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International audience; Species in the stem borer noctuid subtribe Sesamiina are notoriously difficult to distinguish because most related species have homogeneous wing patterns and almost indistinguishable genitalia. The latter is potentially problematic because this group includes several important pest species that are usually baregly distinguishable from non-pest species. In this study we focus on the Mediterranean corn stalk borer Sesamia nonagrioides (Lefèbvre), an important pest of maize with a wide area of distribution that covers most of Africa and extends to the south of Europe and western Asia. According to a recent study, this pest consists of three allopatric populations that were formerly considered as distinct species or subspecies. Here we rely on recent collections of 5470 specimens (sampled in 17 countries and 175 localities) that putatively belong to S. nonagrioides. Integrative taxonomy studies allowed us to unravel the existence of six new species that are closely related to S. nonagrioides and described in this paper. In contrast to S. nonagrioides these new species have more specific ecological preferences, as they are associated with a limited number of plant species and habitats. Dating and population genetic analyses carried out on 100 S. nonagrioides specimens also indicate a more complex population structure than previously thought for S. nonagrioides, which can probably be accounted for by late Cenozoic environmental changes.

The aim of this study was to review the species of Conicofrontia Hampson, a small genus of noctuid stem borers (Noctuidae, Apameini) that is distributed in East and Southeastern Africa. We review the morphology of species in this group and provide new diagnoses and ecological data for five species. The following taxonomic changes are proposed: Hygrostola dallolmoi (Berio, 1973) (= Conicofrontia dallolmoi Berio, 1973) comb. n. and Conicofrontia bipartita (Hampson, 1910) (= Phragmatiphila bipartita Hampson, 1910) comb. n., stat. rev. One new species is also described: C. lilomwa, sp. n. from Tanzania. Wing patterns as well as male and female genitalia of the five species are described and illustrated. Finally we carried out molecular phylogenetic and molecular species delimitation analyses on a multi-marker dataset of 31 specimens and 15 species, including the five mentioned species. The results of molecular analyses provide a clear support for the proposed taxonomical changes

Conicobruchus flabellicornis (Boheman, 1829) Bruchus flabellicornis Boheman, 1829: 105 Bruchus antonioi Pic, 1943: 6 (synonymy in Decelle, 1975: 18) Conicobruchus flabellicornis: Decelle, 1951: 181 Material examined. Holotype (female) of Bruchus antonioi: Angola: San Antonio (now Soyo), 1929 (H. Brauns) [MNHN]. Other material: Kenya: 3 ♂, 4 ♀, Suam, 01�� 11.982 ���N 34 �� 49.538 E, 1951m, 12.xi. 2008, ex Crotalaria incana [2 ♂ 0 1209, 0 2312, specimen GK 200 used for DNA extraction] (B. Le Ru) [CBGP]; 1 ♀, Kakamega, 00�� 10.990 ���N 34 �� 58.137 E, 1693m, 11.xi. 2008, ex Crotalaria incana purpurascens [specimen GK 243 used for DNA extraction] (B. Le Ru) [CBGP]; 1 ♂, 3 ♀, Burguret, 00��04.926���N 37 �� 02.203E, 1915m, 21.xi. 2011, ex Crotalaria incana purpurascens [1 ♂ 0 2112, specimen GK 452 used for DNA extraction] (B. Le Ru) [CBGP]; 2 ♂, 1 ♀, same data, but ex Crotalaria brevidens intermedia [1 ♂ 0 2412, specimen GK 442 used for DNA extraction] (B. Le Ru) [CBGP]. Togo: 1 ♀, Lom��, 28.viii. 1995, ex Crotalaria goreensis (I.A. Glitho) [MNHN]. Black, but Kenyan specimens often with elytral disc dark red. A pod sample collected in Suam yielded two black and five ���red��� specimens, whereas one sample from Burguret yielded only ���red��� specimens. Color is apparently unrelated with sex. Other major morphological traits are as follows: pronotum 1.37 to 1.43 times wider at base than long, with oblique lateral impression, constricted anteriorly into a wide neck, strongly rugose-punctate, with several white hair spots. These were described in detail by F��hraeus (1839: 4): a small spot in the ante-scutellar fossa, a short line just before apex, at posterior angles, and a lateral spot on each side; an additional pair of white spots is often visible on sides of basal lobes, scutellum with similar white setation. Elytral striae strong, with small punctures, interstriae flat, with series of shallow circular punctures. Male genitalia. Median lobe (Fig. 7) of moderate length, stout (maximum width excluding basal hood / total length = 0.17), widened apically, basal hood widened, concave posteriorly; ventral valve large, subtriangular, with apex acute, bearing a few sensillae and two lateral groups of 3���5 setae; no hinge sclerite; internal sac without central column of tubercules, but with rather dense hyaline scales and tubercles, ending posteriorly in a dorsal mass of small, poorly sclerotized teeth, blending into a short series of well sclerotized teeth of variable length; a pair of strong ventro-lateral dented rods; between these a few isolated teeth; posteriorly two poorly defined groups of dented masses; apical ampoule devoid of any ornamentation, gonopore unsclerotized; basal strut narrow, without