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Research Article
Biting the bullet: revisionary notes on the Oraseminae of the Old World (Hymenoptera, Chalcidoidea, Eucharitidae)
expand article infoRoger A. Burks, John M. Heraty§, Jason Mottern|, Chrysalyn Dominguez§, Scott Heacox§
‡ University of California, Riverside, United States of America
§ University of California, Riverside, Riverside, United States of America
| USDA Systematic Entomology Laboratory, Washington, United States of America
Open Access

Abstract

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.

Parasitoid, Formicidae, morphology, revision, Palaeotropical

Keywords

Parasitoid, Formicidae, morphology, revision, Palaeotropical

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 northern-most records in India (New Delhi), Taiwan, Iriomote Island of Japan, and the southwestern highlands of Yemen (Heraty 1994, 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, 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 in either case, Orasema is never monophyletic. Character support for a monophyletic Orasema s.l. as presented in Heraty (1994) is weak. Of the characters discussed, only the basal petiolar flange is likely derived, but this has proven to be highly variable in the Old World taxa although not as prominent as can be observed in the New World taxa. Heraty (2000) proposed only a single unambiguous morphological synapomorphy (#52:0) for Orasema, the first-instar larvae without a tergopleural line, the absence of which is now regarded as plesiomorphic for the family as based on outgroup comparison.

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.

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 (1994, 2002) and Heraty (2017). Asterisk (*) denotes type species of genus. Australian refers to Australia including Tasmania. NR is new host record.

Genus Species Distribution Host genus
Australosema gen. n. (=valgius group) A. politurae sp. n. Australia
A. synempora (Heraty, 1994), comb. n. Australia
A. valgius (Walker, 1839), comb. n. * Australia Pheidole
A. verghetta sp. n. Australia
Cymosema gen. n. C. capelina sp. n. Australia
C. waterworthae sp. n. * Australia
Hayatosema gen. n.
(=assectator group)
H. assectator (Kerrich, 1963)* Indo-Pacific Pheidole
H. delhiensis (Narendran & G. Kumar, 2005) Indo-Pacific
H. initiator (Kerrich, 1963) Indo-Pacific
H. kailashi (G. Kumar & Sureshan, 2015) Indo-Pacific
H. nigra (Heraty) Ethiopian
H. nirupama (G. Kumar & Narendran, 2007) Indo-Pacific
H. plicator sp. n. Indo-Pacific
H. siruvanica (G. Kumar & Sureshan, 2015) Indo-Pacific
Ibitya gen. n. (=communis group) I. communis (Risbec, 1952), comb. n. * Madagascar Pheidole NR
I. seyrigi (Risbec, 1952), comb. n. Madagascar
Indosema I. indica Husain & Agarwal, 1983 * India & Kenya
Ivieosema gen. n. (=striatosoma group) I. confluens sp. n. Madagascar
I. limula sp. n. Madagascar
I. fraudulenta (Reichensperger), comb. n. Ethiopian Pheidole
I. striatosoma (Heraty), comb. n. * Ethiopian
Leiosema gen. n. (=glabra group) L. glabra (Heraty, 1994), comb. n. * Ethiopian
L. lesiolouna sp. n. Ethiopian
Losbanus rev. stat. (=uichancoi group) L. bouceki (Boucek), comb. n. New Guinea
L. ishii (Heraty), comb. n. Taiwan
L. promecea (Heraty), comb. n. New Guinea
L. rugulosa (Heraty), comb. n. New Guinea
L. uichancoi Watanabe comb. n. * Philippines
Matantas gen. n. (=koghisiana group) M. koghisiana (Heraty), comb. n. * New Caledonia
Orasemorpha O. didentata (Girault, 1940) Australia
O. eribotes (Walker, 1839)* Australian Pheidole
O. goethei (Girault, 1934) Australia
O. myrmicae (Girault, 1936) Australia Pheidole
O. pytallus (Walker, 1846) Australia
O. sparsepilosa Heraty, 1994 Australia
O. tridentata (Girault, 1915) Australia Pheidole
O. varidentata (Girault, 1936) Australian
O. xeniades (Walker, 1839) Australian Pheidole
Timioderus T. acuminatus Heraty, 1994 Ethiopian Pheidole
T. coronula Heraty, 1994 Ethiopian
T. peridentatus Heraty, 1994 Ethiopian
T. coronula Heraty, 1994 Ethiopian
T. refringens Waterston, 1916 * Ethiopian
Zuparka gen. n. (=monomoria group) Z. fisheri sp. n. Madagascar
Z. monomoria (Heraty, 2000), comb. n. * Madagascar Monomorium

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).

In the following taxonomic treatments, we provide only new nomenclatorial acts or new information. We do not revisit the genera Indosema, Orasemorpha and Timioderus, which were revised in Heraty (1994). The identification key includes treatments only for the newly decribed species. Two undescribed genera are included along with illustrations that have only been collected as pharate adults (Figs 85, 86).

