Zootaxa,Redescription and placement of the Oriental ...

Zootaxa 1654: 55–60 (2007) www.mapress.com / zootaxa/

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Redescription and placement of the Oriental Scaphepyris rufus Kieffer (Hymenoptera: Bethylidae) G. O. LANES & C. O. AZEVEDO Universidade Federal do Espírito Santo, Departamento de Biologia, Av. Marechal Campos 1468, Maruípe, 29040-090 Vitória ES, Brazil. E-mail: [email protected], [email protected]

Abstract The Sclerodermini (Bethylidae: Epyrinae) genus Scaphepyris Kieffer and its single species, S. rufus Kieffer, are known only from Indonesia, and are revised. The genus is characterized by having the clypeus produced medially with the apical margin subangulate, the mesopleuron with a large, long upper groove, the ocelli and wings absent, the mid tibiae strongly spinose, the eyes extremely reduced, and a constriction between the propodeum and the mesonotum. These two latter conditions place it in Pristocerinae. A key for the Pristocerinae genera where females are known is provided. Key words: Pristocerinae; Epyrinae; Sclerodermini

Introduction Scaphepyris was proposed by Kieffer (1905) and placed in subfamily Bethylinae (now Bethylidae), which was considered to belong to Proctotrypidae (now Proctotrupidae) at that time. Kieffer (1914) established the family Bethylidae for what is considered now to be Bethylidae, Dryinidae, Embolemidae and Sclerogibbidae together. He also established five tribes in Bethylinae, namely Pristocerini, Mesitiini, Bethylini, Epyrini, and Sclerodermini. He placed Scaphepyris on the latter tribe. Berland (1928) raised Bethylidae to superfamily rank and, thus, the five tribes established by Kieffer (1905) were raised to subfamily. When Evans (1964) revised the American Bethylidae he split Scleroderminae (sensu Berland) into two tribes (Sclerodermini and Cephalonomiini) and transferred them to Epyrinae, even though he had never studied some genera from the Old World. Subsequent authors have treated the Indonesian genus Scaphepyris as Sclerodermini. Terayama (1995) proposed a phylogeny for the Sclerodermini, but he excluded Scaphepyris, because he was not able to find the holotype. According to him, the holotype was presumably lost during the World Wars. But recently while making a cladistic analysis of Sclerodermini we were able to find the holotype of Scaphepyris deposited at Museo Cívico di Storia Naturale Giacomo Doria, and we verified that this genus belongs to Pristocerinae.

Material and methods The examined material was borrowed from Museo Cívico di Storia Naturale Giacomo Doria, Genova, Italy by Roberto Poggi (MCSN). The measurements and indices used in this study are as follow: LH, head length, measured in dorsal view,

Accepted by J. Pitts: 7 Nov. 2007; published: 5 Dec. 2007

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from the crest of the vertex to the median apical margin of the clypeus; WH, head width, measured in dorsal view, its maximum width; WF, width of frons, measured in dorsal view, its minimum width; HE, height of eye, measured in dorsal view, its maximum height (length); VOL, vertex-ocular line, measured in dorsal view, the distance from the top of the eye to vertex crest. The terms for structures follow Evans (1964) and those for integument sculpture follow Harris (1979).

Key to genera of Pristocerinae (Female) The key is modified from Evans (1964) and Terayama (2003). It includes three genera, Proasapenesia Kieffer based on P. lacteipennis Kieffer, which is the only species of this genus with known female described by Krombein (1989); Caloapenesia Terayama based on C. brevis Azevedo, the only species of this genus with known female described by Azevedo (2004), and Scaphepyris based on S. rufus Kieffer described below. The females of Pristocera Klug and Acrepyris Kieffer are indistinguishable in this key. Yasumatsu (1955) proposed Neopristocera as a subgenus of Pristocera with P. japonica Yasumatsu as the type species. Evans (1963) accepted Yasumatsu’s classification for Pristocera, but considered Neopristocera as a junior synonym of Acrepyris because P. japonica is very similar to the species of Acrepyris. According to Evans (1963), males of Acrepyris differ from those of Pristocera mainly by hypopygium and genitalia characters, and other characters such as the number of aedeagus valves, and digiti shape. Yasumatsu (1955) and Evans (1963) were not able to find characters to separate the females of these taxa. Terayama (1996) re-established the status of Acrepyris based on cladistic analysis, but he analyzed only male specimens. Characters to separate females of these genera are still unknown. Currently only geographic distribution has been used to separate females of these genera; Acrepyris is mostly distributed in the New World with a few species in the Oriental and southeast Palaearctic regions, whereas Pristocera predominantly occurs in the Ethiopian and the Oriental regions. So, we considered it appropriate not to separate these genera in the key. 1 -

