European Journal of Taxonomy 4: 1-12

http://dx.doi.Org/10.5852/ejt.2011.4

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2011 • Isabel D.C.C. Alencar & Celso O. Azevedo

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Research article

Review of Neurepyris Kieffer, 1905 (Hymenoptera, Bethylidae),

a new synonym of Pristocera King, 1808

Alencar I.D.C.C. & Azevedo C.O. 2011. Review of Neurepyris Kieffer, 1905 (Hymenoptera, Bethylidae), a new

synonym of Pristocera YAug, 1808. European Journal of Taxonomy 1-12. http://dx.doi.Org/10.5852/ejt.2011.4

Isabel D.C.C. ALENCARi^ & Celso O. AZEVEDO^

Tnstituto Federal de Eduea 9 ao, Cieneia e Teenologia do Espirito Santo, Campus Santa Teresa,

Rodovia ES-080, km 21, 29660-000 Santa Teresa, ES, Brasil.

^Universidade Federal do Espirito Santo, Departamento de Biologia,

Av. Mareehal Campos, 1468, 29.040-090, Vitoria, ES, Brasil

^Email: [email protected]

Abstract. The genus Neurepyris Kieffer, 1905 is revised. The adult male holotypes N. rufiventer

Kieffer, 1913 from Eritrea and N. tagala (Ashmead, 1905) from the Philippines are redeseribed and

illustrated. Both speeies are transferred from the subfamily Epyrinae to Pristoeerinae beeause they have

the metanotum well developed medially. Neurepyris rufiventer is transferred to Pristocera King, 1808

beeause the hypopygium is deeply divided into two apieal lobes, the subdiseoidal and eubital veins do

not reaeh the border of the forewing, the pronotal dise has the anterior region slightly elevate medially,

and the stigma is elongate. Neurepyris tagala is transferred to Apenesia Westwood, 1874 beeause the

basal tooth of mandible is not eurved inward, the median lobe of elypeus is not depressed near the

antennal insertions, and the aedeagus eonsists of one lamina. The genus Neurepyris is eonsidered a

junior synonym of Pristocera beeause its type speeies is N. rufiventer.

Key words. Epyrinae, Pristoeerinae, Apenesia, Afrotropieal region. Oriental region.

Introduction

Kieffer (1905) ereated Neurepyris Kieffer, 1905 to aeeommodate a speeies of Epyrini, whieh has a

longitudinal earina on the propodeal dise absent, unidentate tarsal elaw and laneeolate stigma. Kieffer &

Marshall's key (1904-1906) indieated that there was one speeies from Eritrea known from one female,

but they did not deseribed in that time. This speeies was formally deseribed years later as N. rufiventer

Kieffer, 1913 by Kieffer (1913). Kieffer (1914a) transferred a doubtful Philippine speeies Rhabdepyris

tagala (Ashmead, 1905) to Neurepyris. Both speeies are known only from their types. However our

initial analyses indieate that both speeies are males of Pristoeerinae beeause both of them have the

metanotal well developed medially, propodeum with eonvex sides, metasoma depressed and genitalia

with eomplex aedeagus.

In this study, we aimed to reeonsider the genus, presenting a new view about both speeies here redeseribed

and illustrated.

European Journal of Taxonomy A\ 1-12 ( 2011 )

Material and methods

The material studied was borrowed from the Museo Civieo di Storia Naturale 'Giaeomo Doria' - MCSN

(F. Penati) and the National Museum of Natural History - USNM (D. Furth).

The nomenelature of integument seulpture follows Harris (1979), and general terms follow Evans (1964)

and Azevedo (1999).

Measurements and indiees used in this study are as follows: body length measured from the apex of

elypeus to the posterior margin of the last metasomal segment; LFW, length of forewing; LH, length

of head, measured in frontal view, from vertex erest to median apieal margin of elypeus; WH, width of

head, measured in frontal view, its maximum width ineluding eyes; WF, width of frons, measured in

frontal view, its minimum width, usually about bottom of eyes; HE, height of eye, measured in lateral

view, aeross its maximum height (length); OOE, oeelli-oeular line, measured in latero-dorsal view, the

shortest distanee from eye top to posterior oeellus; WOT, width of oeellar triangle, measured in frontal

view, maximum width, ineluding oeelli; DAO, diameter of anterior oeellus, measured in frontal view;

distanee of oeellar triangle to vertex: measured in dorso-posterior view, distanee from posterior oeellus

to vertex erest; VOE, vertex-oeular line, measured in dorsal view, distanee from eye top to vertex erest.

