ROYAL ONTARIO ML ^

LIFE SCIENCES
CONTRIBUTIONS

157

,^

>^

^,

I Classification AND
Evolution OF THE ,4

OrASEMINAE IN THE \

^ Old World, it.

Including Revisions [
OF Two Closely -,yJ
Related Genera OF 3

EUCHARITINAE . .j'

(HYMENOPTERA : -^

Eucharitidae) '^

John Michael Heraty -.1=^

' .'-«■'«,

ROM

Digitized by the Internet Archive

in 2011 with funding from

Royal Ontario Museum

http://www.archive.org/details/classificationevOOhera

iA ^> T *-

Classification and Evolution

OF THE ORASEMINAE

IN THE Old World

Habitus of Orasema hoiiceki, 6 .

ROYAL ONTARIO MUSEUM

LIFE SCIENCES
CONTRIBUTIONS

157

Classification and Evolution of the

Oraseminae in the Old World, Including

Revisions of Two Closely Related Genera

of eucharitinae

(HYMENOPTERA: EUCHARITIDAE)

John Michael Heraty

e- 3'^v;

©IW4 Royal Ontario Museum

All riiihts reserved. No pari ot this publication may be reproduced, stored in a retrieval
system or data base, or transmitted, in any form or by any means, electronic, mechani-
cal, photocopying, or otherwise, without the prior written consent of the publisher.

First published in 1994 by the Royal Ontario Museum. !()() Queens Park. Toronto.
Ontario M.^S 2C6.

Publication date: 1 November 1994
ISBN 0-88854-4 12-X
ISSN 0384-8159

Canadian Cataloguing in Publication Data

Heraty, John Michael

Classification and evolution of the Oraseminae
in the old world, including revisions of two closely
related genera of Eucharitinae (Hymenoptera:
Eucharitidae)

(Life sciences contributions; no. 157)
Includes bibliographical references.
ISBN 0-88854-4 12-X

1 . Eucharitidae - Classification. 2. Insects -
Classification. I. Royal Ontario Museum. II. Title.
III. Series.

QL568.E75H4 1994 595.79 C94-93 1709-8

ROYAL ONTARIO MUSEUM PUBLICATIONS IN LIFE SCIENCES
The Royal Ontario Museum publishes books on a variety of subjects in the life sci-
ences, including Life Sciences Contributions, a numbered series of original scientific
publications. All manuscripts considered for publication are subject to the scrutiny
and editorial policies of the Life Sciences Editorial Board, and to independent
refereeing by two or more persons, other than Museum staff, who are authorities in
the particular field involved.

LIFE SCIENCES EDITORIAL BOARD

Senior Editor: K. A. Coates

Editor: D. H. Collins

Editor: R. D. James

External Editor: C. S. Churcher

Manuscript Editor: K. A. Coates
Production Editor: Andrea Gallagher Ellis

Dr. John Michael Heraty is currently a postdoctoral fellow at the Smithsonian
Institution. Washington. D.C.. and a research associate of the Royal Ontario Museum.
Toronto. Correspondence should be addressed to the author 7o Department of
Entomology, University of California, Riverside, California, U.S.A. 92521

PS

— • The Royal Ontario Museum is an agency of the Ontario Ministry of Culture, Tourism

^ ^ and Recreation.

Printed and bound in Canada

Contents

Abstract 1
Introduction 2
Materials and Methods 5
Descriptive Format 5
Terms 6

Measurements and Abbreviations 6
Setation 6
Sculpture 6
Colour 7
Head 7
Antennae 7
Mesosoma 7
Wings 8
Metasoma 8
Sexual Dimorphism 8
Biogeographic Regions 8
Phylogenetic Analysis 9
Analytical Methods 10
Character Analysis 1 1
Parsimony Analysis 19
Discussion of Phylogeny 19
Eucharitidae 20
Oraseminae 20
Eucharitinae 21
Conclusions on Phylogeny 23
Biology 23

Biology and Immature Stages of Orasemorpha 24
Biology and Immature Stages of Orasema 24
Biology and Immature Stages oi Neoloshanus 26
Biology and Immature Stages of Eucharitini 27
Host-Ant Relationships of Eucharitidae 28
Conclusions on Biology and Immature Stages 30
Revision of Oraseminae and Psilocharitini (Eucharitinae) 3 1

Key to Genera of Oraseminae and Psilocharitini 3 1
Revision of the Old World Oraseminae 32
Timioderus Watcrston 32

Key to Species of Timioderus 33
T. peridentatus sp. nov. 34
T. rcfrini^cns Waterston 35
T. acuminatus sp. nov. 36

T. coronula sp. nov. 37
T. ramosus sp. nov. 37

Indosema Husain and Agarwal 38

/. indica Husain and Agarwal 39

Orasemorpha Boucek 40

Key to Species of Orasemorpha 42
O. xeniades (Walker) 43
O. myrmicae (Girault) 44
O. tridentata (Girault) 45
O. goethei (Girault) 46
O. pyttalus (Walker) 47
O. sparsepilosa sp. nov. 48
O. varidentata (Girault) 49
O. didentata (Girault) 50
O. erihotes (Walker) 5 1

Orasema Cameron 54

Key to Old World Species of Orasema 57
O. communis Risbec 59
O. seyrigi Risbec 61
O. iiichancoi (Ishii) 62
O. ishii sp. nov. 64
O. houceki sp. nov. 65
O. promecea sp. nov. 66
O. rugulosa sp. nov. 67
O. striatosoma sp. nov. 68
O . fraudulenta (Reichensperger) 70
O. koghisiana sp. nov. 71
O. glabra sp. nov. 72
O. assectator Kerrich 74
O. nigra sp. nov. 75
O. initiator Kerrich 76
O. synempora sp. nov. 77
O. valgius (Walker) 79
Revision of Psilocharitini (Eucharitinae) 8 1

Psilocharis gen. nov. 81

Key to Species of Psilocharis 83
P. aenigma sp. nov. 84
P. monilicera sp. nov. 85
P. pentella sp. nov. 85
P. pacifica sp. nov. 86
P. dahmsi sp. nov. 87

VI

p. joanneae sp.nov. 88

P. theocles (Walker) 89

P. afra sp. no v. 90

P. hypena sp. nov. 91
Neoloshanus gen. nov. 93

Key to Species of Neoloshanus 96

A^. gemma (Girault) 98

N. wusheanus sp. nov. 99

N. apoanus sp. nov. 100

N. storeyi sp. nov. 101

A^. watanabei sp. nov. 102

N. purpureoventris (Cameron) 103

N. townesi sp. now. 104

A^. laeviceps (Gahan) 105

N. taiwanensis sp. nov. 106

A^. pilosus sp. nov. 107

N. nepalensis sp. nov. 108

N.palgravei (Gimuh) 109

N. gressitti (Watanabe) 1 14

N. kokureanus sp. nov. 114

N. a impetus sp. nov. 115

N. violaceus sp. nov. 1 16
Summary 118
Acknowledgements 120
Literature Cited 121
Figures 127

Appendix 1 : Character Matrix Used for Phylogenetic Analyses 173
Appendix 2: New Synonymies 174
Appendix 3: New Combinations 174
Appendix 4: Lectotypes Designated 174
Appendix 5: New Taxa Described 174

Vll

Classification and Evolution of the Oraseminae in the Old World,
Including Revisions of Two Closely Related Genera of Eucharitinae

(Hymenoptera: Eucharitidae)

Abstract

The Oraseminae and Eucharitinae are hypothesized to form a monophyletic group in
Eucharitidae (Hymenoptera: Chalcidoidea). These 2 subfamihes are redefined to incorpo-
rate new information from species found in the Old World. Phylogenetic methods were
used to analyse morphological characters of adults and immature stages, and behavioural
characters. The Oraseminae was determined to be a monophyletic group; however, charac-
ter states used previously to define the group were found to be homoplastic and resulted in
misclassification of some species. New characters derived from the ovipositor and imma-
ture stages are used to define the subfamily. Monophyly of the Eucharitinae is supported by
adult and larval features. The limits of Eucharitinae are expanded to include 2 new genera
with an independent prepectus that were formerly regarded as Oraseminae, and the subfam-
ily is subdivided into 2 new tribes based on the presence of an independent prepectus
(Psilocharitini) or a fused prepectus (Eucharitini).