keel; lateral lobes cleft to 85 % their length; apex modified, with two lips, the dorsal one bearing about twelve long setae, the ventral one with a dense group of short sensillae. Biology. Examined material was reared from pods of Crotalaria goreensis and Crotalaria incana subsp. purpurascens. This constitutes the first verified record on its biology. Distribution. Angola, Burundi (Decelle, 1956), Democratic Republic of Congo (Decelle, 1951), Kenya, Sierra Leone (F��hraeus, 1839) and Togo. Discussion. Conicobruchus flabellicornis can be distinguished form all other members of the group based on the presence of brushes and spots of dense white setae on the pronotum (present in C. flabellicornis, absent in other species)(Fig. 1)., Published as part of Le Ru, Bruno P., Delobel, Alex, Gy��rgy, Zolt��n, Genson, Gwena��lle & Kergoat, Gael J., 2014, Taxonomy, host-plant associations and phylogeny of African Crotalaria - feeding seed beetles (Coleoptera, Chrysomelidae, Bruchinae): the Conicobruchus strangulatus (F��hraeus) species group, pp. 238-256 in Zootaxa 3895 (2) on pages 247-249, DOI: 10.11646/zootaxa.3895.2.6, http://zenodo.org/record/226173, {"references":["Boheman, C. H. (1829) Novae Coleopterum species. Memoires de la Societe Imperiale des Naturalistes de Moscou, 7, 101 - 133.","Pic, M. (1943) Opuscula martialis. L'echange, numero special, 10, 1 - 16.","Decelle, J. (1975) Les Coleopteres Bruchides d'Angola. Publicacoes Culturais Cia Diamantes Angola, 89, 13 - 32.","Decelle, J. (1951) Contribution a l'etude des Bruchidae du Congo Belge (Col. Phytophaga). Revue de zoologie et de botanique africaines, 45, 172 - 192.","Fahraeus, O. I. (1839) In: Schonherr, C. J. (Ed.), Genera et species Curculionidum, cum synonymia hujus, familiae. Roret, Paris, Fleischer, Stockholm, pp. 1 - 495.","Decelle, J. (1956) Contributions a l'etude de la faune entomologique du Rwanda Urundi (Mission P. Basilewsky, 1953). XCIX. Coleoptera: Bruchidae. Annales du Musee Royal du Congo Belge. Sciences Zoologiques. Tervuren, 51, 423 - 426."]}

Conicobruchus decoratus (F��hraeus, 1871) Bruchus decoratus F��hraeus, 1871: 448 Bruchus decoratus atrodorsalis Decelle, 1960: 137, syn. nov. Bruchus nigromaculatus Pic, 1929 (synonymy in Decelle 1961: 8) Bruchidius decoratus Decelle, 1975: 21 Conicobruchus decoratus: Kergoat et al., 2011: 756 Material examined. Type (male) of Bruchus nigromaculatus: Republic of South Africa: Port St John, v. 1923, R.E. Turner [dissected, genitalia on card in DMHF]. Other material: Botswana: 1 ♀, Lerib��, 1923 (R. Ellenberger) [MNHN]. Republic of Mozambique: 1 ♂, Nova Chupanga, v. 1928 (P. Lesne) [MNHN]. Tanzania: 2 ♂ (C. Conrads) [MNHN]. Zambia: 1 ♂, 2 ♀, Samununga, 13 �� 37.156 ���S 24 ��07.349���E, 1091m, 19.iii. 2012, ex Crotalaria aculeata claessensii, [1 ♂ 0 2012, specimen GK 446 used for DNA extraction] (B. Le Ru) [CBGP]. Body black, with large red parts; head (including antennae), thoracic sternites and legs always black, pronotum and abdomen always red. Elytra entirely black in some specimens, but usually red with more or less extensive black markings (Fig. 1): usually base, humerus, a round spot at basal fourth in interstriae 2 to 4, apical fourth to third of elytra, often also along suture. Other important morphological traits are as follows: pronotum 1.5 times wider at base than long, its sides almost straight, slightly sinuated, not constricted anteriorly into a neck; moderately lobed posteriorly and laterally, with dense white setation on lobes; strial punctures small, circular, closely spaced, interstriae finely imbricate, shining, with minute punctures; male antennae serrate. Male genitalia. Median lobe (Fig. 5) of moderate length, stout (maximum width excluding basal hood / total length = 0.15), not widened apically, basal hood moderately widened, not concave posteriorly; ventral valve large, subtriangular, bearing apically numerous sensillae and basally two lateral groups 9���12 setae; no hinge sclerite; internal sac anteriorly with minute spinules and scales, then densely lined with strands of hyaline and sclerotized spines; apical ampoule devoid of any ornamentation, gonopore not sclerotized; basal strut (Fig. 6) narrow, without keel; lateral lobes cleft to about 90 % their length; apex modified, with two lips, the dorsal one bearing about 15 setae, the ventral one densely lined with long setation. Biology. Zambian material was reared from pods of Crotalaria aculeata subsp. claessensii. This constitutes the first report on its biology. Distribution. Botswana, Democratic Republic of Congo (Decelle, 1960), Republic of Mozambique, Republic of South Africa, Tanzania and Zambia. Discussion. Conicobruchus decoratus can be distinguished from other members of the C. strangulatus species group by the shape of the pronotum and structure of male genitalia. Contrary to the other species of the group, in C. decoratus the pronotum is short, its side non-concave and it is not compressed anteriorly into a neck (Fig. 1). Male genitalia are also very distinctive, being less widened anteriorly and the basal strut not concave posteriorly (Fig. 5). Specimens with black elytra were considered as a subspecies (atrodorsalis) by Decelle (1960). However the pattern of elytral coloration