Biting the bullet: revisionary notes on the Oraseminae of the Old World (Hymenoptera, Chalcidoidea, Eucharitidae)

Twelve genera of Oraseminae (Hymenoptera: Eucharitidae) are recognized in the Old World. The genus Orasema Cameron is now considered as found only in the New World, and the Old World species, previously treated as species groups, are now treated as distinct genera. Eight new genera are proposed: Australosema gen. n., Cymosema gen. n., Hayatosema gen. n., Ibitya gen. n., Ivieosema gen. n., Leiosema gen. n., Matantas gen. n., and Zuparka gen. n. The genus Losbanus Watanabe is given revised status from Orasema. Nine new species are proposed: Australosema politurae sp. n. (Australia), A. verghetta sp. n. (Australia), Cymosema capelina sp. n. (Australia), C. waterworthae sp. n. (Australia), Hayatosema plicator sp. n. (Vietnam), Ivieosema confluens sp. n. (Madagascar), Iv. limula sp. n. (Madagascar), Leiosema lesiolouna sp. n. (Republic of Congo), and Zuparka fisheri sp. n. (Madagascar). New combinations (from Orasema) include Australosema valgius (Walker, 1839), A. synempora (Heraty, 1994), Hayatosema assectator (Kerrich, 1963), H. delhiensis (Narendran & G. Kumar, 2005), H. initiator (Kerrich, 1963), H. kailashi (G. Kumar & Sureshan, 2015), H. nigra (Heraty, 1994), H. nirupama (G. Kumar & Narendran, 2007), H. siruvanica (G. Kumar & Sureshan, 2015), Ibitya communis (Risbec, 1952), Ib. seyrigi (Risbec, 1952), Ivieosema fraudulenta (Reichensperger, 1913), Iv. striatosoma (Heraty, 1994), Leiosema glabra (Heraty, 1994), Losbanus bouceki (Heraty, 1994), L. ishii (Heraty, 1994), L. promecea (Heraty, 1994), L. rugulosa (Heraty, 1994), Matantas koghisiana (Heraty, 1994), and Zuparka monomoria (Heraty, 2000). Losbanus uichancoi Ishii, 1932 is given revised status from Orasema. Distribution records, a new host association of Pheidole (Myrmicinae) for Ibitya communis, and an identification key to the Old World genera and species are provided. JHR 55: 139–188 (2017) doi: 10.3897/jhr.55.11482 http://jhr.pensoft.net Copyright Roger A. Burks et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. RESEARCH ARTICLE Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139–188 (2017) 140


Introduction
Oraseminae constitutes one of the major lineages of the family Eucharitidae, with the estimated number of species approaching 200 (Heraty 2002), all of which are parasitoids of ants (Hymenoptera: Formicidae).There are verified host records of Oraseminae attacking six genera in the ant subfamily Myrmicinae and dubious records on Ecitoninae and Formicinae (Heraty 1994, 2002, Lachaud and Pérez-Lachaud 2012).Hosts for the Old World Oraseminae are species of the myrmicine genera Monomorium Mayr and Pheidole Westwood (Table 1).While Orasema Cameron is present throughout the New World with a northern-most record from southern Canada, Old World Oraseminae are known only from the tropics and subtropics, with northernmost records in India (New Delhi), Taiwan, Iriomote Island of Japan, and the southwestern highlands of Yemen (Heraty 1994(Heraty , 2002)).
The Old World species of Oraseminae were previously revised by Bouček (1988) and Heraty (1994), with a new species of Orasema from Madagascar added by Heraty (2000).A total of 36 species of Old World Oraseminae were placed into four genera: Indosema Husain & Agarwal, Orasema Cameron, Orasemorpha Bouček, and Timioderus Waterston.The Old World species of Orasema were grouped into six species groups by Heraty (1994), with two new groups added by Heraty (2000).Based on an analysis of 62 morphological and behavioral characters, Indosema, Timioderus and Orasemorpha were monophyletic and sister to Orasema, with the Old World species groups of Orasema paraphyletic to the New World Orasema (Heraty 2000).Among these genera, Timioderus and Indosema were considered distinct because of fusion of the mesosoma across the transscutal articulation between the axillae, and because they have eggs that are cylindrical instead of stalked (Heraty 1994(Heraty , 2000)); all other known eggs of Eucharitidae are stalked with the exception of Gollumiellinae (Heraty et al. 2004).Both Timioderus and Orasemorpha are the only Oraseminae known to deposit multiple eggs into a single oviposition puncture (unknown in Indosema), whereas all known Orasema sensu lato deposit a single stalked egg into each oviposition puncture made by the specialized ovipositor (Das 1963, Heraty 1994, 2000, Ishii 1932).All Oraseminae can be defined by the presence of an enlarged, toothed and usually anteriorly curved ovipositor that is used to enlarge a hole in plant tissue to deposit their eggs.
When using molecular data, the New World species of Orasema are consistently monophyletic (Heraty et al. 2004, Murray et al. 2013).The Old World Oraseminae are either monophyletic (Heraty 1994, Heraty et al. 2004) or paraphyletic (Murray et al. 2013) to the New World Orasema.Monophyly of Orasema s.l.worldwide is never supported.More recent unpublished analyses consistently support either monophyly of the Old World Oraseminae (ribosomal analyses with 18S, 28S, COI), or their paraphyly (Anchored Enrichment, 366-411 loci) with respect to the New World Orasema, but Table 1.List of Old World Oraseminae, with their geographic distribution and ant host genus.For a more complete list of synonymies and host information see Heraty (1994Heraty ( , 2002) ) and Heraty (2017).Asterisk (*) denotes type species of genus.Australian refers to Australia including Tasmania.NR is new host record.
