re

JHR 55: 139-188 (2017) JOURNAL OF "ewe sna
doi: 10.3897/jhr.55.1 1482 ME Hymenopter a

http://jhr.pensoft.net The narational Society of Hymenoptersts. RESEARCH

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

Roger A. Burks', John M. Heraty', Jason Mottern'?,
Chrysalyn Dominguez', Scott Heacox'

| Department of Entomology, University of California, Riverside, CA 925212 USDA Systematic Entomology
Laboratory, Washington, DC 20013-7012

Corresponding author: John M. Heraty (john.heraty@ucr.edu)

Academic editor: P Jansta | Received 14 December 2016 | Accepted 24 March 2017 | Published 28 April 2017
http.//zoobank. org/OE1BOA14-F871-4CD8-9F29-10B255677621

Citation: Burks RA, Heraty JM, Mottern J, Dominguez C, Heacox S (2017) Biting the bullet: revisionary notes on
the Oraseminae of the Old World (Hymenoptera, Chalcidoidea, Eucharitidae). Journal of Hymenoptera Research 55:
139-188. https://doi.org/10.3897/jhr.55.11482

Abstract

Twelve genera of Oraseminae (Hymenoptera: Eucharitidae) are recognized in the Old World. The ge-
nus 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., [bitya gen. n., [vieosema 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. (Aus-
tralia), Cymosema capelina sp. n. (Australia), C. waterworthae sp. n. (Australia), Hayatosema plicator sp. n.
(Vietnam), vieosema 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 Ovasema)
include Australosema valgius (Walker, 1839), A. synempora (Heraty, 1994), Hayatosema assectator (Ker-
rich, 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), lbitya communis (Risbec, 1952), Ib. seyrigi (Risbec, 1952),
Ivieosema fraudulenta (Reichensperger, 1913), lv. 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 [bitya communis, and an identification key to the Old World genera and

species are provided.

Copyright Roger A. Burks et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

140 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

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 Ora-
seminae 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 Monomo-
rium 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 south-
western highlands of Yemen (Heraty 1994, 2002).

The Old World species of Oraseminae were previously revised by Boucek (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 Boucek, and Timi-
oderus 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 Ovasema (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 /ndosema), 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 anteri-
orly 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.1. 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

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 141

Table |. 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 Host genus
Australosema gen. n.
(-valgius group) Pheidole

A. verghetta sp. n. Australia

C. capelina sp. n. Australia

Cymosema gen. n. ;
C. waterworthae sp. n. * Australia

H. assectator (Kerrich, 1963)* Indo-Pacific Pheidole
H. delhiensis (Narendran & G. Kumar, 2005) Indo-Pacific

H.. initiator (Kerrich, 1963) Indo-Pacific
Hayatosema gen. n. H. kailashi (G. Kumar & Sureshan, 2015) Indo-Pacific
(=assectator group) H. nigra (Heraty) Ethiopian

H. nirupama (G. Kumar & Narendran, 2007) Indo-Pacific
LH. plicator sp. n. Indo-Pacific
H. siruvanica (G. Kumar & Sureshan, 2015) Indo-Pacific
Ibitya gen. n. I, communis (Risbec, 1952), comb. n. * Madagascar Pheidole NR

(=communis group) L. seyrigi (Risbec, 1952), comb. n. Madagascar
Indosema L. indica Husain & Agarwal, 1983 * India & Kenya
L. confluens sp. n. Madagascar

Ivieosema gen. n. L. limula sp. n. Madagascar

(=striatosoma group) | I. fraudulenta (Reichensperger), comb. n. Ethiopian Pheidole
L. striatosoma (Heraty), comb. n. * Ethiopian

Leiosema gen. n. L. glabra (Heraty, 1994), comb. n. * Ethiopian

(=glabra group) L. lesiolouna sp. n. Ethiopian

L. bouceki (Boucek), comb. n. New Guinea

L. ishii (Heraty), comb. n. Taiwan
Losbanus rev. stat. :
L. promecea (Heraty), comb. n. New Guinea

(=uichancoi group)
L. rugulosa (Heraty), comb. n. New Guinea

L. uichancoi Watanabe comb. n. * Philippines

Ae M. koghisiana (Heraty), comb. n. * New Caledonia
O. didentata (Girault, 1940) Australia
O. eribotes (Walker, 1839)* Australian Pheidole
O. goethei (Girault, 1934) Australia
O. myrmicae (Girault, 1936) Australia Pheidole
Orasemorpha 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

T. acuminatus Heraty, 1994 Ethiopian Pheidole
T. coronula Heraty, 1994 Ethiopian

Timioderus T. peridentatus Heraty, 1994 Ethiopian
T. coronula Heraty, 1994 Ethiopian

T. refringens Waterston, 1916 * Ethiopian
Zuparka gen. n. Z. fisheri sp. n. Madagascar

(=monomoria group) |Z. monomoria (Heraty, 2000), comb. n. * Madagascar Monomorium

142 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

in either case, Orasema is never monophyletic. Character support for a monophyletic
Orasema s.\. 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 Ovasema. Given the wealth of morphological
differences in the three other genera of Oraseminae, the associated biological differ-
ences, 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 gen-
era and recognize only the monophyletic New World clade as Orasema sensu stricto.
Herein we recognize 12 Old World genera, of which eight are new and one, Losbanus,
is given revised status. Several new species are described, bringing the number of Old
World species to 45, which is far overshadowed by the incredible diversity of Orasema
in the New World.

Materials and methods

Material was examined from ANIC: Australian National Insect Collection, Canberra;
BMNH: Museum of Natural History, London, UK; BPBM: Bernice P. Bishop Mu-
seum, 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 Insti-
tute, 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 Mu-
seum 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

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 143

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 [ndosema, Orasemorpha and Timi-
oderus, which were revised in Heraty (1994). The identification key includes treat-
ments 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 ante-
rior margin broadly rounded posterior to stigmal vein making fore wing ap-
pear ovate (Figs 15, 27). Petiole narrowed gradually into foramen (Fig. 20),
lacking anterior petiolar flange (Australian; males unknown).......... eee
cut Gainer aavaet wants les og ietuc Geert uv eshere debe stan eee dusk chan ees 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).
PEC Halle Wat aD bee ciy op vests vest on csaetrane sist extenrie the sea cnstecestoe sisi ou icons teow ostoncicana teaser? 3
Mandibles minute and lacking distinct teeth (Fig. 16, md). Mesosomal dor-
SumuenitirelyssmG Oth ie MU)» 63..035 a. ssSsnsteasneceedeoneseeutt 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.
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) (Beii@ pianeand Wala CaS) ka.s.ssis, vobeacesmsestasehunenduteamtp Mae attnouag stbavnsnmurhs
Py ee Ivieosema gen. n. (striatosoma-group of Heraty 1994)...4
Upper frons without dorsal pit. Other characters variable... eee eeeeeeeeeees vi
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 articu-
lation fused medially. Fore wing with very short setae; with distinct longitu-
dinal swelling posterior to submarginal vein (Figs 44, 55) (Malagasy)......... 6
Mesoscutellum reticulate dorsally... Ivieosema fraudulenta (Reichensperger)
Mesoscutellum longitudinally striate.......... Ivieosema striatosoma (Heraty)
Face smooth. Scape brown. Upper frons with two depressions between eye
and antennal scrobe (Figs 39, 47) ......scseseseeee Ivieosema confluens sp. n.