Results

GQxms PristoceraYAug, 1808

NeurepyrisYiQffQX, 1905: 29. Type-speeies: Neurepyris rufiventer Kieffer, by subsequent monotypy;

syn. nov.

Pristocera rufiventer (Kieffer, 1913) eomb. nov.

Figs 1-3

Neurepyris rufiventer Kieffer, 1913: 107.

Neurepyris rufiventer - Kieffer 1914a: 370-371. — Gordh & Moezar 1990: 135.

Material examined

Holotype S\ label 1: 380 (3107360?) // label 2: Epyris rufiventer H label 3: Neurepyris rufiventer Kieffer //

label 4: v. Boll. Eab. Z. Postiei (Portiei) 7, 1913, p. 107 // label 5: typus // Eabel 6: Holotypus, Neurepyris

rufiventer, J.J. Kieffer., 1913 //Eabel 7: Museo Genova, eoll. P. magretti (dono 1913).

Penati's note: There is a rhomboid label margined by blue band writing n. 310 attaehed to loeality label

(MCSN).

Type-locality

Eritrea.

Description

Holotype f (Fig. lA). Body length 5.8 mm. EFW 4.5 mm.

Color. Head and mesosoma blaek, exeept pronotal eollar and posterior end of pronotal dise dark

eastaneous, antenna, mandible, tegula and metasoma light eastaneous, petiole dark eastaneous, legs dark

eastaneous to light eastaneous distally, veins eastaneous, pterostigma dark eastaneous, wing membrane

subhyaline.

ALENCAR I.D.C.C. & AZEVEDO C.O., Review oiNeurepyris Kieffer

Fig. 1. Pristocera rufiventer comb, nov., holotype, S- A. Habitus. B. Head, dorsal view. C. Mandible,

frontal view. D. Mesosoma, dorsal view. E. Mesopleuron, lateral view. F. Mesosoma, ventral view (bars

0.5 mm).

European Journal of Taxonomy A\ 1-12 ( 2011 )

Head. (Fig. IB). Mandible with five sharp teeth, upper two redueed (Fig. 1C). Clypeus with tmneate

median lobe, 0.4x as long as wide; median earina eonspieuous, high and arehed in profile. Antenna short,

1.9 mm, reaehing posterior end of pronotum; seape arehed, widened distad; pubeseenee sub-appressed

with some setae ereet, setae long. First four antennomeres in ratio of ~8:2:5:4; segment XI 2.5x as long

as wide. Frons polished, shining, punetate-punetulate. Frontal groove absent. Oeellar triangle elevate,

its frontal angle aeute, oeelli large. Anterior oeellus surpassing anterad imaginary line of eye top. LH

Fix WH; WF 0.6x WH; WF 0.9x HE; OOL 1. lx WOT; DAO 0.2x WOT; posterior oeellus distant from

vertex erest 3.5x DAO. Profile of vertex erest eonvex in dorsal view, eorner rounded; VOL Fix HE.

Temple profiles almost parallel in dorsal view. Oeeipital earina present ventrally and dorsally.

Mesosoma. (Fig. ID-F). Thorax dorsum shinning, punetulate, setae long. Pronotal dise trapezoidal;

side slightly eoneave; anterior margin earinate; anterior region slightly elevate medially with rounded

top. Notaulus eonspieuous, ineomplete anteriorly, eonvergent posteriorly, gradually wider and deeper

posteriorly, foveolate inside, not reaehing posterior margin of mesoseutum. Parapsidal furrow ineomplete

and ineonspieuous anteriorly, straight, narrow, very shallow. Lateral area of mesoseutum slightly elevate

without fovea anteriorly. Seutellar groove deep, narrow, eoneave, dilated laterally. Metanotum with

large median elevation; median fovea sub-trapezoidal with anterior base F3x wider than posterior one;

metanotal groove foveolate laterally, first fovea very large and triangular, punetulate inside, others