In support of several upcoming phylogenetic studies, we have decided to recognize the problems inherent in the Old World Orasema.Given the wealth of morphological differences in the three other genera of Oraseminae, the associated biological differences, and the lack of morphological and molecular support for Orasema s.l., the best approach is to fragment the Old World species groups of Orasema into different genera and recognize only the monophyletic New World clade as Orasema sensu stricto.Herein we recognize 12 Old World genera, of which eight are new and one, Losbanus, is given revised status.Several new species are described, bringing the number of Old World species to 45, which is far overshadowed by the incredible diversity of Orasema in the New World.

Materials and methods
Material was examined from ANIC: Australian National Insect Collection, Canberra; BMNH: Museum of Natural History, London, UK; BPBM: Bernice P. Bishop Museum, Honolulu, HI, USA; CASC: California Academy of Sciences, San Francisco, CA, USA; CNCI: Canadian National Collection of Insects, Ottawa, Canada; MNHP: Muséum National d'Histoire Naturelle, Paris, France; NMPC: National Museum (Natural History), Prague, Czech Republic; PPRI: Plant Protection Research Institute, Pretoria, South Africa; ROME: Royal Ontario Museum, Toronto, ON, Canada; TMP: Ditsong National Museum of Natural History (formerly Transvaal Museum), Pretoria, South Africa; UCRC: UCR Entomology Research Museum, Riverside, CA, USA; UNMO: University of Montana, Bozeman, MT, USA; USNM: National Museum of Natural History, Washington, DC, USA.Each specimen is labeled with a UCRCENT or CASENT barcode specimen identification number; UCRCENT codes include an additional reference to the museum of deposition.Georeferenced points for specimens estimated from Google Earth are italicized.Photographs were taken using a Leica Imaging System with a Z16 APO A microscope, and stacked using Zerene Stacker (version 1.04, Zerene Systems, LLC).
Morphological terms follow Heraty et al. (2013) and Burks et al. (2015).All terms were verified using the Hymenoptera Anatomy Ontology (http://portal.hymao.org/)when they are represented in that resource, and variations from their preferred terms are discussed here: palpomere formula follows a maxillary:labial format; mandibular tooth count follows a right:left format; surface sculptural terms follow illustrations provided by Harris (1979).Important features include the base of the petiole, which can either be tapered to the articulating condyle within the propodeal foramen (Figs 20,25) or truncate basally external to the foramen (Figs 5,11,67,75), and if truncate then with an anterior transverse basal carina (apc,Figs 5,67,75).

Key to Oraseminae of the
Discussion.Defined by Heraty (1994) as the Orasema valgius group for two species from Australia, and was distinguished from the glabra and koghisiana groups in part by an at least partly reticulate face and a labrum with 4 digits (Heraty, 1994).Australosema politurae and related species expand the definition of the group to include an almost or completely smooth face.Originally recognized to include two species, Australosema synempora (Heraty) and A. valgius (Walker).Herein we add two new species, A. politurae and A. verghetta.
Diagnosis.Separated from Orasema s.s. by a relatively weak or absent basal petiolar flange, and smooth lateral areas on the propodeal disc in combination with a rugoseareolate or reticulate median channel.Distinguished from other Old World genera by the head subtriangular in frontal view (Fig. 2); scrobal depression evenly impressed and lacking parallel channels or dorsal depressions; dorsal occipital margin abrupt and rounded or carinate; funicle 7-segmented; labrum with 4-digits; mesonotum with only minute setae; mesoscutal lateral lobes and frenum sculptured or smooth; transscutal articulation complete and relatively straight; propodeal disc with lateral areas at most weakly sculptured (Fig. 5), but usually smooth (Fig. 11), and with a broad sculptured median channel; prepectus foveate and tightly articulated with pronotum ventrally (Fig. 12); fore wing with basal area bare and specular area either bare or pilose (Figs 1, 7), wing disc with very short setae; postmarginal vein much longer than stigmal vein and reaching about half distance to wing apex; petiole base truncate with weak basal flange (Fig. 5); antecostal sulcus present and smooth, at most weakly foveate laterally; first valvula of ovipositor with 3-4 lateral teeth and robust oblique subapical carina.Male scape lacking ventral pores.
Similar genera that have a smooth face and smooth lateral regions on the propodeal disc include Leiosema, Matantas, and Zuparka.Australosema is distinguished from Leiosema and Zuparka by several features, including the absence of a parallel channeled scrobes with dorsal depressions, 3-4 large teeth on the ventral (first) valvula of the ovipositor instead of 9-10 minute teeth, and absence of ventral pores on the male scape.It differs from Matantas primarily by having a 4-digitate rather than 6-8 digitate labrum.
Diagnosis.Differs from most Australosema by having an almost entirely smooth face, with the frons not swollen and the dorsal occipital carina weak.It differs from A. synempora in several features, including having a completely sculptured axilla and mesoscutal lateral lobe and a less densely setose fore wing.Matantas koghisiana also has a smooth face and frenum, but A. politurae has four labral digits and a more elongate mesosoma.Additional features include the lateral lobe of mesoscutum almost entirely rugose-reticulate, smooth near posterior edge; axilla reticulate (Fig. 4); mesepisternum not swollen anterior to mesocoxae (Fig. 1).Propodeum sublaterally nearly smooth (Fig. 5), and in the female the F2 is longer than F3 and expanding apically (Fig. 3).
Metasoma.Petiole 4.0-4.8×as long as broad, 1.6-1.7×as long as metacoxa, finely longitudinally rugose-reticulate, with some incomplete longitudinal carinae laterally and ventrally (Fig. 5).Antecostal sulcus of Gs 1 smooth, very shallow medially; acrosternite smooth, uniformly rounded or with a transverse elevation near midlength.Cercus with a curved seta that is much longer than the others.Hypopygium with a few short setae and a slightly longer posterolateral seta on each side.Ovipositor yellow medially, orange laterally, dorsal valve with 7-8 medially interrupted sawtooth-like annuli; ventral valve with 3-4 weakly defined teeth along lateral carina, and with oblique lateral carina immediately basal to toothed apex (as in Fig. 7).