144

9 (7)

10 (9)

11 (9)

Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

Face smooth to shallowly reticulate. Scape white. Upper frons with single
transverse depression between eye and antennal scrobe (Fig. 50)... eee
aaa eee Sessa Aiea sas eae aco at a ee pane cas a oak Ivieosema limula sp. n.
Transscutal articulation (between mesoscutum and axilla) obliterated medi-
allivecWiaxillaryraid Waly tal paliot-abSetit oss. 5ncte8 le ctanete ted Meee BS, fo ee 8
Transscutal articulation complete with a distinct transverse suture (Fig. 4).
Masa llaeysang labial spate lepresetit 5, ..sanera ee Gt, .o rade aR eee eo reso anal 9
Body metallic blue or green. Head and mesosoma strongly sculptured and ei-
ther coriaceous or rugulose. Anellus present. Mandibles truncate or spatulate,
maxilla small. Petiole not fused ventrally. Ovipositor curved ventrally. Body
lengehrar leasti3:5 samme Ethiopian) \.....cs3sesee beets odeed Timioderus Waterston
Body black or brown. Head and mesosoma almost smooth. Anellus absent.
Mandibles absent, maxilla enlarged and lobate. Petiole fused ventrally. Ovi-
positor long and relatively straight. Body length about 2 mm (Kenya [New
Record: UCRCENT 00478780, Olorgesailie National Monument, R. Cope-
laid | cand litte seas ec gover cebbeeese cocgseecde sions Indosema Husain & Agarwal
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 (Gs,). Base of petiole lack-
inp aneancerioraorsal carina’ (AUStraltan)*.ccm sale eaotanceouogtes oenasutoonttelap hepesr ne 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 (Gs,) meeting apex of petiole
(Figs 64, 86). Base of petiole usually truncate and usually with an anterior
transverse-carina: (Ris. 75) (Old W orld tropics) se «2th: Gece. ac stasncotaencetebe aye 11
Antennal flagellomeres cylindrical. Base of petiole narrowed to condyle........
SAAN NE RR eO TGR RO ORR Rae Ne Ae WEL Orasemorpha Boutéek
Antennal flagellomeres lobate (Fig. 85). Base of petiole truncate... eee
ohare Bayh ae emae 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).
Lower face lateral to clypeus finely reticulate (Fig. 29). Frenum and propo-
deum evenly sculptured, propodeum lacking a distinctly differentiated nar-
row median band of areolate sculpture. Prepectus evenly reticulate or rugose,
dorsal margin without a raised rim and overlapped by ventral margin of mes-
oscutum. 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
qilfvurber atekal sheet int Ghilie. oy herecrse: eke a eee et ee cee eB ee alae
SER ee Hayatosema gen. n. (assectator-group of Heraty, 1994)
Face variously sculptured, but if finely reticulate then propodeum with dis-
tinctly differentiated median channel (Fig. 11), prepectus broadly foveate
with a distinctly raised dorsal margin (Fig. 12) or callus with dense patch of

1OACIuk

13 (12)

14 (12)

1523}

16 (15)

17 (14)

18. (17)

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 145

long hairs. Both sexes with 7—9 funiculars. Ventral valve of ovipositor with
od: laverall tec rnetbiog hay Ge. 28. cele eee cel ee RA, ON ke 12
Antenna with 8—9 funiculars, if 7, then labrum with more than 7 digits,
body black, and propodeal disc evenly sculptured (Jb. seyrigi, Madagascar).
Ocellar-ocular sulcus usually distinct. Propodeal disc evenly sculptured, rare-
ly smooth. Fore wing often without speculum oo... eee eeeeseeseeeeeeeeeeeee 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: Pore wine Stations VALIAD eve steerecderaalte sux. ssducvecssnwenasthoeccos splences puna 14
Face smooth laterally (Fig. 36). Labrum with more than 9 digits (one unde-
scribed specimen with 5 digits). Antenna with 7—8 funiculars (Madagascar)...
ser Ibitya gen. n. (communis-group of Heraty, 2000; 2 species described)
Face evenly sculptured laterally. Labrum with 4 digits (one undescribed Austra-
lian species with 6). Antenna with 8—9 funiculars (Paleotropical excluding Mada-
OASCAL) seman Losbanus tev. stat. (uichancoi-group of Heraty, 1994; 5 species)
Propodealkdisccevierily sculptured ses sceczene acceeeirnscdeost teh ieee arta tc ns! 15
Propodeal disc smooth laterally, usually with a distinct median longitudinal
channel with strong areolate sculpture (Figs 11, 73), rarely weakly reticulate
pave hall] ys TMB) i toc ha Ped Rete Sy ace cea Saar hove Daal doe da tae San ci 17
Face and frenum completely smooth (Figs 57, 58, 62) (Ethiopian)...............
SRR Ct A Br Leiosema gen. n. (glabra-group of Heraty, 1994)...16
Face completely and evenly reticulate; scutellar disc and frenum rugose-retic-
Gate CP 1c BO) Me ISU PAT Morn. Meus hiuwcnstiunls Satta pbacetns caubteiueaited ohak Tel tan bhiakl
pt EG ee dash 5 ty Ba Undescribed and unplaced teneral male (new genus 2)
Antenna encased in exuvia, no ant host information, close to Australose-
ma but very distinct from other species (Australia: QLD: Thornton Peak,
QM:UCRCENT00241816).