is extremely variable, ranging from red with few black markings to entirely black. Because of this high level of variability we chose to establish the synonymy Bruchus decoratus atrodorsalis Decelle = Conicobruchus decoratus (F��hraeus)., Published as part of Le Ru, Bruno P., Delobel, Alex, Gy��rgy, Zolt��n, Genson, Gwena��lle & Kergoat, Gael J., 2014, Taxonomy, host-plant associations and phylogeny of African Crotalaria - feeding seed beetles (Coleoptera, Chrysomelidae, Bruchinae): the Conicobruchus strangulatus (F��hraeus) species group, pp. 238-256 in Zootaxa 3895 (2) on pages 246-247, DOI: 10.11646/zootaxa.3895.2.6, http://zenodo.org/record/226173, {"references":["Fahraeus, O. I. (1871) Coleoptera Caffrariae, family Brenthidae, Anthribidae et Bruchidae. Ofversigt af Kongliga Vetenskaps- Akademiens Forhandlingar, Stockholm, 28, 433 - 452.","Decelle, J. (1960) Bruchidae (Coleoptera: Phytophagoidea). Exploration du parc National de l'Upemba I. Mission G. F. de Witte), 59, 135 - 143. [1946 - 1949]","Pic, M. (1929) Nouveautes diverses. Melanges exotico-entomologiques, 53, 1 - 27.","Decelle, J. (1975) Les Coleopteres Bruchides d'Angola. Publicacoes Culturais Cia Diamantes Angola, 89, 13 - 32.","Kergoat, G. J., Le Ru, B. P., Genson, G., Cruaud, C., Couloux, A. & Delobel, A. (2011) Phylogenetics, species boundaries and timing of resource tracking in a highly specialized group of seed beetle (Coleoptera: Chrysomelidae: Bruchinae). Molecular Phylogenetics and Evolution, 59, 746 - 760. http: // dx. doi. org / 10.1016 / j. ympev. 2011.03.014"]}

Le Ru, Bruno P., Delobel, Alex, György, Zoltán, Genson, Gwenaëlle, Kergoat, Gael J. (2014): Taxonomy, host-plant associations and phylogeny of African Crotalaria - feeding seed beetles (Coleoptera, Chrysomelidae, Bruchinae): the Conicobruchus strangulatus (Fåhraeus) species group. Zootaxa 3895 (2): 238-256, DOI: http://dx.doi.org/10.11646/zootaxa.3895.2.6

Conicobruchus atrosuturalis (Pic, 1939) Bruchus atrosuturalis Pic, 1939: 31. Conicobruchus atrosuturalis: Decelle 1951: 181. Material examined. Paratype (male) of Bruchus atrosuturalis var. Pic: Ethiopia: ix. 1936 (genitalia on card in drop of DMHF) [MNHN] Other material: Kenya: 1 ♂, Chyulu Hills, vii. 1938 [1 ♂ 00514] Coryndon Museum /Expdt Chyulu Hills / July 38 alt. 5600, Imp. Inst./Entom [MNHN]; 1 ♂, Kakamega, 00�� 10.930 N 34 �� 56.137 E, 1693m, ex Crotalaria incana purpurescens, 0011. xi. 2008 [1 ♂ 01109] (B. Le Ru) [MNHN]; 1 ♂, 2 ♀, Magadi, 01�� 46.142 S 36 �� 20.657 E, 856m, i. 2007, ex Crotalaria uguenensis [1 ♂ 0 2307, specimen GK 35 used for DNA extraction] (B. Le Ru) [CBGP]. Body black, except elytra bright yellowish-red with black markings along suture (Fig. 1), also along apex of elytra in some specimens; ventrites dark reddish brown to black in typical form, red in paratype and Kenyan specimens reared from Crotalaria uguenensis seeds; last visible tergite red or black. Major morphological traits are as follows: pronotum 1.3 times wider at base than long, its sides straight basally, strongly convergent anteriorly, with a distinct neck; lack of white hair spots on pronotum (setation uniformly pale greyish or yellowish); elytral striae with small, closely spaced punctuation, interstriae with punctures varying from strong to shallow. Male genitalia. Median lobe (Fig. 2) similar with that of Conicobruchus strangulatus (Fig. 10), stout (maximum width excluding basal hood / total length = 0.19), slightly widened apically, basal hood moderately widened, concave posteriorly; ventral valve large, subtriangular, with apex acute, bearing numerous sensillae and two lateral groups of 5 to 8 setae; no hinge sclerite. Internal sac densely lined anteriorly with minute spinules (Fig. 2 A) and ctenoid scales (Fig. 2 B), followed by an area almost devoid of ornamentation, then dense scales and spines blending dorsally into a short series of well sclerotized small teeth (Fig. 2 C); a pair of strong ventro-lateral dented rods (Fig. 2 D) similar to those observed in Conicobruchus nodieri and C. astragalinae, and between them zero to three isolated teeth (Fig. 2 E); posteriorly a zone with a few strong, isolated teeth (Fig. 2 E) and a number of setae; then again two or more groups of dented sticks or masses (Fig. 2 F); apical ampoule (Fig. 2 G) devoid of any ornamentation, gonopore not sclerotized; basal strut narrow, without keel. Lateral lobes cleft to nearly 90 % of their length; apex modified, with two lips, the dorsal one bearing about 12-15 long setae, the ventral one with a dense group of short sensillae. Biology. Examined material was reared from pods of Crotalaria uguenensis and C. incana subsp. purpurescens in Kenya. Distribution. Ethiopia and Kenya. Discussion. Conicobruchus atrosuturalis has a very specific color pattern with a black body and bright orange elytra (Fig. 1). This pattern can only be found in red specimens of C. flabellicornis; the two species can be easily distinguished based on the presence of brushes and spots of dense white setae on the pronotum (absent in C. atrosuturalis, present in C. flabellicornis)., Published as part of Le Ru, Bruno P., Delobel, Alex, Gy��rgy, Zolt��n, Genson, Gwena��lle & Kergoat, Gael J., 2014, Taxonomy, host-plant associations and phylogeny of African Crotalaria - feeding seed beetles (Coleoptera, Chrysomelidae, Bruchinae): the Conicobruchus strangulatus (F��hraeus) species group, pp. 238-256 in Zootaxa 3895 (2) on pages 242-244, DOI: 10.11646/zootaxa.3895.2.6, http://zenodo.org/record/226173, {"references":["Pic, M. (1939) Nouveaux coleopteres, principalement phytophages, de l'Ethiopie et Somalie italienne. Memorie della Societa Entomologica Italiana, 17, 31 - 37.","Decelle, J. (1951) Contribution a l'etude des Bruchidae du Congo Belge (Col. Phytophaga). Revue de zoologie et de botanique africaines, 45, 172 - 192."]}

