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Taxonomy of Allobethylus Kieffer (Hymenoptera: Bethylidae) from southeastern Asia

Diego N. Barbosa ⁎, Celso O. AzevedoUniversidade Federal do Espírito Santo, Departamento de Biologia, Av. Marechal Campos 1468, Maruípe, 29.040-090 Vitória ES, Brazil

a b s t r a c ta r t i c l e i n f o

Article history:Received 2 June 2010Revised 21 September 2010Accepted 28 September 2010Available online 13 November 2010

Keywords:EpyrinaeScleroderminiNew speciesOriental regionNew key

Two new species of Allobethylus Kieffer, 1905 are described and illustrated. Allobethylus korystus sp. nov. fromThailand is characterized as having a mandible with four apical teeth, a clypeus with a trapezoidal frontalprofile and a pronotal declivity with a conspicuous crest. The last two traits are new characteristics to thegenus. Allobethylus lamprus sp. nov. from Myanmar is characterized as having a mandible with three apicalteeth, a clypeus with a triangular frontal profile and a bidentate median lobe. Additionally, A. multicolor Kiefferis recorded for the first time from Thailand. An updated world key to the species of genus is also provided.

© Korean Society of Applied Entomology, Taiwan Entomological Society and Malaysian Plant ProtectionSociety, 2010. Published by Elsevier B.V. All rights reserved.

Introduction

The genus Allobethylus Kieffer, 1905 belongs to Sclerodermini(Epyrinae). Before now, there were eight species: two from theAustralian region, three from the Nearctic region, one from theNeotropical region and one from the Palaearctic region. The genus ischaracterized as having an elongate and rectangular head, elongatemandibles, fully developed forewings with three closed cells, and wellpigmented, tubular radial veins of the forewings (Azevedo, 2005).

Kieffer, 1905 first described the genus but did not place any speciesinto the genus at that time. Later, Kieffer (1908) describedA. multicolor from Papua New Guinea, which is now the type-speciesof Allobethylus. Terayama (1995) described the phylogenetic relation-ships of the tribe Sclerodermini and placed Allobethylus into this tribe.He also synonymized Nesepyris Bridwell, 1920 with Allobethylus andtransferred the four species from the former to the later genus. Lastly,Terayama (1999) described Allobethylus tomoae based on a series offemales from Japan.

In this work we describe two new species: A. korystus sp. nov. fromThailand and A. lamprus sp. nov. from Myanmar. We also recordA. multicolor Kieffer from the Oriental region (Thailand).

Materials and methods

The specimens examined belong to the Queen Sirikit BotanicalGarden (QSBG) in Thailand (Wichai Srisuka). Specimens werecollected by the “Thailand Inventory Group for EntomologicalResearch” project (grant #DEB-0542864) coordinated by MichaelSharkey from University of Kentucky (U.S.A.).

Measurements and indices used in this study are as follows:body length was measured from the apex of the clypeus to theposterior margin of the last metasomal segment, excluding malegenitalia or female sting; LH, length of head, was measured infrontal view, from vertex crest to median apical margin of clypeus;WH, width of head, was measured in frontal view, its maximumwidth including eyes; WF, width of frons, was measured in frontalview, its minimum width usually about the bottom of eyes; HE,height of eye, was measured in lateral view, across its maximumheight (length); OOL, ocello-ocular line, was measured in latero-dorsal view, the shortest distance from eye top to posteriorocellus; WOT, width of ocellar triangle, was measured in frontalview, maximum width including ocelli; DAO, diameter of anteriorocellus, was measured in frontal view; distance of ocellar triangleto vertex was measured in dorso-posterior view, distance fromposterior ocellus to vertex crest; VOL, vertex-ocular line, wasmeasured in dorsal view, distance from eye top to vertex crest;and LFW, Length of Forewing, measured in dorsal view from tegulato extremity.

The nomenclature of integument sculpture follows Harris (1979),and general terms follow Evans (1964) and Azevedo (1999).

The descriptions were elaborated with DELTA (DescriptiveLanguage for Taxonomy) as proposed by Dallwitz et al. (1993) and

Journal of Asia-Pacific Entomology 14 (2011) 89–94

⁎ Correponding author. Tel.: +55 27 3335 7497; fax: +55 27 3335 7250.E-mail addresses: barbosa.laelius@gmail.com (D.N. Barbosa),

bethylidae@gmail.com (C.O. Azevedo).