Male.Length 2.1-2.6 mm.Antennal flagellum (Fig. 6) densely covered with long decumbent setae, with longitudinal sensilla extending beyond funicular apices, funiculars separated by stalk-like constrictions; F7 as long or longer than F6, shorter than F8; scape and pedicel yellow, flagellum brown.Mesoscutal midlobe transversely rugosereticulate; mesoscutellum rugose-areolate with underlying reticulate sculpture.Petiole 5.7-6.8× as long as broad, 1.9-2.4×as long as metacoxa, narrowest slightly beyond midlength, finely reticulate but becoming smooth posteriorly.Similar species.A single specimen (gaster missing, likely female; UCR-CENT0092152, D0161) from southeast QLD is molecularly similar and is placed as the sister of A. politurae.It is distinct by having a completely smooth face with a strongly swollen frons.Two other series (all males from QLD) likely represent another two species that share similar features to the D0161 specimen.All specimens have a 4-digitate labrum, 7 funiculars, a completely smooth face, and a strongly carinate dorsal occipital margin.Etymology.Italian noun used in heraldry to refer to a line along the middle, referring to the longitudinal scutellar groove present in females.

Australosema verghetta
Diagnosis.Differs from other Australosema by the presence of a longitudinal median groove on the frenum of females, and similar to some of the included species, the frons is rugose-reticulate and the lower face smooth.The female also has F2 much narrower than F4, not longer than F3, with few longitudinal sensilla (Fig. 9).Males are very similar to those of A. valgius (Walker), but differ in having a mostly or entirely dark brown scape (Fig. 8), while that of A. valgius is almost entirely yellow.
Female.Length 2.1-2.6 mm.Head and mesosoma metallic green to blue-green, metasoma brown with metallic green luster.Scape and pedicel dark brown; anellus pale brown; flagellum dark brown.Coxae metallic green basally, becoming brownish to yellowish apically, remainder of leg yellowish except femora mostly dark brown and with yellowish tips.Fore wing venation brown, setae dark.Gaster metallic green.
Male.Length 2.4-2.6 mm.Scape dark brown for most or all of its length (Fig. 14), yellowish in only a small basal area if at all; antennal flagellum densely covered with short decumbent setae, with longitudinal sensilla extending beyond funicular apices; F7 shorter than F6 and F8; entire antenna dark brown and scape with metallic luster.Frenum without longitudinal groove; axillula with reticulate sculpture and longitudinal carinae.Metatibia dark brown except at apices.Petiole 5.9-7.4× as long as broad, 2.2-2.6× as long as metacoxa, narrow at midlength, reticulate.Etymology.Based on the Greek κύμα (kyma) for wave and referring to the sinuate transscutal articulation; gender feminine.
Discussion.The two species in this group are each known from single specimens collected in Australia, both among the smallest known Eucharitidae.They differ from previously described species of Orasema by having a cylindrical ventrally fused petiole that tapers basally to the articulating condyle (Figs 20,25), reduced mouthparts (Figs 16,21), and advanced axillae (Figs 19,24).The mesoscutum and mesoscutellum are separated by a distinct transscutal articulation, but the sclerites are tightly associated and the axillae are advanced (thus somewhat resembling the state in Timioderus and Indosema).Based on morphological (reduced anellus, reduced mouthparts, smooth head and mesosoma) and molecular evidence, they are the potential sister group of Indosema.The fore wing shape is unique within Eucharitidae, with the anterior leading edge of the fore wing anteriorly curved at the stigmal vein giving the wing an oval-shaped appearance (Fig. 27); in other Eucharitidae the wing margin beyond the stigmal vein is straight (Fig. 7).Male unknown.
Diagnosis.Recognized from most Oraseminae by the sinuate closely associated transscutal articulation, minute discoidal anellus, reduced mouthparts that are at most bidentate and medially directed (not falcate), basally tapered petiole and elongate slightly curved ovipositor.Distinguished from other Old World genera by the head transverse in frontal view and mostly smooth (Figs 16,21); scrobal depression strongly impressed and forming weak parallel channels, but without dorsal foveae; dorsal occipital margin abrupt and rounded; funicle 7-segmented; mandibles, chisel-shaped and slightly broader than long or bidentate with both teeth projecting medially (not falcate as in other Eucharitidae); labrum appears to be membranous and without digits; palpi greatly reduced; mesonotum appearing bare, at most with minute setae; mesoscutal lateral lobes entirely smooth or weakly reticulate laterally; frenum smooth; transscutal articulation sinuate and distinct but sclerites closely associated and appearing fused; callus bare; propodeal disc smooth (Fig. 20) or weakly reticulate with single median carina (Fig. 24); prepectus foveate and loosely articulated with pronotum ventrally (Fig. 18); fore wing with basal area and specular area evenly covered with minute setae, and anterior margin of fore wing disc curved at stigmal vein (Fig. 27); postmarginal vein short and about 1.5× as long as stigmal vein; petiole base tapered and without transverse flange (Figs 20,25); petiole fused ventrally, basally tapering to condyle and without basal flange; acrosternite swollen and finely reticulate, antecostal sulcus not apparent, but a distinct transition to the smooth posterior region of Gs 1 ; ovipositor long and slightly curved anteriorly (Fig. 26); first valvula with 3-4 lateral teeth and with a strong oblique subapical ridge.
Host association.Unknown.Distribution.Two species.Australia (Queensland and Western Australia).Female.Length 1.3 mm.Head and mesosoma dark brown with very faint metallic luster.Scape, pedicel and anellus pale brown, funiculars and clava brown.Leg brown to milky brown, with slightly darker areas on the coxa basally, femur except at its tips, tibia subbasally, and last two tarsomeres.Fore wing hyaline; venation pale brown, setae dark.Gaster brown with faint metallic luster.
