3032: 17 32 (2011) Article ZOOTAXA ...sharkeylab.org/sharkeylab/docs/posts/web/Stein_etal_2011_Epyris... · MCSN Museo Civico di Storia Naturale “Giacomo Doria”, Italy (Roberto

Accepted by A.S. Lelej: 22 Aug. 2011; published: 19 Sep. 2011

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Zootaxa 3032: 17–32 (2011) www.mapress.com/zootaxa/ Article

17

Revision of Epyris afer (Magretti, 1884), comb. rev. (Hymenoptera: Bethylidae) with new synonymy and description of two new species

PAULO R.W. STEIN1,3, ISABEL D.C.C. ALENCAR2, DIEGO N. BARBOSA1 & CELSO O. AZEVEDO1

1Universidade Federal do Espírito Santo, Departamento de Biologia, Av. Marechal Campos 1468, Maruípe, 29.040-090 Vitória ES, Brazil. E-mail: [email protected]; [email protected]; [email protected] Federal de Educação, Ciência e Tecnologia do Espírito Santo – IFES, Campus Santa Teresa, Rodovia ES 080, Km 21, São João de Petrópolis, 29660-000, Santa Teresa, ES, Brazil. E-mail: [email protected] author. E-mail: [email protected]

Abstract

Pristocera afra Magretti, 1884 which currently belongs to genus Epyris Westwood is revised and redescribed. The lecto-type of P. afra Magretti is designated. New synonymy is proposed for Epyris afer (Magretti, 1884): Epyris pilosipes Ki-effer, 1904, syn. nov.; Epyris analis Kieffer, 1905, syn. nov.; Epyris secundus Brues, 1906, syn. nov.; Epyris rugicollisCameron, 1906, syn. nov.; Epyris plurilineata Turner, 1928, syn. nov. Two new species Epyris enerterus Stein & Azeve-do, sp. nov. (Myanmar) and Epyris penatii Stein & Azevedo, sp. nov. (Sudan) are described and illustrated. Epyris afer(Magretti) is newly recorded from Yemen, United Arab Emirates, Thailand, and Vietnam.

Key words: Epyrinae, Disepyris, Lytepyris, Pristocera, new synonymy, lectotype designation

Introduction

Magretti (1884) described Pristocera afra based on five syntypes collected on flowers of Asclepiadoideae fromSudan and Ethiopia. According to him, the main diagnostic characters to identify this species were the five propo-deal discal carinae, the metasoma black basally and reddish apically and the legs brown-reddish. In Magretti’swork there is no evidence of comparison between this species with similar species, nor any character that justifiesits classification as Pristocera Klug.

Magretti (1897) observed, among the bethylids from Myanmar collected by Leonardo Fea, specimens that fithis concept of P. afra. In that study, he analyzed the concept of this species and transferred it to Epyris Westwood.Magretti op. cit. reported the similarity of this species with E. piceiventris Westwood, 1874 (presently in Rhab-depyris Kieffer) and E. lathrobioides Westwood, 1874 (presently in Apenesia Westwood), both Australian species.

Kieffer (1913) created the genus Lytepyris Kieffer, which had as its main diagnostic character, the radial veinshorter than basal vein. The species E. afer and Trachepyris biscrensis Kieffer & Marshall, 1906 were included inthis genus. Kieffer (1914) designated L. biscrensis as type-species of the genus.

While studying the type-species of Lytepyris Kieffer and Disepyris Kieffer, Terayama (2004) concluded theyhad no significant generic differences and proposed the former as junior synonym of the latter. He also transferredthe two Lytepyris species to Disepyris.

According to Gordh and Móczár (1990) the type depository of Disepyris afer is dubious. However, recently,the syntypes series and some additional specimens of this taxon were found in the Museo Civico di Storia Naturale“Giacomo Doria” (MCSN), Genoa (Italy), by Fabio Penati.

Our research show that none of the specimens of Disepyris afer belong to Disepyris but rather to Epyris, basedon the scutellar pits are not connected by any groove, the setae of the protarsomeres are not stout, the eyes are notlarge, and the stigma is not large. We observed a wide range of structural variation representing multiple specieswithin the material identified as D. afer. Therefore, the aim of this study is to review the taxonomic identity of D.afer, designate a lectotype, propose new synonyms, and describe new species previously confused under this name.

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STEIN ET AL.18 · Zootaxa 3032 © 2011 Magnolia Press

Material and methods

Material was provided by the following institutions:

MCSN Museo Civico di Storia Naturale “Giacomo Doria”, Italy (Roberto Poggi), BMNH The Natural History Museum, United Kingdon (David Notton), CNCI Canadian National Collection of Insects, Canada (John Huber), QSBG Queen Sirikit Botanical Garden, Thailand (Wichai Srisuka), the Thai material was collected under the

scope of “Thailand Inventory Group for Entomological Research” project (grant #DEB-0542864)coordinated by Michael Sharkey from University of Kentucky (U.S.A.),

RMNH Rijksmuseum van Natuurlijke Historie, Netherlands (Guido Keijl),UFES Universidade Federal do Espírito Santo, Brazil (Marcelo T. Tavares),MNHN Muséum National d’Histoire Naturelle, Paris (Claire Villemant).

The specimens studied here were compared with species of genus Epyris. We analyzed 172 species of Epyris, 117of which were interpreted by direct observation of specimens, photographs or illustration of holotypes, another 55species were compared by descriptions present in the literature.

The terms of body structures, measurements and indices used in this study follows Evans (1964) and Azevedo(1999). The nomenclature of leg surface was interpreted as in Aguiar and Gibson (2010). Those related to integu-ment sculpture follows Harris (1979).

Abbreviations and measurements used in this study are as follows: body length is measured: from the apex ofclypeus to the posterior margin of the last metasomal segment, excluding the male genitalia or the female sting;(LH) length of head in frontal view: from the vertex crest to the median apical margin of clypeus; (WH) width ofhead in frontal view: maximum width including eyes; (WF) width of frons in frontal view: minimum width usuallyat a virtual line that crosses the anterior margin of compound eyes; (HE) height of eye in lateral view: across itsmaximum length; (OOL) ocellar-ocular line in laterodorsal view: the shortest distance from posterior margin ofcompound eye to posterior ocellus; (WOT) width of ocellar triangle in frontal view: maximum width, includingocelli; (DAO) diameter of anterior ocellus in frontal view; (VOL) vertex-ocular line in dorsal view: distance fromeye top to vertex crest; (LFW) length of forewing in dorsal view.