Mandibles 3:3 dentate (Fig. 62). Scutellar disc with prominent reticulate
sculpture (Fig. 61). Pronotal neck extended anteriorly and reticulate (Fig.
O'l).Periiale. tia Owe. penta: esSamibatrseet Bokeh eosnth! 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 59-GO) «0.0... .eeeeeseeeeeee Leiosema glabra (Heraty)
Labrum with 6-8 digits. Face smooth (Papua New Guinea [undescribed] and
Wews@aledonial) i Arcus clean win anne waresu Bie dotacn hi dtnibsertedsotialedt Matantas
gen. n. (koghisiana-group of Heraty, 1994), M. koghisiana (Heraty)
Labrum with'4. digits. Pace-smootmor reticulate: o...cfai0dcdin Aoitats 18
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

146 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

seapesclands: presented Malagasy cab Mets scassees entones ocean Manet sateen at deed dadenne acd.
Mes decsbnnattueses Zuparka gen. n. (monomoria-group of Heraty, 2000)...22
193CLS) Face entirelyssmmOoth (Fite 2) 2. cesar covsecccoreetncovscivesrseuescedet A. politurae sp. n.
= Face with at least frons with reticulate sculpture (Fig. 8)... cece cece 20
20 (19) Frenum with median longitudinal groove (Fig. 10). Scape dark metallic
brownvimbochisexds.® 2... ucottiesn concer. vast phhonia ea aerossaaee A. verghetta sp. n.
i Frenum without median longitudinal groove. Scape yellow or dark brown.....21
21 (20) Lateral lobe of mesoscutum and axilla smooth and polished ........ eee
sony eed LES! § tot a be eee Dew Ae hh tee eae eB, A. synempora (Heraty)
= Lateral lobe and axilla coriaceous to rugose, axilla weakly carinate with sur-
face imbricate to reticulate. Scape yellow to dark brown (see Heraty 1994 for
CLISCUISSI OM Ole VAL LAIUES Foece vas cts sewosttave sk cevesabeet A ctivoovetae A, valgius (Walker)
22 (18) Face entirely smooth (Figs 69, 79). Propodeal disc with lateral areas smooth
GEIB: 7D) iastsorce cate tonngatea senso en tobnsude fea viisent data, tues e neha’ Z. fisheri sp. n.
— Frons weakly coriaceous (Fig. 81). Propodeal disc with lateral areas weakly
SGU PIMA a cee ccce see eset dnt re Rea nnn eee baits hoa Z. monomoria (Heraty)

Australosema Heraty & Burks, gen. n.
http://zoobank.org/9A5268EF-48B5-4293-9562-46A528062AAE
Figs 1-14

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; gen-
der feminine.

Discussion. Defined by Heraty (1994) as the Orasema valgius group for two spe-
cies 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 spe-
cies, 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

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 147

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 propo-
deal 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 ovi-
positor 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: P/heidole (Girault 1913[175], Heraty 2000).

Distribution. Four species. Australia including Tasmania.

Australosema politurae Burks, sp. n.
http://zoobank.org/D4F9B53D-B548-4676-832B-68 DD3FD08CC8
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, fu-
niculars 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), 1.1-1.2x 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 cari-
nate posterior to ocellar triangle. Labrum with 4 digits. Palpal formula 3:3. Flagellum

148 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

Figures 1-6. Australosema politurae. Paratype female: | 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; Ilm; mesoscutal lateral
lobe; lpr = lateral propodeal disc; mlm = mesoscutal midlobe; mpr = median propodeal disc; pet = petiole;

not = notauli; scd = scutellar disc.

1.1-1.3x head height, with 7 funiculars; anellus transverse; F2 2.0—2.8x as long as
broad, F2 1.4—1.6x as long as F3.

Mesosoma (Figs 4-5). Mesoscutal midlobe densely reticulate, lateral lobe shallowly
transversely rugose-reticulate and becoming smooth posteriorly. Axilla rugose-reticu-
late, with longitudinal carinae posteriorly that extend across the scutoscutellar suture;

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 149

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, poster-
oventrally 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-re-
ticulate dorsolaterally. Fore wing 2.3—2.4x as long as broad; costal cell with 1 un-
interrupted 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—2.0x as long as broad, only slightly expanded
at uncus; marginal vein about 6x as long as stigmal vein. Hind wing costal cell with a
few ventral setae in apical third.

Metasoma. Petiole 4.0—4.8x as long as broad, 1.6—1.7x as long as metacoxa, finely
longitudinally rugose-reticulate, with some incomplete longitudinal carinae laterally
and ventrally (Fig. 5). Antecostal sulcus of Gs, smooth, very shallow medially; acros-
ternite smooth, uniformly rounded or with a transverse elevation near midlength. Cer-
cus 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 an-
nuli; 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, funicu-
lars 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—6.8x as long as broad, 1.9-2.4x as long as metacoxa, narrowest slightly beyond
midlength, finely reticulate but becoming smooth posteriorly.

Material examined. Holotype: Australias QLD: Mt. Glorious, 27°19'54"S,
152°45'29"E, 7-13.i1.1998, N. Power, Malaise trap [19, ANIC: UCR-
CENT00091415]. 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
[12, UCRC: UCRCENT00312128]. Mt. Glorious, 27°19'54"S, 152°45'29"E,
10-16.1.1997, N. Power, Malaise Trap [19, 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: UCR-
CENT00312060-62]. Mt. Glorious, 27°19'54"S, 152°45'29"E, 3-9.i.1998, T. Hill-
er, MT 2 [12, UCRC: UCRCENT003 12063].

150 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

Similar species. A single specimen (gaster missing, likely female; UCR-
CENT0092152, D0161) from southeast QLD is molecularly similar and is placed
as the sister of A. politurae. It is distinct by having a completely smooth face with a
strongly swollen frons. Two other series (all males from QLD) likely represent another
two species that share similar features to the D0O161 specimen. All specimens have
a 4-digitate labrum, 7 funiculars, a completely smooth face, and a strongly carinate
dorsal occipital margin.

Australosema verghetta Burks, sp. n.
http://zoobank.org/F282 DE42-2C64-4FF4-B131-0752F56D00BA
Figs 7-14

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 me-
dian 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—1.3x as broad as high. Face mostly reticulate, smooth ven-
trally 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 cl-
ypeus 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—1.2x height of head, with 7 funiculars, dark brown; anel-
lus transverse; F2 1.7—2.3x as long as broad, 0.7—1.0x 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-retic-
ulate except peripherally with many subparallel carinae; frenal line posteriorly cari-
nate, frenum rugose-reticulate and with median longitudinal groove; axillular sulcus
foveate, axillula with reticulate sculpture laterally, and with subparallel longitudi-
nal carinae. Mesepisternum anterodorsally smooth and overlapping posterior lower

Biting the bullet: revisionary notes on the Oraseminae of the Old World... hot

Figures 7-12. Australosema verghetta. Paratype female: 7 habitus, lateral 8 head, anterior 9 antennae,

dorsal 10 mesosoma, dorsal; 1 1 propodeum and petiole, dorsal 12 mesosoma, lateral. acs = antecostal sul-

cus; apc = anterior petiolar carina; axlg = axiullar groove; cal = callus; F = flagellomere; frg = frenal groove;

Ipr = lateral propodeal disc; not = notaulix; SSS = scutoscutellar sulcus; tsa = transscutal articulation.

margin of prepectus, posteroventral surface with horizontal shelf that is shorter than
mesepimeron but swollen and smooth anterior to mesocoxa; upper and lower mese-
pimeron 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—2.5x as long as broad; costal cell ventrally setose in apical half; basal cell

152 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

0.15 mm

Figures 13-14. Australosema verghetta. Paratype female: 13 ovipositor, lateral. Paratype male: 14 an-
tenna, ventral. Fu = funicle; dv = dorsal valve; lhs = lateral hypopygial seta; ocv = oblique carina of ventral

valvula; vv = ventral valve; vvlt = ventral valve lateral teeth.