Mesosoma .Mesoscutum smooth and glossy, notauli evenly impressed and smooth.Axilla strongly advanced, smooth; scutoscutellar suture finely foveate; mesoscutellar disc and axillula smooth, without axillular sulcus; frenum distinguished by a slight change in elevation at frenal line, smooth.Propodeal disc with median carina, weakly sculptured laterally; callus bare, separated from propodeal disc by wrinkle-like longitudinal carinae.Metacoxa smooth to shallowly reticulate.Fore wing spatulate, 2.6× as long as broad; costal cell with 1 complete row of ventral setae; submarginal vein with several long hairs; basal cell setose dorsally and ventrally; speculum absent; cubital and subcubital folds setose; marginal fringe present; stigmal vein 1.5× as long as broad; dorsal setae of disc much longer than ventral setae; postmarginal vein about 1.5× as long as stigmal vein.Hind wing costal cell with a few setae in apical third.
Metasoma.Petiole 0.7× as long as metacoxa, 1.8× as long as broad, smooth except for longitudinal carinae, including a median carina dorsally, narrowing anteriorly to abruptly expand at the anterior condyle, without anterior transverse carina, ventrally smooth (Fig. 20).Antecostal sulcus of Gs 1 crossed by many longitudinal carinae; acrosternite smooth.Cercus with all setae subequal in length.Hypopygium bare.Ovipositor only slightly exerted in holotype and features not distinguished.
Metasoma .Petiole 1.6× as long as broad, 1.0× as long as metacoxa, finely longitudinally rugose-reticulate, with a pair of complete lateral longitudinal carinae and some additional incomplete longitudinal carinae anteriorly, narrowing anteriorly to abruptly expand anteriorly at the condyle, without anterior transverse carina, ventrally with a longitudinal groove that expands at apices.Acrosternite with irregular raised sculpture, swollen and differentiated across indistinct antecostal sulcus from remaining smooth Gs 1 .Cercus with all setae subequal in length.Hypopygium bare.Ovipositor elongate and slightly recurved, dorsal valve with 7 medially interrupted annuli; ventral valve with 5-6 teeth, and with oblique carina ending in a notch immediately basal to toothed apex.
Discussion.This group and three of its seven previously described species were characterized as the Orasema assectator-group by Heraty (1994: 73-74).The four Indian species described in Narendran and Girish Kumar (2005), Girish Kumar and Narendran (2007) and Girish Kumar and Sureshan (2015) are readily placed in the assectator-group and hence in this genus (Table 1).Hayatosema is morphologically similar to species in the New World placed in the cockerelli and bakeri species groups.They were not resolved from these taxa based on the morphological analyses of Heraty (1994Heraty ( , 2000)), and molecular data clearly place them as distinct and placed with the other Old World genera (Murray et al. 2013, Heraty unpublished).The only consistent differentiating feature from Orasema is that the lateral aspect of the mesepisternum is reticulate with the ventral aspect smooth and shining in all Hayatosema, whereas if it is reticulate laterally in Orasema s.s., it is at least weakly sculptured ventrally.The antennal funicular count is unusual in the African species, with O. nigra Heraty having from 7-8 funiculars in females and 8-9 funiculars in males, all from the same collecting event.The Indo-Pacific species all have 7 funiculars in both sexes.Notably, molecular analyses fail to support the monophyly of the African and Indo-Pacific species, although sampling is currently very poor.

Etymology.
A noun meaning "one who folds clothing," derived from the Latin verb plicare (to fold).Refers to the rough propodeal and petiolar surface sculpture.
Female.Length 3.5 mm.Head, mesosoma, and petiole black with slight metallic blue to purple luster.Antenna pale yellow.Coxae black to dark brown, with faint metallic purple luster; femora dark brown except white at tips; tibiae and tarsi pale yellow.Fore wing venation yellowish-white.Gaster dark brown.
Mesosoma .Pronotum with small protrusion ventral to mesothoracic spiracle (Fig. 32: protrusion).Mesoscutum finely rugose-reticulate, posterior margin sinuate and midlobe strongly convex posteromedially, mesoscutal midlobe abruptly elevated above lateral lobes; notauli broadly and deeply impressed anteriorly.Axilla finely rugose-reticulate; mesoscutellar disc finely rugose-reticulate; frenal line as a deep groove with a carinate edge posteriorly; frenum finely rugose-reticulate; axillular sulcus not indicated, axillula finely rugose-reticulate.Mesepisternum anteriorly with a smooth area dorsolaterally, laterally finely reticulate and ventrally smooth; not overlapping prepectus; ventrally smooth and forming a rounded horizontal surface anterior to mesocoxae; upper mesepimeron elevated and smooth; transepimeral sulcus foveate; lower mesepimeron glossy, shallowly irregularly sculptured.Propodeal disc medially areolate with some underlying irregular sculpture, with irregular median carina that is present only for a short distance near propodeal midlength, sublaterally rugose-reticulate; callus bare, smooth over most of surface but coarsely areolate to foveate peripherally, including foveate depression separating metapleuron from hind wing base.Metacoxa rugose-reticulate, glossy posteriorly.Fore wing 2.4× as long as broad; costal cell with ventral row of setae apically, bare basally; basal cell with 2 ventral setae anteriorly; speculum present, partially crossed by a patch of ventral setae near midlength of fore wing; cubital fold with only a single ventral seta posterior to basal cell, bare posterior to speculum; subcubital fold bare posterior to basal cell, with only some ventral setae posterior to speculum; wing disc covered by dense long setae (Fig. 33); stigmal vein 2.0× as long as broad, only slightly expanded at uncus; postmarginal vein about 3.0× as long as stigmal vein.Hind wing costal cell with only a few ventral setae apically.