The information included in the material examined section strictly follows those contained in the label of spec-imen analyzed. The complementary information is given in square brackets, the updated and corrected informationis provided in curly brackets.

The descriptions were performed with DELTA (Descriptive Language for Taxonomy) developed by Dallwitz(1980) and Dallwitz et al. (1999). The photographs were obtained through the Entovision® system and imageswere combined by Helicon Focus®.

Taxonomy

Epyris afer (Magretti, 1884), comb. rev.(Figs .1, 4, 7–12, 19, 20, 23, 24, 27, 29, 31–33)

Pristocera afra Magretti 1884: 533–534, pl. 1 fig. 1, ♀ ♂, lectotype (designated here), ♀, examined; Dalla Torre 1898: 561(catalog); Gordh & Móczár 1990: 133 (catalog).

Epyris afer: Magretti 1897: 319 (part.); Kieffer 1904: 396 (key); Kieffer 1908: 27 (catalog). Lytepyris afer: Kieffer 1913: 108; Kieffer 1914: 416–417 (key, redescription); Gordh & Móczár 1990: 133 (catalog).Disepyris afer: Terayama 2004: 97–98.Epyris pilosipes Kieffer 1904: 396, 404–405 (key, description), ♀ (syntype examined); Kieffer 1908: 28 (catalog); Kieffer

1914: 310, 330, fig. 140 (key, redescription); Kurian 1954: 269 (catalog); Richards 1955: 357, 358 (distribution); Gordh &Móczár 1990: 101 (catalog). Syn. nov.

Epyris analis Kieffer 1905: 111-112, ♀ (holotype examined), nom. praeocc., non Cresson 1872: 193. Syn. nov.Epyris secundus Brues 1906: 143 (new name for E. analis Kieffer 1905: 111-112); Kieffer 1908: 27 (catalog); Kieffer 1914:

312, 326-327 (key, redescription); Kurian 1954: 270 (catalog); Kurian 1955: 90 (key). Syn. nov.Epyris rugicollis Cameron 1906: 288, ♀ (holotype examined); Gordh & Móczár 1990: 127 (catalog). Syn. nov.


Zootaxa 3032 © 2011 Magnolia Press · 19REVISION OF EPYRIS AFER (MAGRETTI, 1884), COMB. REV.

Isobrachium rugicolle: Kieffer 1908: 26 (catalog); Kieffer 1914: 368, 369 (key, redescription); Kurian 1954: 273 (catalog). Isobrachium rugicollis: Gordh & Móczár 1990: 127 (catalog). Epyris plurilineata Turner 1928: 130–131, ♀ (holotype examined); Gordh & Móczár 1990: 102 (catalog). Syn. nov.Epyris plurilineatus: Benoit 1957: 14 (distribution, variation); Gordh & Móczár 1990: 102 (catalog).

Type material examined. LECTOTYPE of Pristocera afra (designated here): ♀, [ETHIOPIA], Metemma,[13.026817, 36.161562], 24.III.1883, P. Magretti col. (MCSN). PARALECTOTYPES of Pristocera afra: [ETHIO-PIA], 2 ♀, Metemma, [13.026817, 36.161562], 24.III.1883, P. Magretti col. (MCSN); 1 ♂, 22.III.1883 (MCSN). 1♀, Bahr el Salaam, rive {= river banks}, [13.85, 36.1333333], 14.III.1883, P. Magretti col (MCSN). SYNTYPE ofEpyris pilosipes: ♀, GHINEA PORTOGHESE {GUINEA-BISSAU}, Bolama, [11.578581, -15.483679], VI–XII.1899, L. Fea col. (MCSN); HOLOTYPE of Epyris plurilineata: ♀, S[OUTHERN] RODESIA {ZIMBABWE},Thabasilitche, {coordinates not found, coordinates of Bulawayo, another occurrence of the species = -20.17,28.58}, 6.VII.1924, R.H.R. Stevenson col. Brit[ish] Mus[eum] 1927-344, B.M.Type Hym. 13302 (BMNH);HOLOTYPE of Epyris analis: ♀, INDES [INDIA], Mahi {=Chennai} [13.083333, 80.283333], Ernest André col-lection 1914, Muséum Paris EY6341 (MNHN). HOLOTYPE of Isobrachium rugicollis: ♀, [PAKISTAN, Bal-ochistan province] Quetta, [30.1872, 67.0125], 6.02.{= VI.1902 ?}, Nurse col. 1915-34 (BMNH).

Afrotropical material (new). UNI[TED] ARAB EMIRATES, 3 ♀, Wadi Wurayah, 25.24N, 56.17E, 12–14.IV.2005, M[alaise] T[rap] & Y[ellow] P[an] T[rap], T. P[ape] col. (UFES). 2 ♀, Sharjah, 25.21N, 55.42E,27.IV–05.V.2005, light tr[ap], A. v[an] Harten col. (UFES no. 30424, no. 30425). 13 ♀, Bithnah, 25.11N, 56.14E,04.VII–12.VII.2006, M[alaise] T[rap], A. van Harten col. (UFES no. 30483); 2 ♀, 12.VIII–09.IX.2006 (UFES no.30422); 2 ♀, 19.X–16.XI.2006 (UFES). 3 ♀, Al Ajban, 24.36N, 55.01E, 26.VI–25.VII.2006, M[alaise] T[rap], A.van Harten col. (UFES no. 30423, no. 30593). YEMEN, 1 ♀, Se[i]yun, [15.94092, 48.780134], VI.2002, Lighttrap, A. van Harten & G. Ba Saheh col. (CNCI). 2 ♀, Al Lahima, [15.4, 43.5333333], 09.IV–05.VI.2001, M[alaise]T[rap], A. van Harten col. (CNCI). 1 ♀ Lahj, [13.066666, 44.883333], IX.2000, M[alaise] T[rap], A. van Harten &A. Sallam col. (CNCI); 1 ♀, XI.2000 (CNCI); 3 ♀, IV–V.2001 (CNCI); 3 ♀, 17.V–15.VI.2001 (CNCI); 2 ♀,07.IX.2001 (CNCI); 2 ♀, 3.V.2002 (CNCI).