Key to Oraseminae of the Old World

1 Transscutal articulation medially sinuate; axillae strongly advanced laterally beyond posterior margin of mesoscutum (Figs 19, 24). Fore wing with anterior margin broadly rounded posterior to stigmal vein making fore wing appear ovate (Figs 15, 27). Petiole narrowed gradually into foramen (Fig. 20), lacking anterior petiolar flange (Australian; males unknown) Cymosema gen. n....2
Transscutal articulation either absent medially (Fig. 42) or present and at most slightly sinuate (usually straight) and axillae not strongly advanced (Figs 4, 10, 32). Fore wing with anterior margin relatively straight (Figs 7, 33). Petiole variable 3
2 (1) Mandibles minute and lacking distinct teeth (Fig. 16, md). Mesosomal dorsum entirely smooth (Fig. 19) C. capelina sp. n.
Mandibles 3/2 dentate with teeth narrow and weakly curved (Figs 21, 22). Mesosoma anteriorly reticulate (Fig. 24) C. waterworthae sp. n.
3 (1) Upper frons with a large pit immediately below median ocellus (Fig. 39, sdp). Antenna with 7 funiculars; anellus transverse and distinct (Fig. 40). Petiole without anterior carina. Metasomal tergite margins laterally sinuate (Figs 45, 46) (Ethiopian and Malagasy) Ivieosema gen. n. (striatosoma-group of Heraty 1994)...4
Upper frons without dorsal pit. Other characters variable 7
4 (3) Face reticulate. Transscutal articulation complete. Fore wing with moderately long setae; without swelling posterior to submarginal vein (Ethiopian) 5
Face smooth (Figs 39, 47) or with fine sculpture (Fig. 50). Transscutal articulation fused medially. Fore wing with very short setae; with distinct longitudinal swelling posterior to submarginal vein (Figs 44, 55) (Malagasy) 6
5 (4) Mesoscutellum reticulate dorsally Ivieosema fraudulenta (Reichensperger)
Mesoscutellum longitudinally striate Ivieosema striatosoma (Heraty)
6 (5) Face smooth. Scape brown. Upper frons with two depressions between eye and antennal scrobe (Figs 39, 47) Ivieosema confluens sp. n.
Face smooth to shallowly reticulate. Scape white. Upper frons with single transverse depression between eye and antennal scrobe (Fig. 50) Ivieosema limula sp. n.
7 (3) Transscutal articulation (between mesoscutum and axilla) obliterated medially. Maxillary and labial palpi absent 8
Transscutal articulation complete with a distinct transverse suture (Fig. 4). Maxillary and labial palpi present 9
8 (7) Body metallic blue or green. Head and mesosoma strongly sculptured and either coriaceous or rugulose. Anellus present. Mandibles truncate or spatulate, maxilla small. Petiole not fused ventrally. Ovipositor curved ventrally. Body length at least 3.5 mm (Ethiopian) Timioderus Waterston
Body black or brown. Head and mesosoma almost smooth. Anellus absent. Mandibles absent, maxilla enlarged and lobate. Petiole fused ventrally. Ovipositor long and relatively straight. Body length about 2 mm (Kenya [New Record: UCRCENT 00478780, Olorgesailie National Monument, R. Copeland] and India) Indosema Husain & Agarwal
9 (7) Petiole of female transverse, at most as broad as long. Petiole of male may be twice as long as broad, but then distal third or half supported ventrally by swollen anterior expansion of first gastral sternite (Gs1). Base of petiole lacking an anterior dorsal carina (Australian) 10
Petiole of both sexes cylindrical and at least as long as broad (Figs 5, 67, 86), but usually much longer and gaster never appearing sessile. Petiole of male with anterior margin of first gastral sternite (Gs1) meeting apex of petiole (Figs 64, 86). Base of petiole usually truncate and usually with an anterior transverse carina (Fig. 75) (Old World tropics) 11
10 (9) Antennal flagellomeres cylindrical. Base of petiole narrowed to condyle Orasemorpha Bouček
Antennal flagellomeres lobate (Fig. 85). Base of petiole truncate Undescribed and unplaced teneral male (new genus 1)
Antenna encased in exuvia, collected from base of ant-infested termite mound, very distinct from other genera and shows some similarities with Timioderus (Australia: QLD: Mareeba; ANIC: UCRCENT00238798).
11 (9) Lower face lateral to clypeus finely reticulate (Fig. 29). Frenum and propodeum evenly sculptured, propodeum lacking a distinctly differentiated narrow median band of areolate sculpture. Prepectus evenly reticulate or rugose, dorsal margin without a raised rim and overlapped by ventral margin of mesoscutum. Callus at most with a few minute setae. Female antenna with 7–8 funiculars, male with 7–9 funiculars. Ventral valve of ovipositor with several minute lateral teeth (Fig. 35) Hayatosema gen. n. (assectator -group of Heraty, 1994)
Face variously sculptured, but if finely reticulate then propodeum with distinctly differentiated median channel (Fig. 11), prepectus broadly foveate with a distinctly raised dorsal margin (Fig. 12) or callus with dense patch of long hairs. Both sexes with 7–9 funiculars. Ventral valve of ovipositor with 3-4 lateral teeth (Figs 13, 76) 12
12 (11) Antenna with 8–9 funiculars, if 7, then labrum with more than 7 digits, body black, and propodeal disc evenly sculptured (Ib. seyrigi, Madagascar). Ocellar-ocular sulcus usually distinct. Propodeal disc evenly sculptured, rarely smooth. Fore wing often without speculum 13
Antenna with 7 funiculars, if labrum with more than 7 digits, then body metallic blue green and propodeal disc smooth laterally. Ocellar-ocular sulcus absent. Fore wing setation variable 14
13 (12) Face smooth laterally (Fig. 36). Labrum with more than 9 digits (one undescribed specimen with 5 digits). Antenna with 7–8 funiculars (Madagascar) Ibitya gen. n. (communis -group of Heraty, 2000; 2 species described)
Face evenly sculptured laterally. Labrum with 4 digits (one undescribed Australian species with 6). Antenna with 8–9 funiculars (Paleotropical excluding Madagascar) Losbanus rev. stat. (uichancoi-group of Heraty, 1994; 5 species)
14 (12) Propodeal disc evenly sculptured 15
Propodeal disc smooth laterally, usually with a distinct median longitudinal channel with strong areolate sculpture (Figs 11, 73), rarely weakly reticulate medially (Fig. 5) 17
15 (13) Face and frenum completely smooth (Figs 57, 58, 62) (Ethiopian) Leiosema gen. n. (glabra-group of Heraty, 1994)...16
Face completely and evenly reticulate; scutellar disc and frenum rugose-reticulate (Fig. 86) (Australian) Undescribed and unplaced teneral male (new genus 2)
Antenna encased in exuvia, no ant host information, close to Australosema but very distinct from other species (Australia: QLD: Thornton Peak, QM:UCRCENT00241816).
16 (15) Mandibles 3:3 dentate (Fig. 62). Scutellar disc with prominent reticulate sculpture (Fig. 61). Pronotal neck extended anteriorly and reticulate (Fig. 61). Female unknown Leiosema lesiolouna sp. n.
Mandibles 3:2 dentate. Pronotal neck not extended and smooth. Scutellar disc at most with shallow reticulate sculpture anteriorly, weakly reticulate or smooth posteriorly (Figs 5960) Leiosema glabra (Heraty)
17 (14) Labrum with 6–8 digits. Face smooth (Papua New Guinea [undescribed] and New Caledonia) Matantas gen. n. (koghisiana-group of Heraty, 1994), M. koghisiana (Heraty)
Labrum with 4 digits. Face smooth or reticulate 18
18 (17) Antecostal sulcus present, dividing first gastral sternite and separating off a distinct acrosternite (Figs 1, 7). Face smooth or with at least some reticulate sculpture. Scrobes without parallel median channels. Male scape glands absent (Australian) Australosema gen. n. (valgius-group of Heraty, 1994)...19
Antecostal sulcus absent (first gastral sternite smooth and uninterrupted). Face entirely smooth. Scrobes with parallel median channels (Fig. 79). Male scape glands present (Malagasy) Zuparka gen. n. (monomoria-group of Heraty, 2000)...22
19 (18) Face entirely smooth (Fig. 2) A. politurae sp. n.
Face with at least frons with reticulate sculpture (Fig. 8) 20
20 (19) Frenum with median longitudinal groove (Fig. 10). Scape dark metallic brown in both sexes A. verghetta sp. n.
_ Frenum without median longitudinal groove. Scape yellow or dark brown 21
21 (20) Lateral lobe of mesoscutum and axilla smooth and polished A. synempora (Heraty)
Lateral lobe and axilla coriaceous to rugose, axilla weakly carinate with surface imbricate to reticulate. Scape yellow to dark brown (see Heraty 1994 for discussion of variants) A. valgius (Walker)
22 (18) Face entirely smooth (Figs 69, 79). Propodeal disc with lateral areas smooth (Fig. 73) Z. fisheri sp. n.
Frons weakly coriaceous (Fig. 81). Propodeal disc with lateral areas weakly sculptured Z. monomoria (Heraty)

Australosema Heraty & Burks, gen. n.

Figs 1–6, 7–12, 13–14

Type species

Eucharis valgius Walker: 412–414. Heraty 1994: 79–81, figs 122, 124, 131, 192, 221, 238, 248, 262–263.

Etymology

Random combination of letters and referring to the distribution; gender feminine.

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 rugose-areolate 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.

Host association

Myrmicinae: Pheidole (Girault 1913[175], Heraty 2000).

Distribution

Four species. Australia including Tasmania.

Australosema politurae Burks, sp. n.

Figs 1–6

Etymology

Latin noun in genitive case meaning “a polishing,” referring to the smooth face.

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).

Female

Length 2.2–3 mm. Head and mesosoma dark metallic green, petiole black, gaster brown with faint metallic luster. Scape, pedicel, anellus yellowish, funiculars and clava brown. Coxae metallic green basally, becoming yellowish apically, femora pale brown except yellowish at tips, remainder of leg yellowish. Fore wing venation pale brown, setae dark. Gaster brown to dark brown, with metallic luster.

Head (Figs 2–3), 1.1–1.2× as broad as high. Face almost entirely smooth except for minute setal pits ventrally, dorsally with shallow irregular sculpture near ocellar triangle; scrobal depression narrow and impressed, without dorsal scrobal foveae; without longitudinal furrow between torulus and eye; supraclypeal area and clypeus smooth, supraclypeal area slightly protruding, anteclypeus present; vertex weakly carinate posterior to ocellar triangle. Labrum with 4 digits. Palpal formula 3:3. Flagellum 1.1–1.3× head height, with 7 funiculars; anellus transverse; F2 2.0–2.8× as long as broad, F2 1.4–1.6× as long as F3.

Mesosoma (Figs 4–5). Mesoscutal midlobe densely reticulate, lateral lobe shallowly transversely rugose-reticulate and becoming smooth posteriorly. Axilla rugose-reticulate, with longitudinal carinae posteriorly that extend across the scutoscutellar suture; mesoscutellar disc finely rugose-reticulate, with some wrinkle-like carinae posteriorly; frenal line carinate, frenum rugose-reticulate; axillular sulcus carinate, axillula glossy and very shallowly sculptured aside from a few longitudinal carinae. Mesepisternum anterodorsally mostly smooth but not overlapping posterior edge of prepectus, posteroventrally smooth, not becoming horizontal anterior to mesocoxa; upper and lower mesepimeron mostly smooth except for foveate area at posterior edge, transepimeral sulcus foveate. Propodeum with broad rugose-reticulate median channel, sublaterally nearly smooth; callus smooth, with a few tiny setae. Metacoxa shallowly rugose-reticulate dorsolaterally. Fore wing 2.3–2.4× as long as broad; costal cell with 1 uninterrupted row of ventral setae; basal cell bare; speculum present; cubital fold with uninterrupted ventral row of setae; subcubital fold setose posterior to speculum, bare posterior to basal cell; stigmal vein 1.5–2.0× as long as broad, only slightly expanded at uncus; marginal vein about 6× as long as stigmal vein. Hind wing costal cell with a few ventral setae in apical third.

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 Gs1 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 rugose-reticulate; 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.

Material examined

Holotype: Australia: QLD: Mt. Glorious, 27°19'54"S, 152°45'29"E, 7-13.ii.1998, N. Power, Malaise trap [1♀, ANIC: UCRCENT00091415]. Paratypes: Australia: QLD: Brisbane Forest Pk, 27°25'04"S, 152°49'48"E, 29.xi-5.xii.1997, N. Power, dry sclerophyll, Malaise Trap 3 [1♂, ANIC: UCRCENT00312131]. Lamington Nat. Park, Binna Burra, 28°11'53"S, 153°11'16"E, 10.xii.2002, George, Hawks, Munro, & Owen, rainforest, sweep [1♀, UCRC: UCRCENT00312128]. Mt. Glorious, 27°19'54"S, 152°45'29"E, 10-16.i.1997, N. Power, Malaise Trap [1♀, UCRC: UCRCENT00312135]. Mt. Glorious, 27°19'54"S, 152°45'29"E, 13-24.iii.1998, N. Power, Malaise Trap [1♂, UCRC: UCRCENT00312133]. Mt. Glorious, 27°19'54"S, 152°45'29"E, 24-30.xi.1997, T. Hiller, MT 3 [3♂, UCRC: UCRCENT00312060 – UCRC: UCRCENT00312060–62]. Mt. Glorious, 27°19'54"S, 152°45'29"E, 3-9.i.1998, T. Hiller, MT 2 [1♀, UCRC: UCRCENT00312063].