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—2.0x as long as broad, only slightly expanded at uncus;
postmarginal vein about 5.0x as long as stigmal vein. Hind wing costal cell setose in
apical third, bare in basal two thirds.

Metasoma (Figs 11, 13). Petiole 0.9-1.1x as long as metacoxa, 2.4—3.1x 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 Gs,
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 medi-
ally 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 longitudi-
nal carinae. Metatibia dark brown except at apices. Petiole 5.9—7.4x as long as broad,
2.2—2.6x as long as metacoxa, narrow at midlength, reticulate.

Material examined. Holotype: Australia: SA: Mt. Barker, 511m, 35°04'00"S,
138°55'17"E, 22-23.1.1999, J. Heraty, Eucalyptus scrub [12, ANIC: UCR-
CENT00091308]. Paratypes: Australia: SA: Mt. Barker, 511m, 35°04'00"S,
138°55 17 EF; 22-234,1999; 1) Yrletaty Eucalyptus scrub [144 199, ANIC: UCR-
CENT00311977-81, UCRC: UCRCENT0009 1307, UCRCENT00091309, UCR-
CENT00311982-89, UCRC: UCRCENT00312042-59].

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 153

Cymosema Heraty & Burks, gen. n.
http://zoobank.org/8A98705 1-99 CE-482E-ABB1-165B9ACDB8FF
Figs 15-27

Type species. Cymosema waterworthae Burks & Mottern sp. n.

Etymology. Based on the Greek xbua (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 /ndosema. 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 Eu-
charitidae); 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 1.5x 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 swol-
len and finely reticulate, antecostal sulcus not apparent, but a distinct transition to the
smooth posterior region of Gs,; 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).

154 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

Cymosema capelina Burks & Mottern, sp. n.
http://zoobank.org/4E96F380-F80F-417C-B154-D49256EABA22
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, 1.4x as broad as high. Head receding ventral to
toruli (Fig. 18); face dorsally smooth, ventrally coriaceous; supraclypeal area and cl-
ypeus medially smooth and laterally shallowly coriaceous, anteclypeus absent; vertex
not carinate. Mandibles truncate and broader than long, broadly separated. Palpal for-
mula 2:1, palpi long. Pedicel broader than F2; anellus small but easily visible; flagellum
1.4x head height, with 7 funiculars that expand apically and have semi-erect long setae
arising from their bases; F2 1.3x as long as broad, 1.0x as long as F3.

Mesosoma (Figs 18-19). Mesoscutum smooth and glossy, notauli evenly im-
pressed and smooth. Axilla strongly advanced, smooth; scutoscutellar suture finely
foveate; mesoscutellar disc and axillula smooth, without axillular sulcus; frenum dis-
tinguished 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, 2.6x as long as broad; costal cell with 1 complete row of ventral
setae; submarginal vein with several long hairs; basal cell setose dorsally and ven-
trally; speculum absent; cubital and subcubital folds setose; marginal fringe present;
stigmal vein 1.5x as long as broad; dorsal setae of disc much longer than ventral se-
tae; postmarginal vein about 1.5x as long as stigmal vein. Hind wing costal cell with
a few setae in apical third.

Metasoma. Petiole 0.7x as long as metacoxa, 1.8x as long as broad, smooth except
for longitudinal carinae, including a median carina dorsally, narrowing anteriorly to
abruptly expand at the anterior condyle, without anterior transverse carina, ventrally
smooth (Fig. 20). Antecostal sulcus of Gs, crossed by many longitudinal carinae; acros-
ternite smooth. Cercus with all setae subequal in length. Hypopygium bare. Oviposi-
tor 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 [12, ANIC:
UCRCENT00238787], deposited in ANIC.

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 155

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 21-27

Etymology. Named in honor of the collector, Rebeccah Waterworth.

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

156 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

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.

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, 1.6x as broad as high. Head receding ventral to to-
ruli; 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 1.5x

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 157

0.1 mm

Figure 27. Cymosema waterworthae. Holotype female: fore wing. arrow points to end of postmarginal

vein.

head height, with 7 parallel-sided funiculars; anellus minute and hardly visible; F2
1.6x as long as broad, F2 1.1x 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 ante-
riorly and laterally; notauli deep anteriorly but very shallow posteriorly. Axilla strongly ad-
vanced, smooth anteriorly, coriaceous posteriorly; mesoscutellar disc smooth to very shal-
lowly 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 eleva-
tion posterolateral to spiracle (Fig. 24, flange). Metacoxa shallowly reticulate dorsolater-
ally. Fore wing spatulate (Fig. 27), 2.7x as long as broad; costal cell and submarginal vein
bare; basal cell ventrally setose, also with a few dorsal setae near basal fold; speculum ab-
sent; cubital and subcubital folds setose; marginal fringe absent; stigmal vein 2.0x as long
as broad; postmarginal vein about 1.5x as long as stigmal vein. Hind wing costal cell bare.

Metasoma (Figs 25-26). Petiole 1.6x as long as broad, 1.0x as long as metacoxa,
finely longitudinally rugose-reticulate, with a pair of complete lateral longitudinal cari-
nae and some additional incomplete longitudinal carinae anteriorly, narrowing ante-
riorly to abruptly expand anteriorly at the condyle, without anterior transverse carina,
ventrally with a longitudinal groove that expands at apices. Acrosternite with irregular
raised sculpture, swollen and differentiated across indistinct antecostal sulcus from
remaining smooth Gs,. 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 im-

mediately basal to toothed apex.
Male. Unknown.

158 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

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. Wa-
terworth, sweep [12, UCRC: UCRCENT00364784], deposited in UCRC. Gaster

mounted separately on card.

Hayatosema Heraty & Burks, gen. n.
http://zoobank.org/D6C6360D-85FB-458C-9EC8-DFAD4F930030
Figs 28-35

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

Etymology. Named after Mohammad Hayat of Aligarh Muslim University; gen-
der 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 In-
dian 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 consist-
ent differentiating feature from Ovasema 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 hav-
ing 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 spe-
cies, 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 oc-
cipital 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; mes-
oscutal 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 ru-
gose 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

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 159

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).

Hayatosema plicator Burks, sp. n.
http://zoobank.org/OF73EB6C-36D2-4BB8-AC07-45 177 DDE900B
Figs 28-35

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; prono-
tum 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 me-
tallic 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) 1.4x 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 1.3x height of head; anellus
transverse; F2 2.0x as long as broad, F2 1.3x as long as F3.