Oriental material (new). [MYANMAR], 1 ♀, Bhamò, [24.2667, 97.2333], VIII.1886, L. Fea col. (MCSN). 1♀, Palon (Pegú), [17.4333, 95.9], VII–IX.1887, L. Fea col (MCSN). THAILAND, 1 ♀, Mae Hong Son, NamtokMae Surin N[ational] P[ark], Walkway top of reservoir, 19°20.893'N, 97°59.005'E, 3–10.II.2008, M[alaise] T[rap],Na-maadkam & M col. T3485 (QSBG). 2 ♀, Loei, Phu Ruea N[ational] P[ark], Nern Pitsawong, 17°29.676'N,101°21.093'E, 1168 m, 19–26.XI.2006, M[alaise] T[rap], Patikhom Tumtip col. T1123 (QSBG). 1 ♀, Phu Kra-dueng N[ational] P[ark], Road to Ta Krong waterfall of Na Noy Forest Unit, 16°48.913'N, 101°47.634'E, 265 m,14–20.XI.2006, M[alaise] T[rap], Suthin Gong-lasae col. T1073 (QSBG). 1 ♀, Ubon Ratchathani, Pha TaemN[ational] P[ark], Don Huay Can, 15°40.016'N, 105°30.502'E, 246 m, 9–15.VI.2007, M[alaise] T[rap], Tongcam& Banlu col. T2208 (QSBG); 2 ♀, Saeng Chan Waterfall, 15°31.985'N, 105°35.774'E, 155 m, 12–20.III.2007,M[alaise] T[rap], Porntip Tonsu & Bunlu Sapsiri col. T2146 (QSBG). 1 ♀, Chaiyaphum, Pa Hin Ngam N[ational]P[ark], car park at Tung Dok Grajeaw, 15°38.438'N, 101°23.576'E, 780 m, 12–18.VII.2006, M[alaise] T[rap], Kra-tae Sa-nog & Buakaw Adnafai col. T333 (QSBG); 2 ♀, creek at Tung Dok Grajeaw, 15°38.391'N, 101°23.609'E,750 m, 24–30.VII.2006, M[alaise] T[rap], Kratae Sa-nog & Buakaw Adnafai col. T338 (QSBG); 1 ♀, ecotonebetween mix deciduous and dipterocarp forest, 15°38.132'N, 101°23.922'E, 698 m, 19–25.II.2007, M[alaise]T[rap], Katae Sa-nog & Buakaw Adnafai col. T1652 (QSBG). 1 ♀, Kanchanaburi, Khuean SrinagarindraN[ational] P[ark], Tourist center, 14°38.136'N, 98°59.837'E, 210 m, 11–18.VII.2008, M[alaise] T[rap], Somboon &Chatchawan col. T3432 (QSBG). 1 ♀, Nakhon Nayok, Khao Yai N[ational] P[ark], Nhong ping khaokeaw,14°23.094'N, 101°23.055'E, 733 m, 12–19.III.2007, M[alaise] T[rap], Wirat Sukho col. T2104 (QSBG); 1 ♀,26.III–2.IV.2007, Pong Sandao col. T2109 (QSBG). 1 ♀, Chanthaburi, Khao Khitchakut N[ational] P[ark], 100 mN/Prabaht Unit, 12°48.842'N, 102°9.144'E, 203 m, 7–14.VII.2008, M[alaise] T[rap], Suthida & Charoenchai col.T2973 (QSBG). 5 ♀, Prachuab Khiri Khan, Khao Sam Roi Yot N[ational] P[ark], Saline wetland/Pa Gwad/N,12°9.2'N, 99°58.298'E, 22–29.III.2009, M[alaise] T[rap], Yai & Amnad col. T4219 (QSBG); 2 ♀, Saline wetland/Pa Gwad/S, 12°9.173'N, 99°58.244'E, T4221 (QSBG); 2 ♀, Nursery, 12°7.58'N, 99°57.478'E, 29.VI–6.VII.2008,M[alaise] T[rap], Amnad & Yai col. T3049 (QSBG); 1 ♀, 30.VI–1.VII.2008, Pan trap, T3031 (QSBG); VIET-NAM, 1 ♀, Dak Lak, Chu Yang Sin N[ational] P[ark], n[ea]r dam, [12.46277, 108.409464], c. 500 m, 3–9.VI.2007,M[alaise] T[rap], C. v. Achterberg & R. de Vries col. RMNH’07 (RMNH). 1 ♀, Ninh Thuân, Núi Chiúa N[ational]P[ark], northeast part, [11.715754, 109.138353], M[alaise] T[rap], 90–150 m, 23–30.V.2007, C. v. Achterberg & R.



de Vries col. RMNH’07 (RMNH). 1 ♀, Dông Nai, Cát Tiên N[ational] P[ark], Bot[anical] garden, [11.470501,107.259089], c. 100 m, 13–20.V.2005, M[alaise] T[rap] 14-19, C. v. Achterberg & R. de Vries col. RMNH’05(RMNH). 2 ♀, Cát Tiên N[ational] P[ark], Dong trail, [11.470501, 107.259089], c. 100 m, 9.IV–19.V.2007,M[alaise] T[rap], M. P. Quy, N. T. Manh & C. v. Achterberg col. RMNH’07 (RMNH). 1 ♀, Cát Tiên N[ational]P[ark], Ficus trail, [11.470501, 107.259089], c. 100 m, 9–30.IV.2007, M[alaise] T[rap], M. P. Quy & N. T. Manhcol. RMNH’07 (RMNH).

Type condition. The lectotype and the three female paralectotypes are in good condition. However the onlyparalectotype male is broken in several parts, but none is missing.