2(1) 3(1)

4(3) 5(4) 6(3) -

Propodeum strongly constricted at its extreme anterior end; mesonotum elongate, mesopleuron does not cover the lateral of propodeum in dorsal view; eye eye absent, or consist of one facet.......................... 2 Propodeum not constricted at anterior end, broadly in contact with mesonotum, constricted at or near spiracles if at all; mesonotum short, mesopleuron cover at least anterior third of lateral of propodeum in dorsal view; eye consist of more than one facet (some exceptions) ........................................................ 3 Mesonotum triangular in dorsal view; eye absent ................................................... Prosapenesia Kieffer Mesonotum with lateral margin more or less parallel in lateral view; eye consist of one facet or sometimes absent..................................................................................................... Pseudisobrachium Kieffer Mesopleuron, seen in dorsal view, very narrow; mesothorax barely wider than across prothorax in dorsal view; sides of propodeum more or less parallel in dorsal view, at most weakly constricted ............... ..............................................................................................................................Dissomphalus Ashmead Mesopleuron, seen in dorsal view, quite large; thorax distinctly wider than elsewhere; propodeum with evident constriction at or near spiracles................................................................................................... 4 Propodeal disc strongly constricted anteriorly, maximum width at least 2.0x constriction width ......... 5 Propodeal disc moderately constricted, maximum width < 1.9x constriction width.............................. 6 Palpal formula 5:3, fore basitarsus strongly curved ........................................... Caloapenesia Terayama Palpal formula 6:3; fore basitarsus not curved .................................. Pristocera Klug/Acrepyris Kieffer Body at most only weakly flat dorsoventrally; mid tibia spinose above; lateral of pronotum weak to moderately arched, moderately visible in dorsal view ........................................................................... 7 Body strongly flattened; mid tibia smooth above; lateral of pronotum strongly arched, broadly visible in dorsal view ............................................................................................................................................. 8

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7(6) Clypeus emarginated, truncate or somewhat produced medially (trapezoid in some species); mesopleuron smooth ................................................................................................................Apenesia Westwood Clypeus strongly produced medially, apical margin subangulate; mesopleuron with large and long upper groove ............................................................................................................. Scaphepyris Kieffer 8(6) Base of pronotum not in contact with mesonotum in dorsal view; mesonotum divided into mesoscutum and scutellum ............................................................................................................ Afgoiogfa Argaman Base of pronotum in contact with base of mesonotum in dorsal view; mesonotum not divided ............. ........................................................................................................................... Parascleroderma Kieffer

Scaphepyris Kieffer, 1905 Scaphepyris Kieffer, 1905: 29. Type species: Scaphepyris rufus Kieffer, 1904. Original designation.

Diagnosis. Female. Body weakly flattened. Eye present. Clypeus with median lobe well developed, apical margin subangulate. Occipital carina complete. Lateral of pronotum moderately arched. Mesonotum triangulate and short; not divided into mesoscutum and scutellum. Propodeal disc weakly constricted anteriorly. Mesopleuron expanded laterally; subtegular groove long. Mid tibia very spinose.