Mesosoma (Figs 31-34). 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 sul-
cus not indicated, axillula finely rugose-reticulate. Mesepisternum anteriorly with a
smooth area dorsolaterally, laterally finely reticulate and ventrally smooth; not overlap-
ping prepectus; ventrally smooth and forming a rounded horizontal surface anterior
to mesocoxae; upper mesepimeron elevated and smooth; transepimeral sulcus foveate;

160 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

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.

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-reticu-
late; callus bare, smooth over most of surface but coarsely areolate to foveate peripher-
ally, including foveate depression separating metapleuron from hind wing base. Meta-
coxa rugose-reticulate, glossy posteriorly. Fore wing 2.4x 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

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 161

E ™ row bs

lateral teeth

Figures 34-37. 34-35 Hayatosema plicator. Holotype female: 34 propodeum and petiole, dorsal
oblique 35 ovipositor, dorsal 36-37: [bitya communis. Female: 36 head, frontal 37 pupa. hex = hind

coxa; pet = petiole.

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
2.0x as long as broad, only slightly expanded at uncus; postmarginal vein about 3.0x
as long as stigmal vein. Hind wing costal cell with only a few ventral setae apically.

Metasoma (Figs 34-35). Petiole 1.7x as long as broad; 0.9x as long as metacoxa,
with sharp anterior transverse carina, areolate and with a complete dorsal median ca-
rina. Antecostal sulcus of Gs, 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) [12, ROME 2001504], deposited in ROME.

162 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

Ibitya Heraty & Burks, gen. n.
http://zoobank.org/F 1DA0653-6359-4DB9-9110-3703880DDB18
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 Jbitya seyrigi
(Risbec) and /. 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 recog-
nize the two currently described communis-group species as the genus /bitya, 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 val-
vula of the ovipositor with sinuate ridges as opposed to distinct lateral teeth. Dis-
tinguished 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: /bitya 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, il-
lustrated); 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 /. communis, 5 female

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 163

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 Ovasema, including
three swollen tubercles over the petiolar region and pronounced linear swellings along
the posterior margins of Mt, .. 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 mesos-
cutellum as well as a single medial posterior tubercle, and with paired lateral swellings
associated with the linear swellings on each of Mt, ..

New host record. Myrmicinae: P/heidole 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 vetera-
trix [13, 12, 6 pupae, 1 5 pupa: CASC: UCRCENT00417452, UCRC: UCR-
CENT00417446-00417451, 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
[13, CASC: CASENT02212472]. Antsiranana Prov.: botanical garden near the en-
trance to Andasibe National Park, 1025m, 18°55'35"S, 48°24'28"E, 1-5.ix.2001, R.
Harin’ Hala, tropical forest, Malaise trap [24, 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 [13, 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 [19,
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’Haala,
mixed tropical forest, Malaise trap [34 12, CASC: UCRCENT00417457-59, UCR-
CENT00417550].

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 accu-
rate 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 1.5x head height;
7 funiculars; F2 1.9x as long as broad, 1.1x as long as F3; clava as long as preceding

164 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

two flagellomeres. Labrum with 9 long digits, each with a long apical seta, digits ar-
ranged 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 la-
brum with 7 digits, and a long petiole about as long as the hind femur. ‘The male shares
many features with J/. 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 (19, CAS:
UCRCENT00417549).

Distribution. Madagascar.

Ivieosema Heraty & Burks, gen. n.
http://zoobank.org/11180142-ED8D-4F39-AE24-6358C767A9D8
Figs 38-54

Type species. Orasema striatosoma Heraty, 1994: 68-70, figs 98-105.

Etymology. Named after Michael Ivie of Montana State University; gender femi-
nine.

Discussion. Defined by Heraty (1994: 68) as the Orasema striatosoma-group. Ivie-
osema 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 sev-
eral features with [vieosema striatosoma (Heraty), and to a lesser degree with [vieosema
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

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 165

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 lat-
eral margin of gastral tergites 1-4 sinuate; first valvula of ovipositor with several fine
lateral teeth. Male scape lacking ventral pores.

Host association. Myrmicinae: [vieosema fraudulenta reared from Pheidole mega-
cephala Fabricius (Reichensperger 1913).

Distribution. Ethiopian and Malagasy regions.

Ivieosema confluens Burks, sp. n.
http://zoobank.org/DD89 153A-F762-43D6-80DE-B823E00EC907
Figs 38-48

Etymology. Latin noun meaning “a place where rivers meet,” referring to the meso-
somal surface sculpture.

Diagnosis. Recognized from other Orasema and African [vieosema by the fore
wing lacking setae and with a medial longitudinal fold. Distinguished from J. 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 depres-
sion 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 fem-
ora 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, 1.4x as broad as long. Face flat and
smooth; scrobal depression broad and shallow, extending laterally well beyond to-
ruli 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 1.0x head height, 7 funiculars; anellus longer than broad (Fig. 40,
F1); F2 1.7x as long as broad, 1.1x as long as F3.

166 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

Figures 38-43. /vicosema 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.

Mesosoma (Figs 41-44). Mesoscutal midlobe finely reticulate, lateral lobe mostly
coriaceous, notauli very shallow and with vague margins; transscutal articulation rep-
resented only by a vague depression medially (Fig. 42, tsa). Axilla anteriorly reticulate,
posteriorly longitudinally striate; mesoscutellum and axillula finely longitudinally stri-
ate; 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

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 167

Figures 44-48. [vicosema 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 (2"*) valvula, F1/an = anellus; wv = ventral (1") valvula.

transepimeral sulcus. Propodeum finely reticulate, callus bare and smooth to shallowly
reticulate. Metacoxa smooth. Fore wing 2.3x 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

168 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

narrowing apically, not separate from wing margin at base of stigmal vein; stigmal vein
4.5x as long as broad, without an uncus; postmarginal vein absent; cubital and subcu-
bital folds bare; marginal fringe absent. Hind wing costal cell bare.

Metasoma (Fig. 45). Petiole 1.7x as long as broad, 0.9x as long as metacoxa, longitu-
dinally shallowly rugose-reticulate. Gt, with a deep longitudinal median depression with
carinate lateral margins equal to length of petiole; shallower smooth depression present
on Gt,. Antecostal sulcus of Gs, smooth, very shallow medially; acrosternite swollen and
with shallow irregular sculpture; posterior Gs, smooth to finely punctate. Cercus with all
setae subequal in length. Hypopygium with a few small subapical setae laterally. Oviposi-
tor dorsal valve with 9 annuli, including a basal annulus that is broadly separated from the
rest; ventral valve with 5-G 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 api-
ces; F7 shorter than F6 and F8; entire antenna brown. Petiole 1.3x as long as broad,
2.5x as long as metacoxa. Gaster with broad, striate longitudinal depression on Gt.
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: UCR-
CENT00435300], deposited in TMP. Paratype: Madagascar: Toliara Prov.: Tsi-
manampetsotsa 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 [19, CASC:
CASENT2022276].