Distribution. Democratic Republic of the Congo, Ethiopia, Guinea-Bissau, India, Israel, Myanmar, Pakistan,South Africa, Sudan and Zimbabwe. It is recorded for the first time from Thailand, United Arab Emirates, Vietnam,and Yemen.

Diagnosis. FEMALE. Mandible with pre-apical lower tooth, sensillae chaetica in lower margin of mandiblepresent. Apex of clypeus with long setae. Head as wide as long. Space between inner and second pair of propodealdiscal carina with a structure resembling a fovea. Metatibia with dense series of short setae in posterior face. Meta-soma dark castaneous nearly black with apex orange.

Redescription. LECTOTYPE. Body length 7.5 mm. LFW 3.8 mm.Color. Head, mesosoma and procoxa dark castaneous nearly black. Metasoma dark castaneous nearly black

with apex orange. Meso and metacoxae and femora dark castaneous. Scape, pedicel, flagellum, mandible, tegula,wing venation, trochanters and tibiae castaneous. Palpi light castaneous. Wings subhyaline.

Head (Fig. 1). Mandible (Figs. 7–9) apex as wide as base, with five rounded apical teeth, two lower wider thanothers, pre-apical lower tooth present; sensillae chaetica in lower margin of mandible present. Clypeus shorter thanwide, median lobe angulate; median carina present; apex of clypeus with long setae; lateral lobe reduced. Transver-sal section of scape cylindrical; first four antennal segments ratio 23:7:7:9, segment III as long as thick; antennalsensillae not visible. Antennal scrobe projected forward, dorsally sulcate. Toruli distant from each other 1.7 × theirdiameter. Eye almost reaching upper mandibular condyle, prominent, glabrous. Frons coriaceous; punctures smalland shallow, separated from each other by 1.5 × their diameter. WH 1.0 × LH; WF 1.3 × HE; WF 0.6 × WH; OOL1.7 × WOT; VOL 0.6 × HE; distance of posterior ocellus to vertex crest 1.0 × DAO. Ocelli small, frontal angle ofocellar triangle acute. Temple parallel in dorsal view. Vertex almost straight in dorsal view, with soft callositymedially. Crest with continuous series of short setae. Occipital carina present dorsally and ventrally. Gena flat-tened. Hypostomal carina straight and polished.

Mesosoma (Fig. 19). Pronotal collar polished, with transversal groove. Pronotal disc 0.9 × longer than wide,coriaceous, subtrapezoidal, sparsely punctate, transversal carina in anterior margin absent. Mesoscutum coria-ceous, lateral slightly elevate posteriorly. Notaulus touching posterior margin of mesoscutum, convergent posteri-orly, gradual anteroposterior enlargement. Parapsidal furrow parallel, shorter and as deep as notaulus, not reachingposterior margin of mesoscutum. Scutellum coriaceous, scutellar groove absent, scutellar pit oval. Propodeal disc0.9 × as long as wide; anterior carina medially narrower than laterally; anterior corner fovea-shaped; five discalcarinae present, median carina complete, inner discal carina complete, space between median and inner discalcarina with longitudinal ridge, second pair of discal carina incomplete and short, space between inner and secondpair of discal carina with structure resembling fovea; lateral margin straight; sublateral carina absent, lateral carinapresent, space between sublateral and lateral carina with series of transversal ridges; posterior margin somewhatconcave; posterior corner with suboval fovea; lateral of propodeum strigate; declivity of propodeum strigate, withmedian carina. Mesopleuron (Fig. 20) coriaceous; subtegular groove elongate, large anteriorly and narrow posteri-orly; anterior fovea closed; sub-anterior fovea closed and small; mesopleural fovea closed; mesopleural pit present;mesopleural elevation present; lower fovea open, groove of inferior margin of lower fovea present throughoutextension of margin; episternal furrow with inner margin striate. Epicnemium lateromedially enlarged, with medialconstriction. Pleurosternum with straight longitudinal groove.

Legs. Profemur 2.4 × longer than wide. Mesotibia spinose. Metatibia with dense series of short setae in poste-rior face. Tarsomeres distally spinose, spines of protarsus short and cylindrical, spines of meso and metatarsi longand flattened. Tarsal claws bidentate, inner tooth as long as apical tooth.

Wing. Forewing with metacarpus present, 0.2 × as long as stigma length; radial vein curved forward, 2.6 × lon-ger than basal; basal vein truncate; transverse median vein convex. Hind wing with two basal and four apical ham-uli.



Metasoma. Ventral surface of petiole (Fig. 27) unsegmented, space between carinae of petiole wide. Tergum IIlonger than others; terga III–VI with line of transverse setae on dorsal surface.

Variation. Hind wing with three basal and five apical hamuli.Diagnosis. MALE. Posterior ocellus distant from vertex crest 0.3 × DAO. Mesoscutum and scutellum pol-

ished. Mesotibia without spines. Protarsomeres without spines. Hind wing with five medial hamuli. Paramerestrongly arched ventrad, narrow and almost 3.0 × longer than basiparamere. Aedeagus inner margin slightlyexpanded subapically. Apodeme base projected laterally, not dilated.

Redescription. PARALECTOTYPE. Body length 4.0 mm. LFW 2.2 mm.Color. Head, mesosoma and coxae dark castaneous nearly black. Metasoma dark castaneous nearly black with

apex castaneous. Wing venation, trochanters and femora dark castaneous. Scape, pedicel, flagellum, mandible, teg-ula and tibiae castaneous. Palpi and tarsi light castaneous. Wings subhyaline.

Head (Fig. 4). Mandible (Figs. 10–12) apex as wide as base, with five rounded apical teeth, two lower widerthan others, pre-apical lower tooth present; sensillae chaetica in lower margin of mandible present. Clypeus shorterthan wide, median lobe angulate; median carina present; apex of clypeus with long setae; lateral lobe reduced.Transversal section of scape elliptical; first four antennal segments ratio 18:5:4:7, segment III 0.6 × longer thanthick; antennal sensillae visible in ventral surface of antenna. Antennal scrobe projected forward, not dorsally cari-nate. Toruli distant from each other 1.5 × their diameter. Eye almost reaching upper mandibular condyle, promi-nent, glabrous. Frons coriaceous; punctures small and shallow, separated from each other by 3.0 × their diameter.WH 0.9 × LH; WF 1.5 × HE; WF 0.7 × WH; OOL 1.9 × WOT; VOL 1.0 × HE; distance of posterior ocellus to ver-tex crest 0.3 × DAO. Ocelli small, frontal angle of ocellar triangle acute. Temple parallel in dorsal view. Vertexstraight in dorsal view. Crest without setae. Occipital carina present dorsally and ventrally. Gena flattened. Hypos-tomal carina straight and polished.