Keys to genera and species, descriptions, and biogeographic information are provided
for the 4 genera of Oraseminae and the 2 genera placed in the Psilocharitini. Descriptions of
immature stages and behaviour are provided. Four genera (with type species in parentheses)
included in the Oraseminae are ludosema Husain and Agarwal, 1 species (/. indica Husain
and Agarwal); Orasema Cameron, 16 species in Old World {O. stramineipes Cameron);
Orasemorpha Boucek, 9 species {Eucharomorpha viridis Girault); and Timioderus
Waterston. 5 species (T. refringens Waterston). Two new genera of Eucharitinae are
described: Neoloshanus, 16 species {Orasema palgravei Girault). and Psilocharis, 9 species
(Orasema theocles Walker). Fifty-six species of Eucharitidae are treated, of which 33 are
described as new. Timioderus includes 4 new species (type locality in parentheses): T.
aciiminatus (Cape Prov., South Africa), T. coromda (Grahamstown, South Africa), T. peri-
dentatus (Mossel Bay, South Africa), and T. ramosus (Aliwal North, South Africa). One
new species of Orasemorpha is described. O. sparsepilosa (Moree, Australia). Orasema
includes 9 new species: O. houceki (Kokoda. Papua New Guinea). O. glabra (Klaserie,
South Africa), O. ishii (Tungpu, Taiwan), O. koghisiana (Koghis Mtns, New Caledonia), O.
nigra (Royal Natal N. P., South Africa), O. promecea (Baiycr Riv., Papua New Guinea), O.
rugulosa (New Britain, Papua New Guinea), O. synempora (Mt Webb, Australia). O. stri-
atosoma (Kampala, Uganda). Psilocharis includes 8 new species: P. aenigma
(Ambohitsitondrona, Madagascar), P. afra (Kalinzu Forest, Uganda), P. dahmsi (Swart
Valley, Papua New Guinea), P. hypena (Tenompok, Sarawak, Malaysia). P. Joanneae (Mt
Webb N. P., Australia). P. monilicera (Grahamstown, South Africa), P. pacifica (Mt
Dalaikoro. Fiji), and P. pentella (Aceh. Sumatra). Neoloshanus includes 1 1 new species: N.
anapetus (Mayoyao. Philippines), N. apoanus (Mt Apo, Philippines), N. kokureanus
(Kokurc, Solomon Islands), N. nepalensis (Godavari, Nepal), N. pilosns (Fyan. Vietnam).
A', storeyi (Nondugl, Papua New Guinea). A', laiwanensis (Wufeng, Taiwan), N. lownesi

(Kassam Pass. Papua New Guinea), A', violaceus (Baiyer Riv.. Papua New Guinea), N.
watanahci (Mayoyao. Philippines), and N. wushcanus (Wushc. Taiwan).

The following new conibinalions are proposed: Oiascinorpha myrmicae (Girault),
Orasema fraudulenta (Reichensperger); Psilocharis theocles (Walker), Neoloshanus
ycninui (Girault), Neoloshanus piirpureoveniris (Walker), Neoloshanus laeviceps
(Gahan), Neoloshanus pali^ravei (Girault), Neoloshanus gressitti (Watanabe), and
Stilhula ranomafanae (Risbec). Lectotypes are designated for 9 species.

The following new synonymies are proposed: Parasemora Gemignani = Orasema
Cameron; Orasema viridicyanea Risbec = Timioderus refringens Waterston;
Euiharomorplia wheeler! Brues = Orasemorpha tridentata (Girault): Eucharomorpha
duhia Girault, E. fuscipes Girault, E. partiglahra Girault, and E. viridis Girault =
Orasemorpha erihotes (Walker); Psilogaster nishidai Ishii and Nagasawa, Loshanus
petersoni Hedqvist, and Orasema indica Snehalatha and Narendran = Neoloshanus
palgravei (Girault).

A parsimony analysis of 36 taxa using 62 characters resulted in 20 trees of 183
steps. Only 16 of 45 genera of Eucharitinae were used in the analysis of relationships
among genera with a free prepectus, but these represent the major lineages that have
some known biological information. The phylogeny of Eucharitidae is correlated with
ant hosts. Myrmicinae are postulated as the ancestral host of Oraseminae. and
Ponerinae as the ancestral host for Eucharitinae. Correlations between ant-host sub-
families with the phylogeny for Eucharitidae suggest adaptation to new hosts through
colonization rather than coevolution. Phylogenetic relationships among species of
Oraseminae and basal Eucharitinae are used to analyse the historic biogeography.

Introduction

The Eucharitidae (Hymenoptera: Chalcidoidea), as based
on the subfamilies Oraseminae and Eucharitinae (sensu
Graham. 1969; Burks, 1979; Heraty, 1985), is a mono-
phyletic group united by characters of adults, including a
digitate labrum with digits arranged in the same plane
(Darling, 1983a, 1988a), falcate or sickle-shaped
mandibles (except in Timioderus Waterston), obliterated
malar sulcus, and a pronotum that is ventral to the meso-
scutum and not visible in dorsal view (Heraty, 1985,
1989). Monophyly of Eucharitidae is also supported by
characters of first-instar larvae, which include reduction
in number of dorsal setae and loss of spiracles (Heraty
and Darling. 1984).

The Oraseminae and Eucharitinae are parasites of ants
(Heraty and Darling, 1984; Darling, 1988a; Heraty,
1985). Three additional subfamilies, Akapalinae,
Echthrodapinae, and Philomidinae, were recognized and
included in the Eucharitidae by Boucek (1978, 1988).
None of these subfamilies is known to be a parasite of
ants, and the close relationships between them and the
other eucharitine subfamilies have been questioned
(Darling, 1988a, 1992; Heraty, 1990). Before these subfa-
milial relationships can be assessed, it is necessary to
understand the bounds and relationships of the
Oraseminae and Eucharitinae (Eucharitidae s.s.). In treat-

ing the Oraseminae of the Old World tropics, it became
apparent that existing definitions of both subfamilies were
not adequate and needed to be reevaluated.