Scaphepyris rufus Kieffer, 1904 (Figs. 1–8) Diagnosis. Female. Antenna compact. Mandible with three apical teeth. Clypeus with high median carina. Pronotal disc separated from mesonotum by large and deep depression. Propodeal disc with only lateral carina defined. Fore basitarsus with distal margin spinose. Description. Body length 4.00 mm. Color. Head, mesosoma and metasoma castaneous; antenna, mandible, palpi and legs light castaneous. Head (Fig. 1). Mandible with three apical teeth (Fig. 2). Clypeus with median lobe projected, apical margin subangulate, median carina high, forming keel in lateral view. Antenna compact (Fig. 3); first four antennal segments in ratio of about 40:8:8:7; segment III 1.25× as wide as long; segment XI 1.71× as wide as long. Frons coriaceous with large punctures separated each other by about 1.0× own diameter. Eye reduced and circular. Ocelli absent. WH 0.73× LH; WF 0.89× WH; WF 9.17× HE; VOL 12.33× HE. Temple convex in dorsal profile. Vertex slightly convex, corner rounded. Occipital carina complete, not visible in dorsal view. Mesosoma (Fig. 4). Thorax coriaceous. Pronotal disc long and separated from mesonotum by large and deep depression; anterior carina absent; pronotal collar large. Mesonotum short, triangular, not divided into mesoscutum and scutellum; without notaulus, parapsidal furrow, groove or pits. Metanotum absent medially. Propodeal disc 1.55× as long as wide; maximum width of propodeum about 1.57× width at constriction; transverse carina and discal carinae absent, lateral carina defined; spiracle circular, located on lateral of propodeum; declivity without median and lateral carinae. Mesopleuron large in dorsal view; weakly coriaceous; upper groove present; episternal furrow, mesopleural pit, anterior fovea and prepectal and postpectal carinae absent (Fig. 5). Tegula and wings absent. Fore femur 2.5× as long as wide; fore tibia not spinose; fore basitarsus with some spines on distal margin (Fig. 6). Mid tibia strongly spinose (Fig. 7). Hind tibia not spinose (Fig. 8). Metasoma. Long, 1.77x mesosoma; not petiolate, coriaceous. Material examined. Holotype, 1 &, [INDONESIA], Sumatra, Si-Rambé, xii.[18]90-iii.[18]91, E. Modigliani col. (MCSN). REDESCRIPTION AND PLACEMENT OF SCAPHEPYRIS

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FIGURES 1–8. Scaphepyris rufus female. 1. Head in dorsal view; 2. Mandible in frontal view; 3. Antenna; 4. Thorax in dorsal view; 5. Mesopleuron in lateral view; 6. Fore leg; 7. Mid leg; 8. Hind leg. (scale bar = 0.5 mm)

Discussion The monotypic genus Scaphepyris is now the 24rd assigned to Pristocerinae and the only one known exclusively from females. The eight genera keyed above are known from both males and females, but the remaining 14 genera are known only from males. So it might be possible that the female of Scaphepyris represents one of these 14 genera since the sexual dimorphism is strong in Pristocerinae, particularly Protisobrachium Benoit and Neoapenesia Terayama because they are the only two genera from Oriental region with female unknown. Scaphepyris rufus was not formally described; Kieffer (1905) just indicated it as a new genus and species in a key. Kieffer (1914) classified it as Sclerodermini because it is wingless and curiously compared it with Pristocera, which, at the time, belonged to another tribe. Scaphepyris is recognized here as belonging to the Pristocerinae. This conclusion is based on the presence of the diagnostic characters shared with females of other genera of this subfamily, such as the eyes being extremely reduced, the constriction between the propodeum and the mesonotum, and the mesopleuron clearly visible in dorsal view. The female of Scaphepyris also lacks ocelli and wings. These conditions also are shared with females of many species of Sclerodermini and probably led Kieffer (1914) to misclassify it as Sclerodermini.