Mesosoma (Fig. 23). Pronotal collar polished, without transversal groove. Pronotal disc 0.6 × longer than wide,coriaceous, trapezoidal, sparsely punctate, transversal carina in anterior margin present. Mesoscutum polished, lat-eral slightly elevate posteriorly. Notaulus touching posterior margin of mesoscutum, convergent posteriorly, grad-ual anteroposterior enlargement. Parapsidal furrow convergent posteriorly, shorter and as deep as notaulus, notreaching posterior margin of mesoscutum. Scutellum polished, scutellar groove absent, scutellar pit circular. Prop-odeal disc 0.5 × as long as wide; anterior carina medially narrower than laterally; anterior corner groove-shaped;five discal carinae present, median carina complete, inner discal carina incomplete, space between median andinner discal carina without sculpture, second pair of discal carina complete, space between inner and second pair ofdiscal carina without ridge; lateral margin sinuous; sublateral carina present and weak, lateral carina present, spacebetween sublateral and lateral carina with series of transversal ridges; posterior margin somewhat concave; poste-rior corner with suboval fovea; lateral of propodeum strigate; declivity of propodeum areolate, with median carina.Mesopleuron (Fig. 24) coriaceous; subtegular groove elongate, large anteriorly and narrow posteriorly; anteriorfovea closed and large; sub-anterior fovea closed and large; mesopleural fovea open; mesopleural pit present;mesopleural elevation present; lower fovea open, groove of inferior margin of lower fovea present only in anteriorportion of margin; episternal furrow with inner margin striate. Epicnemium lateromedially enlarged. Pleurosternumwith lozenge-shaped longitudinal groove.

Legs. Profemur 2.8 × longer than wide. Mesotibia without spines. Metatibia with sparse series of short setae inposterior face. Protarsomeres without spines. Meso and metatarsomeres distally spinose, spines long and flattened.Tarsal claws bidentate, inner tooth as long as apical tooth.

Wing. Forewing with metacarpus present, 0.7 × as long as stigma length; radial vein curved forward, 3.0 × lon-ger than basal; basal vein slightly concave; transverse median vein convex. Hind wing with five medial hamuli.

Metasoma. Ventral surface of petiole (Fig. 29) unsegmented, space between carinae of petiole narrow. TergumII as long as others; terga III–VI with sparse setae on dorsal surface.

Genitalia (Figs 31–33). Paramere positioned dorsally, arched strongly ventrad, narrow, very elongate, almost3.0 × longer than basiparamere, apical margin convex, membranous expansion large and stout; cuspis elongate, itsapex surpassing apical half of paramere length, divided into two arms only at apex, its ventral base with roundedprojection, ventral arm slightly longer and narrower than dorsal arm; digitus with apical margin denticulate, itsapex not reaching cuspis apex; aedeagus bottle-shaped, short, its apex not reaching cuspis length, apex with con-cavity inside, inner margin slightly expanded subapically; inner lobe membranous; apodeme not extending beyondgenital ring, base projected lateral and not dilated.



Discussion. This species does not belong to Disepyris because the scutellar pits are not connected by anygroove, the setae of the protarsomeres are not stout, the eyes are not large, and the stigma is not large. Besides, ithas the pronotal disc ecarinate, wider posteriorly and the anterior corners not acutely angled, the mesoscutum hasnotauli and the antennae are not spinose. This combination of characteristics corresponds to Epyris and because ofthat we revalidated the combination to Epyris as proposed by Magretti (1897).

The lectotype designation was necessary because there were three different species in the type series. The spec-imen selected represents Magretti’s original idea for the name of E. afer.

Three species are considered junior subjective synonyms of E. afer, namely E. pilosipes, E. plurilineatus andE. analis (=secundus) because they have the mandible with five rounded apical teeth, the two lower ones widerthan the upper ones; the presence of long setae in the apex of median clypeal lobe; the presence of occipital carinaventrally, and the propodeal anterior corner foveolate.

Another species, Isobrachium rugicollis is also considered as junior subjective synonym of E. afer. It wasdescribed in Epyris by Cameron (1906) based on one single female. However Kieffer (1908) transferred it to Iso-brachium Förster, 1856, although he did not explain the reason for this nomenclatural act. Isobrachium has as maindiagnostic character the lack of notauli, which is absent in this species, according to the original description. Cam-eron’s description may have led Kieffer to propose the transference to Isobrachium. However, our observationsshowed the presence of notauli in the holotype, so, the inclusion of this species in Isobrachium became unjustifiedand we revalidate the name established by Cameron (1906) and propose it as an Epyris species. Nevertheless, whenwe transferred the species to Epyris, we realized that it is too similar to E. afer exactly by same reasons explainedabove to E. pilosipes, E. secundus and E. plurilineatus. Thus we considered it as a junior subjective synonym of E.afer.

Epyris afer has sexual dimorphism. The main differences between both sexes are: the females have VOL 0.6 ×HE, the distance of posterior ocellus to vertex crest 1.0 × DAO, the scutellar pits oval, the propodeal disc 0.9 × aslong as wide with anterior corner foveolate, the mesotibiae spinose and the forewings with metacarpus 0.2 × aslong as stigma length, whereas the male has VOL 1.0 × HE, the distance of posterior ocellus to vertex crest 0.3 ×DAO, the scutellar pits circular, the propodeal disc 0.5 × as long as wide with anterior corner groove-shaped, themesotibiae without spines, the forewing with metacarpus 0.7 × as long as stigma length.

