounded laterally, flattened in lateral view, smooth and punctate. Subalar groove areolate. Mesopleuron areolate, with short, sparse setae. Precoxal sulcus absent. Surface of mesosternum weakly punctate. Propodeum 0.29x as long as mesosoma (in dorsal view), entirely areolate, carinated areas present at least basally (medio-longitudinal carina dividing in two and enclosing medio- triangular area). Propodeum vertical posteriorly, with short, sparse setae. Propodeal transverse carina with two spines laterally and two angles medially. Median tubercles present and short (as long as wide), rather smooth at their apices. Distance between median tubercles 0.6x distance between median and lateral tubercle. Lateral spines twice as long as their transverse width, straight, with rounded apex. Surface of metapleuron areolate, with short, sparse setae. Metapleural flange absent. Wings (Fig. 39): Pterostigma 4.50x as long as wide. Marginal cell of fore wing closed distally. Vein r 0.63x as long as vein 3-SR, 0.17x as long as vein SR1, 0.36x as long as vein 2-SR. Vein 2-SR of fore wing present. Vein 1-SR+M absent. Vein r-m of fore wing with wide bulla. Vein cu-a of fore wing postfurcal. Vein CU1a (relative to cubital vein (2-CU1) of fore wing) not interstitial, arising behind middle of distal vein of subdiscal cell (vein CU1a present basally as short sclerotized stub). Vein CU1b postfurcal with m-cu. First subdiscal cell of fore wing closed distally (vein 2cu-a present). Vein 2–1A long. Hind wing vein 1-SC+R present. Subbasal cell of hind wing distinctly enlarged. Hind wing vein M+CU 1x as long as vein 1-M. Hind wing vein r absent, m-cu absent. Basal cell of hind wing closed. Hind wing vein cu-a present. Legs: Femora not swollen. Hind coxa shortened, dorsally and ventrally smooth. Hind femur 4.4x as long as wide. Hind femur with sparse setae, its surface dorsally and ventrally smooth. Hind tibia 5.87x as long as wide, 2.60x as long as hind basitarsus, apically without specialized patch of setae on its inner side. Hind basitarsus 6x as long as wide, 0.72 as long as tarsomeres 2–5. Metasoma (Fig. 41). Metasoma without visible sutures on carapace. Carapace anteriorly longitudinally rugose- areolate (two medio-longitudinal rugae clearly indicated basally, surface rather mostly punctate between longitudinal rugae), posteriorly longitudinally rugulose (in frontal view). Carapace without apical opening, with short setae. Ovipositor sheath 0.13x as long as metasoma. Ovipositor sheath apically weakly spatulate, straight, with short, thin setae.

14 · Zootaxa 3385 © 2012 Magnolia Press BRAET ET AL. FIGURES 37–42. Chelonus (Microchelonus) merdicus Rousse & Braet sp. nov., Holotype female. FIGURES 43–44. Phanerotomella erena Braet sp. nov., Holotype male. 37, 43. full imago, laterally ; 38. head, face ; 39. fore and hind wings; 40. head and mesosoma, lateral; 41. metasoma, dorsal ; 42. head and anterior part of mesosoma, dorsal; 44. full imago dorsally.

CHELONINAE WITH SPINED PROPODEA Zootaxa 3385 © 2012 Magnolia Press · 15 FIGURES 45–47. Chelonus (Microchelonus) curvimaculatus Cameron, 1906 (specimen fromMadagascar, CAS). FIGURES 48–51. Odontosphaeropyx flavifasciatus (Zettel, 1990a) (specimen from DRC, MRAC). Figure 52. Phanerotomella spinosa Granger, 1949 (specimen from Madagascar, CAS). Figure 53. Phanerotomella annulicornis Granger, 1949 (specimen from Madagascar, CAS). 45, 52, 53. full imago, laterally; 46. apex of metasoma; 47. full imago, dorsally; 48. metasoma, lateral; 49. metasoma, dorsal; 50. head and anterior part of mesosoma, lateral ; 51. head and anterior part of mesosoma, lateral.

16 · Zootaxa 3385 © 2012 Magnolia Press BRAET ET AL. FIGURES 54–58. Phanerotomella antennata Granger, 1949 (specimen from Madagascar, CAS). FIGURES 59–60. Phanerotomella gigantea Granger, 1949 (specimen from Madagascar, CAS). FIGURES 61–63. Phanerotomella seyrigi Granger, 1949 (specimen from Madagascar, CAS). 54, 59. full imago, lateral; 55. head and mesosoma, lateral; 56. propodeum with spines and metasoma, lateral; 57, 63. wings; 58, 62. full imago, dorsal; 60. metasoma, dorsal; 61. full imago, lateral.

Color. Scapus yellowish but pedicel and inner posterior part of scapus infuscate to brownish, flagellomeres dark-brownish. Head blackish, mandible (except apex) and palpi yellowish, apex of mandible brownish, scapus yellowish brown with the posterior inner side brownish to infuscate. Mesosoma blackish. Fore wing hyaline, veins yellowish-brown, stigma yellowish. Coxae, trochanters, femora and fore tibia yellowish; outer sides of mid and

CHELONINAE WITH SPINED PROPODEA Zootaxa 3385 © 2012 Magnolia Press · 17 hind tibia mostly infuscate and inner sides yellowish; all tarsi strongly infuscate; hind tibial spurs pale whitish-yellow. Carapace mainly blackish apically with long, oval patch covering medio-anterior 2/3 of carapace. Ovipositor sheath blackish. Hosts records. None. Comments. The dimensions of the first five flagellomeres are similar to those of species in the subgenus C. (Baculonus). This is the main character separating this species from its closely related species C. (M.) matilei sp. nov. See C. matilei sp. nov. for additional comments. The two paratypes housed in the MNHN collections (box 45, under the label Chelonus spinosus Sigwalt) have probably been studied by the late Dr Sigwalt but he never published the description. Etymology. “merdicus” is an entomological term in Reunion referring to any small insect (Martiré & Rochat 2008). Distribution record. Ethiopian (La Réunion). Type material. Holotype: & (coll. P. Rousse): "Reunion Plaine des Palmistes / Piton Bébour, forêt primaire de Bébour-Bellouve, alt. 1475m, XII.2010, tente Malaise". Paratypes: % (MHNP), "La Reunion, Cilaos, R.F." "Insti- tut scientifique de Madagascar" "hand writing label: C. spinosissimus (probably by Sigwalt)" ; % (MHNP), "La Reunion, Cilaos, XI.(19)55-R.P" "Institut scientifique Madagascar" "type".

Odontosphaeropyx gracilis Braet, sp. nov. (Figs 69–75)

FEMALE (Holotype). Body length 5 mm, fore wing length 4.5 mm. Head (Figs 71-74). Head width 1.04 x its median length (in lateral view). Antennal sockets inserted close top of eye. Antenna with 36 flagellomeres (right antenna broken apically). Scapus 1.75x as long as wide. Scapus longer than first flagellomere, without apical lobe. First flagellomere length equal to the second, 2.5x as long as wide. Penultimate flagellomere 2x as long as wide, 0.4x as long as first segment, shorter than apical segment. Eye length 1.43x length of temple (in dorsal view), 1.47x as high as broad, without setae. Maxillary palpus shortened (at most reaching the anterior coxa). Clypeus flat in lateral view, without carina/lamella on lower margin. Clypeus with one large apical acute tooth medially, finelly punctate. Malar suture absent. Malar space 0.87x basal width of mandible, 0.25x eye height. Face straight, in lateral view, smooth, punctate, with dense, short, fine setae. Temple strongly swollen, in dorsal view, smooth and finely punctate, with sparse, short setae. Frons smooth, flat, with carina between antennal sockets: lateral carina of frons long and meeting the lateral ocelli. POL as long as ocellar diameter, 0.30x OOL. Vertex smooth and sparsely punctate, with sparse, short setae. Occipital carina complete, joining hypostomal carina at mandible. Mesosoma (Figs 72-73). Pronotum ventrally striate/costate, punctuate-areolate elsewhere. Pronotal furrow absent. Mesosoma 1.56x as long as wide in lateral view, 1.62x as long as wide in dorsal view. Mesoscutum sharply raised anteriorly, at right angle with pronotum, median lobe without median groove anteriorly, mesocutum with sparse, long setae. Notauli complete, rather scrobiculate anteriorly, long and meeting each other medially. Surface of mesoscutum medially, near scutellar sulcus mostly punctate. Scutellar sulcus transversally long and narrow, 5x as wide as long, 2x as wide as scutellum length. Scutellar sulcus smooth, with 6 transverse carina. Scutellum flattened in lateral view, smooth and punctate, rounded, smooth and shining laterally. Epicnemial carina present. Sub- alar groove scrobiculate-punctate. Mesopleuron largely smooth, with sparse and short setae. Precoxal sulcus anteriorly scrobiculate, medially punctuate and posteriorly absent. Surface of mesopleuron glabrous around precoxal sulcus. Mesosternum smooth. Propodeum 0.26x as long as mesosoma (in dorsal view). Propodeum entirely rugose, with one medio-longtudinal ruga and several transversal. Propodeum sloping from base to apex, with long and sparse setae. Propodeum without apophyses. Metapleuron largely areolate-punctate, with long and sparse setae. Metapleural flange absent. Wings (Fig. 69): Pterostigma 5x as long as height. Marginal cell of fore wing closed distally. Vein r 1.25x as long as 3-SR, 0.21x as long as SR1, 0.4x as long as 2-SR. Vein 1-SR+M straigth. First discal cell 3.24x as long as wide (measured perpendicular to vein 1-CU1+2-CU1). Vein r-m of fore wing present, mostly tubular, with large posterior bulla. Vein m-cu of fore wing antefurcal. Vein cu-a of fore wing postfurcal. Vein CU1a (relative to vein 2-CU1 of fore wing) not interstitial, arising behind middle of distal vein of brachial cell. Vein CU1b postfurcal with m-cu. First subdiscal cell of fore wing closed distally (vein 2cu-a present). Vein 2–1A long. Hind wing three distal hamuli. Hind wing vein 1-SC+R present. Subbasal cell of hind wing small, hind-