Conicobruchus cicatricosus (F��hraeus, 1839) Bruchus cicatricosus F��hraeus, 1839: 39. Bruchus cicatricosus pallidioripennis Pic, 1941: 12, syn. nov. Conicobruchus cicatricosus: Kergoat et al., 2011: 756 Material examined. Kenya: 2 ♂, Josa Wundanyi, 03�� 25.863 S 31 �� 21.314 E, 1378m, 13.iv. 2012, ex Crotalaria laburnifolia tenuicarpa [1 ♂ 0 2212, specimen GK 448 used for DNA extraction] (B. Le Ru) [CBGP]. Tanzania: 1 ♂, Kidevu, 03�� 09.402S 36 �� 41.058 E, 1818m, 25.ii. 2008, ex Crotalaria sp. [specimen GK 195 used for DNA extraction] (B. Le Ru) [CBGP]. Republic of South Africa: 1 ♂, 1 ♀, Port Elisabeth [1 ♂ 04908]; 1 ♀, East London, ix. 1915 (R. Ellenberger) [MNHN]. Tanzania: 1 ♂, xii 7 n fv, ex Crotalaria sp. [1 ♂ 00508] (C. Conrads) [MNHN]. Zimbabwe: 1 ♂, 1 ♀, Masvingo, 22.i. 1998 [1 ♂ 19007, specimen GK 171 used for DNA extraction] (M. Halada) [O��LM]. Body entirely black, or black with elytra and legs more or less light reddish brown (Fig. 1). The black form corresponds to variety A of F��hraeus, the paler form to his variety B, as well as to Pic���s variety pallidioripennis. These are mere colour forms of the same species. Other important morphological traits are as follows: pronotum 1.2 times wider at base than long, its sides straight basally, strongly convergent anteriorly, then constricted into a distinct neck; lack of white hair spots on pronotum (setation uniformly yellowish); elytral striae thin and narrow, interstriae alternating greyish setose spots with bare areas surrounding large ocellate punctures, elytral disc thus showing a checkered pattern, less visible laterally and posteriorly. Male antennae strongly pectinate. Male genitalia. Median lobe (Fig. 3) of moderate length, stout (maximum width excluding basal hood / total length = 0.20), slightly widened apically, basal hood moderately widened, concave posteriorly; ventral valve large, subtriangular, bearing apically numerous sensillae and basally two lateral groups of 6 to 9 setae; no hinge sclerite; internal sac densely lined anteriorly with minute spinules and ctenoid scales, followed by strands of hyaline spicules and tubercles, progressively transformed laterally into a pair of groups of very dense setae; posteriorly numerous strong teeth, associated or not in two to four dented sticks or masses; apical ampoule devoid of any ornamentation, gonopore not sclerotized; basal strut (Fig. 4) narrow, without keel; lateral lobes cleft to about 90 % their length; apex modified, with two lips, the dorsal one bearing 15-20 long setae, the ventral one densely lined with long setation. Biology. The type series was reared from Crotalaria volubilis seeds in ���Caffraria��� (now Republic of South Africa). The name C. volubilis Thunberg is not valid (ILDIS 2014; TROPICOS 2014), so that the actual host remains unknown (but C. capensis according to De Luca, 1965). Examined material was reared from pods of Crotalaria laburnifolia subsp. tenuicarpa in Kenya and of Crotalaria sp. in Tanzania. Distribution. Kenya, Republic of South Africa, Tanzania and Zimbabwe. Discussion. Conicobruchus cicatricosus can be distinguished from other members of the group by its elytral setation (Fig. 1), in which bare spots alternate with densely setose patches (in other species elytra are uniformly covered with setae). Contrary to C. decoratus and C. rubricollis, the male genitalia of C. cicatricosus includes large sclerites. However, in contrast with C. atrosuturalis, C. flabellicornis and C. strangulatus, proximal sclerites are absent., Published as part of Le Ru, Bruno P., Delobel, Alex, Gy��rgy, Zolt��n, Genson, Gwena��lle & Kergoat, Gael J., 2014, Taxonomy, host-plant associations and phylogeny of African Crotalaria - feeding seed beetles (Coleoptera, Chrysomelidae, Bruchinae): the Conicobruchus strangulatus (F��hraeus) species group, pp. 238-256 in Zootaxa 3895 (2) on pages 244-245, DOI: 10.11646/zootaxa.3895.2.6, http://zenodo.org/record/226173, {"references":["Fahraeus, O. I. (1839) In: Schonherr, C. J. (Ed.), Genera et species Curculionidum, cum synonymia hujus, familiae. Roret, Paris, Fleischer, Stockholm, pp. 1 - 495.","Pic, M. (1941) Coleopteres du globe. L'echange, 57, 13 - 16.","Kergoat, G. J., Le Ru, B. P., Genson, G., Cruaud, C., Couloux, A. & Delobel, A. (2011) Phylogenetics, species boundaries and timing of resource tracking in a highly specialized group of seed beetle (Coleoptera: Chrysomelidae: Bruchinae). Molecular Phylogenetics and Evolution, 59, 746 - 760. http: // dx. doi. org / 10.1016 / j. ympev. 2011.03.014","ILDIS (2014) International Legume Database and Information Service. Legume Web. Available from: http: // www. ildis. org (accessed 26 Nov 2014)","TROPICOS (2014) Tropicos. org. Missouri Botanical Garden. Available from: http: // www. tropicos. org (accessed 18 Aug. 2014)","De Luca, Y. (1965) Catalogue de metazoaires parasites et predateurs de Bruchides (Coleopteres). Journal of Stored Products Research, 1, 51 - 98. http: // dx. doi. org / 10.1016 / 0022 - 474 X (65) 90007 - X"]}