Similar species

A single specimen (gaster missing, likely female; UCRCENT0092152, 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.

Figures 1–6.

Australosema politurae. Paratype female: 1 habitus, lateral 2 head, anterior 3 antennae, ventral 4 mesosoma, dorsal 5 propodeum and petiole, oblique dorsal. Paratype male: 6 antennae, lateral. acl = anteclypeus; acr = acrosternite; acs = antecostal sulcus; apc = anterior petiolar carina; ax = axilla; cal = callus; cly = clypeus; F = flagellomere; fr = frons; Fu = funicular; F1/an = anellus; llm = mesoscutal lateral lobe; lpr = lateral propodeal disc; mlm = mesoscutal midlobe; mpr = median propodeal disc; pet = petiole; not = notauli; scd = scutellar disc.

Australosema verghetta Burks, sp. n.

Figs 7–12, 13–14

Etymology

Italian noun used in heraldry to refer to a line along the middle, referring to the longitudinal scutellar groove present in females.

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.

Head (Figs 8–9) 1.2–1.3× as broad as high. Face mostly reticulate, smooth ventrally adjacent to mouthparts; vertex with weak carina posterior to ocellar triangle; scrobal depression narrow and impressed, without dorsal scrobal foveae; longitudinal furrow present amidst sculpture between torulus and eye; supraclypeal area and clypeus smooth and separated by only a very shallow depression, with short decumbent setae, supraclypeal area protruding, anteclypeus present. Labrum with 4 digits. Palpal formula 3:3. Flagellum 1.1–1.2× height of head, with 7 funiculars, dark brown; anellus transverse; F2 1.7–2.3× as long as broad, 0.7–1.0× as long as F3, nearly parallel-sided and having few longitudinal sensilla (Fig. 9, F2).

Mesosoma (Figs 10–12). Mesoscutum transversely rugose-reticulate, lateral lobe nearly smooth at posterior margin. Axilla rugose-reticulate and with longitudinal carinae that extend across the scutoscutellar suture; mesoscutellar disc rugose-reticulate except peripherally with many subparallel carinae; frenal line posteriorly carinate, frenum rugose-reticulate and with median longitudinal groove; axillular sulcus foveate, axillula with reticulate sculpture laterally, and with subparallel longitudinal carinae. Mesepisternum anterodorsally smooth and overlapping posterior lower margin of prepectus, posteroventral surface with horizontal shelf that is shorter than mesepimeron but swollen and smooth anterior to mesocoxa; upper and lower mesepimeron smooth, transepimeral sulcus foveate. Propodeal disc with a meandering median carina crossed by short transverse carinae, sublaterally smooth; callus smooth and with very short setae. Metacoxa basally rugose-reticulate, apically smooth. Fore wing 2.4–2.5× as long as broad; costal cell ventrally setose in apical half; basal cell bare; speculum present; cubital fold setose ventrally posterior to basal cell but bare posterior to speculum; subcubital fold setose along speculum but bare posterior to basal cell; stigmal vein 1.5–2.0× as long as broad, only slightly expanded at uncus; postmarginal vein about 5.0× as long as stigmal vein. Hind wing costal cell setose in apical third, bare in basal two thirds.

Metasoma (Figs 11, 13). Petiole 0.9–1.1× as long as metacoxa, 2.4–3.1× as long as broad, transversely rugose-reticulate and with a few short longitudinal wrinkle-like carinae; basally truncate with at most a weak basal flange. Antecostal sulcus of Gs1 smooth to finely sculptured laterally; acrosternite smooth. Cercus with 1 curved seta that is much longer than the others. Hypopygium with a few short setae and 1 longer posterolateral seta on each side (Fig. 13, lhs). Ovipositor dorsal valve with 7–8 medially interrupted annuli; ventral valve with 4–5 weakly defined teeth, and with oblique subapical carina immediately basal to toothed apex (Fig. 13, ocv).

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.

Material examined

Holotype: Australia: SA: Mt. Barker, 511m, 35°04'00"S, 138°55'17"E, 22-23.i.1999, J. Heraty, Eucalyptus scrub [1♀, ANIC: UCRCENT00091308]. Paratypes: Australia: SA: Mt. Barker, 511m, 35°04'00"S, 138°55'17"E, 22-23.i.1999, J. Heraty, Eucalyptus scrub [14♂ 19♀, ANIC: UCRCENT00311977–81, UCRC: UCRCENT00091307, UCRCENT00091309, UCRCENT00311982–89, UCRC: UCRCENT00312042–59].

Figures 7–12.

Australosema verghetta. Paratype female: 7 habitus, lateral 8 head, anterior 9 antennae, dorsal 10 mesosoma, dorsal; 11 propodeum and petiole, dorsal 12 mesosoma, lateral. acs = antecostal sulcus; apc = anterior petiolar carina; axlg = axiullar groove; cal = callus; F = flagellomere; frg = frenal groove; lpr = lateral propodeal disc; not = notaulix; SSS = scutoscutellar sulcus; tsa = transscutal articulation.

Cymosema Heraty & Burks, gen. n.

Figs 15–20, 21–26, 27

Type species

Cymosema waterworthae Burks & Mottern sp. n.

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 Gs1; 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).

Figures 13–14.

Australosema verghetta. Paratype female: 13 ovipositor, lateral. Paratype male: 14 antenna, ventral. Fu = funicle; dv = dorsal valve; lhs = lateral hypopygial seta; ocv = oblique carina of ventral valvula; vv = ventral valve; vvlt = ventral valve lateral teeth.

Cymosema capelina Burks & Mottern, sp. n.

Figs 15–20

Etymology

Noun from Spanish, meaning wide-brimmed floppy hat, referring to the head shape.

Diagnosis

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.

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.

Head (Figs 16–17) transverse, 1.4× as broad as high. Head receding ventral to toruli (Fig. 18); face dorsally smooth, ventrally coriaceous; supraclypeal area and clypeus medially smooth and laterally shallowly coriaceous, anteclypeus absent; vertex not carinate. Mandibles truncate and broader than long, broadly separated. Palpal formula 2:1, palpi long. Pedicel broader than F2; anellus small but easily visible; flagellum 1.4× head height, with 7 funiculars that expand apically and have semi-erect long setae arising from their bases; F2 1.3× as long as broad, 1.0× as long as F3.

Mesosoma (Figs 18–19). 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 Gs1 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.

Male

Unknown.

Material examined

Holotype: Australia: Queensland: Heathlands, 11°27'00"S, 142°21'00"E, 25.vii-18.viii.1992, P. Zborowski, J. Cardale, Malaise trap [1♀, ANIC: UCRCENT00238787], deposited in ANIC.

Figures 15–20.

Cymosema capelina. Holotype female: 15 habitus, lateral 16 head, anterior 17 antennae, lateral 18 mesosoma, lateral 19 mesosoma, dorsal 20 propodeum and petiole, oblique dorsal. ax = axilla; F = flagellomere; md = mandible; pet = petiole; prp = propodeum; tsa = transscutal articulation.

Cymosema waterworthae Burks & Mottern, sp. n.

Figs 21–26, 27

Etymology

Named in honor of the collector, Rebeccah Waterworth.

Diagnosis

Mandibles bidentate and overlapping, teeth of equal length and projecting medially. Funiculars cylindrical. Frons weakly reticulate. Submarginal vein bare, fore wing without marginal fringe.

Female

Length 1.4 mm. Head, mesosoma, and metasoma brown with metallic luster. Scape yellowish-brown, pedicel and flagellum brown. Legs brown, with tibial apices, and first four tarsomeres pale brown. Fore wing hyaline; venation pale brown, setae dark. Gaster brown with faint metallic luster.

Head (Figs 21–22) transverse, 1.6× as broad as high. Head receding ventral to toruli; face reticulate, including supraclypeal area; clypeus shallowly transversely rugose-reticulate, anteclypeus present; vertex not carinate. Mandibles bidentate, both teeth projecting medially, narrow and only weakly curved (Fig. 21). Palpal formula 2:1, palpi minute. Pedicel broader than F2; scape and pedicel reticulate; flagellum 1.5× head height, with 7 parallel-sided funiculars; anellus minute and hardly visible; F2 1.6× as long as broad, F2 1.1× as long as F3.