Discussion.Heraty (1994) proposed a uichancoi-group that included Ibitya seyrigi (Risbec) and I. communis (Risbec) as well as five other Indo-Pacific species of Orasema (now Losbanus).Based largely on all of the Malagasy species having more than 8 labral digits (Fig. 36), Heraty (2000) proposed a separate communis-group.Herein we recognize the two currently described communis-group species as the genus Ibitya, although other potential species from Madagascar have been identified.Illustrations and a key to species, as Orasema, are provided in Heraty (1994).
Diagnosis.Separated from other Oraseminae by having the apex of the first valvula of the ovipositor with sinuate ridges as opposed to distinct lateral teeth.Distinguished from other Old World genera by the head subtriangular in frontal view; face entirely smooth laterally, scrobal depression evenly impressed and lacking parallel channels and dorsal foveae; dorsal occipital margin abrupt and rounded or carinate; antenna with 7 or 8 funiculars (11-12 antennal segments); labrum with 8-10 digits (rarely 5) (Fig. 36); mesonotum appearing bare, at most with minute setae; mesoscutal lateral lobes and frenum evenly sculptured and never smooth and shining; transscutal articulation complete; propodeal disc evenly sculptured; prepectus foveate and tightly articulated with pronotum ventrally; fore wing with basal area pilose or bare, and with specular area pilose, wing disc with dense long setae; postmarginal vein much longer than stigmal vein and reaching beyond half distance to wing apex; petiole base truncate with weak basal flange; antecostal sulcus smooth and distinctly impressed; first valvula of ovipositor with 3-4 sinuate lateral ridges.Male scape lacking ventral pores.
Discussion.A new host record is made for Pheidole veteratrix Forel based on a single collection that included a soldier gripping the wing of an adult I. communis, 5 female and 1 male pupae, and two unparasitized ant larvae.The pupa has the basic features of an Orasema, but with additional pustulate swellings not found in other taxa.Pupa (Fig. 37).Length 5.1-5.5 mm.Typical morphology for Orasema, including three swollen tubercles over the petiolar region and pronounced linear swellings along the posterior margins of Mt 1-5 .Additional atypical tubercular swellings occur over each anellus, each side of the occiput, on the dorsal margin of each flagellomere, laterally on the anterior face of the mesoscutal midlobe, axilla, laterally (paired) on the mesoscutellum as well as a single medial posterior tubercle, and with paired lateral swellings associated with the linear swellings on each of Mt 1-5 .
New Discussion.The redescription and illustrations provided by Heraty (2002) are accurate for this species.The holotype was missing the antennae and the labrum was not visible.Based on the new material: antenna with 11 segments, all flagellomeres beyond anellus densely pilose with numerous longitudinal sensilla; flagellum 1.5× head height; 7 funiculars; F2 1.9× as long as broad, 1.1× as long as F3; clava as long as preceding two flagellomeres.Labrum with 9 long digits, each with a long apical seta, digits arranged in clusters of 1-1-3-3-1.Ovipositor as described for O. communis Risbec in Heraty (2002: fig. 82), but distinctly curved cephalad.
A single male of a different species (Reserve Betampona, from sifted litter; CAS, UCRCENT 00417456) also has an antenna with 11 segments and 7 funiculars, a labrum with 7 digits, and a long petiole about as long as the hind femur.The male shares many features with I. seyrigi, and may be the same species, however there are dramatic differences in especially the coarsely rugose scutellar disc.
New material examined.Madagascar: Fianarantsoa Prov.: Parc National Ranomafanae, radio tower at forest edge, 1130m, 21°15'03"S, 47°24'26"E, 6-17. vii.2003Discussion.Defined by Heraty (1994: 68) as the Orasema striatosoma-group.Ivieosema previously contained two mainland African species, which Heraty (1994) noted as being very different from other Orasema.The two new Malagasy species that we include are even more different from other Oraseminae in many ways, but share several features with Ivieosema striatosoma (Heraty), and to a lesser degree with Ivieosema fraudulenta (Reichensperger): a large pit immediately ventral to the median ocellus (Fig. 39), continuous striae extending from the axillae to the mesoscutellar disc (Figs 42, 52), propodeal shape and sculpture (Figs 41,43,52), and the laterally sinuate posterior margin of the gastral tergites 49).They differ from the African I. striatosoma and I. fraudulenta in several major ways, including the presence of two additional pairs of depressions between the eye and antennal scrobe, prepectus slightly swollen rather than weakly foveate, lack of any indication of an anterior transverse petiolar carina, and presence of a thickened basal fold in the fore wing.They also exhibit further reductions in surface sculpture, mouthparts and wing setation.All of the species have a closely appressed pronotum and prepectus that appear fused in the Malagasy species, a nearly quadrate anellus, and a thickened marginal vein.Notably the anterior margin of the fore wing beyond the stigmal vein is rounded similar to that of Cymosema.
Diagnosis.Separated from other Oraseminae by the deep sharply impressed pit just below the median ocellus (Fig. 39).Distinguished from other Old World genera by the head subtriangular in frontal view (Figs 39,50); scrobal depression below pit broadly impressed, lacking parallel channels or dorsal scrobal foveae; dorsal occipital margin abrupt and rounded; funicle 7-segmented; labrum with 4-digits; mesonotum appearing bare, at most with minute setae; mesoscutal lateral lobes and frenum finely sculptured; transscutal articulation complete or fused medially; propodeal disc evenly and finely sculptured (Figs 43,52); prepectus with upper panel broad and flat, at most with a slightly raised posterior margin, very tightly appressed to posterior margin of pronotum (Figs 41,52); fore wing with basal area pilose or bare (Figs 44,55), wing disc with dense long setae (African species) or lacking setae but with sockets apparent (Malagasy species); marginal vein thickened along entire length; postmarginal vein absent or not much longer than stigmal vein; petiole base narrowed into foramen and lacking basal flange (Figs 45,52); antecostal sulcus present and smooth; posterior lateral margin of gastral tergites 1 -4 sinuate; first valvula of ovipositor with several fine lateral teeth.Male scape lacking ventral pores.