The Eucharitidae is by far the largest and most diverse
group of hymenopteran parasitoids that attack eusocial
insects. Forty-seven genera and 394 species of
Eucharitidae are known from every zoogeographical
region of the world. Eucharitids are notably absent from
New Zealand and a few of the smaller oceanic islands,
and are relatively poorly represented throughout the
Palaearctic region. The family is most abundant in num-
bers both of individuals and of species in tropical regions
(Heraty, 1985). No genera are shared between the
Nearctic and Palaearctic regions and only a few New
World genera {Orasema Cameron, Kapala Fabricius, and
Pseudochalcura Ashmead) have species in the Old World
tropics also (Heraty, 1985, 1986).

Species of Eucharitidae s.s. attack a wide range of ant
hosts, and recent taxonomic changes in the group (cf.
Boucek, 1988) indicate a general congruence between ant
hosts and eucharitid genera. Adult females deposit their
eggs away from the host in plant tissue using a wide vari-
ety of methods (Clausen, 1940a. 1940b. 1940c. 1941).
The first-instar larva is the active stage that is responsible
for gaining access to the ant host.

Brief taxonomic reviews of the Eucharitidae are pre-
sented in Heraty (1985) and Boucek (1988). Walker
(1862) first recognized the family and proposed the name
Eucharidae. It was divided into 2 subfamilies when Kirby
(1886) erected a second subfamily Eucharissinae for what
are now recognized as derived members of the
Eucharitinae (Boucek, 1988). Boucek (1978) suggested
inclusion of the Philomidinae within the Eucharitidae,
and Burks (1979) erected the subfamily Oraseminae to
include the single genus Orasema Cameron. Oraseminae
was later defined by Heraty (1985) and Boucek (1988).
Boucek (1988) recognized 5 subfamilies of Eucharitidae:
Philomidinae, Echthrodapinae. Akapalinae, Oraseminae,
and Eucharitinae. The Echthrodapinae and Akapalinae
were described as new subfamilies to encompass 2 genera
(Boucek, 1988). The Echthrodapinae and Philomidinae
are parasitoids of stem- or ground-nesting bees, respec-
tively (Michener, 1969; Boucek, 1988; Darling, 1992).
Based on morphology of immature stages, Philomidinae
have been placed as sister group to the Perilampidae -i-
Eucharitidae, although this relationship is not supported
by adult characters (Darling 1988a, 1992). The biology of
Akapalinae is unknown. Only Oraseminae and
Eucharitinae are known to be parasites of ants.

Oraseminae has been defined by several symple-
siomorphic character states which include a freely articu-
lating prepectus, antenna with an annular basal flagellom-
ere or ring segment (Boucek, 1988; Heraty, 1985), and
the first phragma ventrally inflected behind the pronotum
(Heraty, 1989). Morphology of the immature stages is
known in detail only for members of Orasema Cameron,
which show alternate states to the Eucharitinae. These
states include separation of tergites I and II, lack of a
complete tergopleural line, and presence of cranial setae
(Heraty and Darling, 1984; Johnson et al., 1986). The
Oraseminae was regarded as a monophyletic group based
solely on possession of a subapically expanded and
strongly ridged ovipositor. The absence of an expanded
ovipositor from some of the species included by Boucek
(1988) within Orasema prompted this study of Old World
Eucharitidae.

According to Steyskal (1980), the correct subfamily
name should be Orascmatinae and all species of Orasema
should have neuter endings to conform to the Latin stem
of sema. Cameron often created euphonic words for gen-
era without regard to their classical origin (Boucek. pers.
comm.. 1991 ). Translation of the Greek ora- (sign, token,
or mark) and the Latin -sema (edge or region) does not
refer to any peculiar attribute of this genus, and Dalla
Torre (1898) referred to the name Orasema as "Eniomol.
ohscurar supporting the lack of any intentional meaning
by Cameron. Of 7 generic names of Eucharitidae pro-
posed by Cameron, none can be translated to mean any
feature of the respective genera. Thus, Orasema should

be treated as an arbitrary combination of letters. The type
species, Orasema stramineipes Cameron, does not have a
gender-specific ending and does not imply Cameron's
intentions. If no gender was attributed or implied by the
author, and if the ending is clearly a natural Latin femi-
nine (as in -ma), then that gender is appropriate to the
ending (Article 30d; ICZN, 1985). By treating Orasema
as being of feminine gender, no changes are required for
previously described species, and the usage of
Oraseminae is maintained.

Heraty (1985) recognized 4 genera in Oraseminae:
Orasema Cameron, Loshanus Ishii. Psilogastrellus
Ghesquiere (in part), and Parasemora Gemignani.
Several species belonging to distantly related genera of
Eucharitidae were incorrectly placed in the genus
Psilogastrellus (= Psilogaster Blanchard. 1840). which is
based on the type species Psilogaster cupreiis Blanchard
(Boucek, 1988). Psilogastrellus fraudulentus
Reichensperger and the genera Parasemora and Loshanus
are here placed in Orasema, and species previously
placed in Loshanus are herein transferred to 3 different
genera. The Oraseminae as recognized here and by
Bou&k (1988) consists of 4 genera: Indosema Husain and
Agarwal (I species), Orasema (16 Old World species,
80-100 New World species), Orasemorpha (9 species),
and Timioderus (5 species).

Orasema is the most speciose genus of Oraseminae
and is the only circumtropical genus. Forty-two species
are described, of which 37 occur in the New World
(Table 1). Orasema was revised by Gahan (1940), who
based his study on fewer than 300 specimens. 1 can segre-
gate the New World species into at least 6 different
species groups, and the Old World species described
herein cannot be directly placed within any of these
groups. However, the range in morphology among Old
World species is equivalent to that among the most dis-
similar species groups of the New World. Considerable
biological information is available for species of Orasema
from around the world and indicates very conservative
morphology and habits of immature stages. Putative
synapomorphies for Oraseminae based on the behaviour
of largely New World Orasema include internal para-
sitism during the first instar, use of a thysanopteran or
homopteran intermediate host to gain access to the ant
host, and parasitism of Myrmicinae (Formicidae).

Monophyly of the subfamily Eucharitinae was sup-
ported previously by the anterior fusion of the prepectus
to the pronotum and complete internal enclosure of the
mcsothoracic spiracle, the loss of the antennal ring seg-
ment, and the formation of the first phragma as a strong
anterior internal ridge above the dorsal margin of the
pronotum (Graham. 1969; Riek. 1970; Heraty. 1985,
1989; Boucek. 1988). Boucek (1988) extended
Eucharitinae to include Anorasenia Boucek and

Table I. List of described species of New World Orasema. and information of geographical region and type locality.