Mesosoma (Figs 23–24, 27). Mesoscutal midlobe coriaceous anteriorly, becoming smooth posteriorly, lateral lobe mostly smooth but with shallow irregular sculpture anteriorly and laterally; notauli deep anteriorly but very shallow posteriorly. Axilla strongly advanced, smooth anteriorly, coriaceous posteriorly; mesoscutellar disc smooth to very shallowly coriaceous; frenal line indicated by narrow groove, frenum smooth; axillular sulcus narrow, axillula coriaceous. Mesepisternum reticulate anterodorsally, becoming smooth ventrally and forming a rounded horizontal shelf anterior to mesocoxae; mesepimeron shallowly reticulate and without a transepimeral sulcus. Propodeal disc finely reticulate, with a weak median carina, sublaterally reticulate and with an oblique elevation anterior to a steep descent towards the broad petiolar foramen; callus bare, with a flange-like elevation posterolateral to spiracle (Fig. 24, flange). Metacoxa shallowly reticulate dorsolaterally. Fore wing spatulate (Fig. 27), 2.7× as long as broad; costal cell and submarginal vein bare; basal cell ventrally setose, also with a few dorsal setae near basal fold; speculum absent; cubital and subcubital folds setose; marginal fringe absent; stigmal vein 2.0× as long as broad; postmarginal vein about 1.5× as long as stigmal vein. Hind wing costal cell bare.

Metasoma (Figs 25–26). 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 Gs1. 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.

Male

Unknown.

Material examined

Holotype: Australia: Western Australia: D’Entrecasteaux N.P., Coastal Survivors Walk, 105m, 34°50'19"S, 116°00'17"E, 8.xii.2009, R. Waterworth, sweep [1♀, UCRC: UCRCENT00364784], deposited in UCRC. Gaster mounted separately on card.

Figures 21–26.

Cymosema waterworthae. Holotype female: 21 head, anterior 22 antennae, lateral 23 mesosoma, lateral 24 mesosoma, dorsal 25 petiole, lateral 26 ovipositor. pet = petiole.

Figure 27.

Cymosema waterworthae. Holotype female: fore wing. arrow points to end of postmarginal vein.

Hayatosema Heraty & Burks, gen. n.

Figs 28–33, 34–35

Type species

Orasema assectator (Kerrich, 1963: 367–368). Heraty, 1994: 74–75, figs 111–112.

Etymology

Named after Mohammad Hayat of Aligarh Muslim University; gender feminine.

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 (1994, 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.

Diagnosis

Distinguished from other Old World genera by the head subtriangular in frontal view (Fig. 29); lower face excluding clypeal region finely reticulate; scrobal depression evenly impressed, lacking parallel channels and dorsal foveae; dorsal occipital margin abrupt with a rounded or carinate margin; antenna with 7-8 funiculars in females (Fig. 30) and 7–9 funiculars in males; labrum with 4-digits; mesonotum appearing bare, at most with minute setae, notauli deeply and broadly impressed; mesoscutal lateral lobes and frenum evenly sculptured; transscutal articulation complete and weakly sinuate, axillae not advanced; propodeal disc evenly sculptured without distinctly differentiated narrow median area (Fig. 34); prepectus finely reticulate or rugose with undifferentiated dorsal margin that is partially overlapped by lateral margin of mesonotum (Fig. 31); tightly articulated with mesepisternum ventrally (Fig. 31); fore wing with basal area and specular area bare (Fig. 33), wing disc with dense long setae; postmarginal vein relatively short, less than 1/3 distance to apex of wing; petiole base truncate with weak basal flange (Fig. 34); antecostal sulcus foveate or irregularly sculptured; first valvula of ovipositor with 7-10 minute lateral teeth (Fig. 35). Male scape lacking ventral pores.

Host association

Hayatosema initiator (Kerrich) reared from Pheidole (Das 1963; Kerrich 1963).

Distribution

Four species. Ethiopian and Indo-Pacific (Heraty, 1994).

Figures 28–33.

Hayatosema plicator. Holotype female: 28 habitus, lateral 29 head, anterior 30 antennae, dorsal 31 mesosoma, lateral 32 mesosoma, dorsal 33 fore wing, dorsal.

Figures 34–37.

34–35 Hayatosema plicator. Holotype female: 34 propodeum and petiole, dorsal oblique 35 ovipositor, dorsal 36–37: Ibitya communis. Female: 36 head, frontal 37 pupa. hcx = hind coxa; pet = petiole.

Hayatosema plicator Burks, sp. n.

Figs 28–33, 34–35

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.

Diagnosis

Flagellum with 7 funiculars (Fig. 30). Prepectus rugose, mostly smooth, and without evenly reticulate sculpture as in all other species. Supraclypeal area and clypeus smooth (Fig. 29). Mesoscutal midlobe strongly elevated anteriorly; pronotum with rounded anterolateral protrusions (Fig. 32). Propodeal disc evenly areolate, with irregular, incomplete median carina (Fig. 34). This species keys to O. initiator in Heraty (1994), but differs by having a proportionally longer antennal flagellum to head height (1.3 versus 1.1–1.2), longer F2 (2.0 versus 1.4-1.5), frenum finely rugose-reticulate (versus reticulate), and a distinctly areolate (versus finely reticulate or nearly smooth) propodeum. It will key to O. assectator in Girish Kumar and Sureshan (2015), but differs most distinctly by the rugose versus reticulate prepectus.

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.

Head (Figs 29–30) 1.4× as broad as high. Face rugose-reticulate; supraclypeal area and clypeus smooth; anteclypeus absent; vertex reticulate. Labrum with 4 digits. Palpal formula 3:3 but with minute 2nd palpomeres. Flagellum 1.3× height of head; anellus transverse; F2 2.0× as long as broad, F2 1.3× as long as F3.

Mesosoma (Figs 3134). 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.

Metasoma (Figs 34–35). Petiole 1.7× as long as broad; 0.9× as long as metacoxa, with sharp anterior transverse carina, areolate and with a complete dorsal median carina. Antecostal sulcus of Gs1 finely sculptured laterally, becoming smooth and very shallow medially; acrosternite smooth. Hypopygium with a few short setae, and 1 slightly longer posterolateral seta on each side. Ovipositor dorsal valve with 7 medially interrupted annuli; ventral valve with 7–8 minute teeth along lateral carina, with only a rounded elevation basal to toothed apex.

Male

Unknown.

Material examined

Holotype: Vietnam: Thua Thien-Hue: Bach Ma Natl. Pk. Parashorea Trail, 9 km from park entrance, 100m, 16°12'02.4"N, 107°50'49.6"E, 26.v-13.vi.2001, D.C. Darling, N. Tatarnic, B. Hubley, Dipterocarp forest-closed canopy, Malaise trap (fine) [1♀, ROME 2001504], deposited in ROME.

Ibitya Heraty & Burks, gen. n.

Figs 36, 37

Type species

Orasema communis Risbec, 1952: 412–414. Heraty 1994: 59–60, figs 80, 82, 84–85.

Etymology

Named after one of the Madagascar collecting localities, Ibity; gender feminine.

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.

Host association

Myrmicinae: Ibitya communis reared from Pheidole veteratrix Forel.

Distribution

Madagascar.

Ibitya communis (Risbec), comb. n.

Fig. 37

Orasema communis Risbec, 1952: 412–414. Type data: Madagascar: Bekily. Lectotype male (examined), by subsequent designation Heraty, 1994: 59. Type depository: MNHP. Citations: Heraty, 1994: 59-61 (subsequent description, both sexes, illustrated); Heraty, 2002: 51 (catalog).

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 Mt1–5. 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 Mt1–5.

New host record

Myrmicinae: Pheidole veteratrix.

New material examined

Madagascar: 45 km S Ambalavao, 785m, 22°13'00"S, 47°01'00"E, 2.x.1993, B.L. Fisher, nest, rotten log rainforest, ex: Pheidole veteratrix [1♂, 1♀, 6♀ pupae, 1 ♂ pupa: CASC: UCRCENT00417452, UCRC: UCRCENT00417446–00417451, UCRC: UCRCENT00417453]. La Mandraka, 18°54'19"S, 47°55'13"E, ii.1944, A. Seyrig [1♂, NMPC: UCRCENT00416474]. Antananarivo Prov.: Ankokoy Forest, 3 km E of Ibity, 1700m, 20°04'03"S, 46°59'58"E, 14-24.xi.2008, M.Irwin, R.Harin’Hala, Uapaca forest, Malaise trap [1♂, CASC: CASENT02212472]. Antsiranana Prov.: botanical garden near the entrance to Andasibe National Park, 1025m, 18°55'35"S, 48°24'28"E, 1-5.ix.2001, R. Harin’ Hala, tropical forest, Malaise trap [2♂, CASC: UCRCENT00091471–72]. Fianarantsoa Prov.: 7 km W Ranomafana, 1100m, 24°34'04"S, 46°50'13"E, 15-21.ix.1988, C. Kremen, montane rainforest, small clearing, Malaise Trap [1♂, EMEC: UCRCENT00404908]. 7 km W Ranomafana, 900m, 21°15'33"S, 47°23'12"E, 23-28.ii.1990, W.E. Steiner, montane rainforest, Malaise trap in small clearing [1♀, USNM: UCRCENT00416660]. Parc National Ranomafanae, radio tower at forest edge, 1130m, 21°15'03"S, 47°24'26"E, 16.x-8.xi.2001, 6-17.vii.2003, R. Harin’Hala, mixed tropical forest, Malaise trap [3♂ 1♀, CASC: UCRCENT00417457–59, UCRCENT00417550].