18 · Zootaxa 3385 © 2012 Magnolia Press BRAET ET AL. wing vein M+CU 0.34x as long as 1M. Vein r of hind wing present, vein m-cu absent. Basal cell of hind wing closed, vein cu-a present (but translucent). Legs: Femora not swollen. Hind coxa shortened, smooth dorsally and ventrally. Hind femur 4.17x as long as wide. Hind femur smooth, finely punctate dorsally and ventrally, with sparse setae. Hind tibia with dense stout and rather short setae, apically without patch of setae on its inner side. Hind tibia 6.5x as long as wide, as long as hind tarsus. Hind basitarsus 6x as long as wide, as long as tarsomeres 2–5.

FIGURE 64. Phanerotomella crassa Zettel, 1989 (specimen from Madagascar, CAS). Figure 65. Phanerotomella subdentata Granger, 1949 (specimen from Madagascar, CAS). FIGURES 66–68. Phanerotomella tristis Granger, 1949 (specimen from Madagascar, CAS). 64–66. full imago, lateral; 67. full body, dorsal; 68. full imago, dorsal.

Metasoma (Figs 70, 75). Metasoma with two sutures on carapace, 2.85x as long as wide. First tergite as long as apical width. First tergite without basolateral process (pointed wing-like, spine-like projections). Medio-lateral carina of first tergite very short (less than the third of petiole length). First tergite longitudinally rugulose areolate. Median length of second tergite 1.14x its basal width, 1.14x length of first tergite. Second metasomal suture weakly curved. Second tergite striate-rugulose, third tergite punctate. Remaining tergites concealed by the metasomal carapace. Ovipositor sheath 0.18x as long as metasoma.Ovipositor sheath with apical setae, glabrous basally Color. Antenna and scapus dark brown. Head dark brown, setae whitish. Mesosoma blackish, propleuron dark brown. Fore wing banded, infuscated but lighter basally and transverse white band medially, stigma brown, hind wing infuscate apically, veins brownish. Legs dark brown, fore and mid coxa white-yellow with faint brown patch basally, fore and mid trochanters white-yellowish (but mid trochantellus brown), tibial spur pale yellowish to whitish. Metasoma dark-brownish, second tergite rather red-brownish. Ovipositor sheath orange basally and dark brownish apically. Hosts records. None.

CHELONINAE WITH SPINED PROPODEA Zootaxa 3385 © 2012 Magnolia Press · 19 FIGURES 69–75. Odontosphaeropyx gracilis Braet, sp. nov., Holotype female. 69. full imago, lateral; 70. metasoma, lateral; 71. frons and vertex; 72. head and mesosoma, lateral; 73. head and mesosoma, dorsal; 74. head, frontal; 75. metasoma, dorsal.

20 · Zootaxa 3385 © 2012 Magnolia Press BRAET ET AL. Morphological variation. The main variations are in the body length (4.9-5.05mm) and fore wing length (4.25-4.5mm) in paratypes. Sculptures of propodeum are more or less pronouced, and sometimes the medilongitu- dinal carina is bifurcate enclosing a narrow medio-longitudinal area Comments. This species is strongly related to the P. leucocoxus Braet, sp. nov. But they can be distinguished as shown in the key. Etymology. From the slender appearance of this species. Distribution record. Ethiopian (Madagascar). Type material. Holotype: &(MHNP): "Madagascar Nord Montagne d'Ambre, les Roussettes 1100m, xi & xii.(19)58, Andria Robinson". Paratypes: 3&& & 4%% (MHNP), "Madagascar Nord Montagne d'Ambre, les Rous- settes 1100m, XI & XII.(19)58, Andria Robinson" "Institut scientifique Madagascar"; & (MHNP), "Mont d'Ambre, XII.(19)48, Inst. Scient. Madagascar, RP"; 2%% (MHNP), "Madagascar Nord, dct(district) Diègo-Suarez, Anala- merana 80m, I.(19)59, Andria Robinson" "Institut scientifique Madagascar"; 2%% (MHNP), "Madagascar Nord, dct(district) Diègo-Suarez, Montagne des Français, II.(19)59, Andria Robinson" "Institut scientifique Madagas- car".

Odontosphaeropyx leucocoxus Braet, sp. nov. (Figs 22–29)

MALE (2 specimens). Body length 6.4-7.4 mm, fore wing length 5.4-5.8 mm. Head (Figs 26, 28–29). Head width 1.0-1.1x its median length (in lateral view). Antennal socket inserted close top of eye. Antenna with 38 flagellomeres (left antenna broken apically). Scapus 2.25x as long as wide. Scapus longer than first flagellomere, without apical lobe. First flagellomere longer than second, 2.2-3.0x as long as wide, without placodes. Penultimate flagellomere 1.5x as long as wide, 0.3x as long as first segment, 0.57x as long as apical segment. Eye length 0.9–1.35x length of temple (in dorsal view), 1.4x as high as broad, without setae. Max- illary palpus shortened (at most reaching anterior coxa). Clypeus flat in lateral view, without carina/lamella on lower margin. Clypeus with one large apical blunt tooth medially, weakly punctate. Malar suture absent. Malar space 1.00-1.10x basal width of mandible, 0.25x eye height. Face straight, in lateral view, smooth, punctate; with dense, long, fine setae. Temple normal, in dorsal view, smooth and punctate, with sparse, long setae. Frons smooth, flat, with carina between antennal socket; lateral carina of frons long and meeting lateral ocellus (carina rather weak but clearly present). POL 0.80-1.00x as long as ocellar diameter, 0.25-0.27x OOL. Vertex smooth and punctate, with sparse, long setae. Occipital carina complete, joining hypostomal carina at mandible. Mesosoma (Figs 27–28). Pronotum smooth and areolate. Pronotal furrow absent. Mesosoma 1.50-1.60x as long as wide in lateral view, 1.8x as long as wide in dorsal view. Mesoscutum sharply raised anteriorly, at right angle with pronotum, median lobe without median groove anteriorly, mesocutum with sparse, long setae. Notauli complete, scrobiculate, long and meeting each other medially. Surface of mesoscutum medially, near scutellar sulcus mostly punctate. Scutellar sulcus transversely long and narrow, 4.00-4.20x as wide as long, 0.85–1.10x as wide as scutellum lengthg. Scutellar sulcus smooth with 3 to 6 full transverse carina. Scutellum flattened in lateral view, smooth and punctate, rounded, smooth and shining laterally. Epicnemial carina present. Subalar groove scrobiculate. Mesopleuron mostly smooth and punctate. Precoxal sulcus, with sparse, long setae. Precoxal sulcus anteriorly scrobiculate, medially and posteriorly absent. Mesosternum smooth. Propodeum 0.46-0.48x as long as mesosoma (in dorsal view). Propodeum entirely areolate. Propodeum convex from base to apex, with long and sparse setae. Propodeal transverse carina without acute angles or apophyses. Metapleuron areolate, with sparse, long setae. Metapleural flange absent. Wings (Fig. 23): Pterostigma 3.75x as long as height. Marginal cell of fore wing closed distally. Vein r 1.75x as long as 3-SR, 0.20x as long as SR1, 0.30x as long as 2-SR. Vein 1-SR+M straight. First discal cell 3x as long as wide (measured perpendicular to vein 1-CU1+2-CU1). Vein r-m of fore wing mostly tubular, with large posterior bulla. Vein m-cu of fore wing antefurcal. Vein cu-a of fore wing postfurcal. Vein CU1a (relative to vein 2-CU1 of fore wing) not interstitial, arising behind middle of distal vein of brachial cell. Vein CU1b postfurcal with m-cu. First subdiscal cell of fore wing closed distally (vein 2cu-a present). Vein 2–1A long. Hind wing vein 1-SC+R present. Subbasal cell of hind wing small, hind wing vein M+CU 0.60x as long as 1-M. Vein r of hind wing present, vein m-cu absent. Basal cell of hind wing closed, vein cu-a present (but nebulous). Legs: Femora not swollen. Hind coxa shortened, smooth dorsally and ventrally. Hind femur 4.20x as long as wide. Hind femur smooth dorsally and ventrally, with sparse setae. Hind tibia with dense, stout and rather short setae, apically