Orascma Cameron, 1884: 1U5

aenea Gdih-M, 1940:443 Neotropical

or^f/jmia Gemignani. 1933:489 Neotropical

aiiii'oviriclis Gahan. 1940:448 Nearctic

hakcri Gdhdn. 1940:452 Nearctic

beameri Gahan. 1940:447 Nearctic

brasiliensis Brethes, 1927:331 Neotropical

cameroni Howard, 1896:133 Neotropical

cockerelli G&han, 1940:453 Nearctic

coloradensis Wheeler, 1907: 1 2 Nearctic

co5ronc-e/j5/.j Wheeler and Wheeler, 1937:164 Neotropical

delicatula (WdilkeT). 1862:377 Neotropical

deltae Gem'ignan\, 1937:161 Neotropical

fesiiva Fahricius, 1804:157 Neotropical

/r^>'c/?e/ (Gemignani), 1933:192 Neotropical
gemignanii DeSanus, 1968:8

{maculata Westwood) [= Oheza, Heraty 1985| Neotropical

m/>iM/a Ashmead, 1888:188 Nearctic

minutissima Howard. 1897:84 Neotropical

neomexicana Gahar\, 1940:450 Nearctic

occidentalis Ashnxead, 1892:355 Nearctic

pireta Heraty, 1993:171 Neotropical

rapo (Walker), 1839:66 Neotropical

rohertsoni Gahan, 1940:451 Nearctic

salehrosa Heraty, 1993:171 Neotropical

simplex Heraty, 1 993 : 1 7 1 Neotropical

simulatrix Gahan, 1940:450 Nearctic

sixaolae Wheeler and Wheeler. 1937: 163 Neotropical

smithi Howard, 1896:134 Neotropical

stramineipes Cameron, 1884:105 Neotropical

.yw^fl/ifl^ Gemignani, 1947:6 Neotropical

/^.va/jo Gahan, 1940:440 Nearctic

tolteca Mann, 1914:183 Nearctic

v;a/?o; Gemignani, 1937:162 Neotropical
violacea k^hmead, 1888:187

(violacea) Gemignani, 1947:8 [= O. gemignanii DeSantis] Nearctic

vvWc/rv Ashmead, 1895:553 Nearctic

H'/?e<?/('7/ Wheeler, 1907:14 Nearctic

M'o/res/m (Girault), 1913a:62 Neotropical

.vw/;//;(^7)//.v (Cameron), 1909:433 Neotropical

Missiones. Argentina

Las Flores, Argentina

Uvalde, Texas. U.S.A.

Fort Collins, Colorado, U.S.A.

Ridgeway, Colorado, U.S.A.

Sao Paulo, Brazil

Grenada. West Indies

Albuquerque, New Mexico. U.S.A.

Colorado, U.S.A.

Zent, Costa Rica

Australia (?) [error]

Tigre. Argentina

Central America

Puerto San Bias. Argentina

Isla Martin Garcia, Argentina

Florida, U.S.A.

St. Vincent. West Indies

Organ Mtns, New Mexico, U.S.A.

Los Angeles, California, U.S.A.

Paraguay

Brazil

South Florida, U.S.A.

Argentina

Argentina

Oracle, Arizona, U.S.A.

Limon Prov.. Costa Rica

Grenada, West Indies

Bugaba, Panama

Prov. Mendoza, Argentina

Denison, Texas, U.S.A.

San Miguel, Mexico

Cruz Colorada, Argentina

East Florida. U.S.A.
Tepic. Mexico
Austin, Texas
Paraguay
Argentina

GoUumiella Hedqvist. which both have a fused prepectus
but have a distinct aneUiform first flagellomere and the
first phragma inflected behind the pronotum (Heraty.
1992). The subfamily Eucharitinae is here redefined to
encompass 2 new genera that have an unfused prepectus:
Neolosbanus gen. nov. (16 species), and Psilocharis gen.
nov. (9 species).

In this paper, four aspects of the systematics of
Oraseminae and Eucharitinae are treated: (1) a phyloge-
netic hypothesis is presented for the Oraseminae,

Psilocharitini. and Eucharitini (for genera with known
host or larval information); (2) the biology of Oraseminae
and Eucharitinae is reviewed, and new information is pre-
sented on the biology and morphology of immature stages
for Orasema and Neolosbanus: (3) revisionary studies are
presented for 6 genera that belong to the Oraseminae and
Psilocharitini; and (4) the biogeography of the
Oraseminae and Psilocharitini is presented and based,
where possible, on the phylogenetic hypotheses.

Materials and Methods

Specimens of Eucharitidae were examined from over 90
museums in North and South America. Europe. Japan.
Australia, and Taiwan. More than 7000 specimens
belonging to the Oraseminae were accumulated, with the
majority of this material represented by New World spec-
imens. The large amount of material examined has
allowed for a relatively thorough treatment of the world
species, which has been useful for defining generic and
geographical boundaries of taxa.

Museum acronyms are based largely on the system
offered by Amett and Samuelson (1986) with some devi-
ations after Heppner and Lamas (1982). Material referred
to in the text was borrowed from the following institu-
tions: American Entomological Institute, (AEI).
Gainesville, United States (H. Townes); American
Museum of Natural History (AMNH), New York, United
States (M. Favreau); Aligarh Muslim University
(AMUA), Aligarh. India (S. Farooqi. no loan); Australian
National Insect Collection (ANIC), Canberra, Australia
(I. Naumann); British Museum (Natural History)
(BMNH). London, England (J. Noyes and Z. Boucek);
Bernice P. Bishop Museum (BPBM), Honolulu, United
States (G. Nishida); California Academy of Sciences
(CAS). San Francisco, United States (W. Pulawski);
Canadian National Collection (CNC), Agriculture
Canada, Ottawa, Canada (G. Gibson and C. Yoshimoto);
Entomological Institute, Faculty of Agriculture. Hokkaido
University (EIHU). Sapporo. Japan (S. Takagi): Indian
Agricultural Research Institute (lARI), New Delhi, India
(S. I. Farooqui); Illinois Natural History Survey (INHS),
Champaign, United States (W. LaBerge); Personal
Collection of John M. Heraty (JMH); Entomological
Laboratory. Faculty of Agriculture. Kyushu University
(KUEC). Fukuoka. Japan (Y. Hirashima); Museum of
Comparative Zoology (MCZ). Cambridge, United States
(S. Shaw); Museum National d"Histoirc Nalurellc
(MNHP), Paris, France (J. Rasplus); Museum of Victoria

(MUV), Victoria, Australia (K. Walker); Museum Zoo-
logicum Bogoriense (MZB), Bogor. Indonesia (M. Amir);
National Institute of Agro-environmental Sciences
(NIAS). Ibaraki. Japan (K. Konishi); Queensland
Museum (QMB), Brisbane. Australia (E. C. Dahms);
Royal Ontario Museum (ROM). Toronto. Canada (D. C.
Darling); South Australian Museum (SAMA). Adelaide.
Australia (E. Matthews); South African Museum
(SAMC). Capetown, South Africa (H. Robertson); Texas
A & M University (TAMU). College Station. United
States (H. R. Burke); Taiwan Agricultural Research
Institute (TARI). Taichung. Taiwan (L.-Y. Chou);
University of Queensland (UQIC). St. Lucia. Australia
(M. Schneider); National Museum of Natural History,
Smithsonian Institution (USNM), Washington, D.C.,
United States (E. E. Grissell); West Australian Museum
(WAM), Perth, Australia (T. Houston); Zoologisches
Forschunginstitut und Museum "Alexander Koenig"
(ZFMK), Bonn. Germany (H. Roer); Zoologisches
Museum. Humboldt-Universitiit (ZMHB). Berlin.
Germany (F. Koch); Zoologisk Museum, Universitets
Zoologiske Museum (ZMUC), Copenhagen, Denmark
(B. Peterson).

DESCRIPTIVE EORMAT

Each description is based on the total number of speci-
mens examined. Mensural character states, where possi-
ble, are based on a representative sample of 10 females
and 10 males over the geographical range of the species,
except for total length of the body, which is based on all
of the examined specimens. Descriptions of the opposite
sex are based only on character states and measurements
that deviate or exceed ranges presented in the description,
which is usually based on females.