The females of Epyris afer are similar to E. hirtipennis Cameron,1909 because both species have the mandiblewith five rounded apical teeth, the two lower ones wider than the upper ones, the pre-apical lower tooth present andthe sensillae chaetica in lower margin, the distance of posterior ocellus to vertex crest 1.0 × DAO, the propodealdisc 0.9 × as long as wide and the protarsomeres with spines short and cylindrical. However, E. afer has the scutel-lar pits oval, the space between inner and second pair of propodeal discal carina with a structure resembling a foveaand the pleurosternum with straight longitudinal groove, whereas E. hirtipennis has the scutellar pits circular, thespace between inner and second pair of propodeal discal carina without fovea-like structure, and the pleurosternumwith elliptical groove.

On the other hand, the male of Epyris afer is similar to E. penatii sp. nov. because both species have the scutel-lar pits circular, the propodeal disc ranging 0.5–0.6 × as long as wide, the anterior fovea of mesopleuron closed andlarge and the parameres positioned dorsally and elongate. However, E. afer has mandibular pre-apical lower toothand the sensillae chaetica present, the eyes glabrous, VOL 1.0 × HE, the protarsomeres without spines and the innerlobe of aedeagus without setae, whereas E. penatii sp. nov. has the mandibular pre-apical lower tooth and the sen-sillae chaetica absent, the eyes weakly pilose, VOL 0.5 × HE, the protarsomeres distally spinose and the inner lobeof aedeagus setose.

Epyris afer was previously known from Israel, Ethiopia, Guinea-Bissau, Democratic Republic of Congo, Zim-babwe, South Africa, Pakistan and Myanmar based on observations of 26 specimens (Fig. 37, red circles). Thelarge gap among these sites suggests that this species has a broad distribution in several sites through Afrotropicaland Oriental regions, and even through Mediterranean region since it occurs in Israel. We have searched for morespecimens within the material from regions where collection have been recently carried out, namely Yemen andUAE (projects coordinated by A. van Harten), Madagascar (project coordinated by B. Fisher), Thailand (projectcoordinated by M. Sharkey) and SE Asia (project coordinated by C. van K. Achterberg). We were able to find 71specimens from Yemen, UAE, Thailand and Vietnam (Fig. 37, blue squares). This confirms that this species iswidely distributed throughout the Afrotropical and Oriental regions. However we were not able to find E. aferamong the 2102 specimens of Epyris identified by Mugrabi and Azevedo (2010) from Madagascar. Also, therewere not any specimens within material from Indonesia and Malaysia.



FIGURES 1–6. Head. 1. Epyris afer ♀, lectotype, dorsal view. 2–3. E. enerterus sp. nov. ♀, holotype. 2. Dorsal view; 3. Ven-tral view. 4. E. afer ♂, paralectotype, dorsal view. 5–6. E. penatii sp. nov. ♂, holotype. 5. Dorsal view; 6. Ventral view. (Scalebar = 200 µm)



FIGURES 7–18. Mandible. 7–9. Epyris afer ♀, lectotype. 7. Frontal view; 8. Ventral view; 9. Dorsal view. 10–12. E. afer ♂,paralectotype. 10. Frontal view; 11. Ventral view; 12. Dorsal view. 13–15. E. enerterus sp. nov. ♀, holotype. 13. Frontal view;14. Ventral view; 15. Dorsal view. 16–18. E. penatii sp. nov. ♂, holotype. 16. Frontal view; 17. Ventral view; 18. Dorsal view.(Scale bar = 200 µm).

Epyris enerterus Stein et Azevedo, sp. nov.(Figs 2–3, 13–15, 21–22, 28)

Epyris afer Magretti 1897, 17: 319, part. (addition of Myanmar specimens to the syntype series of Epyris afer).

Material examined. HOLOTYPE: ♀. BIRMANIA {MYANMAR}. Bhamò, [24.2667, 97.2333], VIII.1880, L.Fea. col. (MCSN). PARATYPE: THAILAND, 1 ♀, Sakon Nakhon, Phu Phan N[ational] P[ark], dry evergreen nearhouse no.1567, 16°48.628'N, 103°53.591'E, 522 m, Pan trap, 4–5.VI.2007, Winlon Kongnara col. T2486 (QSBG).

Type condition. The right proleg of holotype is missing.Distribution. Myanmar and Thailand.Diagnosis. FEMALE. Mandible with pre-apical lower tooth, sensillae chaetica in lower margin of mandible

present. Frons nearly 1.7 × wider than eye length. Head longer than wide. Distance of posterior ocellus to vertexcrest 3.0 × DAO. Gena depressed. Notaulus with abrupt anteroposterior enlargement. MALE unknown.

Description. FEMALE. Body length 6.9 mm. LFW 3.8 mm.Color. Head, mesosoma and procoxa dark castaneous nearly black. Scape dark castaneous with apex castane-

ous. Mandible, meso and metacoxae, trochanters, femora and metasoma dark castaneous. Pedicel, flagellum, palpi,tegula, tibiae and tarsi castaneous. Wing venation light castaneous. Wings subhyaline.

Head (Figs 2, 3). Mandible (Figs. 13–15) apex as wide as base, with four sharpened apical teeth, three lowerwider than last one, pre-apical lower tooth present; sensillae chaetica in lower margin of mandible present. Clypeusshorter than wide, median lobe angulate; median carina present; apex of clypeus without setae; lateral lobereduced. Transversal section of scape elliptical; first four antennal segments ratio 23:7:6:9, segment III 0.7 × longer



than thick; antennal sensillae not visible. Antennal scrobe projected forward, dorsally sulcate. Toruli distant fromeach other 2.3 × their diameter. Eye almost reaching upper mandibular condyle, prominent, glabrous. Frons coria-ceous; punctures large and deep, separated from each other by 1.0 × their diameter. WH 0.8 × LH; WF 1.7 × HE;WF 0.7 × WH; OOL 1.5 × WOT; VOL 1.4 × HE; distance of posterior ocellus to vertex crest 3.0 × DAO. Ocellismall, frontal angle of ocellar triangle acute. Temple parallel in dorsal view. Vertex straight in dorsal view. Crestwithout setae. Occipital carina present dorsally. Gena depressed (Fig. 3). Hypostomal carina sinuous and polished.