1226-8615/$ – see front matter © Korean Society of Applied Entomology, Taiwan Entomological Society and Malaysian Plant Protection Society, 2010. Published by Elsevier B.V.All rights reserved.doi:10.1016/j.aspen.2010.09.005

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Journal of Asia-Pacific Entomology

j ourna l homepage: www.e lsev ie r.com/ locate / jape

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the specimens were photographed with the EntoVision system(GTVision, Hagerstown, Maryland).

Our key is modified from Azevedo (2005) and includes these twonew species.

Key for females

1. Clypeuswithmedian lobe tridentate; sides of head slightly concavein dorsal view; notaulus progressively widened backward;Japan ................................................................................... A. tomoae Terayama- Clypeus with median lobe trapezoidal or nearly so; sides of headstraight in dorsal view; notaulus evenly narrow ……………… 2

2(1). Mandible with 3 to 4 apical teeth; notaulus strong and completeor nearly so …............................................................................................. 3

- Mandible with 2 apical teeth; notaulus absent or weak andincomplete .................................................................................................... 5

3(2). Mandible elongate, overlapping more than the apical half of theoppositemandible, with 4 apical teeth .............................................. 7

- Mandible of regular size, overlapping less than the apical half ofthe opposite mandible, with 3 apical teeth; propodeal discwithout any lateral constriction………............................................... 4

4(3). Forewing with radial vein pigmented only on median third;Papua New Guinea and Thailand..………….. A. multicolor Kieffer

- Forewing with radial vein tubular and entirely pigmented ……

…………………...………………….………………….………………. 85(2). Head with sides converging posterior the eyes; Dominican

Republic ............................................................... A. antelleanus (Evans)- Headwith sides parallel behind the eyes ............................................ 6

6(5). Head ~0.67× as wide as long; median lobe of clypeusemarginated; transverse median vein of forewing slightlyoblique; Florida (U.S.A.) andMexico ...…… A. floridanus (Evans)

- Head ~0.75× as wide as long; median lobe of clypeus truncate;transverse median vein of forewing slightly straight and erect;Eastern U.S.A ........................................................... A. virginianus (Evans)

7(3). Clypeus subtrapezoidal with apical margin sub-angulate(Plate 1, panel 2); pronotal declivity with crest (Plate 1,panels 3, 4); propodeal disc not emarginated laterally (Plate 1,panel 3); Thailand ....……………………………A. korystus sp. nov.

- Clypeus truncate with apical margin emarginated; pronotaldeclivity without crest; propodeal disc emarginated laterally;Australia…………………………………… A. binconcavus Azevedo

8(4). Clypeus with median lobe bidentate (Plate 2, panel 7); eyewithout hairs; antenna with pedicel shorter than first flagello-mere and pronotal disc without hairs (Plate 1, panels 7, 8);Myanmar ….........………………………………. A. lamprus sp. nov.

- Clypeus with median lobe thickened; eye with hairs; antennawith pedicel as long as first flagellomere and pronotal disc withhairs; Hawaii and Brazil ...……………………… A. ewa (Bridwell)

Allobethylus korystus Barbosa & Azevedo sp. nov. (Plate 1)

Diagnosis. Head elongated and rectangular; mandible with fourapical teeth; clypeus subtrapezoidal with apical margin sub-angulate,frontal profile trapezoidal in shape and without clypeal carina;pronotum with pronotal declivity with crest, pronotal collar withanterior expansion; scutellum with scutellar groove, with extremityswollen; propodeal disc with median carina incomplete, sublateralcarina present, propodeal declivity without median carina; meso-pleuron without episternal furrow; hindwing with four distal hamuli,separated in pair at median region.

Description of female. Holotype (Plate 1, panel 1). Body 3.10 mm.LFW 1.8 mm.

Colors. Body dark castaneous, nearly black; scape and pedicellight castaneous; flagellum castaneous; mandible castaneous withapex lighter; palpi light castaneous; tegula castaneous; wings hyaline;wing venation castaneous; legs castaneous, except coxae; trochanters,femora, tibiae castaneous; tarsi light castaneous.