CHELONINAE WITH SPINED PROPODEA Zootaxa 3385 © 2012 Magnolia Press · 21 FIGURES 76–79. Odontosphaeropyx ruficeps Cameron, 1910 Holotype female. 76. detail of tarsal claws of mid leg; 77. full imago, lateral and its labels; 78. mesosoma, dorsal; 79; metasoma, dorsal.

without specialized patch of setae on its inner side. Hind tibia 7.00x as long as wide, 0.78–0.95x hind tarsus. Hind basitarsus flattened laterally, 12.00-12.80x as long as wide, 0.89-0.90x as long as tarsomeres 2–5. Metasoma (Figs 24–25). Metasoma with two sutures on carapace, 4.30x as long as wide. First tergite 1.20x as long as apical width. First tergite without basolateral process (pointed flange-like or spine-like projections). Medio-

22 · Zootaxa 3385 © 2012 Magnolia Press BRAET ET AL. lateral carina of first tergite longer than half length of petiole. First tergite longitudinally striate (or costate-reticulate) over entire surface. Median length of second tergite 1.16-1.26x its basal width, 1x length of first tergite. Sec- ond metasomal suture straight. Second and third tergites densely reticulate-punctate. Remaining tergites concealed by metasomal carapace. Setae on metasoma fine and short. Color. Antenna and scapus dark brown. Head dark brown, mandible orange, setae whitish. Mesosoma dark brown, setae rather whitish. Fore wing banded, fore wing infuscate but hyaline basally and with transverse white band medially, hind wing infuscate apically, veins brownish. Legs dark brown. Fore and mid coxa, fore and mid trochanter, tibial spur pale yellowish to whitish. Metasoma dark-brownish, except for semicircular white patch medially at base of second tergite. Hosts records. None. Comments. This distinctive species can be distinguished from all other Pachychelonus by having a long metasoma (more than 4x as long as wide vs. at most 2.3x for other species), the presence of white spots on its dark metasoma and its biogeographic location. Etymology. From the white color of fore and mid coxae. Distribution record. Ethiopian (Madagascar). Type material. Holotype: % (MRAC): "Coll. Mus. Congo, Madagascar, Rogez, for. Analandraraka, VI-37, A. Seyrig"; in good state. Paratype: % (MRAC), "Coll. Mus. Congo, Madagascar, Rogez, V-1931, A. Seyrig"; in good state.

Phanerotomella erena Braet, sp. nov. (Figs 43–44)

MALE (Holotype). Body length 4.5 mm, fore wing length 4.2 mm. Head. Head as wide as medial height. Antennal socket inserted near middle level of eye. Antenna with 45 flagellomeres. Scapus twice as long as wide, 1.1x first flagellomere, without apical lobe. First flagellomere 4.5x as long as wide, longer than second. Penultimate flagellomere 2.7x as long as wide, 0.2x as long as first segment, shorter than apical segment. Eye length 1.2x length of temple (in dorsal view), 1.36x as high as broad, without setae. Maxillary palpus of normal length (reaching between fore and middle coxae). Clypeus flattened in lateral view, with slightly convex lower margin, without apical teeth, weakly punctate. Malar suture absent. Malar space 1.7x basal width of mandible, 0.43x eye height. Face straight in lateral view,flattened with small bump in its middle, close to antennal socket, weakly punctate and longitudinally weakly rugose, with sparse, long setae. Temple not swollen in dorsal view, coarsely punctate, with sparse, long setae. Frons costate/striate, concave, without carina between antennal socket (a short ruga running between antennal socket to ocellus), without lateral carina. POL 1x ocellar diameter, 0.4x OOL. Vertex striate, with faint and poorly defined transverse striate sculpture, with short, sparse setae. Occipital carina complete, joining hypostomal carina at mandible. Mesosoma. Pronotum mostly punctate. Pronotal furrow shortened, laterally scrobiculate. Mesosoma 1.47x as long as wide in lateral view, 1.74x as long as wide in dorsal view. Mesoscutum sharply raised anteriorly, at right angle with pronotum, medio-anteriorly areolate, median lobe without median groove anteriorly. Mesoscutum with short, sparse setae. Surface of mesoscutum medially mostly punctate or areolate near scutellar sulcus. Notaulus entirely absent. Scutellar sulcus 10x as wide as long, 1.3x as long as scutellum, smooth. Scutellar sulcus with 3 complete carina. Scutellum rounded laterally, flattened in lateral view, punctate. Subalar groove areolate. Mesopleuron entirely areolate, with short, sparse setae. Precoxal sulcus absent. Surface of mesosternum weakly punctate. Propodeum 0.46x as long as mesosoma (in dorsal view), entirely sculptured, areolate, carinated areas completely absent. Propodeum rather vertical posteriorly, with short, sparse setae. Propodeal transverse carina not apparent, not distinguishable from sculpturing. Propodeum with two long spine-like apophyses laterally, median tubercles absent. Propodeal lateral spine 5x its transverse width, straight, with rounded apex. Surface of metapleuron areolate, with short, sparse setae. Metapleural flange absent. Wings: Pterostigma 5x as long as wide. Radial (marginal) cell of fore wing closed distally. Vein r 4x as long as vein 3-SR, 0.17x as long as vein SR1, 0.25x as long as vein 2-SR. Vein 2-SR of fore wing present. Vein 1-SR+M straight. First discal cell of fore wing 1.74x as long as wide (measured perpendicular to vein 1-CU1+2-CU1). Vein r-m of fore wing mostly tubular, with posterior bulla. Vein m-cu postfurcal ((RS+M)b absent, second marginal cell not petiolate. Vein cu-a of fore wing present, nervel- lus (cu-a) of fore wing postfurcal. Vein CU1a (relative to cubital vein, 2-CU1, of fore wing) not interstitial, arising

CHELONINAE WITH SPINED PROPODEA Zootaxa 3385 © 2012 Magnolia Press · 23 apical to middle of distal vein of subdiscal cell. Vein CU1b absent. First subdiscal cell of fore wing open (vein 2cu- a absent or only present as faint brownish spot). Vein 2–1A long. Hind wing vein 1-SC+R present. Subbasal cell of hind wing distinctly enlarged. Hind wing vein M+CU 0.81x as long as vein 1-M. Hind wing vein r absent, m-cu absent. Basal cell of hind wing closed. Hind wing vein cu-a present. Legs: Fore femur and mid femur not swollen. Hind coxa large (more than half length of metasoma), dorsally and ventrally smooth. Hind femur not swollen, 4.93x as long as wide. Hind femur with sparse setae, dorsally and ventrally punctate. Hind tibia 5.77x as long as wide, 2.2x hind basitarsus, with dense, stout and rather short setae, presence of sparse, stout spines on outer side of hind tibia. Hind basitarsus 7x as long as wide, equal in length to tarsomeres 2–5. Metasoma. Metasoma with two sutures on carapace. Carapace without apical opening, anteriorly and posteriorly entirely areolate. . First tergite less than 1.5x as long as apical width, without basolateral process near base, median carina present as two short carina and very short. Median length of second tergite 0.9x its basal width, equal to length of first tergite. Second metasomal suture curved. Third tergite areolate-reticulate. Fourth to sixth tergites concealed by metasomal carapace. Color. Scapus brownish, flagellum dark. Head yellowish; median part of frons, stemmaticum, and posterior part of vertex medially mostly dark-brownish; mandible yellowish but apically light brown. Mesosoma bicolored. Pronotum blackish medially and partly blackish laterally. Mesopleuron anteriorly and ventrally, mesosternum entirely, metapleuron ventrally, mesonotum anteriorly and laterally, metanotum laterally, and propodeum laterally mostly brown; other parts yellowish. Fore wing lightly infuscate, veins and stigma brownish. Legs yellowish. Fore tibia apically, fore tarsus entirely, mid tibia apically, mid tarsus mostly, hind trochantellus strongly, hind tibia apically, and hind tarsus entirely brownish. Carapace black with pale patches medially on first and second tergites. Hosts records. None. Comments. This species may be distinguished from other Phanerotomella by the presence of various dark patches on its body. It is closely related to Phanerotomella seyrigi Granger from which it can be separated by having long lateral spines on the propodeum (vs. short). Etymology. From the Greek 'ειρήνη (eirenè)' meaning "the peace" and in honour of my girlfriend who share my study of insects. Distribution record. Ethiopian (Madagascar). Type material. Holotype: % (CAS): "CASENT 2184137""Madagascar, Province Fianarantsoa, PN Ranoma- fana, radio tower at forest edge, elev. 1130m, 14–24.VI.2002, Collector: R. Harin'Hala""21°15'05''S-47°24'43''E, California Acad of Sciences, Malaise trap, mixed tropical forest, MA-02–098–34, CASLOT 024096"; in good state.