Mesosoma (Fig. 21). Pronotal collar coriaceous, without transversal groove. Pronotal disc 1.3 × longer thanwide, coriaceous, subtrapezoidal, densely punctate, transversal carina in anterior margin absent. Mesoscutum cori-aceous, lateral slightly elevate posteriorly. Notaulus not touching posterior margin of mesoscutum, convergent pos-teriorly, abrupt anteroposterior enlargement. Parapsidal furrow convergent posteriorly, shorter and as deep asnotaulus, not reaching posterior margin of mesoscutum. Scutellum coriaceous, scutellar groove absent, scutellar pitoval. Propodeal disc 0.9 × as long as wide; anterior carina medially narrower than laterally; anterior corner fovea-shaped; five discal carinae present, median carina complete, inner discal carina complete, space between medianand inner discal carina with longitudinal ridge, second pair of discal carina incomplete and short, space betweeninner and second pair of discal carina without ridge; lateral margin straight; sublateral carina absent, lateral carinapresent, space between sublateral and lateral carina with series of transversal ridges; posterior margin straight; pos-terior corner with suboval fovea; lateral of propodeum strigulate; declivity of propodeum coriaceous, with mediancarina. Mesopleuron (Fig. 22) coriaceous; subtegular groove elongate, large anteriorly and narrow posteriorly;anterior fovea closed; sub-anterior fovea closed and small; mesopleural fovea closed; mesopleural pit present;mesopleural elevation present; lower fovea open, groove of inferior margin of lower fovea present throughoutextension of margin; episternal furrow with inner margin striate. Epicnemium lateromedially enlarged. Pleuroster-num with straight longitudinal groove.

Legs. Profemur 2.3 × longer than wide. Mesotibia spinose. tibia with sparse series of short setae in posteriorface. Tarsomeres distally spinose, spines of protarsus short and cylindrical, spines of meso and metatarsi long andflattened. Tarsal claws bidentate, inner tooth shorter than apical tooth.

Wing. Forewing with metacarpus present, 0.1 × as long as stigma length; radial vein curved forward, 1.9 × lon-ger than basal; basal vein slightly concave; transverse median vein angulated inferiorly. Hind wing with two basaland four apical hamuli.

Metasoma. Ventral surface of petiole (Fig. 28) unsegmented, space between carinae of petiole wide. Tergum IIlonger than others; terga III–VI with sparse setae on dorsal surface.

Etymology. The specific Greek epithet refers to the shape of gena, which is depressed. Discussion. This female specimen was identified by Magretti (1897) as E. afer. Indeed, they share some char-

acters like the presence of pre-apical lower tooth and the sensillae chaetica in the lower margin of mandible, theratio of propodeal disc 0.9 × as long as wide and the protarsomeres spines short and cylindrical. However, thisspecimen has VOL 1.4 × HE, the distance of posterior ocellus to vertex crest 3.0 × DAO and the gena depressed,whereas E. afer has VOL 0.6 × HE, the distance of posterior ocellus to vertex crest 1.0 × DAO and the gena flat-tened. Because of that we considered a different species.

Historically, the ventral region of body was unexplored in Bethylidae taxonomy. For this reason, similar spe-cies in dorsal and lateral view, but different in ventral view are considered the same. This was what happened to E.enerterus sp. nov. and E. afer. They share some dorsal and lateral similarities (Figs. 19–22), however, when thehead is examined ventrally (Fig. 3), it is possible to observe the depressed gena. This character was not cited toEpyris until now, being observed only in species of Chlorepyris Kieffer, like C. subangulatus (Rosmann &Azevedo, 2005).

Epyris penatii Stein et Azevedo, sp. nov.(Figs 5–6, 16–18, 25–26, 30, 34–36)

Epyris afer Magretti 1897, 17: 319, part. (addition of Myanmar specimens to the syntype series of Epyris afer).

Material examined. HOLOTYPE: ♂. [SUDAN], Kassala, [15.4558, 36.3989], 15.II.1883, P. Magretti. col.(MCSN).



FIGURES 19–22. Mesosoma in dorsal view and mesopleuron. 19–20. Epyris afer ♀, lectotype. 19. Mesosoma; 20. Mesopleu-ron. 21–22. E. enerterus sp. nov. ♀, holotype. 21. Mesosoma; 22. Mesopleuron. (Scale bar = 200 µm)



FIGURES 23–26. Mesosoma in dorsal view and mesopleuron. 23–24. Epyris afer ♂, paralectotype. 23. Mesosoma; 24.Mesopleuron. 25–26. E. penatii sp. nov. ♂, holotype. 25. Mesosoma; 26. Mesopleuron. (Scale bar = 200 µm)



FIGURES 27–30. Petiole in ventral view. 27. Epyris afer ♀, lectotype. 28. E. enerterus sp. nov. ♀, holotype. 29. E. afer ♂,paralectotype. 30. E. penatii sp. nov. ♂, holotype. (Scale bar = 200 µm)

Type condition. The metasoma of holotype is separated of the body and was glued to triangle.Distribution. Sudan.Diagnosis. MALE. Mandible without pre-apical lower tooth, sensillae chaetica in lower margin of mandible

absent. Antennal scrobe not projected forward. Eye weakly pilose. Ocelli large. Vertex convex in dorsal view.Parapsidal furrow shallower than notaulus. Lateral of propodeum areolate. Paramere arched slightly ventrad, largeand 2.0 × longer than basiparamere. Aedeagus inner margin almost straight and inner lobe membranous and setose.Apodeme base rounded, not dilated. FEMALE unknown.

Description. Male. Body length 5.4 mm. LFW 3.1 mm.Color. Head, mesosoma and coxae dark castaneous nearly black. Trochanters, fore and metafemora dark casta-

neous. Metasoma dark castaneous with apex castaneous. Scape, pedicel, flagellum, mandible, tegula, protibia, tarsicastaneous. Palpi and wing venation light castaneous. Wings subhyaline.