Distribution

Madagascar.

Ibitya seyrigi Risbec, comb. n.

Orasema seyrigi Risbec, 1952: 414–416. Type data: Madagascar: Bekily. Holotype female (examined), by monotypy. Type depository: MNHN. Citations: Heraty 1994: 61–62 (subsequent description of female, illustrated); Heraty 2002: 55 (catalog).

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.2003, R. Harin’Hala & M. Irwin, mixed tropical forest, Malaise trap (1♀, CAS: UCRCENT00417549).

Distribution

Madagascar.

Ivieosema Heraty & Burks, gen. n.

Figs 38–43, 44–48, 49–54

Type species

Orasema striatosoma Heraty, 1994: 68–70, figs 98–105.

Etymology

Named after Michael Ivie of Montana State University; gender feminine.

Discussion

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 (Figs 45–46, 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.

Host association

Myrmicinae: Ivieosema fraudulenta reared from Pheidole megacephala Fabricius (Reichensperger 1913).

Distribution

Ethiopian and Malagasy regions.

Figures 38–43.

Ivieosema confluens. Paratype female: 38 head and mesosoma, lateral 39 head, anterior 40 antenna, medial 41 mesosoma, lateral 42 mesosoma, dorsal 43 mesosoma, posterior. dfd = dorsal frontal depression; F1/an = anellus; lg = longitudinal groove; sod = subocellar depression; tsa = transscutal articulation.

Figures 44–48.

Ivieosema confluens. Paratype female: 44 fore wing 45 metasoma, lateral, lateral view of ovipositor in top left corner. Holotype male: 46 habitus, lateral 47 head, anterior 48 antennae, dorsal. dv = dorsal (2nd) valvula, F1/an = anellus; vv = ventral (1st) valvula.

Figures 49–54.

Ivieosema limula. Holotype female: 49 habitus, lateral 50 head, anterior 51 antenna, lateral 52 mesosoma, lateral 53 mesosoma, dorsal 54 ovipositor, lateral. dfd = dorsal frontal depression; F1/an = anellus; lg = longitudinal groove; mts = metascutellum; tsa = transscutal articulation.

Ivieosema confluens Burks, sp. n.

Figs 38–43, 44–48

Etymology

Latin noun meaning “a place where rivers meet,” referring to the mesosomal surface sculpture.

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.

Mesosoma (Figs 4144). Mesoscutal midlobe finely reticulate, lateral lobe mostly coriaceous, notauli very shallow and with vague margins; transscutal articulation represented only by a vague depression medially (Fig. 42, tsa). Axilla anteriorly reticulate, posteriorly longitudinally striate; mesoscutellum and axillula finely longitudinally striate; frenal line as a smooth raised strip. Metascutellum protruding, not separated from rest of metanotum. Mesepisternum reticulate dorsally, smooth ventrally, forming a short rounded horizontal area anterior to mesocoxae; mesepimeron reticulate, without 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. Gt1 with a deep longitudinal median depression with carinate lateral margins equal to length of petiole; shallower smooth depression present on Gt2. Antecostal sulcus of Gs1 smooth, very shallow medially; acrosternite swollen and with shallow irregular sculpture; posterior Gs1 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.

Male

(Figs 46–48). Length 2.5 mm. Antennal flagellum densely covered with long decumbent setae, with only a few longitudinal sensilla extending beyond funicular apices; F7 shorter than F6 and F8; entire antenna brown. Petiole 1.3× as long as broad, 2.5× as long as metacoxa. Gaster with broad, striate longitudinal depression on Gt1. Digitus with 6 spines; parameres present.

Holotype

Madagascar: Toliara Prov.: 18 km NNW Betroke, 825m, 23°09'48"S, 45°58'07"E, 24.x-4.xii.1994, M.A.Ivie & D.A.Pollock, FIT [1♂, UNMO: UCRCENT00435300], deposited in TMP. Paratype: Madagascar: Toliara Prov.: Tsimanampetsotsa National Park, Mitoho Forest, 120m, 24°02'55"S, 43°45'08"E, 24-29.x.2008, M. Irwin & R. Harin’Hala, transitional forest, Malaise trap [1♀, CASC: CASENT2022276].

Ivieosema limula Burks, sp. n.

Figs 49–54, 55

Etymology

Latin noun meaning “small file (carpenter’s or sculptor’s tool),” referring to the finely striate mesosomal surface sculpture.

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.

Head (Figs 50–51) 1.4× as broad as long. Face flat and smooth; medial scrobal depression absent, but frons with paired dorsal frontal depressions lateral to scrobal area (Fig. 50, dfd); lower frons with a distinct vertical groove (Fig. 50, lg); scrobal area with a single medial large pit just below median ocellus; supraclypeal area smooth, clypeus shallowly irregularly sculptured, anterior tentorial pits very deep and broad, anteclypeus very short; scrobal depression broadly and shallowly impressed; vertex not carinate. Labrum with 4 digits. Maxillary and labial palps absent. Flagellum 1.2× head height, 7 funiculars; anellus slightly broader than long, expanding apically (Fig. 51, F1); F2 2.0× as long as broad, F2 1.1× as long as F3.

Mesosoma (Figs 52–53, 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. Gt1 with a deep longitudinal median depression extending length of tergite and with smooth lateral margins. Antecostal sulcus of Gs1 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.

Male

Unknown.

Material examined

Holotype: Madagascar: Toliara Prov: Manderano, 70m, 23°31'39"S, 44°05'18"E, 23.vi-28.vii.2002, Frontier Wilderness Project, gallery forest at the edge of marsh, 5m from road, Malaise trap [1♀, CAS: CASENT2022276], deposited in CAS.

Figure 55.

Ivieosema limula. Holotype female: fore wing.

Leiosema Heraty & Burks, gen. n.

Figs 56, , , , , , , , , , , 67

Type species

Orasema glabra Heraty, 1994: 72–73, figs 119, 125–128, 130, 132.

Etymology

Combination of the Greek λεῖος (leios) for smooth and a contraction of Orasema, referring to the smooth and polished head and frenum; gender feminine.

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 5758, 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.

Diagnosis

Separated from Orasema by the presence of male scape glands (pores) (Fig. 56, vp), broadly impressed rugulose antecostal sulcus and the combination of smooth head, 4-digitate labrum, and smooth frenum (Figs 5961). Distinguished from the other Old World genera by having the head smooth and subcircular in frontal view (Figs 5762); scrobal depression with parallel channels and dorsal foveae (Fig. 57); dorsal occipital margin rounded; funicle 7-segmented; labrum with 4 digits; mesonotum with scattered short setae; mesoscutal lateral lobes and frenum smooth; transscutal articulation complete; propodeal disc evenly rugose; prepectus foveate and loosely articulated with pronotum ventrally (Fig. 63); fore wing with basal area and specular area bare (Fig. 65), with dense and relatively long wing disc setae; postmarginal vein 1.5–3.2× as long as stigmal vein; petiole base truncate with prominent basal flange (Fig. 67); antecostal sulcus broad and reticulate; first valvula of ovipositor with 6–10 minute lateral teeth. Male scape with distinct ventral pores (glands) set in circular depressions across entire ventral surface (Fig. 56).

Distribution

Sub-saharan Africa (Democratic Republic of the Congo, South Africa, Tanzania, Zimbabwe), and Yemen.

Leiosema lesiolouna Heraty & Burks, sp. n.

Figs 61, 62, 64, 66

Etymology

Named for the collection locality of the holotype.

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.

Head (Figs 6162) 1.2× as broad as high; inter-ocular distance 1.2× eye height; malar space 0.4× eye height; supraclypeal area longer than broad, smooth; clypeus smooth; epistomal sulcus distinct and sharply defined; anteclypeus distinct, straight. Mandibular formula 3:3, apical tooth elongate. Maxilla and labium of normal size, palpal formula 3:2. Occiput striate, shallowly emarginate in dorsal view, dorsal margin evenly rounded, temples present, rounded and coriaceous. Antenna 11-segmented; scape narrow and cylindrical, not reaching median ocellus; pedicel subconical, as broad as F1; flagellum 1.4× head height; anellus minute, difficult to distinguish, F2 1.2× as long as broad, 1.2× as long as F3; following funiculars subequal in length, equal in width; clava subcylindrical, longer than preceding 2 funiculars, clava without distinct separation of fused clavomeres.