Distribution.Ethiopian and Malagasy regions.Etymology.Latin noun meaning "a place where rivers meet," referring to the mesosomal surface sculpture.

Ivieosema confluens
Diagnosis.Recognized from other Orasema and African Ivieosema by the fore wing lacking setae and with a medial longitudinal fold.Distinguished from I. limulus by the smooth face, frons with 2 depressions between eye and antennal scrobe (Fig. 39), and the basal gastral tergite with a laterally carinate median longitudinal depression extending the equivalent length of the petiole.
Female.Length 2.8 mm.Head and mesosoma dark brown.Scape, pedicel, and anellus light brown, rest of flagellum darker brown.Coxae, trochanters, most of femora and last tarsomere dark brown; femoral apices, tibiae and tarsi pale white yellow with last tarsomere brown.Fore wing venation glossy and brown.Gaster brown.
Head (Figs 39-40) broadly subtriangular, 1.4× as broad as long.Face flat and smooth; scrobal depression broad and shallow, extending laterally well beyond toruli and forming a single dorsal frontal depression (Fig. 39, dfd); lower frons with a vague longitudinal groove; scrobal area with a single medial large pit just below median ocellus; supraclypeal area smooth, clypeus with shallow transverse sculpture, anterior tentorial pits very deep and broad, anteclypeus short laterally and absent medially; scrobal depression shallow and hardly evident; vertex not carinate.Labrum with 4 digits.Maxillary and labial palps absent.Mandibular formula 3:2, mandibles small but falcate.Flagellum 1.0× head height, 7 funiculars; anellus longer than broad (Fig. 40,  F1); F2 1.7× as long as broad, 1.1× as long as F3.transepimeral sulcus.Propodeum finely reticulate, callus bare and smooth to shallowly reticulate.Metacoxa smooth.Fore wing 2.3× as long as broad; slightly infuscate along cubital fold and posterior to marginal and submarginal veins, and between the oblique folds in the speculum; venation with only a few tiny setae, including on submarginal vein; costal cell bare; basal cell bare; speculum present, with two oblique folds (Fig. 44, fold) that converge towards stigma and basal fold; marginal vein very thick basally and narrowing apically, not separate from wing margin at base of stigmal vein; stigmal vein 4.5× as long as broad, without an uncus; postmarginal vein absent; cubital and subcubital folds bare; marginal fringe absent.Hind wing costal cell bare.
Metasoma (Fig. 45).Petiole 1.7× as long as broad, 0.9× as long as metacoxa, longitudinally shallowly rugose-reticulate.Gt 1 with a deep longitudinal median depression with carinate lateral margins equal to length of petiole; shallower smooth depression present on Gt 2 .Antecostal sulcus of Gs 1 smooth, very shallow medially; acrosternite swollen and with shallow irregular sculpture; posterior Gs 1 smooth to finely punctate.Cercus with all setae subequal in length.Hypopygium with a few small subapical setae laterally.Ovipositor dorsal valve with 9 annuli, including a basal annulus that is broadly separated from the rest; ventral valve with 5-6 lateral teeth and with transverse carina basal to toothed apex.
Diagnosis.Recognized from other Orasema and African Ivieosema by the fore wing lacking setae and presence of a medial longitudinal fold.Distinguished from I. confluens by the reticulate facial sculpture, frons with a single depression between eye and antennal scrobe (Fig. 50), and the basal gastral tergite with a laterally smooth median longitudinal depression extending the entire length of the tergite.
Female.Length 3.1 mm.Head and mesosoma almost entirely dark brown, without metallic luster; pronotum ventrally, prepectus, and acropleuron yellowish-brown.Scape white; pedicel and flagellum pale brown.Coxae and most of femora pale brown; trochanters, femoral apices, tibiae, and tarsi white.Fore wing venation glossy and milky brown.Gaster brown to dark brown.
Mesosoma 55).Mesoscutum finely reticulate, with a shallow median depression; notauli finely reticulate in their troughs, with vague margins and not foveate, merging posteriorly as a transverse depression anterior to the transscutal articulation, which is present medially (Fig. 53, tsa).Axilla anteriorly reticulate, posteriorly longitudinally striate; mesoscutellum and axillula finely longitudinally striate; frenal line as a pale strip slightly elevated above frenum.Metascutellum narrow and protruding, not separated from rest of metanotum but with fine reticulate sculpture anteromedially.Mesepisternum finely reticulate dorsally and anteriorly, smooth posteroventrally, forming a short protruding horizontal surface anterior to mesocoxa.Propodeum finely reticulate, with a shallow and vague median carina; callus bare, glossy and more shallowly reticulate than propodeal disc.Metacoxa smooth.Fore wing (Fig. 55) 2.2× as long as broad; venation with only a few tiny setae, including on submarginal vein; costal cell bare; basal cell bare; speculum present, with two oblique folds that converge anteriorly towards stigma and diverge posteriorly; marginal vein very thick basally and narrowing apically, separating from wing margin to smoothly form the stigmal vein; stigmal vein 1.4× as long as broad; postmarginal vein absent; cubital and subcubital folds with only a few tiny setae beyond speculum; marginal fringe absent.Hind wing costal cell bare.
Metasoma (Fig. 54).Petiole 1.7× as long as broad, 0.9× as long as metacoxa, nearly smooth medially but longitudinally rugose-reticulate laterally, without transverse anterior carina and narrowest anteriorly immediately posterior to the condyle.Gt 1 with a deep longitudinal median depression extending length of tergite and with smooth lateral margins.Antecostal sulcus of Gs 1 smooth; acrosternite smooth to very shallowly reticulate.Cercus with all setae subequal in length.Hypopygium with a few setae submedially.Ovipositor dorsal valve with 8 medially interrupted annuli, including a basal annulus that is broadly separated from the rest; ventral valve with 4 lateral teeth and with a vague transverse elevation immediately basal to toothed apex.