Head (Plate 1, panel 2). Head elongate and rectangular. Mandiblelong, overlapping more than apical half of opposite mandible; withfour sharpened apical teeth, with two inferior teeth longer thanothers; ventral margin with superior corner serrate and with mediantooth at inferior corner. Clypeus long, polished, extending backwardinto frons, median lobe well projected and subtrapezoidal, apicalmargin sub-angulate and trapezoidal in frontal profile, median clypealcarina absent, lateral lobe absent. Inter-torular space more thantorulus diameter. Antenna with insertion lower than clypeal eleva-tion, with sub-erect pubescence, dense, short setae; scape curved,widening to apex; pedicel longer than wide, longer than firstflagellomere. Eye glabrous, oval, on dorsal and anterior half of head.Frons coriaceous, hairy, with slight median crest. First five antennalsegments in ratio of approx. 22:9:5:6:6. LH 1.36× WH. WF 0.86× HE.WF 0.54× WH. OOL 2× WOT. VOL 0.43× HE. Distance of posteriorocellus to vertex crest 2.0× DAO. Frontal angle of ocellar triangleacute; ocellus small; ocellar triangle compact. Temple parallel. Vertexconcave. Corner rounded. Occipital carina absent; occiput with ventralmargin angulate. Gena coreaceous, completely separated from eachother by a median carina. Hypostomal carina sinuous. Palpalformula 4:2, segments with flattened cross section, widened distally,apices of segments with some short setae.

Mesosoma (Plate 1, panels 3–5). Pronotal disc 1.0× as long as wide,coriaceous, trapezoidal, side slightly concave; declivity with mediantransverse crest projected dorsally. Mesoscutum coriaceous; notauluswide, incomplete, convergent posteriorly, well impressed; parapsidalfurrow present, incomplete, parallel, weakly impressed. Scutellumcoriaceous, touching propodeal disc; scutellar groove narrow, arched,with extremity swollen. Propodeal disc 1.0× as long as wide; mediancarina incomplete posteriorly; sublateral carina present but inconspic-uous; lateral carinapresent and conspicuous; posterior carina complete;spiracle elliptical, placed above lateral carina anteriorly; declivity ofpropodeum coriaceous, without median carina; lateral surface ofpropodeum coriaceous. Mesopleuron with subtegular fovea elongated;anterior fovea present and fused with subtegular fovea; mesopleuralfovea shallow; central pit inconspicuous; episternal furrow absent.Propleuron rectangular in ventral view,withmedian carina. Prosternumsmall. Pleurosterna separated from each other by narrow groove.

Wings. Forewing elongate, 2.57× long as its maximum width,dorsal face setose, margin densely fringed, with three closed cells;costal cell very narrow and longitudinally folded and wider in distalpart; costal vein as narrow as subcostal vein; median cell 1.0× as longas submedian cell; stigma prostigma and pterostigma separated fromeach other, sub-developed; radial vein slightly curved forward.Hindwing with costal vein short; dorsal face setose, densely fringedat lower margin; with four distal hamuli, separated into pairs atmedian region.

Legs. Femora dilated; trochanters strong; hindtibia elongate; spur offoretibia strong; tarsi elongate; tarsal claw simple and arched, apex sharp.

Metasoma.Metasoma2.0× longer thanwide; not petiolate, spiraclesof first and second tergites I and II conspicuous; tergum II longer thanothers; segments III–VI with few setae at posterior margin, with amedian constriction at posterior margin. Sting not serrulate.

Variations. Body 3.10 to 3.45 mm. LFW 1.8–1.97 mm. First fiveantennal segments in ratio of approx. 25:8:6:8:8. LH 1.35× WH. WF0.9× HE. WF 0.52× WH. OOL 1.38× WOT. VOL 0.5× HE. Distance ofposterior ocellus to vertex crest 2.0× DAO. Forewing elongate, 2.57 to2.84× long as its maximum width.

Plate 1. Allobethylus korystus sp. nov. Holotype. 1. Habitus; 2. Head in dorsal view; 3. Mesosoma in dorsal view; 4. Mesosoma in lateral view; 5. Mesosoma in ventral view(scale bars 300 μm).