Ivieosema limula Burks, sp. n.
http://zoobank.org/2985A405-15F7-4588-A216-E1C5C76FC5D6
Figs 49-55

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 I[vieosema by the fore
wing lacking setae and presence of a medial longitudinal fold. Distinguished from /,
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 me-
dian longitudinal depression extending the entire length of the tergite.

Female. Length 3.1 mm. Head and mesosoma almost entirely dark brown, with-
out 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) 1.4x as broad as long. Face flat and smooth; medial scrobal

depression absent, but frons with paired dorsal frontal depressions lateral to scrobal

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 169

Figures 49-54. [vicosema 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.

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 1.2x head
height, 7 funiculars; anellus slightly broader than long, expanding apically (Fig. 51,
F1); F2 2.0x as long as broad, F2 1.1x as long as F3.

170 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

Figure 55. /vieosema limula. Holotype female: fore wing.

Mesosoma (Figs 52-53, 55). Mesoscutum finely reticulate, with a shallow median
depression; notauli finely reticulate in their troughs, with vague margins and not fove-
ate, merging posteriorly as a transverse depression anterior to the transscutal articula-
tion, 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 protrud-
ing, not separated from rest of metanotum but with fine reticulate sculpture antero-
medially. Mesepisternum finely reticulate dorsally and anteriorly, smooth posteroven-
trally, 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) 2.2x
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 1.4x 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 1.7x as long as broad, 0.9x as long as metacoxa, nearly
smooth medially but longitudinally rugose-reticulate laterally, without transverse an-
terior carina and narrowest anteriorly immediately posterior to the condyle. Gt, with
a deep longitudinal median depression extending length of tergite and with smooth
lateral margins. Antecostal sulcus of Gs, smooth; acrosternite smooth to very shallowly
reticulate. Cercus with all setae subequal in length. Hypopygium with a few setae sub-
medially. 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.

Biting the bullet: revisionary notes on the Oraseminae of the Old World... FAY

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 for-
est at the edge of marsh, 5m from road, Malaise trap [12, CAS: CASENT2022276],
deposited in CAS.

Leiosema Heraty & Burks, gen. n.
http://zoobank.org/F7181858-49AC-49 1A-ABD2-6E035D26DBFF
Figs 56-67

Type species. Orasema glabra Heraty, 1994: 72-73, figs 119, 125-128, 130, 132.

Etymology. Combination of the Greek Xetoc (/eios) for smooth and a contraction
of Orasema, referring to the smooth and polished head and frenum; gender feminine.

Discussion. Originally treated within Ovasema as the glabra group by Heraty
(1994: p. 72). Heraty (1994) speculated that some of the characteristic features were
shared with species in the New World coloradensis group. However, there is a greater
similarity with Zuparka (Madagascar), including the entirely smooth face (Figs 57-58,
62), scrobal channels with dorsal foveae (Fig. 57), labrum with 4 digits (Figs 57, 62),
and fore wing with bare basal area and speculum (Fig. 65). However, Zuparka has a
laterally smooth or very weakly sculptured propodeal disc (Figs 73, 82) and the prono-
tum and prepectus are more rigidly associated ventrally (Fig. 82). Australosema synem-
pora 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 Leiosemaz.

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 59-61). Distinguished
from the other Old World genera by having the head smooth and subcircular in
frontal view (Figs 57-62); 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; postmar-
ginal vein 1.5—3.2x 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 Af

rica, Tanzania, Zimbabwe), and Yemen.

172: Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

Leiosema lesiolouna Heraty & Burks, sp. n.
http://zoobank.org/9AA354CA-AF78-4523-ACB0-170734056A65
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 4.7x 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 61-62) 1.2x as broad as high; inter-ocular distance 1.2x eye height;
malar space 0.4x 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 1.4x head height; anellus minute, difficult to distinguish, F2 1.2x as
long as broad, 1.2x 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, dor-
sally 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 articula-
tion; scutellar disc about 1.5x 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 later-
ally, smooth ventrally, broadly rounded anterior to mid coxa. Upper mesepimeron
smooth; lower mesepimeron smooth; transepimeral sulcus distinct. Propleuron con-
vex, smooth. Postpectal carina prominent. Hind coxa 1.8x as long as broad, smooth
with very weak reticulate sculpture; hind femur 5.7x as long as broad, with even cover
of appressed long setae; hind tibia densely setose. Fore wing 2.4x as long as broad;
basal cell and speculum bare, costal cell and wing disc densely setose, wing disc setae

Biting the bullet: revisionary notes on the Oraseminae of the Old World... N75

‘a a

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 pro-
podeum 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; Ibr = labrum; no, = pronotum; scp = scape; vp = ventral pores.

long; marginal fringe relatively long; submarginal vein with several long setae; mar-
ginal vein with minute setae; stigmal vein more than 2.5x as long as broad, slightly
angled, slightly narrowed basally and capitate apically (Fig. 66); uncus absent; stigma

174 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

with 3 sensilla in a straight line; postmarginal vein 1.9x as long as stigmal vein. Hind
wing costal cell with a broad bare area.

Metasoma. Petiole cylindrical, linear in profile, 4.6x as long as broad, 1.7x as long
as hind coxa, reticulate, anterior carina strong, lateral margin rounded, ventral sul-
cus 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 [13, UCRC:
UCRCENT00408466].

Leiosema glabra (Heraty), comb. n.

Figs 56-60, 63, 65-67

Orasema glabra Heraty, 1994: 72-73. Holotype, South Africa, Transvaal [2, CNCI].
Mistakenly figured in Heraty (1994) with 6 funicular segments (cf. Fig. 126) in-
stead of 7.

Diagnosis. Both sexes with mandibles 3:2 dentate, mesoscutal midlobe and scutellar disc
weakly reticulate, becoming smooth posteriorly (Figs 59-60). The pronotal neck is not
extended, and is reticulate only along the anterior margin (Fig. 60). Male petiole 2.6-
2.9x as long as broad, female petiole 1.5—2.1x 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 geo-
graphic 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 pat-
terns 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

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 175

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 acros-
ternite same as mesosoma; gaster dark metallic.

Head (Figs 61-62) 1.1-1.2x as broad as high; inter-ocular distance 1.4—1.5x eye
height; malar space 0.3—0.7x 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. Flagel-
lum 1.0—1.2x head height. F2 1.2—1.3x as long as broad, 1.2—1.3x as long as F3; clava
subconical, longer than preceding two funiculars and weakly segmented.