Leiosema
Discussion.Originally treated within Orasema as the glabra group by Heraty (1994: p. 72).Heraty (1994) speculated that some of the characteristic features were shared with species in the New World coloradensis group.However, there is a greater similarity with Zuparka (Madagascar), including the entirely smooth face (Figs 57-58, 62), scrobal channels with dorsal foveae (Fig. 57), labrum with 4 digits (Figs 57,62), and fore wing with bare basal area and speculum (Fig. 65).However, Zuparka has a laterally smooth or very weakly sculptured propodeal disc (Figs 73,82) and the pronotum and prepectus are more rigidly associated ventrally (Fig. 82).Australosema synempora Heraty also has the mesosomal sidelobes, axillae and frenum smooth, but the face is reticulate and the propodeum has lateral glabrous areas.Matantas also has a similar smooth face, axilla and frenum, but the face is distinctly subtriangular, the labrum has 6-8 digits and the propodeum is laterally glabrous.Each of these other genera have only 3 or 4 lateral subapical teeth on the first valvula in contrast to the 6-10 minute lateral teeth of Leiosema.
Distribution.Sub-saharan Africa (Democratic Republic of the Congo, South Africa, Tanzania, Zimbabwe), and Yemen.Diagnosis.The female is unknown, but the male is easily distinguished by the 3:3 dentate mandibles (Fig. 62), strongly reticulate mesoscutal midlobe and scutellar disc, and all tibiae yellow.It can be further differentiated from L. glabra by the anteriorly extended reticulate pronotal neck (Fig. 61), and petiole 4.7× as long as broad, with a strong transverse basal carina (flange) but without a lateral carina.
Male.Length 1.6 mm.Head and mesosoma blue.Scape brown; pedicel brown; anellus yellowish brown; flagellum brown.Mandible yellow, ventral mandibular setae acute; maxilla and labium yellowish brown.Coxae dark blue; fore and mid femur mostly brown, apically yellowish; hind femur mostly dark brown to black, apically yellow; tibiae yellow, but with hind tibia slightly darker basally.Fore wing hyaline; venation clear.Petiole same color as mesosoma; gaster dark brown; acrosternite same color as petiole.
Variation.The few specimens that are known were collected across a broad geographic range from Yemen to South Africa.Females from South Africa, Yemen and Zimbabwe all have a pale brown scape and a completely yellow fore and mid tibia, whereas the female from Tanzania has an almost black scape and all tibiae are basally dark brown.Males generally have the scape and tibiae dark brown, but again these are much darker in specimens from Tanzania.In the description below, the primary measurements are for what appear to be typical L. glabra specimens from South Africa and Zimbabwe.There are a few minor ratio differences for specimens from Tanzania and Yemen (included in parentheses) that are outside of the range of variation from the typical L. glabra specimens.While these are currently considered to be one species, as more material becomes available, additional species will likely need to be recognized.An additional collection from Gambia shows variation that is likely representative of another species that has a more distinctly reticulate mesoscutellum, but we do not treat this as there seems to be a need to collect more material across Africa to assess the patterns of variation.
Male.Length 1.8-2.4mm.Head and mesosoma dark blue, violet or green.Scape dark brown to black; pedicel black; anellus brown; flagellum black.Mandibles dark brown; maxilla and labium brown.Coxae dark blue; femora mostly dark brown to black, with tips pale; fore and mid tibia yellow to almost completely black with pale apex; hind femur dark brown to black with metallic reflections; hind tibia mostly dark brown to black, tips pale.Fore wing hyaline; venation pale brown.Petiole and acrosternite same as mesosoma; gaster dark metallic.
Diagnosis.Separated from most Orasema and other Oraseminae by having 8-9 funiculars versus 7, a weak basal petiolar flange, usually a distinct ocellar-ocular sulcus, 4-7 digitate labrum, and the callar region usually with several long setae (African male with a single long seta).Distinguished from other Old World genera by the head broadly subtriangular to transverse in frontal view; face smooth or sculptured; presence of an ocellar-ocular sulcus in all but the Australian species; scrobal depression evenly impressed and lacking parallel channels and dorsal foveae; dorsal occipital margin abrupt and rounded or carinate; funicle 8-9 segmented; labrum with 4-7 digits (usually 4); mesonotum appearing bare, at most with minute setae; mesoscutal lateral lobes and frenum sculptured or smooth; transscutal articulation complete and only slightly sinuate; propodeal disc evenly sculptured, smooth laterally only in the Australian species; prepectus foveate and tightly articulated with pronotum ventrally; fore wing with basal area and specular area pilose or bare, wing disc with dense long setae; postmarginal vein much longer than stigmal vein and reaching over half distance to wing apex; petiole base truncate with weak basal flange; antecostal sulcus broad and smooth or foveate; first valvula of ovipositor with 3-5 lateral teeth, subapical teeth sometimes formed by oblique ridges.Male scape lacking ventral pores.
Biology.The oviposition habits and morphology of the egg and first instar were described by Ishii (1932) for L. uichancoi.The egg is stalked as in most Eucharitidae.A revised interpretation of the planidum was presented in Heraty (1994) and larvae are the same as for Orasema, with tergites I and II separated and not fused as illustrated by Ishii.Oviposition was made in short parallel rows into incisions made into the undersurface of leaves of Celtis philippinensis Blanco (Cannabaceae) and Leucaena glauca Benth.(Fabaceae).
Discussion.Defined by Heraty (1994: 71) as the Orasema koghisiana-group.No genetic sequence data are available for the group.Matantas koghisiana (Heraty) comb.n. is the only species described for the genus.