Identification keys to Ethiopian species

A. Phanerotomella (modified from Zettel, 1989):

1 Propodeum with long spines laterally (Figs 43, 52, 55, 56); vein 3-SR of fore wing reduced or absent; body length greater than 4 mm; Madagascar ...... 2 - Propodeum lacking long spines laterally, sometimes with short apophyses or short spines laterally, usually triangular when present (Figs 45, 53, 59); vein 3-SR of fore wing variable; body length variable; mainland Africa and Madagascar...... 5 2(1) Second submarginal cell slightly petiolate basally (vein 2-SR+M present) (Fig. 57); wing with transverse brownish stripe under stigma (Fig. 57); head blackish, basal third of antenna whitish (Figs 54, 58); body mainly orange; body length 6.9–7.9 mm...... Ph. antennata Granger, 1949 - Second submarginal cell not petiolate basally (vein 2-SR+M absent); wing without brownish stripe, otherwise colored; head color and body length variable ...... 3 3(2) Body orange; frons laterally carinate, median carina strong; length of malar space 0.75x vertical length of eye; 5.2–6.5 mm (Fig. 52)...... Ph. spinosa Granger, 1949 - Body with large blackish patches; frons nearly without carina, median carina weak; length of malar space 0.66x or less vertical length of eye; body length less than 5 mm ...... 4 4(3) Body entirely blackish; wings strongly infuscate; lateral propodeal spines short (at most 2x as long as their basal diameter); body length 4.3 mm (Figs 61–63) ...... Ph. seyrigi Granger, 1949 - Body blackish with large, light, yellowish stripes and patches; wings weakly infuscate; lateral propodeal spines long (more than 4x their basal diameter); body length 4.5 mm...... Ph. erena Braet sp. nov. 5(1) Body length more than 5 mm (mostly from Madagascar) ...... 6 - Body length less or equal to 5 mm ...... 7

24 · Zootaxa 3385 © 2012 Magnolia Press BRAET ET AL. 6(5) Body blackish; carapace rather elongate; second submarginal cell of fore wing petiolate (vein 3-SR absent and vein r-m start- ing on 2-SR); body length 5.6–6.2 mm (Figs 59–60) ...... Ph. gigantea Granger, 1949 - Body yellowish; carapace rather oval-rounded in dorsal view; vein 3-SR present but short, thus second submarginal cell of fore wing not petiolate; body length 5.1–5.4 mm (Fig. 65) ...... Ph. subdentata Granger, 1949 7(5) Body mostly yellowish-orange ...... 8 - Body mostly blackish ...... 12 8(7) Carapace oval in dorsal view, less than 1.75x as long as wide; Madagascar ...... 9 - Carapace elongate in dorsal view, more than 1.75x as long as wide (character unknown for Ph. lutea); mainland Africa. . . . 10 9(8) Vertex, sides of mesoscutum and patch on carapace brown to dark-brown; second submarginal cell of fore wing strongly petiolate (vein 3-SR absent); antenna with 38–40 segments; body length 3–3.7 mm ...... Ph. punctata Zettel, 1989 - Vertex, mesoscutum and carapace entirely yellowish; second submarginal cell of fore wing weakly or not petiolate (vein 3-SR sometimes present); antenna with 41–45 segments; body length 3.7–4.1 mm (Fig. 64) ...... Ph. crassa Zettel, 1989 10(8) Body entirely yellowish, stigma yellowish; body length 4 mm; Central Africa...... Ph. lutea Szépligeti, 1922 - Color variable but stigma and large lateral patch on scutellum dark-brown; body length less than 4mm; South Africa. . . . . 11 11(10) Face equal or wider than length of face plus clypeus, face and temple shining; body entirely bright yellow; length of outer tibial spur of mid leg more than 0.5x basitarsus...... Ph. aurea Zettel, 1989 - Face narrow, face and temple weakly shining; body yellowish but blackish around scutellum; length of outer tibial spur of mid leg less or equal to 0.5x basitarsus ...... Ph. capensis Zettel, 1989 12(7) First flagellomeres whitish ...... 13 - Flagellum mainly dark-brown (scapus sometimes paler) ...... 14 13(12) Vein R1 of fore wing shorter than length of stigma; body entirely dark-brown to black; Madagascar (Fig. 53). . . . . Ph. annulicornis Granger, 1949 - Vein R1 of fore wing longer than length of stigma; first tergite with yellow patch; D.R. Congo . . . Ph. juliae De Saeger, 1948 14(12) Carapace thin in lateral view; mesopleuron densely and finely punctate; mesopleuron shining; second submarginal cell of fore wing strongly petiolate (vein 3-SR absent); body length 2.7–4.5 mm; Madagascar ...... 15 - Carapace shape unmodified and weakly arched in lateral view; mesopleuron coarsely punctate (punctations often meeting each others); body sculpture usually coarse and large; second submarginal cell of fore wing variable (vein 3-SR sometimes present); body length variable ...... 16 15(14) Scapus whitish (continental Africa) to mainly darkish with only short white stripe (Madagascar), in all cases scapus apically infuscate; mesoscutum regularly punctate; sometimes first tergite with testaceous patch in male, female blackish (Figs 66–68) ...... Ph. tristis Granger, 1949 - Scapus entirely whitish with small infuscation apically; mesoscutum finely punctate on lateral lobes, longitudinally rugose areolate medially; color of first tergite mostly whitish (excepted baso-lateral corners) on female (male unknown) ...... Ph. spB 16(14) Carapace basally blackish; carapace without obvious teeth postero-laterally ...... 17 - Color of base of carapace variable; postero-lateral teeth of carapace variable...... 18 17(16) Carapace sculpture rather reticulate; second submarginal cell with small petiole (vein 3-SR absent); m-cu interstitial with vein 2-SR of fore wing; antenna with 36 segments; body length 3.0–3.3 mm; D.R. Congo ...... Ph. atrata De Saeger, 1948 - Carapace longitudinally coarsely striate; second submarginal cell of fore wing strongly petiolate (vein 3-SR absent); m-cu slightly postfurcal to interstitial with vein 2-SR of fore wing; antenna with 33 segments; body length 2.9 mm; D.R. of Congo ...... Ph. lepida De Saeger, 1948 18(16) Carapace with distinct teeth postero-laterally; vein 3-SR of fore wing very short but present; scapus yellowish; base of carapace black (Madagascar) to yellowish (continental species); D.R. of Congo, Madagascar . . . . Ph. mucronata De Saeger, 1948 - Carapace without distinct teeth postero-laterally; vein 3-SR of fore wing absent, second submarginal cell petiolate; scapus brown; Madagascar ...... Ph. spA