TKRMS

Tonus IoIIdu Hcnily (1^85) \\n c\icvm\\ characters and
(.k'lails c»r length measures, and Heraty (1989) lor internal
morphology of the mesosoma. Terms for immature stages
lollou Heraty and Darling (1984) and Heraty and Barber
(1990). Some modifications of terms for structures are
adopted from Boucx'k (1988). and Gibson (1989), which
are discussed in the following section on descriptive mor-
pholoev. There have been several works in the past few
years dealing uith morphological terms and attributes of
Chalcidoidea (Schauff, 1984; Heraty, 1985, 1986, 1989;
Gibson, 1985. 1986. 1989; LaSalle. 1987; Woolley, 1988;
Boucek. 1988; Huber, 1988). In particular, Gibson
(1986), Huber (1988), and Boucek (1988) provide in-
depth discussions of characters and previously used terms
in an attempt to standardize terms for use in descriptive
analysis. Unfortunately, there are still differences of opin-
ion, even with respect to major terms such as the use of
mesosoma or thorax. One hopes for the emergence of a
standard nomenclature for the field that will minimize
future discussions of morphology. Reference to certain
abbreviations and length measures are summarized in
Figures 6, 27. 31-32, 43, 48, 63, 198, 21 1, and 217-219.

Measurements and Abbreviations

The following abbreviations are used in the text: Fl-Fl 1,
antennal flagellomeres following pedicel and including
the anellus; MPS, multiporous plate sensilla; LOL, lateral
ocellar line — minimum distance between lateral (posteri-
or) ocelli; OOL, ocellar-ocular line — minimum distance
between lateral ocellus and dorsal margin of eye; SSS
(Fig. 63), scutoscutellar sulcus; TSA (Fig. 63), transscutal
articulation. Head height was measured from the top of
the median ocellus to the apex of the clypeus. Interocular
distance is the minimum separation of the eyes at their
inner dorsal margin. Length of the malar space (MS) is
the minimum distance between the ventral margin of the
eye and the lateral margin of the oral fossa (Fig. 32a).
Forewing length was measured from the apex of the
humeral plate to the wing apex. Width of the forewing
was measured as the maximum width perpendicular to the
forewing margin at the stigmal vein. Mesosomal width
was measured across the mesoscutum at the transscutal
articulation and does not include the lateral flange of the
mesoscutum just anterior to the tegula. Mesothoracic
length is measured from the anterior margin of the mesos-
cutum to the posterior ventral margin of the frenum.

Setation

The term microtrichia is reserved for setae that are barely
visible using light microscopy at high magnification
(75x), whereas hair is used for a long, narrow, and
observably flexible seta. The term pilose refers to dense,
shorter setae as found on the disc of the forewing. Setae

may be adpressed (decumbent), suhdecumhcnt . semi-
erect, or r/rr; (Bou&k, 1988).

Sculpture

Sculpture has a wide range of interpretation within the lit-
erature on Hymenoptera. This makes it difficult to
express similar features across groups (Gibson. 1989).
Recent attempts have been made to standardize sculpture
terminology in Hymenoptera (Eady. 1968; Harris. 1979).
and more specifically in Chalcidoidea (LaSalle. 1987;
Boucek, 1988; Gibson, 1989). LaSalle (1987) and Harris
(1979) presented excellent scanning electron microscope
(SEM) photographs of various sculpture types; however,
until a general atlas of microsculpture is available for
Chalcidoidea, it will be difficult to define accurately vari-
ous patterns of sculpture (Bou&k, 1988).

The most commonly used terms in this work are as
follows: aciculate (ac. Fig. 223): finely striate as if
scratched by needle; alveolate: having a regular network
of broad depressions with slightly rounded, narrow septa;
areolate (ar. Fig. 240): having an irregular network of
broad depressions with sightly rounded narrow septa; car-
inate: having one or more raised, longitudinal ridges;
coriaceous (Fig. 33): leatherlike with minute septa;
cremdate (cr. Fig. 256): scalloped and evenly rounded
with sharp dividing septa; foveate (fo. Fig. 232): having
deep, regular depressions or pits with irregular dividing
septa (finer sculpture h foveolate); glahrate (gl. Fig.
189): smooth or nearly so, shiny and almost devoid of all
sculpture, allows for isolated and minute sculpture and/or
scattered fine setae; glabrous (gs. Fig. 197): completely
smooth and polished and devoid of any hairs or setae;
granulate: minutely verrucose and appearing like sandpa-
per even under high magnification: imbricate (im. Fig.
222): having overlapping sculpture like shingles or scales
of fish; piliferous punctate (pp. Fig. 208): having circular
to oval indentations with broad interstices, each depres-
sion associated with small seta; reticulate (re. Fig. 191):
having a fine, regular network of raised septa (minute
alveolate); ribbed: having longitudinal, parallel, and
strongly raised ridges; rugose (ru. Fig. 187): having
rough, raised sculpture with smooth septa and no regular
patterns evident, dividing septa may be prominent
(strongly rugose) or hardly raised above surface (weakly
rugose) (finer sculpture is rugulose): scabrous: having
rough, sharp-walled septa in irregular pattern, septa often
raised in one direction and rasplike (finer sculpture is
scabriculous); strigate: having fine, longitudinal raised
lines; verrucose: covered with fine, irregularly shaped
spots which could be lobes or wartlike protuberances or
small circular depressions (hard to tell).

Mixed sculpture types have 2 descriptors linked by a
hyphen, with the dominant sculpture type first. Specific
terms used in descriptions can be correlated with the illus-

trations provided. Relative terms such as weak, fine,
prominent, or strong are used within descriptions to refer
to carinae, separating walls of areolae, or other sculpture
types. Definitions of this type are for comparative purpos-
es only and are avoided whenever possible as distinguish-
ing features in key couplets. Fine is usually used in refer-
ence to weak carinae or to areolae that are closely set with
narrow interstices.

Colour

The use of colour in delimiting taxa can have severe
drawbacks due to the effects of different preservation
techniques and specimen age on the nature of the colour
(Huber, 1988). There can be marked differences in colour
of specimens depending on whether they are air-dried or
critical-point dried. Colour change caused by long-term
alcohol storage can be slowed considerably by storage at
low temperatures (Huber. 1988). Although quality of
colour (hue or intensity) can change with time, the pattern
and base colour remain largely unchanged and can be
useful descriptive tools. This is particularly evident in
colour patterns of the scape, coxae, femur, and hind tibia.
The head and mesosoma range from black to a strong
metallic lustre of green, blue, or violet, and sometimes
have iridescent reflections (combinations of colours).