Mesosoma (Figs 61, 64). Mesosoma 1.5—1.7x as long as high. Mesoscutal lateral
lobe smooth and shining dorsally, weak rugulose laterally. Scutellar disc 1.5—1.7x 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—2.0x as long as broad, smooth to
coriaceous; hind femur 4.2—5.4x as long as broad, with even cover of elongate, semi-
erect setae. Fore wing 2.1—2.3x as long as broad; marginal vein with long setae; stigmal
vein 1.5—2.6x 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
1.1-3.2x as long as stigmal vein. Hind wing costal cell bare.

Metasoma. Petiole 2.6—2.9x as long as broad, 1.2—1.4x 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 \.0—1.1x as broad as high; eyes separated by 1.3—1.5x eye height; malar space
0.4—0.6x eye height. Flagellum length 0.8—1.0x head height (0.9-1.0 in type mate-
rial); flagellum with 7 funiculars, anellus disc-shaped, F2 1.1—1.4x as long as broad,
1.1—1.2x as long as F3.

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

Metasoma. Petiole 1.5—2.1x as long as broad, 0.9-1.3x 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 [12, NMPC: UCRCENT00416471]. Transvaal: Blyderiviers-
poort, 1200m, 24°53'00"S, 30°45'00"E, iii.1981, G. L. Prinsloo [14, PPRI: UCR-

176 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

CENT00298795]. Waterberg nr Nylstroom [Modimolle], 1200m, 24°48'00"S,
28°25'51"E, i.1980, G. L. Prinsloo [1¢, PPRI: UCRCENT00298796]. Strij-
dom Tunnel, 730 m, 24°27'47"S, 30°36'31"E, 24-25.xi.1991, G.L. Prinsloo [19,
NMPC: UCRCENT00416472]. Tanzania: Tanga: West Usambara, Megwashi
env., 1460m, 4°45'47"S, 38°28'30"E, 31.i-1.ii.2015, P. Jansta & J. Straka [34 19
2?, UCRC: UCRCENT00479260-65]. Zimbabwe: Harare: Chishawasha, 1416m,
17°45'46"S, 31°12'31"E, A. Watsham [22, NMPC: UCRCENT00416454, UCR-
CENT00416456]. Zimbabwe: Mashondland: Mazowe, 125m, 17°30'32"S,
30°58'19"E, i.1975, A. Watsham [62, NMPC: UCRCENT00416459-61, UCR-
CENT00416466-67, UCRCENT00416519]. Makumbi Miss., 17°31'12"S,
31°15'22"E, A. Watsham [14, NMPC: UCRCENT00416462].

Orasema near glabra: Gambia: Banjul: Kotu Stream, 3m, 13°27'41"N,
16°42'13"W, 14.xi.1981, K-J. Hedqvist [54 42, BMNH: UCRCENT00309860-68].
Kotu Stream, 4m, 13°27'42"N, 16°42'13"W, 18.xi.1981, K-J. Hedqvist [192, BMNH:
UCRCENT00309871]. North Bank Division: Upper Baddibu, Farafenni, 13m,
13°34'03"N, 15°44'56"W, 1-2.xi.1977, Michael S6derlund, Meadow (mostly resorts)
with baobab [14, BMNH: UCRCENT00309873]. Western Division: Kombo St
Mary, Bakau, 10m, 13°28'47"N, 16°40'14"W, 28-29.x.1977, Michael Séderlund,
shore meadow [14, BMNH: UCRCENT00309872]. Kombo St Mary, Sara Job
Kunda, 25m, 13°26'40"N, 16°42'46"E, 29.x.1977, Michael Sdderlund, grassland
with palms [146 BMNH: UCRCENT00309859].

Losbanus Ishii, rev. stat.

Losbanus Ishii, 1932: 210. Type species: Losbanus uichancoi Ishii 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, Boucéek, 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. Boucek
(1988) incorrectly used this as a senior synonym of Gollumiella. Heraty (1992) trans-
ferred 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 characteris-

Biting the bullet: revisionary notes on the Oraseminae of the Old World... Ks

tics 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 Ovasema and other Oraseminae by having 8-9
funiculars versus 7, a weak basal petiolar flange, usually a distinct ocellar-ocular sul-
cus, 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 depres-
sion 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 un-
dersurface of leaves of Celtis philippinensis Blanco (Cannabaceae) and Leucaena glauca
Benth. (Fabaceae).

Host association. Unknown.

Distribution. Five species described from southeast Asia (Heraty, 1994). Paleo-
tropical (Australia: Queensland (undescribed); Republic of Congo (undescribed), Pap-

ua New Guinea, Philippines, Taiwan).

Matantas Heraty & Burks, gen. n.
http://zoobank.org/6556925F-B592-4324-B97C-110902691CF2

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

178 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

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 oc-
cipital margin abrupt and rounded, at most with a weak dorsal carina; funicle 7-seg-
mented; labrum with 6-8 digits; mesonotum appearing bare, at most with minute se-
tae; 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 val-
vula of ovipositor with 3 or 4 lateral teeth. Male scape lacking ventral pores.

Host association. Unknown.

Distribution. One species described. New Caledonia (another undescribed spe-

cies from Vanuatu, BPBM: UCRCENT00422296).

Zuparka Heraty & Burks, gen. n.
http://zoobank.org/17C35918-BF49-424D-BFE0-F3541C898EBB
Figs 68-86

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 recent-
ly emerged teneral female. Another distinctive species is recognized herein, Z. fisheri
sp. n. [he two species are morphologically distinct from each other, and the divergence
in their ribosomal and mitochondrial data is generally what is expected between dif-
ferent genera in other Oraseminae. However, morphological features support recogni-
tion 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).

Biting the bullet: revisionary notes on the Oraseminae of the Old World... ee

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 peti-
olar 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 chan-
nels 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; prep-
ectus 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.

Zuparka fisheri Heacox & Dominguez, sp. n.
http://zoobank.org/A2294EE0-09AF-4A98-B9AA-B8D207366A13
Figs 68-80

Etymology. Named after Brian Fisher of the California Academy of Sciences for his
tremendous collecting efforts that have contributed to the understanding of the fauna
of Madagascar.

Diagnosis. Face entirely smooth. Scutellum vaulted anterior to frenum and medi-
ally depressed. Propodeal disc laterally smooth. ‘This species resembles Zuparka mono-
moria, 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 mar-
gins 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. Flagel-
lum 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.

180 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

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; Ipr; lateral propodeal disc; mts = metascutellum; sct = mesoscu-

tellum; tsa = transscutal articulation.

Head (Figs 69, 77-79) 2.2—2.6x 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—1.4x eye height; malar space 0.5—0.7x eye height; distance
from anteclypeus to ventral margin of toruli 0.3—0.4x 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.