B. Odontosphaeropyx (modified from Zettel, 1990)

1 Vein 3-SR of fore wing subequal or longer than vein r; carapace oval in dorsal view, less than 2x longer than wide ...... 2 - Vein 3-SR of fore wing shorter than vein r; carapace elongate in dorsal view, more than 2x longer than wide ...... 4 2(1) Vein 3-SR of fore wing longer than vein r; carapace 1.4x to 1.7x longer than wide ...... 3 - Vein 3-SR of fore wing subequal to vein r; carapace 1.7x longer than wide. 11.7 mm; Kenya ...... O. maximus (Zettel, 2002) comb. nov. 3(2) Mesosoma blackish and carapace yellowish, 1.4x longer than wide. 8.5 mm; South Africa, Zambia, Zimbabwe ...... O. fulviventris (Brues, 1924) comb. nov. - Mesosoma and carapace yellowish, 1.7x longer than wide. 8.3 mm; South Africa (Figs 76–79) . . . O. ruficeps Cameron, 1910 4(1) Thorax rufous-orange, coxae brown to dark-brown; carapace black and second tergite with large transverse patch basally, first tergite basally rugulose-reticulate to finely reticulate apically, covered by dense short whitish setae, carapace 2.3x longer than wide; Nigeria, D.R. of Congo (Figs 48–51) ...... O. flavifasciatus (Zettel, 1990a) comb. nov. - Thorax black, fore and mid coxa mainly whitish, fore coxa black; carapace black, color of second tergite variable, sculptures on first tergite variable, length of carapace more than 2.5x longer than wide; Madagascar...... 5 5(4) Fore and mid coxa and trochanters white (Fig. 27); first tergite 1.2x as long as apical width, with two long dorso-lateral carina (reaching the middle of tergum), longitudinally rugose (Fig. 25); vein 2-SC+R of hind wing short; clypeus ventrally with blunt median tooth (Fig. 26); second tergite with small lateral and one median semicircular patch basally, carapace 4.3x longer than

CHELONINAE WITH SPINED PROPODEA Zootaxa 3385 © 2012 Magnolia Press · 25 wide (Fig. 25) ...... O. leucocoxus Braet, sp. nov. - Fore and mid coxa whitish, more or less largely brownish basally, fore and mid trochanters whitish (Fig. 69); first tergite as long as apically wide, with two short dorso-lateral carina, rather areolate rugulose; vein 2-SC+R of hind wing long; at most; clypeus ventrally with acute median tooth (Fig. 74); second tergite red-brownish, carapace 2.8x longer than wide (Fig. 75) ...... O. gracilis Braet, sp. nov.

Additional illustrations of O. fulviventris and O. flavifaciatus can be viewed on the WaspWeb web site housed at the Iziko institution (http://www.waspweb.org/) (accessed 31 May 2012).

New data about species of Cheloninae from Ethiopian region (New localities are indicated with an asterisk *)

Ascogaster breviventris Granger 1949

Ascogaster breviventris Granger 1949: 199 (key), 201 (descr. &, Madagascar). Ascogaster breviventris Granger, 1949: Shenefelt 1973: 818 (cat.).

Distribution: Madagascar: Prov. Antananarivo: Massif de l'Ankaratra – Prov. Toliara: Antanimora – Province Fian- arantsoa*: NP Ranomafana. Endemic. New data: %, “MADAGASCAR: Province Fianarantsoa, Parc National Ranomafana, radio tower at forest edge, elev 1130 m, 5–15.VII.2002; 21°15.05’S - 47°24.43’E; coll: M. Irwin, R. Harin'Hala” “California Acad of Sciences, malaise, mixed tropical forest, MA-02-09B-34”.

Chelonus (Microchelonus) curvimaculatus Cameron 1906 (Figs 45–47)

Chelonella curvimaculata (Cameron 1906): Wilkinson 1932: 9 (fig. 2b, biol., Mauritius), 10 (key). Chelonella curvimaculata (Cameron 1906): Moutia & Mamet 1947: 28 (cat., Mauritius). Chelonus curvimaculatus Cameron 1906: Granger 1949: 192 (fig. 196), 193 (key, descr. % & &, Madagascar), 427 (biol.), 428 (biol.). Chelonus (Neochelonella) curvimaculatus Cameron, 1906: Anonymous 1960: 339 (biol., Madagascar). Chelonus curvimaculatus Cameron 1906: Anonymous (det. Fischer) 1963: 338 (biol., Madagascar), 362 (biol.). Chelonus curvimaculatus Cameron 1906: Brénière 1965: 346 (cat., Madagascar). Chelonus curvimaculatus Cameron 1906: Appert, Betbeder-Matibet & Ranaivosoa 1969 Microchelonus curvimaculatus Cameron 1906: Shenefelt 1973: 881 (tax., cat.). Chelonus curvimaculatus Cameron 1906: Williams. 1974: 3 (cat., Mauritius). Chelonus curvimaculatus Cameron 1906: Williams. 1980: 4 (cat., Mauritius). Chelonus curvimaculatus Cameron 1906: van Achterberg & Polaszek 1996: 53 (key, tax., descr. % & &, Madagascar, Mauri- tius), 102 (fig. 235), 105 (figs. 263, 264). Chelonus curvimaculatus Cameron 1906: Ganeshan & Williams. 2001: 4 (cat., Mauritius).

Hosts: Gelechiidae: Phthorimaea operculella (Zeller, 1873), Scrobipalpa heliopa (Lower, 1900), Pectinophora gossypiella (Saunders, 1843) – Plutellidae: Plutella xylostella (Linnaeus, 1767). Distribution: Madagascar: Prov. Mahajanga: Maromandia – Prov. Antananarivo: Ampefy – Prov. Toliara: Behara, Bekily, Fort-Dauphin, Manombo, Ranomafana, Andohahela NP* – Mauritius: Mauritius. Réunion*: Petite Ile / Piton Bloc, St Pierre / Bassin Martin. Known from the Ethiopian Region. New data: 3 && & %, “MADAGASCAR: Tulear Province, Andohahela National Park, Tsimelahy, Parcelle II; 24°56.21’S - 46°37.60’E; 15-28.I.2004, California Acad of Sciences“ “colls M. Irwin, F. Parker, R. Harin’Hala, elev 180 m, malaise trap in transitional forest, MA-02-20-52“; &, “Petite Ile / Piton Bloc, bordure de parcelled en zone maraichère, alt. 850m, IV.2007, D-Vac, coll. Cirad PVBMT”; &, “St Pierre / Bassin Martin, Station expéri- mentale de l’Armeflhor, parcelle AB, alt. 290m, XII.2010, tente Malaise, coll. Cirad PVBMT” ; 2%%, “St Pierre, jardin de centre ville, alt. 40m, XII.2010, piège jaune, coll. T. Ramage”.

Phanerotomella annulicornis Granger 1949: 211 (key), 213 (descr. &, Madagascar). Phanerotomella annulicornis Granger 1949: Shenefelt 1973: 929 (cat.). Phanerotomella annulicornis Granger 1949: Zettel 1989: 70 (key), 86 (tax., descr. &, Madagascar), 142 (figs. 289-293).

Distribution: Madagascar: Prov. Toliara: Bekily – Province Fianarantsoa*: Manombo Special Reserve. Endemic. New data: &, “MADAGASCAR: Province Fianarantsoa, Manombo Special Reserve camp site, 32 km SSE of Farafangana, 20.I–II.2005; 23°01.31’S - 47°43.20’E; California Acad of Sciences“ “colls M. Irwin, R. Harin'Hala, malaise trap, lowland rain-forest, elev 36m, MA-28-11”.

Phanerotomella antennata Granger 1949

Phanerotomella antennata Granger 1949: 211 (key, descr. % & &). Phanerotomella antennata Granger 1949: Shenefelt 1973: 929 (cat.). Phanerotomella antennata Granger 1949: Zettel 1989: 68 (key), 71 (tax., descr. % & &, Madagascar), 129 (figs. 216-221).

Distribution: Madagascar: Prov. Mahajanga: Andriba – Prov. Toamasina: Andekaleka (Rogez) – Province Fian- arantsoa*: Ranomafana NP, Manombo Special Reserve. Endemic. New data: 4 && & 2 %%, “MADAGASCAR: Province Fianarantsoa, Parc National Ranomafana, radio tower at forest edge, elev 1130 m; 5 – 15.VII.2002; 21°15.05’ S-47° 24.43’ E; coll: M. Irwin, R. Harin'Hala. California Acad of Sciences, malaise trap, mixed tropical forest. MA-02-09B-34“; 2 %%, “MADAGASCAR: Province Fian- arantsoa, Manombo Special Reserve camp site 32 km SSE of Farafangana, 20 Jan - 2 Feb 2005; 23° 01.31’ S-47° 43.20’ E; California Acad of Sciences, coll: M. Irwin, R. Harin'Hala malaise trap, lowland rainforest, elev 36m MA-28-11”.

Phanerotomella crassa Zettel 1989

Phanerotomella crassa Zettel 1989: 69 (key), 78 (descr. % & &, Madagascar), 135 (figs. 240-254).

Distribution: Madagascar: Prov. Toliara: Berenty Reserve, Andohahela NP*. Endemic. New data: %, “MADAGASCAR: Tulear Province, Andohahela National Park, Tsimelahy, Parcelle II; 24°56.21’S - 46°37.60’E; 15-28.I.2004, California Acad of Sciences“ “colls M. Irwin, F. Parker, R. Harin’Hala, elev 180 m, malaise trap in transitional forest, MA-02-20-52“.