Head

Boucek (1988) provided an extensive description of the
structural features of the chalcidoid head. Some features
peculiar to Eucharitidae are listed below. In some taxa, a
narrow transverse depression on the vertex extends from
the lateral ocellus to the dorsal margin of the eye, here
termed the ocellar-ocular groove (OCG, Fig. 92). The
malar space may be spanned by a narrow depression
termed the malar depression (MD, Figs. 95, 223). This
depression is not associated with a fine groove or line as
in other Chalcidoidea and is often poorly defined when
present. The dorsal margin of the occiput is often cari-
nate. Homology of this carina with a similar carina of
Torymidae is uncertain but the term applied is the same
(occipital carina). The clypcus is margined dorsally by the
epistomal sulcus, and laterally by the clypeogenal suture
(E. C. Fig. 205; Richards, \911). The apical margin of the
clypeus (CLY) has a distinct glabrous flange or ventral
projection called the antcclypeus (A. Fig. 190) after
Snodgrass (1935) and Habu (1960). The region dorsal to
the clypeus is often swollen and may be bordered laterally
by the frontogenal sulcus (F, Fig. 205; Richards, 1977;
Matsuda. 1980). The labrum is usually flattened and digi-
tate with an elongate seta at the apex ol' each digit (Figs.
187-205; Darling. 1988a). The gcna extends behind the
mandible to the hypostomal carina. Medial to the hypos-
lomal carina, the hypostoma may project as a small lobe
behind each mandible (cl. Richards, 1977). The projec-

tions of the hypostoma are not considered homologous
with the postgenal extensions found in the Eucharitinae
(Heraty, 1985, 1986). Mandibular dentition is presented
as number of teeth on the specimen's right and left
mandible, respectively; for example, 3/2 dentate means
right mandible with 3 teeth and left mandible with 2 teeth.

Antennae

The antennae of Oraseminae and Psilocharitini are almost
all strongly geniculate with an elongate scape and a well-
defined pedicel. Antennal segments (or flagellomeres) are
numbered Fl-Fll, including the basal anellus or ring
segment as Fl. The first segment (Fl) is either anelliform
or absent from all members of the Oraseminae; if present
it can be recognized by its absence of multiporous plate
sensilla (Boucek, 1988; Gibson, 1989). Funicular seg-
ments are between the anellus (Fl) and clava, and in
some taxa may include F2 to FIO. The terminal clava may
be comprised of 1 to 3 fused segments (F9-F1 1 ). In most
cases individual segments of the clava cannot be identi-
fied; if apparent they are not counted individually in the
total of antennal segments (clava counts as 1 segment).
The term clava or ciuh is somewhat erroneous within the
Eucharitidae as it is often not distinguished from the pre-
ceding segments. Fused segments of the clava can usually
be viewed with transmitted light, and 1 1 segments can be
distinguished in most species of Oraseminae. Because of
variability in the degree of fusion of segments in the
clava, counts of funicular segments are more reliable in
the determination of reductions in antennal segments.

Mesosoma

Terms for the mesosoma follow Heraty (1985, 1989) and
Gibson (1986). Mesosoma refers to the combined thorax
and first abdominal segment found in the Apocrita. The
term lateral lobe (Fig. 63) refers to the mesoscutal lateral
lobe (Gibson, 1986, Heraty. 1989). side lobe (Heraty,
1985). or scapulae (Boucek. 1988). Axilla refers only to
the dorsal axillar surface (Fig. 63; Gibson. 1986; Heraty,
1989). Scut el him refers only to the dorsal area defined by
the sculoscutellar sulcus anteriorly, frenal line posteriorly,
and axillular sulci laterally or maximum dorsal width if
the axillar sulci are absent (Fig. 63). The posterior region
of the scutellum may be separated by a frenal line to form
a frenal area. Descriptions of the frenal line refer only to
the dorsal aspect (Fig. 63). The central region of the
propodeum delimited by the postspiracular sulci laterally
is termed the propodeal disc (Heraty. 1985). The supra-
coxal flange is a sharp ridge that extends between the
postspiracular sulcus and the propodeal loranicn above
the hind coxa (Boucek. 1988). The callus is a usually
swollen area between the postspiracular sulcus and the
metepimeron and may have a distinct patch of elongate
hairs; a small pointed "nib" is found on the dorsal medial

aspeci of the callus of some laxa (CN, Fig. 224). The nib
is ilifficult lo observe and (he callus must be viewed at an
oblique angle to highlight the protuberance. The meso-
plcuron is divided posteriorly into an upper and lower
mesepimeron separated by a iranscpimeral sulcus
(Gibson, 1^X6). The anterior region of the incsopleuron is
the niesepisternum. A wedge-shaped sternaular area
marked by areolate-rugose or foveate sculpture is often
found anterior and ventral to the femoral groove (SA. Fig.
224). This is not considered homologous to the stemaulus
of Ichneumonoidea, even though it is in a similar loca-
tion. The pronotum is usually irregularly sculptured later-
ally with a broadly impressed median longitudinal sulcus,
here termed the pronotal sulcus (Fig. 219).

Wings

Terms follow Boucek (1988) for nomenclature and vein
definitions (Fig. 43). The speculum (SP) is recognized as
a bare area just posterior to the parastigmal vein and base
of the marginal vein (Fig. 122). Absence of the speculum
refers to an even pilosity of this region. Because the apex
of the postmarginal vein is difficult to resolve, measures
of length are prone to error; therefore, only large differ-
ences in length were considered as diagnostic.

Metasoma

The metasoma of Eucharitidae consists of a distinct peti-
ole, comprised of the first metasomal tergite, and the
gaster (Boucek, 1988; Gibson, 1989). The petiole is well
developed in most genera and is fused ventrally in all
except females of Timioderus Waterston. Numbering of
gastral tergites follows Gibson (1989); Mt, to Mt,; Mt,^
and Mtg are fused into a syntergum. The first metasomal
stemite is absent (see discussion of Ms., in Phylogenetic
Analysis). The second metasomal (= first gastral) stemite
(Ms,) is constricted at the base in some taxa by a trans-
verse sulcus which is often crenulate and deeply
impressed (Fig. 256). Five gastral stemites are present in
females, designated Ms, to Ms^, with the last sternite
termed the hypopygium. Males have 7 gastral sternites
designated Ms., to Ms^^.

Terms of female genitalia follows Boucek (1988). The
ovipositor stylets are formed by the paired first valvulae
and fused second valvulae (Fig. 31). The ovipositor
sheaths are usually elongate and subtriangular and the
apical gonostylus may be separated from the basal second
valvifer by an oblique suture (Figs. 51, 257). In many
orascmine taxa. the first and second valvulae are expand-
ed and strongly ridged for piercing plant tissue. Notable
features of the first valvulae are the ventral subapical
ridge (observed in profile), and the longitudinal ridge of
fine teeth that occurs laterally between the apex and sub-
apical ridge (SAR, LT, Figs. 3 1 , 262). The apex of the
second valvula may have several strong transverse ridges.

or lateral teeth and a smooth median dorsal area. The
ovipositor in some laxa may be strongly curved forward
but dorsal and ventral aspects are used as if the ovipositor
is straight (i.e., second valvula is always dorsal).

Genitalia of males are generally not useful for identifi-
cation at the species level although some taxa show spe-
cialized features. Terms follow Heraty (1985, 1986; Fig.
103). Genitalia are referred to as being "typical" if the
genitalic capsule is narrow, a small median process is pre-
sent, the parameres are narrow and moderately elongate
(length is about 0.6x width of basiparameres), the digitus
is fiat and disclike with 3 to 5 marginal teeth (strong coni-
cal structures in marginal sockets), and the aedeagus is
subovate apically (Fig. 274).

Sexual Dimorphism

Sexes are readily distinguishable based on the relative
length of the petiole, the shape of the gaster, sometimes
the number of antennal segments, and the presence or
absence of an ovipositor or genitalia that usually are visi-
ble externally. More subtle differences can also be
observed. Males are usually less robust than females: the
mesosoma is slightly smaller in proportion to body size,
sculpture may be more prominent, coloration is darker, and
metallic reflections, if present, tend to be more extensive.