Head (Figs 5, 6). Mandible (Figs. 16–18) apex wider than base, with five sharpened apical teeth, two lowerwider than others, pre-apical lower tooth absent; sensillae chaetica in lower margin of mandible absent. Clypeuslonger than wide, median lobe angulate; median carina present; apex of clypeus without setae; lateral lobe reduced.Transversal section of scape cylindrical; first four antennal segments ratio 16:6:10:11, segment III 1.7 × longer thanthick; antennal sensillae not visible. Antennal scrobe not projected, dorsally carinate. Toruli distant from each other



FIGURES 31–36. Genitalia. 31–33. Epyris afer ♂, paralectotype. 31. Dorsal view; 32. Ventral view. 33. Lateral view. 34–36.E. penatii sp. nov. ♂, holotype. 34. Dorsal view; 35. Ventral view. 36. Lateral view. (Scale bar = 250 µm)



FIGURE 37. Current known distribution of Epyris afer. Red circles represent the previous data and blue squares represent thenew data of species.

2.2 × their diameter. Eye almost reaching upper mandibular condyle, prominent, weakly pilose. Frons coriaceous;punctures large and deep, separated from each other by 1.0 × their diameter. WH 1.1 × LH; WF 1.3 × HE; WF 0.6× WH; OOL 0.8 × WOT; VOL 0.5 × HE; distance of posterior ocellus to vertex crest 0.5 × DAO. Ocelli large, fron-tal angle of ocellar triangle right. Temple posteriorly convergent in dorsal view. Vertex slightly convex in dorsalview. Crest without setae. Occipital carina present dorsally and ventrally. Gena flattened (Fig. 6). Hypostomalcarina triangular and polished.

Mesosoma (Fig. 25). Pronotal collar polished, with shallow transversal groove. Pronotal disc 0.6 × longer thanwide, coriaceous, trapezoidal, sparsely punctate, transversal carina in anterior margin absent. Mesoscutum coria-ceous, lateral slightly elevate posteriorly. Notaulus touching posterior margin of mesoscutum, parallel, gradualanteroposterior enlargement. Parapsidal furrow parallel, shorter and shallower than notaulus, not reaching posteriormargin of mesoscutum. Scutellum coriaceous, scutellar groove absent, scutellar pit circular. Propodeal disc 0.6 × aslong as wide; anterior carina medially narrower than laterally; anterior corner groove-shaped; five discal carinaepresent, median carina complete, inner discal carina incomplete, space between median and inner discal carinawithout sculpture, second pair of discal carina complete, space between inner and second pair of discal carina with-out ridge; lateral margin sinuous; sublateral carina present, lateral carina present, space between sublateral and lat-eral carina with series of transversal ridges; posterior margin somewhat concave; posterior corner with subovalfovea; lateral of propodeum areolate; declivity of propodeum areolate, with median carina. Mesopleuron (Fig. 26)



coriaceous; subtegular groove elongate, large anteriorly and narrow posteriorly; anterior fovea closed and large;sub-anterior fovea closed and small; mesopleural fovea open; mesopleural pit present; mesopleural elevation pres-ent; lower fovea open, groove of inferior margin of lower fovea present only in anterior portion of margin; epister-nal furrow with inner margin striate. Epicnemium lateromedially enlarged. Pleurosternum with lozenge-shapedlongitudinal groove.

Legs. Profemur 2.3 × longer than wide. Protarsomeres distally spinose, spines long and flattened. Tarsal clawsbidentate, inner tooth shorter than apical tooth.

Wing. Forewing with metacarpus present, 0.4 × as long as stigma length; radial vein curved forward, 2.4 × lon-ger than basal; basal vein slightly concave; transverse median vein convex. Hind wing with three basal and fiveapical hamuli.

Metasoma. Ventral surface of petiole (Fig. 30) unsegmented, space between carinae of petiole narrow. TergumII as long as others; terga III–VI with sparse setae on dorsal surface.

Genitalia (Figs 34–36). Paramere positioned dorsally, arched slightly ventrad, large, elongate, 2.15 × longerthan basiparamere, apical margin convex, slightly larger medially in lateral view; cuspis elongate, its apex surpass-ing apical half of paramere length, divided into two arms only at apex, arms almost equally sized; digitus with api-cal margin denticulate, its apex not reaching cuspis apex; aedeagus bottle-shaped, short, its apex not reachingcuspis length, apex with concavity inside, inner margin almost straight; inner lobe membranous and setose; apo-deme not extending beyond genital ring, base rounded and not dilated.

Etymology. The specific epithet is given in honour to Fabio Penati, curator of MCSN.Discussion. This male specimen was identified by Magretti (1897) as E. afer. However, as compared to the

male of E. afer above, they do not represent the same species. This species has the mandibular pre-apical lowertooth and the sensillae chaetica absent, the eyes weakly pilose, VOL 0.5 × HE, the protarsomeres distally spinoseand the inner lobe of aedeagus setose, whereas E. afer has the mandibular pre-apical lower tooth and the sensillaechaetica present, the eyes glabrous, VOL 1.0 × HE, the protarsomeres without spines and the inner lobe of aedea-gus without setae.

Acknowledgements

We are grateful to Fabio Penati, David Notton, John Huber and Michael Sharkey for loaning the material studied;Agnièle Touret-Alby from MNHN for providing the photographs of some holotypes deposited there; to AlexandreP. Aguiar, Elaine D. G. Soares and Antônio C. C. Macedo for the revision of text and valuable contribution; toGeane O. Lanes by the confection of the map; to NSF grant #DEB-0542864 for the financial support of TIGERproject (Thailand Inventory for Entomological Research); to FAPES grant #003/2009 for the Master Sciencesscholarship to the first author, to CNPq grants #303216/2004-2, #306231/2007-7, #502656/2007-7 and 301669/2010-4 for the fellowships provided to fourth author, to CNPq grants #563953/05-5, 474116/2003-4, 473386/2008-9, 473386/2008-9, 502958/2008-1and 501185/2010-0 and FAPES grants #39353842/07, #41106407/08 and#45429065/2009 for the financial support; to Programa de Taxonomia CNPq grant #563953/05-5, to ProgramaCasadinho CNPq grants #620064/2006-4 and 620068/2008-6. We thank two anonymous reviewers for their valua-ble comments and suggestions.

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