HYM. RES. J. Vol. 15(2), 2006, pp. 317-347 The Species of Sternanlopius Fischer (Hymenoptera: Braconidae, Opiinae) and the Braconid Sternaulus Robert A. Wharton Department of Entomology, Texas A&M University, College Station, TX 77843, USA email: [email protected] — Abstract. The opiinebraconid genus Sternaulopius Fischer is recognized as valid, redefined, and one new species, Sternaulopius duplicatus, from Madagascar, is described. The type species, Sternaulopius bisternaulicus Fischer, is fully redescribed,withnewhostand distribution records. The only other previously included species, Sternaulopius beieri Fischer, is placed as a junior subjective synonym of Opius bajulus Haliday, new synonym. Opius bajulus is also redescribed and the genus group name Biophthora Foerster is resurrected for this species. Opius rossicus Szepligeti, is transferred tothegenus Biophthora (new combination) and Opius castaneusGranger, typespeciesof Frekius Fischer, is transferred to Utetes Foerster (new combination). Thus, Frekius is a junior subjective synonym of Utetes, new synonym, but is retained as a valid subgenus. Biophthora and Sternaulopius are compared to Xynobius Foerster (where Opius bajulus has frequently been placed), and Xynobius is redefined and treated as a subgenus of Eurytenes Foerster. Stigmatopoea Fischer is also recognized as a valid subgenus of Eurytenes. Characters used todefine these genus-group taxa are discussed in detail, with emphasis on venation, placement of metasomal spiracles, and sculptural details of the body. Use of the term sternaulus for a longitudinal groove on the ventral mesopleuron in Ichneumonoidea is reviewed, and it is shown that the sternaulus in cryptine and mesochorine Ichneumonidae is not homologous to the precoxal sulcus in Braconidae based on dissections of associated thoracic musculature. A true sternaulus, defined internally as the ridge supporting the origin of the mesopleural-basalare muscle, is rarely present in Braconidae. The genus Sternaulopius Fischer, 1965 Opiinae is still unsettled. Wharton (1988) was described to accomodate a single observed that, except for the double ster- species from the Democratic Republic of naulus, the two described species more the Congo. Subsequently, Fischer (1968) closely resembled other species within described a second species from Germany. Opius Wesmael s. /. than they did each No additional species have been described, other. On this relatively limited evidence, and only five specimens have been re- Wharton (1988) transferred both species to corded (Fischer 1965, 1972, Quicke et al. Opius, thus treating Sternaulopius as a syn- 1997). The two species that have been onym of Opius while at the same time included in Sternaulopius (Figs 1-20) have noting that Opius s. I. was not demonstra- two distinct grooves (sternauli) on each bly monophyletic. Quicke et al. (1997) side of the mesopleuron but in nearly all recognized six genera that were formerly other braconid wasps only a single groove treated by Fischer (1972, 1977, 1987) as is present, or the groove is completely lost subgenera of Opius, and reported that the (exceptions include Trigastrotheca laikipien- venom apparatus of a specimen of Sternau- sis Quicke and some species of Pambolus lopius beieri Fischer resembled that ofsome, Haliday). These two species of Sternnulo- but not all of the species that they included pius are thus distinctive, though their in one of these genera, Xynobius Foerster. placement in the classification of the Nothing else has been published on Ster- 318 Journalof Hymenoptera Research Figs 1-4. Sternaulopius bisternaulicus reared from Strombosia fruits: 1, face. 2, lateral view of head. 3, dorsal view ofhead. 4, right mandible, with oblique views ofclypeus and labrum. naulopius, undoubtedly due to the paucity the mesopleuron of Braconidae, while of individuals available for study. Van other braconid specialists have used the Achterberg (2004) recently re-characterized name precoxal suture/sulcus (notably van Xynobius, but made no mention of Sternau- Achterberg 1975, 1993, Shaw and Hud- lopius. dleston 1991) or simply longitudinal fur- The term sternaulus has long been used row of mesopleuron (Muesebeck 1970). by students of Ichneumonidae to describe The term precoxal sulcus has often been a longitudinal groove on the lower part of applied more widely in the Hymenoptera, the mesopleuron extending posteriorly though not necessary consistently (com- from the ventral-lateral region of the pare Richards 1956 with Bohart and epicnemial (= prepectal) carina towards Menke 1976). Gibson et al. (1998) treat the coxifer (or pleural coxal process) the sternaulus as a synonym of the (Viereck 1916, Richards 1956, Townes transepisternal line/sulcus (though their 1969). Several workers (e.g. Granger focus is on Chalcidoidea). Fischer (1972) 1949, Fischer 1958, Mason 1964, Marsh noted that it was probably incorrect to 1971, Wharton et al. 1997) have also equate the sternaulus with the precoxal applied this term to a similar groove on sulcus in Braconidae, and van Achterberg Volume 15, Number 2, 2006 319 Figs 5-10. Stemaulopius bisternaulicus reared from Strombosia fruits: 5, left side of mesosoma, left arrow = propleural flange, upper rightarrow = precoxal sulcus, lower rightarrow = sternaulus. 