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Material Examined. Holotype, female. THAILAND, Phetchabun, KhaoKho NP Mixed deciduous forest, 16 32′N 101 2′E, 537 m, Malaise trap,26.xii.2006–2.i.2007, Somchai Chachumnan & Saink Singtong leg.,T1187 (QSBG). Paratypes, THAILAND, Phetchabun, Khao Kho NP, 1female, mixed deciduous forest near office, 16 39.55′N 101 8.123E,230 m, Malaise trap, 12–19.ii.2007, Somchai Chatchumnan & SainkSingtong leg., T1604 (QSBG); 1 female, Savana at nursery, 16 52′N 1018′E, 520 m, Malaise trap, 5–12.x.2006, Somchai Chatchumnan andSaink Singtong leg., T804 (UFES).

Distribution. Thailand.Etymogeny. The specific epithet korystus, from Latinized Greek,

refers to crest at pronotal declivity.Variation. Head with frons without slight median crest.Remarks. This is species is similar to A. biconcavus Azevedo, 2005 by

having the mandibles very long, overlapping more than apical half ofopposite mandible and tetradentate; the clypeus with elevation atventral margin; the mesoscutum with notauli well impressed andconvergent posteriorly; the scutellum with scutellar groove narrowand with extremity swollen. However, A. korystus sp. nov. has theclypeus long andwith apical margin sub-angulate; the pronotumwithpronotal collar with anterior expansion and pronotal declivity with acrest and the propodeal disc not strongly bi-concave in dorsal view,whereas A. biconcavus has the clypeus short and with apical marginemarginated; the pronotum with pronotal collar without anteriorexpansion and pronotal declivity without a crest and the propodealdisc strongly bi-concave in dorsal view.

Allobethylus lamprus Barbosa & Azevedo sp. nov. 2 (Plate 2)

Diagnosis.Headelongatedandrectangular;mandiblewith threeapicalteeth, one inferior longer than others; clypeus subtrapezoidal, frontalprofile triangular in shape and with clypeal carina; mesoscutum withnotauli narrow incomplete and well impressed; scutellumwith scutellargroove without extremity swollen; propodeal disc with median carinaincomplete, sublateral carina absent, propodeal declivitywithoutmediancarina; mesopleuron with episternal furrow; femora and tarsi short.

Description. Holotype (Plate 2, panel 6). Unknown. Body 1.82 mm.LFW 1.58 mm.

Colors. Body dark castaneous; scape and pedicel light castaneous;flagellum light castaneous; mandible castaneous; palpi light casta-neous; tegula castaneous; wings hyaline; wing venation lightcastaneous; coxae dark castaneous; trochanters and femora casta-neous; tibiae and tarsi light castaneous.

Head (Plate 2, panel 7). Head elongated and rectangular. Mandibleshort, overlapping less thanapical half of oppositemandible, slender,withthree sharpened apical teeth, with one inferior tooth longer than others.Clypeus short, extendingbackward into frons,median lobewell projectedand subtrapezoidal, apical margin bidentate, triangular in frontal profile,median clypeal carina present, lateral lobe absent. Inter-torular spacemore than torulus diameter. Antenna with insertion lower than clypealcarina, pubescence sub-erect, sparse, short; scape curved, same diameteraround your extension; pedicel longer than wide, longer than firstflagellomere. Eyeglabrous, oval,most placedondorsal andanteriorhalf ofhead. Frons coriaceous, hairy. First five antennal segments in ratio ofapprox. 22:10:4:6:6. LH 1.55× WH. WF 0.8× HE. WF 0.44× WH. OOL2.27×WOT.VOL1.0×HE.Distanceof posteriorocellus tovertex crest 1.5×DAO. Frontal angle of ocellar triangle acute; ocellus small; ocellar trianglecompact. Temple parallel. Vertex straight. Corner rounded. Occipitalcarina absent; occiput with ventral margin angulate. Gena coriaceous,completely separated from each other by median carina. Hypostomalcarina arched. Palpal formula 4:2, segments with flattened cross section,widened distally, apices of segments with some short setae.