LANES & AZEVEDO

Scaphepyris runs to Apenesia in the keys proposed by Evans (1964) and Terayama (2003) by having the propodeal disc weakly constricted, the mid tibiae very spinose, and the mesopleuron large in dorsal view, but Scaphepyris has the mesopleuron with an elongate upper groove, the clypeus with median lobe very projected and its apical margin subangulate, whereas Apenesia has the mesopleuron smooth, clypeus with median lobe short or somewhat projected and its apical margin emarginated or straight. Scaphepyris females differ from Pseudisobrachium and Prosapenesia females by having the eyes with more than one facet, propodeal constriction placed at or near the spiracles, and a short mesonotum, whereas Pseudisobrachium and Prosapenesia have the eyes absent or with a single facet, the propodeal constriction at anterior end, and an elongate mesonotum. Scaphepyris females differ from Dissomphalus females by having the propodeum with an evident constriction at the spiracles, and the mesopleuron expanded laterally, whereas Dissomphalus has the propodeum more or less parallel sided and the mesopleuron not expanded laterally. Scaphepyris females differ from Acrepyris, Pristocera and Caloapenesia females by having the propodeal disc weak to moderately constrict, whereas these latter genera have strong anterior constriction in propodeal disc. They also differ from females of Afgoiogfa and Parascleroderma by having the body weakly flattened, the lateral margins of the pronotum weak to moderately arched, and the mid tibia spinose above, whereas Afgoiogfa and Parascleroderma have the body strongly flattened, the lateral margins of the pronotum strongly arched, and the mid tibia smooth above. All species of the bethylid subfamily Pristocerinae have small wingless females (Krombein 1989), which makes the association between females and males a difficult task, not only at species level, but also at the generic level. So, collecting couples in copula has been most useful in clarifying the sex associations in this subfamily. The systematics of Pristocerinae has been based strongly on males, because females have reduction of many morphological characters. Females, however, can contribute to an understanding of the generic concept in Pristocerinae, and are important to the systematics of the group even though they are poorly known and rarely collected (Azevedo 2004).

Acknowledgments We are most grateful to Roberto Poggi (MCSN) for the loan of the material, to Paula Hebling Dutra for revising the English; to CNPq for the M.Sc. scholarship from Taxonomy Program to the first author, and for fellowship to the second author and financial support (CNPq grant # 303216/2004-2).

References Azevedo, C.O. (2004) A new species of Caloapenesia (Insecta, Hymenoptera, Bethylidae) from Vietnam, with discovery of the female of the genus. Spixiana, 27, 143–146. Berland, L. (1928) Bethylidae. In Fauna de France 19 - Hyménoptères vespiformes II. Office Central de Faunistique, Paris. pp. 1–206 Evans, H.E. (1963) A revision of the genus Pristocera in the Americas (Hymenoptera, Bethylidae). Bulletin of the Museum of Comparative Zoology, 129(4), 241–90. Evans, H.E. (1964) A synopsis of the American Bethylidae (Hymenoptera, Aculeata). Bulletin Museum of Comparative Zoology, 132, 1–222. Harris, R.A. (1979) A glossary of surface sculpturing. Occasional Papers in Entomology, 28, 1–31. Kieffer, J.J. (1905) Description de nouveaux Proctotrypides exotiques. Annales de la Société scientifique de Bruxelles, 29, 95–142. Kieffer, J.J. (1914) Bethylidae. Das Tierreich, 41, 228–595. Krombein, K.V. (1989) Systematic notes on some Bethylidae from Botswana Pristocerinae (Hymenoptera: Aculeata). Proceedings of the Entomological Society of Washington, 91, 620–631. Terayama, M. (1995) The Phylogeny of the Bethylid Wasp Tribe Sclerodermini (Hymenoptera, Bethylidae). Proceedins

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of the Japanese Society of Systematic Zoology, 54, 65–73. Terayama, M. (1996) The Phylogeny of the Bethylid wasp subfamily Pristocerinae (Hymenoptera, Bethylidae). Japanese Journal of Entomology, 64 (3), 587–601. Terayama, M. (2003) Phylogenetic Systematics of the Family Bethylidae (Insecta: Hymenoptera) Part II. Keys to subfamilies, tribes and genera in the world. The Academic Reports of the Faculty of Engineering of Tokyo Polytechnic University, 26, 16–29. Yasumatsu, K. (1955) Taxonomic notes on three wireworm parasites of the genus Pristocera from the Far East (Hymenoptera: Bethylidae). Journal of the Faculty of Agriculture Kyushu University, 10, 233–249.


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