6, ventral-lateral view of mesopleuron, upper arrow = precoxal sulcus, lower arrow = sternaulus. 7, dorsal view of mesosoma. 8, dorsal viewofpronotum,upperarrow = obliquegrooveon propleuron,lowerarrow = leftnotaulus.9,dorsal view of metanotum and propodeum. 10, dorsal view ofpetiole. (1993) illustrated the two as separate be observed in some Opiinae (undoubt- structures for braconids. In a newsletter, edly referring to Stemaulopius), and Pam- van Achterberg (1994) stated more specif- bolinae where "the sternaulus is clearly ically that the difference between the bent downwards anteriorly and situated sternaulus and the precoxal sulcus can more ventrad." 320 Journalof Hymenoptera Research Figs 11-12. Sternaulopius bisternaulicus reared from Strombosia fruits: 11, lateral view of metasoma. 12, same showingdorsope (leftarrow) and spiracle on second tergum (right arrow). There are no published host records for MATERIALS AND METHODS Sternaulopius (with the exception of in- formation I have recently included on Specimens of Sternaulopius (sensu a website: hymenoptera.tamu.edu/parof- Fischer 1972, 1987) were either reared from fit). However, examination of material fruits infested with Tephritidae or bor- reared from fruit in both Cameroon and rowed from museums. Rearing methods, Kenya (Steck et al. 1983, Copeland et al. localities, and methods of identification of 2002), indicates that tephritid fruit flies flies and plants are described in Steck et al. (Diptera) are the hosts of at least some of (1983) and Copeland et al. (2002). Material the species of Sternaulopius. This reared was borrowed from the Koninklijk Muse- um material forms the basis for the present voor Midden-Afrika, Tervuren, Bel- treatment, including a preliminary exami- gium (MRAC), Museum fur Naturkunde nation of the nature of the sternaulus in der Humboldt-Universitaet, Berlin, Ger- Ichneumonoidea. many (ZMHB), Hungarian Natural History Volume 15, Number 2, 2006 321 Fig. 13. Sternaulopius bisternaulicus reared from Strombosia fruits, ovipositor and ovipositor sheaths. Museum, Budapest (HNHM), Museum Texas A&M University were dissected to National d'Histoire Naturelle, Paris examine the internal musculature of the (MNHN), and the U. S. National Museum mesothorax. All dissected material was of Natural History (USNM). Additional initially stored in 70-80% ethanol, then specimens examined are in the Naturhis- either air dried following transfer through torisches Museum Wien, Austria (NHMVV) 95% ethanol and 99% amyl acetate or and the National Museum of Ireland, dissected while still in alcohol. Specimens Dublin (NMID). of Apis mellifera L. and several other Fly puparia were individually isolated in Apocrita collected in central Texas with Kenya by R. Copeland from one heavily- Malaise traps were also dissected to ascer- infested sampleofStrombosia scheffleri Engl. tain the pattern of general thoracic muscu- (Olacaceae) fruit, collected in Kakamega lature, using the works of Daly (1964) and Forest, Kenya on 30th April 2000. Associ- Gibson (1985) for orientation and identifi- ation ofwasps with hosts was made by the cation of major muscles. All identifica- author, based on characteristics of the tions were made by the author; voucher A&M puparia. All otherhost records listed below specimens are deposited as Texas are based on wasps that emerged from University voucher number 656 in the A&M tephritid puparia that were not individu- Texas University Insect Collection ally isolated. (TAMU). Numerous specimens of Braconidae and Measurements are as in Wharton (1977, Ichneumonidae from material housed at 1986); terminology for descriptions of 322 Journalof Hymenoptera Research V 14 15 Figs 14-16. 14, 15, Fore and hind wings of Sternaulopius bisternaulicus. 16, Hind wing of Biophthora bajulus, arrow = 2-1A. venation and external features of the body precoxal sulcus (following van Achterberg follows Sharkey and Wharton (1997), ex- 1993). Details of the mesothorax associated cept that in the present paper a distinction with the musculature are provided in the is made between the sternaulus and the results and discussion where additional Volume 15, Number 2, 2006 323 Figs 17-20. Sternaulopius beieriholotypefemale (= B. bajulus): 17, lateral habitus. 18, face. 19, dorsal-posterior viewofposteriorpartofmesosomashowingmesonotal midpitandsculptureofscutellumand propodeum. 20, lateral obliqueview ofmesosoma, upper arrow = precoxal sulcus, lower arrow = sternaulus. terms, when used, are specifically defined. also apply to many other taxa both within Thoracic muscles relevant to discussions of and outside apocritan Hymenoptera, as the sternaulus are noted in Figs 21-32. Of evidenced by the detailed studies of Daly particular focus are various axillary mus- (1964) and Gibson (1985, 1986a, b, 1993). cles and especially the mesopleural-basa- The major muscles discussed below for the lare muscle (sensu Gibson 1985). The latter Ichneumonoidea were roughly similar in originates on the wall of the mesopleuron their points of origin and insertion in the and inserts on the basalare sclerite near the few aculeates and non-aculeate apocritans base of the wing, at least in all the that were dissected (namely A. mellifera, Ichneumonoidea (22 genera) and Aculeata Khopalosoma nearcticum Brues, and unde- (4 genera) examined (Table 1). termined species of Gasteruption Latreille and Polities Latreille). Many of the details RESULTS AND DISCUSSION of thoracic musculature found in such excellent works as those of Gibson (1985, Musculature and associated 1993) have been omitted here since the external features major interest is to provide a better char- The observations made here refer specif- acterization of the sternaulus. ically to the Ichneumonoidea (Ichneumo- On the mesopleuron, the sternaulus and nidae + Braconidae). However, they will various pits, depressions, or other changes 324 Journalof Hymenoptera Research Volume 15, Number 2, 2006 325 in elevation are, to a greater or lesser recent taxonomic treatments), rather than extent, the external representation of the parapsidal furrows. attachment of the thoracic muscles. In- Immediately laterad the dorsoventral ternally, the mesothorax, at least in ichneu- indirect flight muscle is the smaller monoids and aculeates, is dominated by (though still fairly large) mesopleural- the massive dorsolongitudinal (Fig. 21) basalare muscle (Figs 23, 25-29, 31). This and dorsoventral (Fig. 22) indirect flight is a distinctive band of muscle fibers muscles. The former originates largely readily identified by the dorsal, sclerotized from the median mesoscutal lobe (and cap that is strongly constricted to form partly from the phragma along the anterior a tendon-like attachment connected to the margin ofthe lobe) and extends posteriorly basalare (Figs 27, 29, 31). In Ichneumonoi- through the middle of the mesothorax. On dea, the mesopleural-basalare originates either side of this median muscle mass is ventrally in the trough of the mesothorax a broad band of dorsoventral muscle fibers just laterad the dorsoventral muscle or it (Fig. 22), which insert on the lateral lobes may arise somewhat higher on the vertical of the mesoscutum and originate ventrally wall of the mesepisternum. The subtegular on the mesothorax. The floor of the (= subalar) ridge, immediately ventrad the mesothorax, where the dorsoventral mus- anterior subalar depression of van Achter- cles originate, is a trough interrupted berg (1988), is the external representation medially by a phragma representing the of the internal pocket in which resides the midventral invagination of the mesoster- sclerotized cap of the mesopleural-basalare num (partially shown in Figs 25 and 29). muscle. The epicnemial carina, when pres- The lateral and ventral sides of the trough ent, delineates the anterior border of this are formed on each side of the body by the muscle mass ventrally, and dorsally it is relatively sharp vertical to horizontal tran- delineated by the pronotal-mesopleural sition in the mesepisternal wall ventrally as suture. it curves towards the ventral midline. The mesopleural-basalare muscle origi- Notauli are perhaps best defined as nates on the sternaulus in those members external grooves on the mesoscutum in- of the Ichneumonidae that possess a ster- dicating the position of the internal ridge naulus (Figs 25, 28), but does not originate or phragma that separates the attachment on the structure that has frequently (e.g. pointofthe dorsolongitudinal muscle mass Sharkey and Wharton 1997) been called the from the attachmentpoints of the right and sternaulus in Braconidae (Figs 29-30). In left dorsoventral muscle bundles (see the the ichneumonid taxa Cryptini and Meso- excellent discussion in Gibson (1985) for chorinae, both of which have a well-de- terminology of mesonotal grooves). In veloped sternaulus ventrolateral^ on the Ichneumonoidea, the position of these mesepisternum, the mesopleural-basalare grooves, when present, relative to the muscle originates on the internal ledge attachment points of these two indirect formed by the anterior portion of the flight muscles confirms that they should be sternaulus as the latter extends posteriorly called notauli (as they are in nearly all from the epicnemial carina. In those mem- mesopleuron, dried specimen, left arrow showing mesopleural-basalare muscle, right arrow showing internal ledge of short sternaulus on which anterior end of mesopleural-basalare originates. 26, Enicospilus sp., Ophioninae, right side of body in ethanol, showing origin of elongate mesopleural-basalare in trough ofmesopleuron. 326 Journalof Hymenoptera Research Figs 27-32. Longitudinal sections through mesosomas of Ichneumonoidea, showing muscles originating on mesopleuron:27-28,Ichneumonidae,Cryptinae,rightsideofbody:27,inethanolshowingmesopleural-basalare muscle(arrow)anditsdorsalcap;28,samespecimen,dried(resultingingapsinmusclebundle),showingorigin of mesopleural-basalare muscle on internal ledge formed by sternaulus (arrow). 29-30, Fopius vandenboschi (Braconidae, Opiinae), dried: 29, ventral and left portions of body, bottom arrow = midventral phragma (partiallybroken in dissection), top arrow = mesopleural-basalaremuscle; 30, rightsideofbody, mesopleural- basalare removed, bottom arrow = region where mesopleural-basalare originates, middle arrow == precoxal sulcusvisiblethroughcuticle,toptwoarrows = musclesinsertingonaxillarysclerites.31,Wroughtoniaferruginea