reetangular and narrower than first one. Propodeal dise F lx as long as wide, dise rugulose; spiraele slightly

arehed, plaeed eompletely at dise. Propodeum rugulose laterally. Propodeal deelivity weak, eonvex in

lateral view, rugulose. Mesopleuron with subtegular groove dilated anteriorly and uniformly narrow

posteriorly, foveolate inside; epistemal groove eontinuous to subtegular one, foveolate; mesopleural

eallus elevate, polished and shinning; anterior region strongly punetuate to punetieulate posteriorly,

median region with irregular punetures (Fig. IE); sub-anterior fovea large and deep. Pleurostemum

with aeetabular earina wide medially followed by large foveae (Fig. IF); median groove irregularly

foveolate; latero-posterior groove narrow, eonvergent medially. Fore femur 4.2x as long as wide. Tarsal

elaws bidentate, teeth sharp.

Wings. (Fig. 2). Forewing with radial vein long and slightly eurved forward apieally; R1 (metaearpus

of Evans) absent; M+Rs (basal vein of Evans) eoneave; eu-a (transverse vein of Evans) angulate; C\xa

(diseoidal vein of Evans) tubular and shortly well pigmented from interseetion with M+Rs; m-eu (first

Fig. 2. Pristocera rufiventer eomb. nov., holotype, S. Forewing, dorsal view (bar 0.5 mm).

ALENCAR I.D.C.C. & AZEVEDO C.O., Review of Neurepyris Kieffer

reeurrent of Evans) and Cue (subdiseoidal vein of Evans) weakly impressed. Hind wing with six apieal

hamuli.

Metasoma. (Fig. 3A-B). Petiole 0.6x as long as wide. Caster shinning, punetieulate to imbrieate-

punetieulate, setose laterally from II tergite. Hypopygium deeply divided (Fig. 3A-B), densely setose

apieally, base very narrow and apex expanded; inner margin with inner fold setose subapieally, base of

fold with inner hook rounded mesad.

Genitalia. (Fig. 3C-D): paramere bilaminar, plaeed ventral-dorsally, 1.5x as long as basiparamere; apex

projeeted upward and arehed subapieally; apieal margin eonvex; ventral margin with basal fold; dorsal

margin slightly eoneave medially. Basiparamere with dorsal margin slightly eoneave. Basivolsella

surpassing ventral margin of basiparamere, margin almost straight. Digitus with apieal margin dentate,

apex sharp and arehed basad. Cuspis large, narrow, apex eoneave. Aedeagus divided into two laminae;

dorsal lamina with apex divided into two pairs of apieal lobes, outer lobe triangular, lateral and dorsal

Fig. 3. Pristocera rufiventer eomb. nov., holotype, S- A. Hypopygium, outer view. B. Hypopygium,

inner view. C. Genitalia, ventral view. D. Genitalia, dorsal view (bars 0.5 mm).

European Journal of Taxonomy A\ 1-12 (2011)

margin converging upward, apex narrow and projeeted ventrad, inner lobe membranous; ventral lamina

slightly shorter than dorsal one, very narrow, lateral margin eonvex with subapieal eonstrietion, inner

margin straight, diverging apiead; ventral margin with basal strap. Genital ring wide laterally. Basal ring

absent. Apodeme not extending beyond genital ring.

Genus Apenesia Westwood, 1874

Apenesia tagala (Ashmead, 1905) eomb. nov.

Figs 4-6

Epyris tagala Ashmead, 1905: 109.

Rhabdepyris (Trichotepyris) tagala Kieffer, 1908: 32.

Neurepyris tagala Kieffer, 1914a: 370-371.

Epyris toga/a-Brown 1906: 68.

Neurepyris tagala - Kieffer 1914b: 285 —Kurian 1954: 273 — Gordh & Moezar 1990: 135.

Material examined

Holotype S\ label 1: Manila // Ph; label 2: Robt Brown // Colleetor; label 3: S Type // No. 8439 //

U.S.N.M.; Label 4 (Ashmead's handwritten): Epyris H tagala H $ Ash. (USNM).

Type-locality

The Philippines, Luzon, Manila.

Description

Holotype f (Fig. 4A). Body length 5.1 mm. LFW 3.7 mm.