Phanerotomella gigantea Granger 1949

Phanerotomella gigantea Granger 1949: 211 (key), 215 (descr. %, Madagascar). Phanerotomella gigantea Granger 1949: Shenefelt 1973: 930 (cat.). Phanerotomella gigantea Granger 1949: Sigwalt 1977: 530 (fig. 3). Phanerotomella gigantea Granger 1949: Zettel 1989: 69 (key), 75 (tax., descr. %, Madagascar), 132 (figs. 232-236).

Distribution: Madagascar: Prov. Toamasina: Andekaleka (Rogez) – Province Fianarantsoa*: NP Ranomafana – Province Antsiranana*: Marojejy NP – Prov. Toliara*: Ivohibe. Endemic. New data: 2&& & 2%%, “MADAGASCAR: Province Fianarantsoa, Parc National Ranomafana, radio tower at forest edge, elev 1130 m, 5–15.VII.2002; 21°15.05’S - 47°24.43’E; coll: M. Irwin, R. Harin'Hala” “California Acad of Sciences, malaise, mixed tropical forest, MA-02-09B-34”; 4 && & %, “MADAGASCAR: Province Fian-

CHELONINAE WITH SPINED PROPODEA Zootaxa 3385 © 2012 Magnolia Press · 27 arantsoa, Parc National Ranomafana, radio tower at forest edge, elev 1130 m, 5–15.VII.2002; 21°15.05’S - 47°24.43’E; coll: M. Irwin, R. Harin'Hala” “California Acad of Sciences, malaise, mixed tropical forest, MA-02- 09B-34”; &, “MADAGASCAR: Toliara, Foret Ivohibe 55km N Tolagnaro, 2–4.XII.2006; 24˚34'08"S - 047˚12'14"E; California Acad of Sciences” “colls B.L.Fisher et al., yellow pan trap, elev 200 m, rainforest, BLF15449”; &, “MADAGASCAR: Province Antsiranana, Marojejy National Park, 5 km W Manantenina village, Camp Mantella, 28.IV – 7.V.2005; 14°26.29'S - 49°46.44'E; California Acad of Sciences” “colls M. Irwin, R. Harin'Hala, malaise trap, low altitude rainforest, elev. 490 m, MA-31-20”. Comments: The body shows a great variation in length (3.5 to 6.2 mm), males are usually smaller than females. The scapus are mostly whitish but sometimes a small blackish patch is present on the outer side. The hind coxae are sometimes entirely whitish.

Phanerotomella mucronata De Saeger, 1948

Phanerotomella mucronata De Saeger 1948: 192 (key), 197 (descr. &, DR. Congo) Phanerotomella mucronata De Saeger 1948 Phanerotomella mucronata De Saeger 1948: Zettel 1989: 70 (key), 83 (tax., descr. &, DR. Congo), 139 (figs. 273-277).

Distribution: D.R. Congo: Prov. Equateur, Bolingo; Madagascar*: Prov. Toliara: Ivohibe. Known from the Ethiopian Region. New data: &, “MADAGASCAR: Toliara Foret Ivohibe 55.0km N Tolagnaro, 2 - 4 Dec 2006, 24˚34'08"- 047˚12'14" E; California Acad of Sciences, coll. B.L.Fisher et al., yellow pan trap, elev 200 m, rainforest, BLF15449“.

Phanerotomella seyrigi Granger, 1949

Phanerotomella seyrigi Granger 1949: 211 (key), 213 (descr. &, Madagascar). Phanerotomella seyrigi Granger 1949: Shenefelt 1973: 931 (cat.). Phanerotomella seyrigi Granger 1949: Zettel 1989: 69 (key), 72 (tax., descr. &, Madagascar), 130 (figs. 222-226).

Distribution: Madagascar: Prov. Toamasina: Ivondro – Prov. Toliara*: Andohahela NP. Endemic. New data: &, “MADAGASCAR: Toliara, Parc National Andohahela, Col de Tanatana, 33.3km NW Tolagnaro, elev 275 m, 22-24.xi.2006; 24˚45'31"S - 046˚51'13"E; California Acad of Sciences” “colls B.L.Fisher et al., Mal- aise trap, rainforest, collection code: BLF15102”.

Phanerotomella spinosa Granger, 1949

Phanerotomella spinosa Granger 1949: 211 (key, fig. 213), 212 (descr. % & &, Madagascar). Phanerotomella spinosa Granger 1949: Docavo Alberti 1960: pl. 16 (fig. 91). Phanerotomella spinosa Granger 1949: Docavo Alberti 1965 Phanerotomella spinosa Granger 1949: Shenefelt 1973: 931 (cat.). Phanerotomella spinosa Granger 1949: Sigwalt 1977: 531 (fig. 5). Phanerotomella spinosa Granger 1949: Zettel 1989: 68 (key), 70 (tax., descr. % & &, Madagascar), 72, 128 (figs. 211-215).

Distribution: Madagascar: Prov. Prov. Antsiranana: Réserve Naturelle Intégrale de Marojejy (Ambasoratra – Prov. Antananarivo: Mandraka – Prov. Toamasina: Andekaleka (Rogez) – Prov. Fianarantsoa: Massif de l'Andringita (Vallée de l'Iantara:Ampasy), Ranomafana NP*. Endemic. New data: 3 %%, “MADAGASCAR: Province Fianarantsoa, Parc National Ranomafana, radio tower at forest edge, elev 1130 m, 5–15.VII.2002; 21°15.05’S - 47°24.43’E; coll: M. Irwin, R. Harin'Hala” “California Acad of Sciences, malaise, mixed tropical forest, MA-02-09B-34”.

Phanerotomella subdentata Granger 1949: 211 (key, fig. 212), 212 (descr. % & &, Madagascar). Phanerotomella subdentata Granger 1949: Shenefelt 1973: 931 (cat.). Phanerotomella subdentata Granger 1949: Zettel 1989: 69 (key), 77 (tax., descr. % & &, Madagascar), 134 (figs. 243-247).

Distribution: Madagascar: Prov. Toliara: Bekily – Province Antsiranana: Marojejy NP*. Endemic. New data: %, “MADAGASCAR: Province Antsiranana, Marojejy National Park, 5 km W Manantenina village, Camp Mantella, 28.IV – 7.V.2005; 14°26.29'S - 49°46.44'E; California Acad of Sciences” “colls M. Irwin, R. Harin'Hala, malaise trap, low altitude rainforest, elev. 490 m, MA-31-20”.

Phanerotomella tristis Granger, 1949

Phanerotomella tristis Granger 1949: 210 (fig. 211), 211 (key), 214 (descr. % & &, Madagascar). Phanerotomella tristis tristis Granger 1949: Shenefelt 1973: 931 (cat.). Phanerotomella tristis Granger 1949: Zettel 1989: 69 (key), 70 (key), 73 (tax., descr. % & &, Madagascar), 131 (figs. 227-231). Phanerotomella tristis ambositrensis: Granger 1949: 214 (descr. % & &). Phanerotomella tristis ambositrensis Granger 1949: Shenefelt 1973: 931 (cat.). Phanerotomella tristis ambositrensis Granger 1949: Zettel 1989: 73 (syn.).

Distribution: Madagascar: Prov. Antananarivo: Mandraka – Prov. Toamasina: Andasibe (Perinet), Andekaleka (Rogez), Ivondro – Prov. Fianarantsoa: Ambositra, Fianarantsoa, Manombo Special Reserve*, NP Ranomafana* – Province Antsiranana*: NP Marojejy. Endemic. New data: & & 4 %%, “MADAGASCAR: Province Fianarantsoa, Manombo Special Reserve camp site, 32 km SSE of Farafangana, 20.I–II.2005; 23°01.31’S - 47°43.20’E; California Acad of Sciences“ “colls M. Irwin, R. Harin'Hala, malaise trap, lowland rain-forest, elev 36m, MA-28-11”; 46 %% & 13 &&, “MADAGASCAR: Prov- ince Fianarantsoa, Parc National Ranomafana, radio tower at forest edge, elev 1130 m, 5–15.VII.2002; 21°15.05’S - 47°24.43’E; coll: M. Irwin, R. Harin'Hala” “California Acad of Sciences, malaise, mixed tropical forest, MA-02- 09B-34”; 2 %%, “MADAGASCAR: Province Antsiranana, Marojejy National Park, 5 km W Manantenina village, Camp Mantella, 28.IV – 7.V.2005; 14°26.29'S - 49°46.44'E; California Acad of Sciences” “colls M. Irwin, R. Harin'Hala, malaise trap, low altitude rainforest, elev. 490 m, MA-31-20”.