Key to the Conicobruchus strangulatus species group based on external morphology of the adults (see Fig. 1) 1. Pronotum short, its sides not concave, not compressed anteriorly into a neck. Mostly red, with dark markings on elytra................................................................................................ C. decoratus - Pronotum long with sides concave, produced anteriorly into a neck.............................................. 2 2 (1). Elytra with bare spots alternating with densely setose patches........................................ C. cicatricosus - Elytra uniformly covered with setae....................................................................... 3 3 (2). Pronotum bearing brushes and spots of dense white setae, quite noticeable on black background........... C. flabellicornis - Lack of white setae spots on pronotum.................................................................... 4 4 (3). Last visible tergite and elytra black or dark reddish-brown......................................... C. strangulatus - Last visible tergite red.................................................................................. 5 5 (4). Body mostly deep reddish brown, elytra mainly black, pronotum with a longitudinal line of whitish setae..... C. rubricollis - Body black, elytra bright orange with suture black............................................... C. atrosuturalis, Published as part of Le Ru, Bruno P., Delobel, Alex, Gy��rgy, Zolt��n, Genson, Gwena��lle & Kergoat, Gael J., 2014, Taxonomy, host-plant associations and phylogeny of African Crotalaria - feeding seed beetles (Coleoptera, Chrysomelidae, Bruchinae): the Conicobruchus strangulatus (F��hraeus) species group, pp. 238-256 in Zootaxa 3895 (2) on page 253, DOI: 10.11646/zootaxa.3895.2.6, http://zenodo.org/record/226173

Conicobruchus strangulatus (F��hraeus, 1839) Bruchus strangulatus F��hraeus, 1839: 4 Bruchus hargreavesi Pic, 1933: 133, syn. nov. Bruchidius hargreavesi: De Luca, 1965: 58 Conicobruchus strangulatus: Decelle, 1951: 181 Bruchidius strangulatus Prevett, 1967: 176 Material examined. Paratype (female) of Bruchus hargreavesi: Uganda: Kampala, ii. 1932, ex pods of Crotalaria [MNHN] Other material: Burkina-Faso: 1 ♂, Bobo-Dioulasso, 1.xi. 2003, light trap [1 ♂ 00714] (H. Perrin) [MNHN]. Mali: 2 ♂, 7 ♀, Bafing, vi. 2008, ex C. cf goreensis [1 ♂ 0 0 614, specimen GK 116 used for DNA extraction] (G.J. Kergoat) [MNHN]. Senegal: 18 ♂, 13 ♀, 5km N. Missira, 25.xi. 1995, ex C. glaucoides [1 ♂ 14495, 1♀ 02210] (H. & A. Delobel) [CBGP]; 4 ♂, 11 ♀, Nianing, xi. 1998, ex C. perrottetii [1 ♂ 19207, 1♀ 11307] (H. & A. Delobel) [CBGP]; 2 ♂, 2 ♀, Dakar, 15.ix. 1999, ex C. podocarpa [1 ♂ 0 2699, specimen Xh 1 used for DNA extraction] (H. & A. Delobel) [CBGP]; 1 ♂, 1 ♀, 5 km N. Missira, 11.xi. 1995, ex C. comosa (H. & A. Delobel) [CBGP]; 1 ♂, Joal - Samba Dia, 19.xi. 1995, ex C. cf goreensis [1 ♂ 14395] (H. & A. Delobel) [CBGP]. Color variable, with two distinct forms: darker specimens can be entirely black whereas lighter specimens are dark reddish brown, with antennae black. Specimens of the lighter form were formerly considered as a distinct species (hargreavesi). Other important morphological traits are as follows: pronotum 1.6���1.7 times wider at base than long; lack of white hair spots on pronotum (setation uniformly pale greyish or yellowish); strong ocellate punctation on interstriae, more or less aligned, particularly deep on elytral base in some specimens; hind femur with small acute tooth. Male genitalia. Median lobe (Fig. 10) of moderate length, stout (maximum width excluding basal hood/ total length = 0.16), strongly widened apically, basal hood moderately widened, concave posteriorly; ventral valve large, subtriangular, with apex acute, bearing numerous sensillae and two lateral groups of 5 to 10 setae; no hinge sclerite; internal sac without central column of tubercles, but with rather dense hyaline scales and tubercles, ending posteriorly in a dorsal mass of small, poorly sclerotized teeth, blending into a short series of well sclerotized teeth of variable length; a pair of strong ventro-lateral dented rods similar to those observed in Conicobruchus nodieri and C. astragalinae, varying in size and shape, from short and straight (Fig 12) to long and crescent (Fig. 13) or rod-shaped (Fig. 14), sometimes very long and articulated (Fig. 15); between these from zero to three isolated teeth; posteriorly a zone with a variable number of strong, isolated sclerotized teeth: absent in specimens from Dakar (Fig. 13) and Bafing (Fig. 15), numerous in male from Missira (Fig. 16); then again two groups of dented sticks or masses, clearly rod-shaped as in Fig. 16 to ill-defined as in male from Nianing (Fig. 17); apical ampoule devoid of any ornamentation, gonopore without sclerotization; basal strut narrow, without keel (Fig. 11); lateral