Mesosoma (Figs 61, 64). Mesoscutal midlobe reticulate, interstices rounded, dorsally with few sparse setae; lateral lobe smooth and shining; notauli deeply impressed. Axilla smooth; dorsally rounded, on roughly same plane as scutellum; scutoscutellar sulcus broad, irregularly foveate. Scutoscutellar sulcus meeting transscutal articulation; scutellar disc about 1.5× as long as broad, reticulate, smooth just anterior to frenal line; frenal line as distinct crenulate groove; axillular sulcus distinct and foveate; axillula very weakly reticulate. Propodeal disc broadly rounded, without depression or carina, rugose-areolate; callus smooth, with several long hairs; callar nib absent. Prepectus evenly triangular, sculpture broadly foveate. Mesepisternum reticulate laterally, smooth ventrally, broadly rounded anterior to mid coxa. Upper mesepimeron smooth; lower mesepimeron smooth; transepimeral sulcus distinct. Propleuron convex, smooth. Postpectal carina prominent. Hind coxa 1.8× as long as broad, smooth with very weak reticulate sculpture; hind femur 5.7× as long as broad, with even cover of appressed long setae; hind tibia densely setose. Fore wing 2.4× as long as broad; basal cell and speculum bare, costal cell and wing disc densely setose, wing disc setae long; marginal fringe relatively long; submarginal vein with several long setae; marginal vein with minute setae; stigmal vein more than 2.5× as long as broad, slightly angled, slightly narrowed basally and capitate apically (Fig. 66); uncus absent; stigma with 3 sensilla in a straight line; postmarginal vein 1.9× as long as stigmal vein. Hind wing costal cell with a broad bare area.

Metasoma. Petiole cylindrical, linear in profile, 4.6× as long as broad, 1.7× as long as hind coxa, reticulate, anterior carina strong, lateral margin rounded, ventral sulcus present with margins narrowly separated. Antecostal sulcus rugulose; acrosternite posteriorly angulate, raised and smooth medially and laterally broadly impressed and reticulate. Genitalia protruding: parameres elongate, digitus broad and flattened with 4 marginal spines.

Female

unknown.

Holotype

Republic of Congo: Pool Dept.: Iboubikro, Lesio-Louna Pk., 340m, 3°16'12"S, 15°28'16"E, 23.vii.2008, Sharkey & Braet, Malaise trap #3 [1♂, UCRC: UCRCENT00408466].

Leiosema glabra (Heraty), comb. n.

Figs 56, , , , 60, 63, 65, , 67

Orasema glabra Heraty, 1994: 72–73. Holotype, South Africa, Transvaal [♀, CNCI]. Mistakenly figured in Heraty (1994) with 6 funicular segments (cf. Fig. 126) instead of 7.

Diagnosis

Both sexes with mandibles 3:2 dentate, mesoscutal midlobe and scutellar disc weakly reticulate, becoming smooth posteriorly (Figs 5960). The pronotal neck is not extended, and is reticulate only along the anterior margin (Fig. 60). Male petiole 2.6–2.9× as long as broad, female petiole 1.5–2.1× as long as broad, with a strong basal carina continuous with a lateral carina and extending 1/3 to 2/3 length of petiole (Fig. 67).

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 1.8–2.4 mm. 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.

Head (Figs 6162) 1.1–1.2× as broad as high; inter-ocular distance 1.4–1.5× eye height; malar space 0.3–0.7× eye height; supraclypeal area slightly broader than long, longer than clypeus, smooth; clypeus smooth; epistomal sulcus distinct and sharply defined; anteclypeus weakly differentiated, straight. Mandibular formula 3:2. Flagellum 1.0–1.2× head height. F2 1.2–1.3× as long as broad, 1.2–1.3× as long as F3; clava subconical, longer than preceding two funiculars and weakly segmented.

Mesosoma (Figs 61, 64). Mesosoma 1.5–1.7× as long as high. Mesoscutal lateral lobe smooth and shining dorsally, weak rugulose laterally. Scutellar disc 1.5–1.7× as long as broad, reticulate; axillula weakly rugulose, nearly smooth. Pronotal neck not extended, at most reticulate along anterior margin. Mesepisternum straight anterior to mid coxa. Propleuron coriaceous. Hind coxa 1.1–2.0× as long as broad, smooth to coriaceous; hind femur 4.2–5.4× as long as broad, with even cover of elongate, semi-erect setae. Fore wing 2.1–2.3× as long as broad; marginal vein with long setae; stigmal vein 1.5–2.6× as long as broad, slightly angled; stigma nearly quadrate but with short apical anterior extension; stigma with 3 sensilla in a straight line; postmarginal vein 1.1–3.2× as long as stigmal vein. Hind wing costal cell bare.

Metasoma. Petiole 2.6–2.9× as long as broad, 1.2–1.4× as long as hind coxa, lateral margin rounded with only short lateral carina in basal 1/4. Acrosternite posteriorly rounded with rugulose sculpture, antecostal sulcus rugulose.

Female

(redescription) (Fig. 59, holotype). Length 2.0–2.6 mm. Scape yellow. Fore femur mostly dark brown to black, tip pale; fore tibia yellow; mid femur mostly dark brown to black, tip pale; mid tibia yellow; hind femur mostly dark brown to black, tip pale; hind tibia pale brown basally, yellow apically.

Head 1.0–1.1x as broad as high; eyes separated by 1.3–1.5× eye height; malar space 0.4–0.6× eye height. Flagellum length 0.8–1.0× head height (0.9–1.0 in type material); flagellum with 7 funiculars, anellus disc-shaped, F2 1.1–1.4× as long as broad, 1.1–1.2× as long as F3.

Mesosoma 1.6–2.1x as long as high. Fore wing 2.3–2.6× as long as broad.

Metasoma. Petiole 1.5–2.1× as long as broad, 0.9–1.3× as long as hind coxa, apical setae of hypopygium with 2–4 long hairs clustered on each side of midline. Ovipositor slightly curved cephalad; first (ventral) valvula with 6–10 small, narrowly separated teeth, second (dorsal) valvula with 6–7 annuli that are broadly separated dorsally by smooth area; subapical carina present; third valvula separated.

Material examined

South Africa: Eastern Cape: Port St. John, 31°37'00"S, 29°32'00"E, i.1974, A. Watsham [1♂, NMPC: UCRCENT00416408]. Orange Free State: Koppies Nat. Res., 26°10'11"S, 27°59'46"E, 22-23.ii.1993, M. Stiller, Acacia karroo, sweep [1♀, NMPC: UCRCENT00416471]. Transvaal: Blyderiviers-poort, 1200m, 24°53'00"S, 30°45'00"E, iii.1981, G. L. Prinsloo [1♂, PPRI: UCRCENT00298795]. Waterberg nr Nylstroom [Modimolle], 1200m, 24°48'00"S, 28°25'51"E, i.1980, G. L. Prinsloo [1♂, PPRI: UCRCENT00298796]. Strijdom Tunnel, 730 m, 24°27'47"S, 30°36'31"E, 24-25.xi.1991, G.L. Prinsloo [1♀, NMPC: UCRCENT00416472]. Tanzania: Tanga: West Usambara, Mgwashi env., 1460m, 4°45'47"S, 38°28'30"E, 31.i-1.ii.2015, P. Jansta & J. Straka [3♂ 1♀ 2?, UCRC: UCRCENT00479260–65]. Zimbabwe: Harare: Chishawasha, 1416m, 17°45'46"S, 31°12'31"E, A. Watsham [2♀, NMPC: UCRCENT00416454, UCRCENT00416456]. Zimbabwe: Mashondland: Mazowe, 125m, 17°30'32"S, 30°58'19"E, i.1975, A. Watsham [6♀, NMPC: UCRCENT00416459–61, UCRCENT00416466–67, UCRCENT00416519]. Makumbi Miss., 17°31'12"S, 31°15'22"E, A. Watsham [1♂, NMPC: UCRCENT00416462].

Orasema near glabra: Gambia: Banjul: Kotu Stream, 3m, 13°27'41"N, 16°42'13"W, 14.xi.1981, K-J. Hedqvist [5♂ 4♀, BMNH: UCRCENT00309860–68]. Kotu Stream, 4m, 13°27'42"N, 16°42'13"W, 18.xi.1981, K-J. Hedqvist [1♀, BMNH: UCRCENT00309871]. North Bank Division: Upper Baddibu, Farafenni, 13m, 13°34'03"N, 15°44'56"W, 1-2.xi.1977, Michael Söderlund, Meadow (mostly resorts) with baobab [1♂, BMNH: UCRCENT00309873]. Western Division: Kombo St Mary, Bakau, 10m, 13°28'47"N, 16°40'14"W, 28-29.x.1977, Michael Söderlund, shore meadow [1♂, BMNH: UCRCENT00309872]. Kombo St Mary, Sara Job Kunda, 25m, 13°26'40"N, 16°42'46"E, 29.x.1977, Michael Söderlund, grassland with palms [1♂, BMNH: UCRCENT00309859].

Losbanus Ishii, rev. stat.

Losbanus Ishii, 1932: 210. Type species: Losbanus uichancoiIshii 1932: 210–211, by monotypy.

Losbanus; Hedqvist, 1978: 229. Short redescription and key to species, most of which are now attributed to Neolosbanus (Eucharitinae).

Losbanus; Watanabe, 1958, 26. Subsequent description.

Gollumiella; Bouček, 1988: 521. Incorrectly placed as senior synonym of Gollumiella.

Orasema; Heraty, 1992: 586. Synonymy based on transfer of L. uichancoi to Orasema.

Orasema; Heraty, 1994: 54–57. Description and treatment as Orasema uichancoi-group.

Orasema; Heraty, 2000: 378. Separation of communis and uichancoi species groups.