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 181

Occiput transversely aciculate, dorsal margin broadly rounded, occipital carina absent;
temples broad. Antenna 1 1-segmented (Fig. 77); scape narrow and cylindrical, slightly
broadening medially and reaching median ocellus, at least 7x as long as broad; pedicel
1.3—2.0x as long as broad, apex broader than anellus; flagellum 0.8—0.9x head height;
funicle 7-segmented; flagellar segments beyond anellus densely setose with numerous
longitudinal sensilla; F2 1.2—1.6x as long as broad, following segments becoming more
transverse; clava subconical, about as long as preceding three segments.

Mesosoma (Figs 70-74). Midlobe of mesoscutum and mesoscutellum finely re-
ticulate; notauli foveate, reaching transscutal articulation (TSA) and broadly sep-
arated 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 fo-
veate scutoscutellar sulcus that is continuous medially; axillular sulcus and frenal
line medially foveate with a weak carina along ventral margin of sulcus, continu-
ing 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—2.9x
as long as mesosoma, 2.4—2.6x 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—0.2x 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—1.9x as long as hind coxa;
finely reticulate; ventral sulcus absent. Gt, as long as or slightly shorter than femur,
smooth and bare; Mt, densely pilose anterior to cercus, cercus with 5 long setae; Gs,
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. Gs, apically strongly convex with sparse ventral hairs; Ms, 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 [12, CASC: UCRCENT00417485].
Paratypes: Madagascar: Antananarivo Prov.: 3 km 41 degrees NE Andranomay,

182 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

0.05 mm

0.1 mm

701002 mm

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 peti-
olar carina; dv = dorsal (2nd) valvula; F = flagellomere; Fl/an = anellus; lhp = petiolar lateral hair patch;

vp = ventral pore of scape; pmv = postmarginal vein; stv = stignal vein; vv = ventral (1st) valvula.

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,

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 183

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.

960m, 12°30'52"S, 49°10'53"E, 12.ii-4.iii.2001, 4-19.iii.2001, R. Harin’ Hala,
Malaise [53 19, CASC: UCRCENT00175169, UCRCENT00278241-45]. Parc
National Montagne dAmbres Wl 25ms 12231137, 49°10'45" Ey 295-1 iti 2001,
5-21.iv.2001, 14-30.v.2001, 30.v-6.vi.2001, R. Harin’Hala, Malaise trap [435 19,
CASC: CASENT02014649, CASENT02015483, CASENT02015488, UCR-
CENT00417343, 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 Rano-
mafana, 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¢ 19, CASC:
UCRCENT00417551, UCRCENT00417553]. Parc National Ranomafana, ra-
dio 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 [24 49,
CASC: UCRCENT00417386—487, UCRCENT00417388, UCRCENT00417489,
UCRCENT00417490, UCRCENT00417492]. Parc National Ranomafana, Vohi-
parara at broken bridge, PT Oms 219133 4°5,047222 1B 21-2 8.542002,-4-12)
ii.2002, M. Irwin and R. Harin’Hala, high altitude rain forest, Malaise trap [13
22, 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 fo-
rest, Malaise Trap [24 22, CASC: UCRCENT00417486, UCRCENT00417497,
UCRCENT00417513, UCRCENT00417538].

Additional material examined. Madagascar: Antsiranana Prov.: Parc National
Montagnad’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 [32 14, 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 [12, CASC: UCRCENT00417338].

184 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

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 rainfo-
rest [12, UCRC: UCRCENT00417370]. Additional material examined: Mada-
gascar: 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 [19,
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, mon-
tane 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.Ha-
rin’ Hala, Uapaca forest, Malaise trap [19, CASC: UCRCENT00352446]. Antsirana-
na 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 [32, CASC: CASENT02015482,
CASENT02015484, CASENT02015487]. Diego Suarez; PN Montange d’Am-
bra, 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 [12, 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 openarea [1 2, 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 [44 219, CASC: CASENT02174558-60,
CASENT02174556, CASENT02174592, CASENT02174594-97, UCRCENT00
417360; UCRC: UCRCENT00417481, UCRCENT00417495-96, UCRCENT0
0417503, UCRCENT00417514, UCRCENT00417531—33, UCRCENT004175
35, 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
[12, CASC: CASENT02174591]. Parc National Ranomafana, radio tower at forest
edge, 1130m, 21°15'03"S, 47°24'26"E, 14-21.1.2002, 28.i-4.i1.2002, 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 [142 34, CASC: UCRCENT00417467, UCRCENT00417371-77,

Biting the bullet: revisionary notes on the Oraseminae of the Old World... 185

G7 h ih yas
, a ua Ne ~
ee oe
ys a a
i ag
vd 02mm al
. ¥ 2

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: VCRCENT00238798
(couplet 10) 86 unknown male Oraseminae, Australia: UCRCENT00241816 (couplet 15).

UCRCENT00417379-85, UCRCENT00417387, UCRCENT00417530]. Parc Na-
tional Ranomafana, Vohiparara at broken bridge, 1110m, 21°13'34"S, 47°22'11"E,
6-15.xii.2001, 14-21.1.2002, 21-28.i1.2002, 4-12.ii.2002, 19-26.ii.2002, 6-15.v.2002,
M. Irwin and R. Harin’Hala, high altitude rain forest, Malaise trap [53 99 32, CAS:
CASENT02174562, CASENT02174583, CASENT02174587-89; UCRC: UC
RCENT00417508, UCRCENT00417511-12, UCRCENT00417525-29, UC

186 Roger A. Burks et al. / Journal of Hymenoptera Research 55: 139-188 (2017)

RCENT00417534, UCRCENT00417465-66, UCRCENT00417468]. Parc Na-
tional 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 [24 192, CASC:
UCRCENT00417443, UCRCENT00417501-02]. Parc National Ranomafanae,
radio tower at forest edge, 1130m, 21°15'03"S, 47°24'26"E, 21-28.i1.2002, 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 [63 72
12, CASC: CASENT02174580-81, UCRCENT00417441, UCRCENT00417488,
UCRCENT00417491; UCRCENT00417493-94, UCRCENT00417498,
UCRCENT00417504—07, UCRCENT00417536, UCRCENT00417554]. Rano-
mafana 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 [12, 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
[12 12, UCRC: UCRCENT00412588-89]. Toamasina Prov.: bot. garden near en-
trance to Andasibe National Park, 1025m, 18°55'35"S, 48°24'28"E, 16-23.xi.2001, R.
Harin’Hala, tropical forest, Malaise Trap [14, CASC: UCRCENT00417494].

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 Tor-
réns 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 com-
mercial products in this publication is solely for the purpose of providing specific in-
formation and does not imply recommendation or endorsement by the USDA; USDA
is an equal opportunity provider and employer.

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