BIOGEOGRAPHIC REGIONS

The 4 major geographical distributions of Oraseminae
and Psilocharitini within the Old World tropics are based
on the presence of unique endemic species or genera;
Ethiopian. Malagasy, Indo-Pacific, and Australian (Fig.
282). These regions are derived from a composite of dis-
tributions for Oraseminae and Eucharitinae (Figs.
275-280).

The Indo-Australasian region is composed of a frag-
mented complex of islands that make it difficult to assign
areas of endemism for the purpose of this study. Two
major regions were recognized from southeast Asia; the
Australian region (Australia excluding north Queens-
land), and the Indo-Pacific region (excluding Australia
except for north Queensland). The 2 regions are apparent
because of several disjunct distributions of species and
genera of Oraseminae and Psilocharitini. The Indo-Pacific
region can be further divided into a number of subregions
as proposed by Grcssitt (1956) and adopted by Schuh and
Stonedahl (1986); Indo-Chinese subregion (mainland
Asia from eastern India to Taiwan and including the
Okinawa Islands, and Japan). Malayan subregion
(Vietnam, southern Thailand. Cambodia, Malaysia.
Borneo, Sumatra, and Java), Philippine subregion
(Philippine Islands). Sulawesi subregion (Sulawesi,
Timor). Papuan subregion (eastern Indonesia, Papua New
Guinea, Caroline Islands. Solomon Islands, and north

Queensland, Australia), and Polynesian subregion
(Pacific islands east of the above regions). Extension of
the Indo-Pacific region to include New Guinea, Cape
York, and the Polynesian islands recognizes a general
similarity in both insect and plant distribution (Gressitt,
1956, 1982; Schuh and Stonedahl, 1986). For purposes of
this study, the Indo-Chinese subregion is shifted slightly
west to accommodate the Indian record of Indosema, and
north to include the Japanese islands of Honshu, Kyushu,
and Tsushima, which are on the marginal distribution of

Schuh and Stonedahl's (1956) Indo-Pacific subregion.
Also, the Malayan subregion is here divided into east and
west to recognize the distinct fauna of Borneo (east
Malayan).

The Ethiopian region was not subdivided. No species
of Oraseminae or Psilocharitini were shared between
Africa and Madagascar, and except for a single record
from Cameroon, all of the Ethiopian taxa are located in
southern and eastern Africa (Figs. 275-277).

Phylogenetic Analysis

There has been an increasing number of attempts to
understand the higher classification of Chalcidoidea
based on analyses of apomorphic character states (see
Heraty and Darling, 1984; Gibson, 1986; Darling, 1988a,
1992; Schauff, 1984, 1991; Woolley, 1988). Within the
Eucharitidae, recent attempts were made to understand
classification above the generic level by means of an
analysis of the larval characters of Philomidinae (Darling,
1992), Perilampidae {sensu Boucek, 1988), and
Eucharitidae (Heraty and Darling, 1984). However, a
phylogenetic treatment of Eucharitidae has been ham-
pered by the need for a thorough treatment of taxa in the
Old World, including the description of several new taxa.
In Boucek's (1988) work on the Australasian
Chalcidoidea, most of the nomenclatural problems were
solved. Several species were reclassified in different
and/or new taxa, which made the relationships of certain
groups clear. Summaries of host relationships for
Eucharitidae published prior to Boucek's reorganization
show very little congruence of genera with host taxa
(Wheeler and Wheeler, 1937; Clausen, 1940c; Johnson,
1988). The revised classification has greatly increased the
correlation of genera of Eucharitidae with the subfamilies
and genera of host Formicidae.

The Oraseminae was originally erected to encompass
only the genus Orascma (Burks, 1979). Heraty (1985)
and Boucek (1988) defined the subfamily based on pres-
ence of a free prepectus, presence of an anellus, constric-
tion of the first gastral sternite, and presence of an
expanded ovipositor. The first 2 character states are plc-
siomorphic within Chalcidoidea and are of no value in
elucidating relationships (Gibson, 1986). The last 2 char-
acter states were considered as unique within the
Chalcidoidea and were thought to be synapomorphies
supporting a monophyletic Oraseminae. However,
Darling (pers. comm.. 1988) observed a similar constric-
tion in Perilampidae. including Chrysolampinac. and the
state is now interpreted as plesiomorphic. Examination of

Old World Oraseminae revealed that an expanded ovipos-
itor is not present in all Oraseminae, as defined above.
Thus, there appeared to be no defining features for the
subfamily.

A study was therefore made of the distribution of char-
acter states within the Oraseminae s.l. (Eucharitidae with
unfused prepectus). Under any hypothesis, the phyloge-
netic relationships of the Oraseminae cannot be tested
without comparison to the more derived Eucharitinae,
especially when the monophyly of the Oraseminae s.l. is
questionable. Genera of Eucharitinae with known biolo-
gies were therefore included in the analysis. Only 16 of
the 45 known genera of Eucharitinae {sensu Boucek,
1988) were treated. However, these genera represent a
diverse sampling of taxa within the Eucharitinae and are
what I believe to be indicative of the major groups.

The outgroup taxa chosen for the Eucharitidae were
Chrysolampinac and Perilampinae, which together have
been regarded as the sister group of the Eucharitidae
(Boucek, 1956; Graham, 1969; Heraty and Darling,
1984). Boucek (1988) questioned this relationship and
alluded to the possibility of a more distant relationship of
these families within the pteromaloid lineage. In any case,
the monophyly of Chrysolampinac + Perilampinae
(exclusive of the Eucharitidae) has not yet been resolved
and various classifications place the Chrysolampinac in
either the Pteromalidae (Riek, 1970; Burks, 1979) or
Perilampidae (Graham, 1969; Boucek, 1988; Darling and
Miller, 1991). In the most recent classification. Boucek
(1988) placed Chrysolampinac within Perilampidae. That
classification is adopted here even though larval charac-
ters suggest that Chrysolampinac is the sister group to
Eucharitidae + Perilampidae (Heraty and Darling. 1984;
Darling, 1992). Character states of the adult that support
Chrysolampinac + Perilampinae include the presence of
external pores on the ventral surface of the male scape
and fusion of the first 2 gastral tcrgites dorsally.
However, both states of the last character occur within

Chrysolanipinae. Rosolution of this conflict in oiitgroup
classification was not an objective ol this analysis. The
polarit) ot character states was evaluated against both the
Pteromalinae (Pieromalidae). which are regarded as the
general oiitgroup lor the Euchariiidae aru! Perilanipidae
(Graham. 1969; Heraty and Darling, 19X4; Boucek,
1988). and a survey of other families of Chalcidoidea.
Most of the polarity decisions were straightforward and
plesiomorphic states were assumed to be those represen-
tative of a generalized pteromalid. The use of any of the
3 other subfamilies included by Boucek (1988) in
Eucharitidae would not affect the polarity of most char-
acters.