Diagnosis.Separated from Orasema by the combination of smooth lateral areas on the propodeal disc in combination with a rugose-areolate or reticulate median channel, smooth face, and labrum with 6-8 digits.Distinguished from other Old World genera by the head subtriangular in frontal view; face smooth and relatively flattened, scrobal depression evenly impressed and lacking parallel channels or dorsal foveae; dorsal occipital margin abrupt and rounded, at most with a weak dorsal carina; funicle 7-segmented; labrum with 6-8 digits; mesonotum appearing bare, at most with minute setae; mesoscutal lateral lobes and frenum smooth; transscutal articulation complete and relatively straight; propodeal disc smooth with a broad sculptured median channel; prepectus foveate and tightly articulated with pronotum ventrally; fore wing with basal area and specular area bare, wing disc setae relatively long and dense; postmarginal vein much longer than stigmal vein and reaching about half distance to wing apex; petiole base truncate with strong basal flange; antecostal sulcus present and smooth; first valvula of ovipositor with 3 or 4 lateral teeth.Male scape lacking ventral pores.
Host association.Unknown.Distribution.One species described.New Caledonia (another undescribed species from Vanuatu, BPBM: UCRCENT00422296).Etymology.Named after Robert Zuparko of the California Academy of Sciences; gender feminine.

Zuparka
Discussion.Defined by Heraty (2000: 375) as the Orasema monomoria group as based on the single species Z. monomoria (Heraty), comb.n. and on a single recently emerged teneral female.Another distinctive species is recognized herein, Z. fisheri sp.n.The two species are morphologically distinct from each other, and the divergence in their ribosomal and mitochondrial data is generally what is expected between different genera in other Oraseminae.However, morphological features support recognition of a single genus.Zuparka is most similar to the African Leiosema, the only other orasemine with ventral pores on the male scape.Based on Z. monomoria, Zuparka is the only genus with a pupa that lacks pronounced tubercles over the petiolar region (c.f.Fig. 37) or laterally on the lateral abdominal region (Heraty, 2000).It is also the only genus known to attack Monomorium (Myrmicinae).
Diagnosis.Separated from other Oraseminae by the presence of ventral pores on the male scape, 4-digitate labrum (Figs 69, 79), parallel scrobal depressions with a dorsal fovea (Fig. 79, dsd), sculptured frenum (Fig. 70, frn), presence of a basal petiolar carina (Fig. 75, apc), and absence of an antecostal sulcus.Distinguished from the other Old World genera by having the head smooth or very weakly coriaceous, and subtriangular in frontal view (Figs 69,79,81); scrobal depression with parallel channels and dorsal fovea (Fig. 79); occiput rounded; funicle 7-segmented; labrum with 4 digits; mesonotum appearing bare, at most with few minute setae; mesosomal lateral lobes and frenum sculptured; transscutal articulation complete and relatively straight; propodeal disc has either smooth or nearly smooth lateral areas on the propodeal disc in combination with a rugose-areolate or reticulate median channel (Fig. 73), or more weakly sculptured propodeum with a weak median channel and a median carina; prepectus foveate dorsally and tightly articulated with pronotum ventrally (Figs 70, 82); fore wing with basal area and specular area bare (Figs 74,83), wing disc with relatively long dense setae; postmarginal vein less than twice as long as stigmal vein; petiole with prominent basal flange; antecostal sulcus absent; first valvula of ovipositor with 3-4 lateral teeth.Male scape with distinct ventral pores (glands) set in circular depressions across entire ventral surface (Fig. 78).
Distribution.Madagascar.Diagnosis.Face entirely smooth.Scutellum vaulted anterior to frenum and medially depressed.Propodeal disc laterally smooth.This species resembles Zuparka monomoria, but can be easily differentiated by the presence of an elevated medial region of the scutellar disc (Fig. 70) with a median depression (Figs 71,72), sculptured axilla (Fig. 71), torulus with ventral margins roughly on the same plane as the ventral margins of the eyes (Figs 69,79).Based on differing structures of the male antenna, what we are currently recognizing as Z. fisheri may ultimately be a complex of species.
Mesosoma .Midlobe of mesoscutum and mesoscutellum finely reticulate; notauli foveate, reaching transscutal articulation (TSA) and broadly separated posteriorly; lateral lobe and axilla broadly rounded; scutellar disc abruptly vaulted anterior to frenum (Fig. 70) and with weak median depression ; scutellar disc about as long as broad, separated from axillae and TSA by deep foveate scutoscutellar sulcus that is continuous medially; axillular sulcus and frenal line medially foveate with a weak carina along ventral margin of sulcus, continuing laterally as a smooth ridge; frenum semicircular and vertical.Propodeal disc rounded, areolate medially, smooth laterally (Fig. 73); callar region smooth with few setae dorsally; metepimeron smooth dorsally and ventrally reticulate.Upper and lower mesepimeron mostly smooth; transepimeral sulcus irregularly foveate; femoral groove obscure.Prepectus foveate dorsally, with a strong posterior flange, ventrally narrow and smooth.Pronotum reticulate laterally and smooth medially.Propleuron imbricate.Fore and mid coxae imbricate; hind coxae reticulate.Fore wing 2.6-2.9× as long as mesosoma, 2.4-2.6× as long as broad; basal cell setose; costal cell ventrally pilose, remainder of wing pilose; stigmal vein as long or slightly longer than wide and perpendicular to wing margin; postmarginal vein 0.1-0.2×length of marginal vein, about twice length of stigmal vein.Hind wing costal cell with several ventral setae in apical half.
Heacox & Dominguez, sp.n. http://zoobank.org/A2294EE0-09AF-4A98-B9AA-B8D207366A13Figs 68-80 Etymology.Named after Brian Fisher of the California Academy of Sciences for his tremendous collecting efforts that have contributed to the understanding of the fauna of Madagascar.

Type species. Eucharis valgius Walker
Mandibles highly reduced, broader than long and truncate.Funiculars tapering to apex.Frons smooth.Submarginal vein with several long setae, fore wing with distinct marginal fringe.