Phanerotomella spA

&, “MADAGASCAR: Province Fianarantsoa, Parc National Ranomafana, radio tower at forest edge, elev 1130 m, 5 – 15 July 2002, 21° 15.05’ S-47° 24.43’ E, coll: M. Irwin, R. Harin'Hala. California Acad of Sciences, malaise trap, mixed tropical forest. MA-02-09B-34”.

Phanerotomella spB

&, “MADAGASCAR: Province Antsiranana, Marojejy Nat'l Park, 5 km W Manantenina village, Camp Mantella, 28 April - 7 May 2005, 14° 26.29' S-49° 46.44' E, California Acad of Sciences, coll: M Irwin, R. Harin'Hala. Mal- aise trap, low altitude rainforest, elev. 490 m, MA-31-20”.

Discussion and conclusions

Braconidae (Hymenoptera, Ichneumonoidea) is a large family present in all parts of the world (except Antarctica) whose members have diverse or even extraordinary morphological characteristics. Among them, long spine-like

CHELONINAE WITH SPINED PROPODEA Zootaxa 3385 © 2012 Magnolia Press · 29 protuberances (teeth) on different parts of the body are not uncommon in several subfamilies. They occur on the head, as in Lasiophorus Haliday, 1838 (Braconinae), on the pronotum, e.g., Spinaria Brullé, 1846 (Rogadinae), on the mesoscutum in Thoracoplites Fischer, 1961 (Rhyssalinae), on the scutellum in Heratemis Walker, 1860 (Alysi- inae), on the propodeum of some Rogadinae, e.g., Conspinaria Schulz, 1906, Pambolinae, e.g., Pambolus Haliday, 1836, and Doryctinae, e.g. Fijispathius Belokobylskij, Iqbal et Austin, 2004; Echinodoryctes Belokobylskij, Iqbal et Austin, 2004; and Ryukyuspathius Belokobylskij, 2008, and finally on the metasoma in Conspinaria Schulz, 1906 (Rogadinae) and Chelonus (Areselonus) Braet, 1999 (Cheloninae) (Braet, 1999; Fisher, 1961; van Achter- berg, 2007). Several Cheloninae genera exhibit a raised transverse carina on propodeum (Chelonus s.l., Asco- gaster). This carina may be a simple ridge, or two (e.g., Chelonus oculator (Fabricius, 1775) to four (e.g., Ascogaster cognata De Saeger, 1948) obtuse to acute apophyses. These may appear as a tubercle, large obtuse triangle, or small pointed teeth. In all cases, these different shapes are pooled under the term apophysis, e.g., triangular apophysis, tubercular apophysis, toothed apophysis. These apophyses are here not considered as spines because they are more or less triangular or blunt and their height is equal to, or less than, their basal diameters. Spines on the propodeum, as defined here, are at least twice (often more) as long as their basal diameter. These spines are only found in a few chelonine taxa and are only present laterally. Lateral protuberances from the carina (apophysis or spines) are sometimes more pronounced than the median ones (e.g., Ascogaster mayamotensis De Saeger, 1948) or the median ones may be absent. Moreover, the relative position of these angles is also variable; either the apophyses are equally spaced on the transverse ridge, or the two median ones are closer one to each other than they are to the outer ones. The observed variations could be useful for future cladistic analyses of the huge genus Chelonus and may be subdivided into eight qualitative characters (here considered as unordered):

State 1: transverse carina of propodeum present/absent. State 2: transverse carina of propodeum lower/higher than the neighboring sculpture. State 3: shape of transverse carina (if present) without angles/with apophyses/with spines. State 4: number of apophyses or spines equal to 0/2/4. State 5: position of apophyses, spines equally spaced/medially tighter. State 6: size of lateral apophyses, spines at most as long as basal diameter/longer than basal diameter. State 7: lateral angles, apophyses, spines (if present) apically blunt/apically acute. State 8: median apophyses absent/as long as lateral ones/longer than lateral ones.

The function(s) of the propodeal apophyses, and spines is unknown. As they are equally developed in both sexes, they probably do not have a sexual function. They could assist in emergence from the host, or in aiding ovi- position into host eggs by stabilizing the abdomen in female. The idea that they may deter predation seems unlikely for those species with smaller apophyses but may have evolved as such in those with long, sharp spines. With the exception of a group of species related to C. (M.) merdicus sp. nov. and C. (M.) matilei sp. nov., all taxa with long spines are restricted to the Malagasy subregion: Madagascar and the Mascarene archipelago (Reunion, Mauritius and Rodrigues islands).

Acknowledgements

We thank Dr. Zuparko (CAS), Dr. Villemant and Ms. Touret-Alby (MNHN), Dr. de Coninck and M. Hannot (MRAC), Dr. Delvare (CIRAD), Dr Franck Koch (MNHU) and Dr. Madl (IFENM) for the loan of specimens. We are grateful to Dr. Madl and to Dr. van Achterberg (Leiden) for bibliographic references extracted from their future catalog of the Malagasy sub-region. We thank Ms. Touret-Alby and M. Hannot for their help in preparing illustrations and photos. Fieldwork in Madagascar could not have been completed without the gracious support of the Malagasy people. The Malaise-trap material was collected and processed in Madagascar through grants from the Schlinger Foundation. We thank Mike Irwin and Eve Schlinger, who planned most of the field trapping program, and Harinhala Rinha, who conducted much of the fieldwork and supervised specimen processing in Antananarivo. We give also many thanks for the collection of Reunion specimens to Dr. Rochat, Ms. Gasnier, Dr. Deguine, MM Ramage, Moutoussamy and Ajaguin-Soleyen. Finally, many thanks to Dr J.T. Jennings and Mss Rebecca Kittel (University of Adelaide, Australia) for their review of our paper.

30 · Zootaxa 3385 © 2012 Magnolia Press BRAET ET AL. This work was partly supported by the Société Entomologique de France (Germaine Cousin Grant to P. Rousse) and by the National Science Foundation (Grants DEB-0072713 to B.L. Fisher and C.E. Griswold and DEB-0344731 to B.L. Fisher and P.S. Ward). We also thank reviewers for their suggestions and improvements.