lobes cleft to 85 % their length; apex modified, with two lips, the dorsal one bearing about twelve long setae, the ventral one with a dense group of short sensillae. Biology. Examined material was reared from pods of Crotalaria comosa, C. glaucoides Baker f., C. cf goreensis, C. perrottetii, C. podocarpa. Recorded from other Crotalaria species: C. calycina Schrank, C. leprieurii Guill. & Perr., C. ochroleuca G. D o n, and C. subcapitata De Wild. (Gillon et al., 1992). Crotalaria pods usually contain a large number of small seeds; Conicobruchus strangulatus larvae are external feeders, at least during the last phases of their development, because their body is much larger than the size of individual seeds; when mature, larvae weave a white cocoon inside the inflated pod, attached to its wall, and emergence occurs through a circular hole. Development of young larvae seems impossible in mature, dry seeds. Distribution. F��hraeus' types are from Senegal and Republic of South Africa; Pic���s typical series of Bruchus hargreavesi is from Uganda; material seen is from Burkina Faso, Mali, Senegal; also reported from Angola (Decelle, 1975), Congo (Decelle, 1951, 1960, Rasplus, 1989), Eritrea (Zampetti, 1988), Ethiopia (Decelle 1971), Ivory Coast (Rasplus, 1989; Gillon et al., 1992), Nigeria (Prevett, 1971), Senegal (Decelle, 1969) and Togo (Woegan et al., 1997). Discussion. Conicobruchus strangulatus cannot be confounded with the three species that have a red last visible tergite (C. atrosuturalis, C. decoratus, C. rubricollis). Differences in elytral setation easily separate C. strangulatus from C. cicatricosus (uniform in C. strangulatus, mix of bare spots and densely setose patches in C. cicatricosus). Finally it can be distinguished from C. flabellicornis based on the presence of brushes and spots of dense white setae on the pronotum (absent in C. strangulatus, present in C. flabellicornis). Conicobruchus strangulatus is the type species of genus Conicobruchus Decelle. Senegalese specimens are either of the lighter (hargreavesi) or of the darker (strangulatus) form. The fact that specimens of both forms emerged from pods of the same host-plant in the same location provides support to establish the synonymy Bruchus hargreavesi Pic = Conicobruchus strangulatus (F��hraeus). Phylogenetic analyses. Maximum likelihood analyses yield a best ML tree with a likelihood score of - 25368.23 (Fig. 18). The genus Conicobruchus is recovered monophyletic with a high support (BV of 97 %). As in the study of Kergoat et al. (2011), Bruchidius biguttatus and B. cisti appear closely related to the genus Conicobruchus (BV of 93 %). Within the genus Conicobruchus the relationships are very similar to those inferred in Kergoat et al. (2011). The six species corresponding to the Conicobruchus strangulatus species group cluster together, with a high bootstrap support (BV of 94 %). The newly sequenced species Conicobruchus indicus is found sister to C. kidevuensis, and appear unrelated to the species of the C. strangulatus species group. Another newly sequenced species, C. medaniensis is found sister to C. albopubens with a high support (BV of 100 %). Within the C. strangulatus species group a clade composed of C. decoratus and C. rubricollis is sister (BV of 94 %) to a wellsupported (BV of 89 %) clade of four species. The latter is encompasses C. cicatricosus on the one hand and a subclade that groups C. flabellicornis sister to C. atrosuturalis and C. strangulatus on the other., Published as part of Le Ru, Bruno P., Delobel, Alex, Gy��rgy, Zolt��n, Genson, Gwena��lle & Kergoat, Gael J., 2014, Taxonomy, host-plant associations and phylogeny of African Crotalaria - feeding seed beetles (Coleoptera, Chrysomelidae, Bruchinae): the Conicobruchus strangulatus (F��hraeus) species group, pp. 238-256 in Zootaxa 3895 (2) on pages 249-253, DOI: 10.11646/zootaxa.3895.2.6, http://zenodo.org/record/226173, {"references":["Fahraeus, O. I. (1839) In: Schonherr, C. J. (Ed.), Genera et species Curculionidum, cum synonymia hujus, familiae. Roret, Paris, Fleischer, Stockholm, pp. 1 - 495.","Pic, M. (1933) Nouveautes diverses. Melanges exotico-entomologiques, 61, 3 - 36.","De Luca, Y. (1965) Catalogue de metazoaires parasites et predateurs de Bruchides (Coleopteres). Journal of Stored Products Research, 1, 51 - 98. http: // dx. doi. org / 10.1016 / 0022 - 474 X (65) 90007 - X","Decelle, J. (1951) Contribution a l'etude des Bruchidae du Congo Belge (Col. Phytophaga). Revue de zoologie et de botanique africaines, 45, 172 - 192.","Prevett, P. F. (1967) Observations on the biology of six species of Bruchidae (Coleoptera) in northern Nigeria. Entomologist's. Monthly Magazine, 102, 174 - 180.","Gillon, Y., Rasplus, J. - Y. & Boughad, A. - M. (1992) Utilisation des graines de Legumineuses par un peuplement de Bruchidae et d'Anthribidae en zone de mosaique foret-savane (Lamto: Cote d'Ivoire). Journal of African Zoology, 106, 421 - 443.","Decelle, J. (1975) Les Coleopteres Bruchides d'Angola. Publicacoes Culturais Cia Diamantes Angola, 89, 13 - 32.","Decelle, J. (1960) Bruchidae (Coleoptera: Phytophagoidea). Exploration du parc National de l'Upemba I. Mission G. F. de Witte), 59, 135 - 143. [1946 - 1949]","Rasplus, J. - Y. (1989) Revision