Mesosoma (Plate 2, panels 8–10). Pronotal disc 0.86× as long aswide, coriaceous, trapezoidal, side straight. Mesoscutum coriaceous;notaulus narrow, incomplete, convergent posteriorly, well impressed;parapsidal furrow incomplete, parallel, inconspicuous. Scutellum

coriaceous, touching propodeal disc; scutellar groove narrow, arched,extremity not swollen. Propodeal disc 1.0× as long as wide; anteriormargin followed by short striae; median carina incomplete; sublateralcarina absent; lateral carina present and conspicuous; posterior carinacomplete; posterior margin followed by short striae; spiracleelliptical, placed above lateral carina anteriorly; declivity of propo-deum coriaceous, without median carina; lateral surface of propo-deum coriaceous. Mesopleuron with subtegular fovea elongated;anterior fovea present and fused with subtegular fovea; mesopleuralfovea shallow; central pit inconspicuous; episternal furrow present.Propleuron rectangular in ventral view, with median carina. Proster-num small. Pleurosterna separated from each other by narrow groove.

Wings. Forewing elongate, 3.0× as long as its maximum width,dorsal face setose, margin densely fringed, with three closed cells;costal cell very narrow, longitudinally folded and swollen in distalpart; costal vein as narrow as subcostal vein; median cell 1.0× as longas submedian cell; stigma prostigma and pterostigma separated eachother, sub-developed; radial vein slightly curved forward.

Legs. Femora dilated; trochanters strong; hindtibia not elongate;spur of foretibia not strong; tarsi not elongate; tarsal claw simple andarched, apex sharp.

Material Examined. Holotype. MYANMAR, Tenasserim, Thagatà,April 1887, leg. L. Fea (MCSN)

Distribution. Myanmar.Etymogeny. The specific epithet lamprus, from Latinized Greek,

refers to body with light color.Remarks.This species is similar toA. ewa (Bridwell, 1920) inhaving the

mandibles short, overlapping less than apical half of opposite mandibleand tridentate; the clypeus carinate and short; the mesoscutum withnotauli narrow and well impressed; the scutellum with scutellar groovenarrow and without extremity swollen. However A. lamprus sp. nov. hasthe median clypeal lobe bidentate; the eyes glabrous; the frons withoutshort line impressed giving rise from the clypeal carina; the antennaewith pedicel shorter than first flagellomere and the pronotal discwithouthairs, whereas A. ewa has the median clypeal lobe trapezoidal; the eyeshairy; the frons with short line impressed giving rise from the clypealcarina; the antennae with pedicel as long as first flagellomere and thepronotal disc with hairs.

Allobethylus multicolor Kieffer, 1906 (Plate 3)

This species was known only from the holotype from Papua NewGuinea. It is now recorded for the first time from Thailand. Althoughthe sites are not very distant from each other geographically, they arein different zoogeographic regions, the Australian and Orientalregions, respectively.

The Thai specimens (Plate 3) differ from the type (see Kieffer (1908)and Móczár (1966), for details) in the following aspects: body length3.07 mm; head 1.60× longer than wide; pedicel 1.57× as long as wide;length of eyes 2.84×distance of eyes frommandibles; eye 0.6× aswide aslong; ratio of antennal segments 1–12 (pedicel and flagellomeres)=7:4:5:5:5:5:5:5:5:5:5:6; length of pronotum (with collar):mesono-tum:scutellum:propodeum=35:13:18:30; mesonotum very short,shorter than scutellum;midtarsi 1.62×as long asmidfemora; hindfemora0.79× as long as hindtarsi; foretibia 1.0× as long as midtibia; breadth offorefemora, midfemora, and hindfemora about equal=10:8:8.

Material Examined. THAILAND, Phetchabun, Khao Kho NP, Mixeddeciduous near office, 16 39.479′N 101 8.105′E, 260 m, Malaise trap, 19–26.ii.2007, Somchai Chachumnan & Saink Singtong leg. T1606 (QSBG).

Discussion

Allobethylus korystus sp. nov. has some unique characters, namely:“pronotal declivity with crest”, “pronotal collar with anteriorexpansion”, “propleuron with subquadrate profile”, and a clypeusstructure not recorded in any Allobethylus species.

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The “pronotal declivitywith crest” (Plate 1, panels 3, 4)wasnever seenin Bethylidae species. It is a conspicuous elevation situated at the upperpart of the pronotal declivity near the anteriormargin of the pronotal disc.Because of that, it gives an impression that the pronotal is longer than itreally is. Therefore, this crest cannot be confused with the anterior carina

of the pronotal of Anisepyris Kieffer. The crest is evident in both the dorsaland lateral views. It is found in both sexes and its function is uncertain.