References

Achterberg, C. van (1984) Essay on the phylogeny of the Braconidae (Hymenoptera: Ichneumonoidea). Entomologisk Tidskrift, 105, 41–58. Achterberg, C. van (1990) Revision of the western Palaearctic Phanerotomini (Hymenoptera: Braconidae). Zoologische Ver- handelingen, 255, 1–106. Achterberg, C. van (1993) Illustrated key to the subfamilies of of the Braconidae (Hymenoptera, Ichneumonoidea). Zoologis- che Verhandelingen Leiden, 283, 1–189. Achterberg, C. van & Polaszek, A. (1996) The parasites of cereal stem borers (Lepidoptera: Cossidae, Crambidae, Noctuidae, Pyralidae) in Africa, belonging to the family Braconidae (Hymenoptera: Ichneumonoidea). Zoologische Verhandelingen, 304, 123 pp. Achterberg. C. van (2007) Revision of the genus Spinaria Brullé (Hymenoptera: Braconidae: Rogadinae), with keys to genera and species of the subtribe Spinariina van Achterberg. Zoologische Mededelingen Leiden, 81(2), 11–83. Baker, C.F. (1926) Braconidae-Cheloninae of the Philippines, Malaya, and Australia. 1. Chelonini (except Chelonus). Philip- pine Journal of Science, 31 451–489. Belokobylskij, S.A. (2008) Rhyukyuspathius, a new peculiar genus of the tribe Spathiini (Hymenoptera: Braconidae: Dorycti- nae) from Japan. Annales Zoologici (Warsaw), 58(4), 761–766. Belokobylskij, S.A., Iqbal, M. & Austin, A.D. (2004) Systematics, distribution and diversity of the Australasian doryctine wasps (Hymenoptera, Braconidae, Doryctinae). Records of the South Australian Museum Monograph Series, 8, 1–150. Braet, Y. & Achterberg, C. van (2001) New species of the genera Foenomorpha Szépligeti (Cenocoeliinae) and Chelonus Pan- zer (Cheloninae) (Hymenoptera: Braconidae), from French Guiana, and Brazil. Zoologische Mededelingen Leiden, 75(6), 103–118. Braet, Y. (1999) Description of new Braconidae (Hymenoptera) from Papua New Guinea. Belgian Journal of Entomology, 1(1), 3–20. Brues, C.T. (1924) Some South African parasitic Hymenoptera of the families Evaniidae, Braconidae, Alysiidae, and Plumarii- dae in the South African Museum with a catalogue of the known species. Annals of the South African Museum, 19, 1–150. Brulle, M.A. (1846) Tome Quatrieme. Des Hymenopteres. Les Ichneumonides. In: Lepeletier de Saint-Fargeau A. 'Histoire Naturelles des Insectes.' Paris. 680 pp. pp. 56-521. Cameron, P. (1910) On some African species of the subfamilies Exothecinae, Aphrastobraconinae, Cheloninae, Doryctinae, Cardiochilinae and Microgasterinae in the Royal Berlin Zoological Museum. Zeitschrift für Naturwissenschaft, 81, 433–450. De Saeger, H. (1942) Espèces nouvelles et notes sur quelques Chelonides (Hym. Braconidae) du Congo belge. Revue de Zoolo- gie et de Botanique Africaines, 36, 8–15. De Saeger, H. (1948) Cardiochilinae et Sigalphinae (Hymenoptea: Apocrita) Fam. Braconidae. Exploration du Parc National Albert. Mission G.F. de Witte, 53, 1–272. Fabricius, J.C. (1775) Systema Entomologiae, sistens Insectorum classes, ordines, genera, species. Flensburgi et Lipsae, 832 pp. Fahringer, J. (1934) Opuscula braconologica, Band 3. Palaearktischen Region. Lieferung 5-8. Fritz Wagner, Wien, 321-594. Fischer, M. (1961) Eine neues Opiinen-Genus aus Afrika (Hymenoptera, Braconidae). Annales Historico-Naturales Musei Nationalis Hungarici, 53, 439–441. Förster, A. (1862) Synopsis der Familien und Gattungen der Braconiden. Verhandlungen des Naturhistorischen Vereins der Preussischen Rheinlande und Westfalens, 19, 225–288. Granger, C. (1949) Braconides de Madagascar. Mémoires de l’Institut scientifique de Madagascar (A), 2, 1–428. Haliday, A.H. (1836) Essay on parasitic Hymenoptera. Entomological Magazine, 4(i), 38–59. Haliday, A.H. (1838) Essay on parasitic Hymenoptera. Entomological Magazine, 5(3), 209–249. He, J.H., Chen, X.X. & Achterberg, C. van (1994) Siniphanerotomella gen. nov., a new genus of the subfamily Cheloninae Nees (Hymenoptera: Braconidae) from China. Zoologische Mededelingen, 68(17), 191–195. He, J.H., Chen, X.X. & Achterberg, C. van (1997) Scabrichelonus, a new subgenus of Chelonus Panzer (Hymenoptera: Bra- conidae: Cheloninae) from China. Zoologische Mededelingen, 71(7), 53–56. Jones, D. (1985) Endocrine interaction between host (Lepidoptera) and parasite (Cheloninae in Hymenoptera): Is the host or the parasite in control? Annals of the entomological Society of America, 78(2), 141–148. Lucas, H. (1849) Exploration scientifique de l'Algérie pendant les années 1840, 1841, 1842. Sciences physiques, Zoologie, III. Insectes. Paris, 527pp. Martiré, D. & Rochat, J. (2008) Les papillons de La Réunion et leurs chenilles. Biotope, Mèze (collection Parthénope). Muséum d'Histoire Naturelle, Paris, France, 496 pp.

CHELONINAE WITH SPINED PROPODEA Zootaxa 3385 © 2012 Magnolia Press · 31 Murphy, N., Banks, J.C., Whitfield, J.B. & Austin, A.D. (2008) Phylogeny of the parasitic microgastroid subfamilies (Hymenoptera: Braconidae) based on sequence data from seven genes, with an improved time estimate of the origin of the lineage. Molecular Phylogenetics and Evolution, 47(1), 378–395. Panzer, G.W.F. (1806) Kritische Revision der Insektenfaune Deutschlands nach dem System bearbeitet II. Nürnberg, 271 pp. Papp, J. (1995) Revision of C. Wesmael's Chelonus species (Hymenoptera Braconidae Cheloninae). Bulletin de l’Institut royal des Sciences naturelles de Belgique, 65, 115–134. Quicke, D.L.J. & Achterberg, C. van (1990) Phylogeny of the subfamilies of the family Braconidae (Hymenoptera: Ichneu- monoidea). Zoologische Verhandelingen Leiden, 258(1), 1–95. Schulz, W.A. (1906) Spolia Hymenopterologica. Paderborn (Junfermann), 356 pp. Shaw, M.R. & Huddleston, T. (1991) Classification and biology of Braconid wasps (Hymenoptera: Braconidae). Handbooks for the Identification of British Insects, 7(11), 126 pp. Shi, M., Chen, X.X. & Achterberg, C. van (2005) Phylogenetic relationships among the Braconidae (Hymenoptera: Ichneu- monoidea) inferred from partial 16S rDNA, 28S rDNA D2, 18S rDNA gene sequences and morphological characters. Molecular Phylogenetics and Evolution, 37, 104–116. Sigwalt, B. (1977) La nomenclature alaire de Comstock-Needham dans l'étude des Braconides: distinction des genres Phanero- toma et Phanerotomella (Hym., Braconidae, Cheloninae). Annales de la Société Entomologique de France, 13(3), 525–534. Sigwalt, B. (1978) Le genre Phanerotomella Szepligeti généralités. Nouvelles espèces du sud-est asiatique (Hym. Braconidae). Annales de la Société Entomologique de France (N.S.), 14(4), 715–725. Szépligeti, G. (1900) Braconiden aus New-Guinea in der Sammlung des ungarischen National Museums. Termeszetrajzi Fuzetek, 23, 49–65. Szépligeti, G. (1908) Braconiden aus der Sammlung des ungarischen National-Museums, 2. Annales Historico-Naturales Musei Nationalis Hungarici, 6, 397–427. Tobias, V.I. (2000) Cheloninae. In: Ler, P.A. (ed.) [Key to the insects of Russian Far East. Vol. IV. Neuropteroidea, Mecoptera, Hymenoptera. Pt 4.] Opredelitel nasekomykh Dalnego Vostoka Rossii. T. IV. Setchatokryloobraznye, skorpionnitsy, pere- ponchatokrylye. Ch. 4. Dalnauka, Vladivostok. pp.426–571. Walker, F. (1860) Characters of some apparently undescribed Ceylon insects. Annals and Magazine of Natural History, (3)5, 304–311. Walker, A.K. & Huddleston, T. (1987) New Zealand chelonine braconid wasps (Hymenoptera). Journal of Natural History, 21(2), 339–361. Wesmael, C. (1835) Monographie des Braconides de Belgique. Nouveaux Memoires de l'Academie Royale des Sciences et Belles-lettres Bruxelles, 9, 1–252. Wesmael, C. (1838) Monographie des Braconides de Belgique. 4. Nouveaux Memoires de l'Academie Royale des Sciences et Belles-lettres de Bruxelles, 11, 1-166. Yu, D.S., Achterberg, C. van & Horstmann, K. (2005) World Ichneumonoidea 2004. Taxonomy, biology, morphology and distribution [Braconidae].― Taxapad 2005 (Scientific names for information management) Interactive electronical catalogue on DVD/CD-ROM. Vancouver. Zettel, V.H. (1989) Die Gattung Phanerotomella Szépligeti (Hymenoptera: Braconidae, Cheloninae). Linzer Biologische Beiträge, 21(1), 15–142. Zettel, V.H. (1990a) Eine Revision der Gattungen der Cheloninae (Hymenoptera, Braconidae) mit Beschreibungen neuer Gat- tungen und Arten. Annalen des Naturhistorischen Museums in Wien, 91(B), 147–196. Zettel, V.H. (1990b) Zwei neue Phanerotoma-Arten aus Südafrica (Hymenoptera, Braconidae: Cheloninae). Linzer Biologische Beiträge, 22, 335–340. Zettel, V.H. (1991) Beiträge zur Kenntnis neotropischer Arten der Gattung Phanerotoma Wesmael: 5. Die Ph. minuta-Gruppe und 6. Die Ph. popovi-Gruppe (Hymenoptera: Braconidae, Cheloninae). Linzer Biologische Beiträge, 23(1), 375–385. Zettel, V.H. (1992a) Beiträge zur Kenntnis neotropischer Arten der Gattung Phanerotoma Wesmael: 7. Die Ph. atriceps- Gruppe (Hymenoptera: Braconidae, Cheloninae). Linzer Biologische Beiträge, 24(2), 663–669. Zettel, V.H. (1992b) Revision der Phanerotoma-Arten Nordamerikas (Hymenoptera: Braconidae, Cheloninae). Linzer Biologis- che Beiträge, 24(1), 275–330. Zettel, V.H. (2002) Pachychelonus maximus sp. n. (Hymenoptera: Braconidae: Cheloninae) aus Kenia. Linzer Biologische Beiträge, 34(1), 195–198.

View publication stats