des especes afro-tropicales du genre Dinarmus (Hymenoptera: Pteromalidae). Annales de la Societe Entomologique de France (N. S.), 25, 135 - 162.","Zampetti, M. F. (1988) Notes on bruchids from East Africa Coleoptera, Bruchidae. Fragmenta Entomologica, 21, 101 - 110.","Decelle, J. (1971) Bruchides (Col.) recoltes en Abyssinie Centrale par l'expedition H. Scott et J. Omer Cooper (Sept 1926 - Jan 1927). Bulletin et Annales de la Societe Royale Entomologique de Belgique, 107, 243 - 259.","Prevett, P. F. (1971) The larvae of some Nigerian Bruchidae (Coleoptera). Transactions of the Royal Entomological Society of London, 123, 247 - 312. http: // dx. doi. org / 10.1111 / j. 1365 - 2311.1971. tb 00845. x","Decelle, J. (1969) Le Parc National du Niokolo-kobo (Senegal). Part III. XVII. Coleoptera: Bruchidae. Memoires de l'Institut Fondamental d'Afrique Noire, 84, 287 - 296.","Woegan, A. Y., Glitho, A. I., Bouchet, F. & Akpagana, K. (1997) Contribution au recensement de quelques legumineuses spontanees et subspontanees, hotes de Bruchidae en zone guineenne au Togo (Coleoptera, Chrysomeloidea). Bulletin de la Societe Entomologique de France, 3, 241 - 250.","Kergoat, G. J., Le Ru, B. P., Genson, G., Cruaud, C., Couloux, A. & Delobel, A. (2011) Phylogenetics, species boundaries and timing of resource tracking in a highly specialized group of seed beetle (Coleoptera: Chrysomelidae: Bruchinae). Molecular Phylogenetics and Evolution, 59, 746 - 760. http: // dx. doi. org / 10.1016 / j. ympev. 2011.03.014"]}

Conicobruchus rubricollis (Pic, 1903) Bruchus rubrithorax Pic, 1903: 169 Bruchus rubricollis Pic, 1913: 45 (name preoccupied) Conicobruchus rubricollis: Kergoat et al., 2011: 756 Material examined. Male paratype, Zimbabwe: Harare (with label ��� Bruchidius rubrithorax Pic ��� handwritten by J. Decelle) [MNHN]. Other material: Kenya: 2 ♂, Thange Town, 02�� 34.738 ���N 37 �� 57.129 ���E, 904m, 24.iv. 2011, ex Crotalaria cf. polysperma [1 ♂ 0 3211, specimens GK 406 and GK 440 used for DNA extraction] (B. Le Ru) [MNHN]; 1 ♂, SE Kenya, Voi (Tsavo), 23.3.- 4.4.1997; 2 ♂, Kenya mer., Tsavo East, Buchuma env., 28.xi. 1997 [specimen GK 196 used for DNA extraction] (M. Snizek) [O��LM]; 1 ♀, Kenya, S. Taita, Mwatate, 30.xi. 1997 (M. Snizek) [O��LM]. Zimbabwe: 1 ♂, Masvingo, 22.i. 1998 [1 ♂ 18907] (M. Halada) [O��LM]. Large species well defined by its mainly red colour, with major part of antennae, elytra, and often disc of mesothoracic sternite, black (Fig. 1); pubescence white, denser on median longitudinal line of pronotum, scutellum, base of elytral suture; pygidium with two lateral spots near base and a longitudinal line. Other major morphological traits are as follows: pronotum 1.5 times wider at base than long, without oblique lateral impression, its sides concave, produced anteriorly into a neck; elytral striae narrow and deep, with small punctures, interstriae wide and flat, with small punctures on shagreened background; antennae serrate. Male genitalia. Median lobe (Fig. 8) of moderate length (maximum width excluding basal hood / total length = 0.23), strongly widened and sclerotized apically; basal hood oval, apically indented; ventral valve large, subtriangular, with apex acute, with numerous sensillae, bearing two lateral groups of 10���12 setae; dorsal valve unsclerotized; no hinge sclerites; anterior part of the internal sac lined with hyaline spinules becoming denser and larger distally, then large ctenoid scales, posterior saccus with numerous sclerotized teeth; apical ampoule unarmed. Tegminal strut (Fig. 9) without keel, triangularly pointed apically, lateral lobes cleft to base; apex of parameres with numerous setae, slightly modified, with dorsal velum. Biology. Specimens from Thange were reared from a sample of Crotalaria sp. near polysperma pods. This constitutes the first report on its biology. Distribution. Kenya and Zimbabwe. Discussion. The red color of the last visible tergite distinguishes Conicobruchus rubricollis from C. cicatricosus, C. flabellicornis and C. strangulatus (Fig. 1). It cannot be confounded with C. decoratus based on differences on the shape of the pronotum. Finally the presence of a longitudinal line of whitish setae on the pronotum separates C. rubricollis from dark specimens of C. atrosuturalis. Male genitalia (Fig. 8) are similar in shape with other members of the group (except C. decoratus) but completely lack large sclerites., Published as part of Le Ru, Bruno P., Delobel, Alex, Gy��rgy, Zolt��n, Genson, Gwena��lle & Kergoat, Gael J., 2014, Taxonomy, host-plant associations and phylogeny of African Crotalaria - feeding seed beetles (Coleoptera, Chrysomelidae, Bruchinae): the Conicobruchus strangulatus (F��hraeus) species group, pp. 238-256 in Zootaxa 3895 (2) on page 249, DOI: 10.11646/zootaxa.3895.2.6, http://zenodo.org/record/226173, {"references":["Pic, M. (1903) Coleopteres de l'Afrique australe. Revue d'Entomologie, 22, 165 - 171.","Pic, M. (1913) Bruchidae, In: Junk, W. (Ed.), Coleopterorum Catalogus. Part 55. Berlin, pp. 1 - 74.","Kergoat, G. J., Le Ru, B. P., Genson, G., Cruaud, C., Couloux, A. & Delobel, A. (2011) Phylogenetics, species boundaries and timing of resource tracking in a highly specialized group of seed beetle (Coleoptera: Chrysomelidae: Bruchinae). Molecular Phylogenetics and Evolution, 59, 746 - 760. http: // dx. doi. org / 10.1016 / j. ympev. 2011.03.014"]}