Color. Head and mesosoma blaek, exeept pronotal eollar and posterior end of pronotal dise dark

eastaneous; metasoma dark eastaneous; antenna eastaneous with flagellomeres progressively darker

distad, palpi light eastaneous; mandible and tegula eastaneous; legs eastaneous with eoxae darker; wing

membrane subhyaline, veins eastaneous.

Head. (Fig. 4B). Mandible with one lower large tooth and upper eutting edge (Fig. 4C). Clypeus projeeting

forward subangulate median lobe, its length 0.19x LH; median earina eonspieuous, slightly high and

arehed in profile. Antenna 1.95 mm long, reaehing posterior end of pronotum; seape arehed, widened

distad; pubeseenee subereet with some ereet setae. First four antennomeres in ratio of -23:6:11:11;

segment XI 2.25x as long as wide. Eye with sparse short hairs. Frons polished, shining, punetures

small, somewhat deep, separated eaeh other by 1.0-3.Ox their diameters. Frontal groove absent. LH

L14x WH; WF 0.52x WH; WF l.Ox HE; frontal angle of oeellar triangle aeute; WOT 3.3x DAO, OOE

L2x WOT; DAO 0.16x WF; posterior oeellus distant from vertex erest 2.3x DAO; anterior margin of

anterior oeellus not reaehing imaginary top eye line. Profile of vertex erest eonvex in dorsal view; eomer

rounded; VOE 0.49x HE. Temple profiles diverging anterad in dorsal view. Oeeipital earina not visible

in dorsal view.

Mesosoma. (Fig. 4D-F). Thorax dorsum shinning, punetures smaller and sparser than those of frons.

Pronotal dise trapezoidal; side straight in dorsal view; anterior margin slightly eonvex, not earinate,

pronotal deelivity vertieal. Notaulus eonspieuous, slightly arehed, eomplete anteriorly, eonvergent

posterad, gradually wider posteriorly, not reaehing posterior margin of mesoseutum. Parapsidal furrow

ineomplete anteriorly, eonspieuous, slightly arehed. Region around parapsidal furrow bulging. Seutellar

groove deep, narrow, arehed, slightly dilated laterally. Metanotum with median elevation; median

fovea semi-eireular; metanotal groove foveolate laterally, first fovea large and triangular, not punetured

ALENCAR I.D.C.C. & AZEVEDO C.O., Review of Neurepyris Kieffer

Fig. 4.Apenesia tagala eomb. nov., holotype, 3 . A. Habitus. B. Head, dorsal view. C. Mandible, frontal

view. D. Mesosoma, dorsal view. E. Mesopleuron, lateral view. F. Mesosoma, ventral view (bars 0.25

mm ).

European Journal of Taxonomy A\ 1-12 (2011)

inside, others reetangular and narrower than first one. Propodeal dise 0.64x as long as wide, median

earina eomplete, dise mgulose; spiraele short, elliptieal, eompletely plaeed at lateral of propodeum,

lateral earina abruptly eurved anteriorly to aeeommodate spiraele; posterior earina little distinguishable.

Propodeum strigulate laterally. Deelivity of propodeum eoarsely mgulose, without median earina,

almost straight in lateral view. Mesopleuron with subtegular groove dilated anteriorly and evenly narrow

posteriorly, foveolate inside; epistemal groove not eontinuous to subtegular one, foveolate; mesopleural

eallus polished and shinning; anterior region strongly punetuate to punetieulate posteriorly (Fig. 4E).

Pleurostemum mostly polished, aeetabular earina wide posterior pit deep (Fig. 4F). Fore femur 3.4x as

long as wide. Tarsal elaws bidentate, teeth sharp.

Wings. (Fig. 5). Forewing with radial vein long and evenly eurved; R1 short than half length of stigma;

M+Rs (basal vein of Evans) eoneave; eu-a (transverse vein of Evans) angulate; C\xa (diseoidal vein of

Evans) tubular for short distant, interseeting M+Rs. Hind wing without basal hamuli, with six equidistant

median hamuli.