The “pronotal collar with anterior expansion” (Plate 1, panels 3, 5)is an expansion placed anteriorly in the pronotal collar producedlaterally in such a way that covers the propleuron dorsally.

Plate 2. Allobethylus lamprus sp. nov. Holotype. 6. Habitus; 7. Head in dorsal view; 8. Mesosoma in dorsal view; 9.Mesosoma in lateral view; 10.Mesosoma in ventral view (scale bars 300 μm).

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The “propleuron with subquadrate profile” (Plate 1, panel 5) wasreported only in Tuberepyris Lanes & Azevedo. According to Lanes andAzevedo (2008), this characteristic configures the main synapomor-phy of Tuberepyris. Considering that both genera are Sclerodermini,the possibility that this characteristic is a homoplasy should beinvestigated in future studies of this tribe.

The bidentate apical margin of the median clypeal lobe ofAllobethylus lamprus sp. nov. is unusual not only for the genus butalso for Sclerodermini. This tribe has a very short clypeus (Evans,1964), so generally these conditions are not presented. Thus, thisbroadens the phenotypic plasticity of the genus.

Acknowledgments

We are grateful to Michael Sharkey for loaning the material studiedand for permission to retain the duplicate; to NSF grant #DEB-0542864for the financial support of TIGER project (Thailand InventoryGroup for Entomological Research); to CNPq grants #303216/2004-2,#306231/2007-7 and #502656/2007-7 for the fellowships provided toboth authors; to CNPq grants #563953/05-5, 474116/2003-4 and473386/2008-9 and FAPES grants #39353842/07, #41106407/08 and#45429065/2009 for financial support; to Programa de Taxonomia —

CNPq grant #563953/05-5; and to Programa Casadinho — CNPq grant#620064/2006-4.

References

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Azevedo, C.O., 2005. A new species of Allobethylus (Hymenoptera, Bethylidae) fromAustralia, with a key to world species. Zootaxa 1064, 25–30.

Bridwell, J.C., 1920. Some notes on hawaiian and other Bethylidae (Hymenoptera) withthe descriptions of a new genus and species. Second paper. Proceedings of theHawaiian Entomological Society 4, 291–314.

Dallwitz, M.J., Paine, T.A., Zurcher, E.J., 1993. User's Guide to the DELTA System: A GeneralSystem for Processing Taxonomic Descriptions, 4th edition. . http://biodiversity.uno.edu/delta/.

Evans, H.E., 1964. A synopsis of the American Bethylidae (Hymenoptera, Aculeata). Bull.Mus. Comp. Zool. 132, 1–222.

Harris, R.A., 1979. A glossary of surface sculpturing. Occas. Pap. Entomol. 28, 1–31.Kieffer, J.J., 1905. Proctotrypides. In: André, E. (Ed.), Species des Hyménoptères

d'Europe & d'Algerie. A. Hermann, Paris, 9, pp. 65–288.Kieffer, J.J., 1908. Description de quelques nouveaux serphides. Bull. Soc. Hist. Nat. Metz

Ser. 3 (25), 1–7.Lanes, G.O., Azevedo, C.O., 2008. Phylogeny and taxonomy of Sclerodermini

(Hymenoptera, Bethylidae, Epyrinae). Insects Syst. Evol. 39, 55–86.Móczár, L., 1966. Remarks on Kieffer's and Marshall's types (Hymenoptera,

Bethylidae) in the Hungarian Natural History Museum. Acta Zool. Acad. Sci.Hung. 12, 339–361.

Terayama, M., 1995. The phylogeny of the bethylid wasps tribe Sclerodermini(Hymenoptera, Bethylidae). Proceedings of the Japanese Society of SystematicZoology 54, 65–73.

Terayama, M., 1999. Descriptions of new species of the family Bethylidae from the Ryukyus,and taxonomic notes on the Japanese species of the genus Sclerodermus. In: Yamane, S.,Ikudome, S., Terayama, M. (Eds.), Identification Guide to the Aculeata of the NanseiIslands, Japan. Hokkaido University Press, Sapporo, pp. 701–725.

Plate 3. Allobethylus multicolor Kieffer. Habitus (scale bar 300 μm).

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