Discussion

Losbanus was first described for a single species, L. uichancoi. Bouček (1988) incorrectly used this as a senior synonym of Gollumiella. Heraty (1992) transferred the type species to Orasema, then used it to represent the Orasema uichancoi-group in Heraty (1994), and in a more restricted sense for all of the Indo-Pacific species in Heraty (2000). Herein we recognize this more restricted uichancoi group (sensu Heraty, 2000) as Losbanus. We have discovered two new species (not described) that expand the geographic and morphological limits of Losbanus. A single male from the Republic of the Congo (UCRC: UCRCENT00434658) fits all of the characteristics of the group and extends the distribution to Africa. Another series of males from northeastern Queensland in Australia (Charmillin Creek and Baldy Mountain Rd; QM: UCRCENT00241810–14) overall fit the description of the genus, but lack the ocellar-ocular sulcus and have a 5-7 digitate labrum, whereas the other species have 4 digits. All species of Losbanus have 8–9 funiculars, a 12-13 segmented antenna, and densely pilose wings. The following species are transferred from Orasema: L. bouceki (Heraty), comb. n., L. ishii (Heraty), comb. n., L. promecea (Heraty), comb. n., L. rugulosa (Heraty), comb. n., and L. uichancoi Ishii comb. n. (key in Heraty, 1994).

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).

Host association

Unknown.

Distribution

Five species described from southeast Asia (Heraty, 1994). Paleotropical (Australia: Queensland (undescribed); Republic of Congo (undescribed), Papua New Guinea, Philippines, Taiwan).

Matantas Heraty & Burks, gen. n.

Type species

Orasema koghisiana Heraty, 1994: 71–72, figs 107–108, 110, 189–190, 211–212, 219, 236, 247.

Etymology

Named after one of the New Caledonia collecting localities, Matanta; gender feminine.

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).

Zuparka Heraty & Burks, gen. n.

Figs 68–73, 74–78, 79–80, 81–86

Type species

Zuparka monomoria Heraty, 2000: 375–378, figs 1–7.

Etymology

Named after Robert Zuparko of the California Academy of Sciences; gender feminine.

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).

Host association

Myrmicinae: Z. monomoria reared from Monomorium sp. (Heraty 2000).

Distribution

Madagascar.

Figs 56–67.

Leiosema glabra. Male: 56 antenna, lateral (Tanzania, TZ) 57 head, frontal, (Yemen) 58 head, oblique (TZ). Female holotype 59 head and mesosoma, dorsal; Male: 60 head and mesosoma, dorsal (South Africa) 63 mesosoma, lateral (SA) 65 wings, dorsal (TZ) 66 stigmal vein (TZ) 67 propodeum and petiole, oblique (TZ). Leiosema lesiolouna. Holotype male: 61 head and mesosoma, dorsal 62 head, oblique 64 head and mesosoma, lateral.acr = acrosternite; apc = anterior petiolar carina; F = flagellomere; frn = frenum; lbr = labrum; no1 = pronotum; scp = scape; vp = ventral pores.

Figures 68–73.

Zuparka fisheri. Paratype female: 68 habitus, lateral 69 head, anterior 70 mesosoma, lateral 71 mesosoma, oblique dorsal 72 mesosoma, anterior 73 propodeum, oblique dorsal. apc = anterior petiolar carina; cal = callus; frn = frenum; lpr; lateral propodeal disc; mts = metascutellum; sct = mesoscutellum; tsa = transscutal articulation.

Figures 74–78.

Zuparka fisheri. Paratype female: 74 fore wing 75 petiolar base 76 ovipositor. Paratype male: 77 antenna 78 pores on scape ventrally. ads = anterodorsal swelling of petiole; apc = anterior petiolar carina; dv = dorsal (2nd) valvula; F = flagellomere; F1/an = anellus; lhp = petiolar lateral hair patch; vp = ventral pore of scape; pmv = postmarginal vein; stv = stignal vein; vv = ventral (1st) valvula.

Figures 79–80.

Zuparka fisheri. Paratype male: 79 head, anterior 80 genitalia and subgenital plate, ventral. acy = anteclypeus; dsd = dorsal scrobal fovea; msd = median scrobal depression; prm = paramere; sgp = subgenital plate; tor = torulus.

Zuparka fisheri Heacox & Dominguez, sp. n.

Figs 68–73, 74–78, 79–80

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.

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.

Female

Length 1.8–2.4 mm. Head and mesosoma, including mid and hind coxae and petiole, dark bluish-black; fore coxae dark bluish-black and yellow apically. Flagellum brown; scape, pedicel and anellus yellow to brown. Legs beyond coxa yellow with femora medially brown and tarsomeres apically pale brown. Gaster dark brown. Wings hyaline, venation pale yellowish-brown.

Head (Figs 69, 7779) 2.2–2.6× as broad as long; face broadly rounded, smooth and polished with faint indication of cellulate sculpture on upper frons and vertex; interocular distance 1.3–1.4× eye height; malar space 0.5–0.7× eye height; distance from anteclypeus to ventral margin of toruli 0.3–0.4× head height; clypeus smooth and with scattered setae, lateral margins and epistomal sulcus strongly impressed; anteclypeus distinct and subtruncate. Mandibular formula 3:2; palpal formula 3:3. Occiput transversely aciculate, dorsal margin broadly rounded, occipital carina absent; temples broad. Antenna 11-segmented (Fig. 77); scape narrow and cylindrical, slightly broadening medially and reaching median ocellus, at least 7× as long as broad; pedicel 1.3–2.0× as long as broad, apex broader than anellus; flagellum 0.8–0.9× head height; funicle 7-segmented; flagellar segments beyond anellus densely setose with numerous longitudinal sensilla; F2 1.2–1.6× as long as broad, following segments becoming more transverse; clava subconical, about as long as preceding three segments.

Mesosoma (Figs 7074). 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 (Figs 71–72); 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.

Metasoma (Figs 75–76) with petiole cylindrical, 1.6–1.9× as long as hind coxa; finely reticulate; ventral sulcus absent. Gt1 as long as or slightly shorter than femur, smooth and bare; Mt7 densely pilose anterior to cercus, cercus with 5 long setae; Gs1 smooth and without antecostal sulcus. Hypopygium with 2 pairs of minute subapical setae. Ovipositor expanded subapically and curved forward; first valvula with 3 lateral teeth and sinuate subapical ridge; second valvula with 8 lateral teeth (Fig. 76).

Male

Length 1.7–1.8 mm. Scape yellow to brown, flagellomeres brown. Fore coxa bluish-black and brown apically; legs yellow to yellowish-brown, hind femur darker with yellow tips, tarsomeres yellow to brown. Scape ventroapically with dense minute pores. Gs8 apically strongly convex with sparse ventral hairs; Ms8 densely setose near cerci. Genitalia with broad flattened parameres with two subapical setae, digiti with 4 lateral spines; aedeagus subapically expanded (Fig. 80).

Holotype

Madagascar: Fianarantsoa Prov.: Parc National Ranomafana, Belle Vue at Talatakely, 1020m, 21°15'59"S, 47°25'07"E, 2-10.i.2002, R. Harin’ Hala, secondary tropical forest, Malaise trap [1♀, CASC: UCRCENT00417485]. Paratypes: Madagascar: Antananarivo Prov.: 3 km 41 degrees NE Andranomay, 11.5 km 147 degrees SSE Anjozorobe, 1300m, 18°28'24"S, 47°57'36"E, 5-13.xii.2000, Fisher, Griswold et al., montane rainforest, Malaise trap [1♂, CASC: UCRCENT00417369]. Antsiranana Prov.: Parc National Montagne d’Ambre, 960m, 12°30'52"S, 49°10'53"E, 12.ii-4.iii.2001, 4-19.iii.2001, R. Harin’Hala, Malaise [5♂ 1♀, CASC: UCRCENT00175169, UCRCENT00278241–45]. Parc National Montagne d’Ambre, 1125m, 12°31'13"S, 49°10'45"E, 29.i-11.ii.2001, 5-21.iv.2001, 14-30.v.2001, 30.v-6.vi.2001, R. Harin’Hala, Malaise trap [4♂ 1♀, CASC: CASENT02014649, CASENT02015483, CASENT02015488, UCRCENT00417343, UCRCENT00417345]. Sakalava Beach, 10m, 12°15'46"S, 49°23'51"E, 16-31.v.2001, R. Harin’ Hala, dwarf littoral forest, sandy trail, Malaise trap [1♂, CASC: UCRCENT00417339]. Fianarantsoa Prov.: Parc National Ranomafana, Belle Vue at Talatakely, 1020m, 21°15'59"S, 47°25'13"E, 28.xi-6.xii.2001, M. Irwin & R. Harin’Hala, secondary tropical forest, Malaise Trap [1♂ 1♀, CASC: UCRCENT00417551, UCRCENT00417553]. Parc National Ranomafana, radio tower at forest edge, 1130m, 21°15'03"S, 47°24'26"E, 16.x-8.xi.2001, 24.v-4.vi.2002, 7-17.v.2003, R. Harin’ Hala, mixed tropical forest, Malaise [2♂ 4♀, CASC: UCRCENT00417386–487, UCRCENT00417388, UCRCENT00417489, UCRCENT00417490, UCRCENT00417492]. Parc National Ranomafana, Vohiparara at broken bridge, 1110m, 21°13'34"S, 47°22'11"E, 21-28.i.2002, 4-12.ii.2002, M. Irwin and R. Harin’Hala, high altitude rain forest, Malaise trap [1♂ 2♀, CASC: UCRCENT00417509–10, UCRCENT00417537]. Toamasina Prov.: bot. garden near entrance to Andasibe National Park, 1025m, 18°55'35"S, 48°24'28"E, 16-24.x.2001, 24.x-1.xi.2001, 1-7.xi.2001, R. Harin’Hala, tropical forest, Malaise Trap [2♂ 2♀, CASC: UCRCENT00417486, UCRCENT00417497, UCRCENT00417513, UCRCENT00417538].