Metasoma. (Fig. 6A-B). Not petiolate; anterior half mostly polished, posterior half mostly very weakly

eoriaeeous; progressively more hairy posterad. Stemite Vll reetangular, without median stalk, lateral

stalk very short, posterior margin with median narrow semieireular eoneavity. Hypopygium (Fig. 6A-

B) strongly eoneave, but not divided into two lobes, inner margin badly eoneave, inner surfaee setose

apieally, median stalk about 0.6x as long as plate, lateral stalk very short, dorsal surfaee short, surrounding

inner margin of lobe, progressively higher basad, posterior margin with perpendieular median lamina

direeted upward.

Genitalia. (Fig. 6C-D): paramere subvertieal, about as long as basiparamere; plaeed dorsally, outer

surfaee mostly hairy, bilaminar, exeept basal-inner quadrant; apex rounded; wide, dorsal margin mueh

developed mesad, espeeially basally, basal half eonvex, apieal half eoneave, with median angulate

eallus; ventral margin straight. Basiparamere with dorsal margin inelined and truneate in lateral view.

Basivolsella eompletely outlined and separated from basiparamere, short, basal half subquadrate. Digitus

laminar, wide, rounded, dorsal surfaee eonvex and dentieulate. Cuspis laminar, base eompletely angled

upward, somewhat narrow, apex rounded. Aedeagus bulging, divided into dorsal and ventral parts,

apex of dorsal one with three pair lobes, dorsal lobe eompletely angled ventrad, tubular, apex deeply

bifiireated; median lobe stout, apieal margin with delieate filament apiead, ventral margin dentieulate,

inner margin membranous and little hairy, ventral lobe with large expansion direeted downward; ventral

surfaee laminar expansion, angled laterad with apex rounded and base with ventral loop of irregular

Fig. 5. Apenesia tagala eomb. nov., holotype, f. Forewing, dorsal view (bar 0.25 mm).

ALENCAR I.D.C.C. & AZEVEDO C.O., Review of Neurepyris Kieffer

margin; ventral part of aedeagus laminar with base wider than apex, inner surfaee vertieal, inner lower

part with ridge, apex divergent. Genital ring subreetangular. Basal ring narrow. Apodeme narrow,

extending beyond genital ring.

Discussion

Aeeording to the most reeent Bethylidae eatalog (Gordh & Moezar 1990), Neurepyris type speeies,

Neurepyris tagala, was eonsidered lost but reeently Fabio Penati found it in the MCSN eolleetion. Thus,

we were able to aeeess it and study both speeies of the genus. Our results eonfirm the initial hypothesis

that both speeies of Neurepyris are Pristoeerinae. Our analysis also indieated that the only two speeies

belong to distinet genera within Pristoeerinae. A historieal review of nomenelatural aets of the genus

indieated that the type speeies designation was wrongly perpetuated by Gordh & Moezar (1990) and its

sex was misidentified.

Fig. 6. Apenesia tagala eomb. nov., holotype, S- A. Hypopygium, outer view. B. Hypopygium, inner

view. C. Genitalia, ventral view. D. Genitalia, dorsal view (bars 0.5 mm).

European Journal of Taxonomy A\ 1-12 (2011)

Kieffer (1913) misidentified the sex of the holotype of Neurepyris rufiventer as a female. However,

the speeimen is elearly a male. There are two identifieation labels attaehed to the type, one is Epyris

rufiventer and the other is Neurepyris rufiventer. However this speeimen has never been formally named

as Epyris rufiventer. That indieates a probable eonfusion about the eoneept of the genus within Epyrini

by the author speeies, although both Neurepyris and Epyris are elassified as Epyrini.

Neurepyris rufiventer is easily reeognized as belonging to Pristoeerinae mostly beeause it has the

metanotum well developed medially. It also has the hypopygium deeply divided into two lobes. Sueh

a eondition is found in five genera of this subfamily: Dicrogenium Stadelmann, 1894, Neodicrogenium

Benoit, 1957, Diepyris Benoit, 1957, Kathepyris Kieffer, 1907 and Pristocera. Among these genera,

the first two have a large spine on the genal area whieh makes them distinet. Diepyris has the mandible

siekle-shaped with only two apieal teeth, whereas Kathepyris and Pristocera have mandibles that are

wide apieally, with four or five apieal teeth. However, Kathepyris has the forewings with subdiseoidal

and eubital veins reaehing the apieal margin of the wing whereas Pristocera does not. Neurepyris

rufiventer should eertainly be assigned to Pristocera beeause it has the eombination of the following

eharaeters: the mandibles wide apieally with five apieal teeth, the genal area without spine, the forewings

with subdiseoidal and eubital veins barely visible and not reaehing apieal margin of the wing, and

the hypopygium divided into two lobes. Beeause of that we propose the transferenee of Neurepyris

rufiventer to Pristocera and the subsequent new eombination {P. rufiventer eomb. nov.).