Busseola fusca (Lepidoptera: Noctuidae) is an important pest of maize and sorghum in sub-Saharan Africa. One century after its first description by Fuller in 1901, inaccurate information based on earlier reports are still propagated on its distribution (e.g., absent from the lower altitudes in East Africa) and host plant range (e.g., feeding on a large range of wild grass species). This review provides updated information on the biology, distribution and genetics of B. fusca with emphasis on insect-plant interactions. Related to this, new avenues of stem borer management are proposed.

Transgenic crops expressing Bacillus thuringiensis (Bt) toxins have been adopted worldwide, notably in developing countries. In spite of their success in controlling target pests while allowing a substantial reduction of insecticide use, the sustainable control of these pest populations is threatened by the evolution of resistance. The implementation of the “high dose/refuge” strategy for managing insect resistance in transgenic crops aims at delaying the evolution of resistance to Bt crops in pest populations by promoting survival of susceptible insects. However, a crucial condition for the “high dose/refuge” strategy to be efficient is that the inheritance of resistance should be functionally recessive. Busseola fusca developed high levels of resistance to the Bt toxin Cry 1Ab expressed in Bt corn in South Africa. To test whether the inheritance of B. fusca resistance to the Bt toxin could be considered recessive we performed controlled crosses with this pest and evaluated its survival on Bt and non-Bt corn. Results show that resistance of B. fusca to Bt corn is dominant, which refutes the hypothesis of recessive inheritance. Survival on Bt corn was not lower than on non-Bt corn for both resistant larvae and the F1 progeny from resistant × susceptible parents. Hence, resistance management strategies of B. fusca to Bt corn must address non-recessive resistance Biosafety South Africa (Grant 08-001) doi:10.1371/journal.pone.0069675

International audience

The noctuid Busseola fusca is the most destructive pest of maize in Cameroon. Various habitat management technologies have been developed and tested but none of them provides a self-sustainable solution. The braconid Cotesia sesamiae, the most common larval endoparasitoid of the B. fusca in eastern Africa, is rare in West Africa and Cameroon. In Cameroon, it is mostly found on non-crop borer species feeding on wild grasses. Thus, two Kenyan C. sesamiae populations and their hybrid were introduced into Cameroon. Suitability studies were carried out with two B. fusca populations, the noctuid Poeonoma serrata, which is commonly found feeding on Napier grass but not maize, and the noctuid Sesamiae cretica, which was recently accidentally introduced into Mauritania, where it attacks sorghum. The performance of the C. sesamiae was influenced by both host species/population and parasitoid population, although the B. fusca population had a minor effect. The reproductive potential of the hybrid was superior over that of its parent populations. The least suitable host was P. serrata, thus it was concluded that it would probably not play a major role in perennating C. sesamiae during the dry season. With S. cretica as host, parasitism was similar but brood size was lower than with either B. fusca population. Release strategies of C. sesamiae and its chances of establishment on B. fusca and S. cretica in the hothumid forest zone and the cool highland of Cameroon and in Mauritania are discussed.

International audience; As a result of an intense host-parasite evolutionary arms race, parasitic wasps frequently display high levels of specialization on very few host species. For instance, in braconid wasps very few generalist species have been described. However, within this family, Cotesia sesamiae is a generalist species that is widespread in sub-Saharan Africa and develops on several lepidopteran hosts. In this study, we tested the hypothesis that C. sesamiae may be a cryptic specialist when examined at the intraspecific level. We sequenced exon 2 of CrV1, a gene of the symbiotic polyDNAvirus that is integrated into the wasp genome and is associated with host immune suppression. We found that CrV1 genotype was more closely associated with the host in which the parasitoid developed than any abiotic environmental factor tested. We also tested a correlation between CrV1 genotype and an infection with Wolbachia bacteria, which are known for their ability to induce reproductive isolation. The Wolbachia bacteria infection polymorphism was also found as a major factor explaining the genetic structure of CrV1, and, in addition, the best model explaining CrV1 genetic structure involved an interaction between Wolbachia infection and host species. We suggest that Wolbachia could act as an agent capable of maintaining advantageous alleles for host specialization in different populations of C. sesamiae. This mechanism could be applicable to other insect models because of the high prevalence of Wolbachia in insects.