Additional material examined

Madagascar: Antsiranana Prov.: Parc National Montagna d’Ambre, 1125m, 12°31'13"S, 49°10'45"E, 12.ii-4.iii.2001, 19.iii-5.iv.2001, 12-14.v.2001, R. Harin’Hala, Malaise trap [3♀ 1♂, CASC: CASENT02015492, UCRCENT00417340, UCRCENT00417344, UCRCENT00417346]. Sakalava Beach, 10m, 12°15'46"S, 49°23'51"E, 16-31.v.2001, R. Harin’ Hala, dwarf littoral forest, sandy trail, Malaise trap [1♀, CASC: UCRCENT00417338].

Zuparka monomoria (Heraty), comb. n.

Figs 81–84

Orasema monomoria Heraty, 2000: 376–378.

Diagnosis

Frons weakly coriaceous. Scutellum evenly rounded anterior to frenum. Propodeal disc laterally weakly sculptured.

Discussion

Previously described from a single female that had recently emerged. The new material examined fits the description of the single female.

Distribution

Madagascar.

Holotype

Madagascar: Fianarantsoa Prov.: Res. Andringitra, 8.5 km SE of Antanitotsy, 1990m, 22°10'00"S, 46°58'00"E, 6.iii.1997, Sylvian, montane rainforest [1♀, UCRC: UCRCENT00417370]. Additional material examined: Madagascar: Majunga Ampijoroa National Park, 160 km N of Maevatanana on RN 04, 43m, 16°19'10"S, 46°48'48"E, Rin’ha Harin’Hala, deciduous forest, Malaise trap [1♀, CASC: UCRCENT00417389]. Antananarivo Prov.: 3km 41d NE Andranomay, 1300m, 18°28'24"S, 47°57'36"E, 5-13.xii.2000, Fisher, Griswold, et al. CASC, montane rainforest, Malaise trap [1♂, CASC: UCRCENT00278242]. Ankokoy Forest, 3 km E of Ibity, 1700m, 20°04'03"S, 46°59'58"E, 12-22.xii.2008, M.Irwin, R.Harin’Hala, Uapaca forest, Malaise trap [1♀, CASC: UCRCENT00352446]. Antsiranana Prov.: Parc National Montagne d’Ambre, 960m, 12°30'52"S, 49°10'53"E, 4-19.iii.2001, 5-21.iv.2001, R. Harin’ Hala, Malaise trap [3♀, CASC: CASENT02015482, CASENT02015484, CASENT02015487]. Diego Suarez: PN Montange d’Ambra, 960m, 12°30'52"S, 49°10'53"E, 21-26.i.2001, M.E. Irwin, E.I. Schlinger & R. Harin‘Hala, Malaise trap [1?, CASC: UCRCENT00417358]. Fianarantsoa Prov.: Miandritsara Forest, 40 km S Ambositra, 825m, 20°47'34"S, 47°10'32"E, 28.xii.2006-4.i.2007, M. Irwin, R. Harin’Hala, low altitude rainforest, Malaise trap [1♀, CASC: CASENT02174550]. near Isalo National Park, at stream E of Interpretive Cntr., 750m, 22°37'36"S, 45°21'29"E, 7-17.v.2003, R. Harin’ Hala, mixed tropical forest, Malaise trap in open area [1♀, CASC: UCRCENT00417378]. Parc Nacional Ranomafana, Belle Vue at Talatalely, 1020m, 21°15'59"S, 47°25'13"E, 22.xi.2001, 6-12.xii.2001, 28.xi-6.xii.2001, 19-26.ii.2002, 10-14.i.2002, 12-19.ii.2002, 23-28.iv.2002, 5-13.v.2002, 15-22.xi.2002, 16-26.ii.2003, 26.ii-10.iii.2003, 10-21.iii.2003, 28.v-6.vi.2003, R. Harin’ Hala, secondary forest, Malaise [4♂ 21♀, CASC: CASENT02174558–60, CASENT02174556, CASENT02174592, CASENT02174594–97, UCRCENT00417360; UCRC: UCRCENT00417481, UCRCENT00417495–96, UCRCENT00417503, UCRCENT00417514, UCRCENT00417531–33, UCRCENT00417535, UCRCENT00417390–93]. Parc National Ranomafana, JIRAMA water works, 690m, 21°14'55"S, 47°27'08"E, 28.i-4.ii.2002, R.Harin Hala, near river, Malaise Trap [1♀, CASC: CASENT02174591]. Parc National Ranomafana, radio tower at forest edge, 1130m, 21°15'03"S, 47°24'26"E, 14-21.i.2002, 28.i-4.ii.2002, 4-12.ii.2002, 7-17.v.2003, 7-18.vi.2003, 27.vi-7.vii.2003, R. Harin’ Hala, mixed tropical forest, Malaise trap [14♀ 3♂, CASC: UCRCENT00417467, UCRCENT00417371–77, UCRCENT00417379–85, UCRCENT00417387, UCRCENT00417530]. Parc National Ranomafana, Vohiparara at broken bridge, 1110m, 21°13'34"S, 47°22'11"E, 6-15.xii.2001, 14-21.i.2002, 21-28.i.2002, 4-12.ii.2002, 19-26.ii.2002, 6-15.v.2002, M. Irwin and R. Harin’Hala, high altitude rain forest, Malaise trap [5♂ 9♀ 3?, CAS: CASENT02174562, CASENT02174583, CASENT02174587–89; UCRC: UCRCENT00417508, UCRCENT00417511–12, UCRCENT00417525–29, UCRCENT00417534, UCRCENT00417465–66, UCRCENT00417468]. Parc National Ranomafanae, radio tower at forest edge, 1130m, 21°15'03"S, 47°24'26"E, 12-19.ii.2002, R. Harin’Hala, mixed tropical forest, Malaise trap [2♂ 1♀, CASC: UCRCENT00417443, UCRCENT00417501–02]. Parc National Ranomafanae, radio tower at forest edge, 1130m, 21°15'03"S, 47°24'26"E, 21-28.i.2002, 4-12.iii.2002, 12-19.iii.2002, 27.ii-9.iii.2003, 20.iii-3.iv.2003, 15-27.iv.2003, 24.v-4.vi.2002, 6-17.vii.2003, R. Harin’Hala, mixed tropical forest, Malaise trap [6♂ 7♀ 1?, CASC: CASENT02174580–81, UCRCENT00417441, UCRCENT00417488, UCRCENT00417491; UCRCENT00417493–94, UCRCENT00417498, UCRCENT00417504–07, UCRCENT00417536, UCRCENT00417554]. Ranomafana National Park Talatakely area, 900m, 21°15'02"S, 47°25'10"E, 9-19.i.2001, D.H. & K.M. Kavanaugh, R.L. Brett, E. Elsom, F. Vargas, mixed tropical forest, flight intercept trap [1♀, CASC: UCRCENT00417347]. Res. Andringitra, 8.5 km SE of Antanitotsy, 1990m, 22°10'00"S, 46°58'00"E, 6.iii.1997, Sylvian, montane rainforest [1♀ 1?, UCRC: UCRCENT00412588–89]. Toamasina Prov.: bot. garden near entrance to Andasibe National Park, 1025m, 18°55'35"S, 48°24'28"E, 16-23.xi.2001, R. Harin’Hala, tropical forest, Malaise Trap [1♂, CASC: UCRCENT00417494].

Figures 81–86.

81–84 Zuparka monomoria. Female: 81 head, frontal 82 mesosoma, lateral 83 habitus, lateral 84 head and mesosoma, dorsal 85 unknown male Oraseminae, Australia: UCRCENT00238798 (couplet 10) 86 unknown male Oraseminae, Australia: UCRCENT00241816 (couplet 15).

Acknowledgements

We would like to thank Gary Gibson and Lisa Bearss for imaging the holotype of Orasema glabra. We would like to thank Austin Baker, Chris Darling and Javier Torréns for comments on earlier drafts of the manuscript. This research was supported by NSF DEB 1257733 and UCR Hatch grants to JMH. Mention of trade names or commercial products in this publication is solely for the purpose of providing specific information and does not imply recommendation or endorsement by the USDA; USDA is an equal opportunity provider and employer.

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