Neurepyris tagala was originally deseribed as a speeies belonging to Epyris Westwood, 1832 (see

Ashmead 1905). However Kieffer (1908) transferred it to Rhabdepyris and indieated a possible

eonfusion about the generie identity of this speeies. Finally it was transferred to Neurepyris by Kieffer

(1914a). Nevertheless, both transferenees remained within Epyrini genera. However, what first ealled

our attention is the faet N. tagala presents the metanotum well developed medially as it oeeurs in all

Pristoeerinae. Aetually, this eondition is pointed out as the main synapomorphy of this subfamily (Sorg

1988; Terayama 1996). Neurepyris tagala has the basal tooth of the mandible not eurved inwardly, the

median lobe of elypeus not depressed near the antennal insertions, and the aedeagus eonsisting of one

lamina. This eombination of eharaeters addresses this speeies to Apenesia Westwood. Beeause of that

we propose the transferenee of Neurepyris tagala and the subsequent new eombination {Apenesia tagala

eomb. nov.).

Aeeording to Gordh & Moezar (1990), N. tagala was designated the type speeies of Neurepyris by

'subsequent monotypy'. However, this assumption does not fit with the nomenelatural history of the

genus.

Neurepyris was established by Kieffer (1905) in a key. He did not provide any deseription nor inelude

any speeies on it. Curiously, Kieffer & Marshall (1904-1906) mentioned that Neurepyris was ereated

to aeeommodate one speeies from Eritrea known only from one single female (see the footnote in

Kieffer & Marshall's key, page 251). However, there was no speeies formally addressed to Neurepyris

in that time. Kieffer (1908) formally deseribed Neurepyris and pointed out that 'd'espece... sur lequel

ce genre est etabli provient de TErythree et n'a pas encore ete decrif (= this genus was established

based one speeies from Eritrea not deseribed yet). Kieffer (1913) formally provided the deseription

of the first speeies of Neurepyris, N. rufiventer, the speeies from Eritrea. Kieffer (1914a) transferred

Rhabdepyris (Trichotepyris) tagala (Ashmead, 1905), previously deseribed in Epyris, to Neurepyris. He

also indieated Neurepyris rufiventer as the type speeies of this genus.

Given this seenario, we offer three eonelusions. First it is elear N. rufiventer is the type speeies by

subsequent monotypy in Kieffer (1913), seeond the indieation of type speeies by Kieffer (1914a)

was unneeessary, and third the eitation of N. tagala as type speeies by Gordh & Moezar (1990) is not

ALENCAR I.D.C.C. & AZEVEDO C.O., Review of Neurepyris Kieffer

adequate. So we assume that the eorreet type-speeies of the genus is Neurepyris rufiventer. Thus it has

to be treated as a new junior synonym of Pristocera.

Acknowledgement

We thank to Fabio Penati (MCSN) and David Furth (USNM) for the loan of the types and other support,

Matthias Buffington for taking the pietures of the genitalia of Neurepyris tagala, Diego N. Barbosa

for taking some pietures of Neurepyris rufiventer, Felipe B. Fraga for his help with the genitalia draws

of Neurepyris rufiventer, CAPES and CNPq (grants #500280/00-8, #303216/2004-2, #306331/2007-

7 and #306231/2007-7) by providing seholarship to the authors and finaneial support; CNPq for the

Programa Casadinho grant #620064/2006^ and #620068/2008-6, and Programa de Taxonomia grant

#563953/05-5 by providing additional finaneial support.

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Manuscript received: 13 August 2011

Manuscript accepted on: 16 November 2011

Published on: 28 December 2011

Topic editor: Malcolm Scoble.

In eomplianee with Artiele 8.6 of the ICZN, printed versions of all papers are deposited in the libraries

of the institutes that are members of the EJT eonsortium.