Nineteen characters were evaluated in a previous study
of phylogenetic relationships between the Eucharitidae.
Perilampinae. and Chrysolampinae (Heraty and Darling,
1984). For that publication, larval characters were known
for only a few species of Orasema and a published
account of Loshaniis (now Orasema) uichancoi. Conflicts
in characters within Oraseminae s.l. suggested that char-
acter states within Orasema were autapomorphic reduc-
tions (Heraty and Darling, 1984). Acquisition of several
new taxa and the opportunity to examine material used in
previous studies by other authors allowed for a reevalua-
tion of character states within Orasema and related taxa.

The character states coded for adults presented several
problems, homoplasy being the largest. For almost every
character state that was evaluated in Oraseminae, the state
was found elsewhere, often within Eucharitinae where it
was clearly a parallel loss or gain. This problem is partly
a result of the diversity of form that occurs in the
Eucharitidae. The large number of taxa and characters
that were evaluated, with all of the confounding homo-
plasy. required computer analysis using parsimony algo-
rithms. I agree with Alexander (1990) that the results of
computer parsimony may not always be a reflection of
evolution because of noise encountered in large data sets.
One way of dealing with this problem is to use the analy-
sis to identify homoplasy and to direct efforts to the eval-
uation of character states that are incongruent with cladis-
tic hypotheses. Noise should not necessarily confound
scenarios that are based on sound character analysis;
rather, it should act as a strong test of the arguments.

ANALYTICAL METHODS

Character states and taxa included in these analyses were
selected during the course of revisionary studies and an
ongoing analysis of the world species of Eucharitidae.
Over the past several years. I have had the opportunity to
examine representatives of almost all of the genera of
Eucharitidae and Perilampidae as well as of several unde-
scribed genera. One theme became evident: the more taxa
that are examined, the more difficult it is to recognize

uniquely derived character states in the Eucharitidae!
Extensive morphological studies will be necessary to
uncover the value of some of these characters.

Morphological studies were carried out on the muscu-
lature of the mesosoma of several eucharitine taxa and
Orasema (Heraty, 1989). In this study, additional infor-
mation was obtained through additional morphological
studies of the mesosoma, internal and external morpholo-
gy of the first gastral stemite (Msj, the scape of males,
and the structure of antennal flagellomeres. Homology of
some structures, such as the pores on the scape of males
in Psilocharitini and Perilampidae (see discussion of char-
acter 3). needs to be determined through more detailed
studies. In order to avoid making a priori assumptions
regarding character states, coding of character states
between taxa was conservative. Therefore, although there
are considerable differences in the morphology of pores
on the scapes of Perilampidae and some Eucharitinae
(e.g., external openings in Perilampidae are not evident in
Eucharitidae), the character states were coded as the
same. This introduced considerable homoplasy into the
data matrix and lowered the consistency index of certain
characters, while not altering the resultant cladogram to a
great degree. Certainly, characters that are shown to be
homoplastic need to be reevaluated for homology.

The analysis includes the genera and species groups
studied in this revisionary work and several species in
each of the genera of Eucharitinae listed in Appendix 1.
The Orasema uichancoi-group was divided into a com-
munis OTU (Operational Taxonomic Unit) which includ-
ed O. communis and O. seyrigi (CO; code for OTU in
data matrix); it also contained a uichancoi OTU (UI),
which included all of the Indo-Pacific species.
Psilocharis was divided into 3 monophyletic groups that
were based on the character analysis of species within
the generic treatment, and included the following
species: P. aenigma (AE), P. monilicera + P. pentella +
P. afra + P. hypena (AF). and P. dahmsi -t- P. pacifica -t-
P. joanneae + P. theocles (TH). The Neoloshanus pal-
gravei-gTOup was divided into 2 OTUs represented by
N. townesi + N. laeviceps (TO), and N. taiwanensis +
N. pilosus + N. palgravei + N. nepalensis (PA). All other
OTUs represent genera or species groups as represented
in the following generic revisions. The Orasema uichan-
coi-group is similar to the O. fesriva-gToup of species in
Central and South America. The majority of New World
Orasema are closest to the O. assectaror-gwup (AS) of
species with respect to the phylogenetic character set. In
Orasema. larval information for characters 50. 56. and 57
is known only for New World species, and this infomia-
tion was included as data for the O. assectator-group.
Character states for Gollumiella and Anorasema are based
on my revisionary study of these genera (Heraty. 1992).
The Eucharitinae s.s. refer to the Eucharitini excluding

10

GoUumiella and Anorasema. The genus Schizaspidia is a
speciose group containing 35 described species, a few of
which exhibit character states of the ovipositor that
directly conflict with those found in the Oraseminae (see
character discussions). Adults of these taxa are otherwise
identical with other species of Schizaspidia and there was
no reason to code them as a different OTU. Chalcura
montana (Ishii) was treated as a separate OTU (CH
MONTANA). I do not consider this species as closely
related to other Chalcura.

Characters were coded as binary variables (0, 1 ) or as
multistate characters (0. 1, 2). Coding of character states
reflects presumed polarity (plesiomorphic = 0), but polar-
ity of all characters was established in the analysis using
outgroup comparisons (Maddison, Donoghue. and
Maddison. 1984). Table 2 lists the plesiomorphic state at
the first ingroup node resulting from the analysis. All
character states were treated as unordered in the analysis
with the exception of the number of male funicular seg-
ments (character 4) and the impression of the femoral
groove (character 29) which was ordered with a user-
defined tree.

CHARACTER ANALYSIS

Characters can be divided into 3 distinct sets: external
adult characters, internal adult characters, and larval char-
acters (including some aspects of behaviour). External
characters of adults were verified in as many species for
each OTU as possible. Examination of internal characters
involved dissections of exemplars from each genus; how-
ever, internal characters are generally conservative
(Heraty, 1989). Complete information on larval morphol-
ogy and behaviour are known for only a small proportion
of Eucharitidae, but representative information is known
for several different taxa. Character state descriptions and
distributions are presented in Table 2 and Appendix 1. An
attempt was made to minimize the number of character
states that were chosen for each character. The following
discussion deals with characters that either cannot be ade-
quately treated in Table 2 or are discussed at greater
length elsewhere (Heraty and Darling, 1984; Heraty,
1985, 1986, 1989; Darling, 1988a; Heraty and Barber,
1990).

Character I: Anelliform first flagellonwrc. The pres-
ence of a distinct ancllus is generally regarded as ple-
siomorphic within the majority of Chalcidoidea
(Konigsmann, 1978; Schauff, 1984; Gibson, 1986). An
anellus is present and distinct in Chrysolampinae and
Pcrilampinac; the loss of an anellus in Eucharitidae is
thus apomorphic. The anellus of most Eucharitidae treat-
ed here is typical for other chalcidoids. In Anorasema. the
ancllus is present (Boucek. 1988; Heraty, 1992). In
Golhtmiclla. the basal nagcllomcrc may be anelliform to

elongate like the following segments, or partially to com-
pletely fused with F2 (Heraty. 1992). The anellus is
absent from all Eucharitinae i\5. (Heraty, 1985; state 1).
In Neolosbanus and GoUumiella, the anellus undergoes
various degrees of fusion with the second flagellomere.
Multiple states for terminal OTUs were generally coded
as anellus present.

Character 2: Length of scape. A geniculate antenna,
with an elongate scape, is regarded as plesiomorphic
within Chalcidoidea (Konigsmann, 1978; Gordh, 1979;
Gibson, 1986). The scape is long and slender in
Chrysolampinae and Perilampinae. The length of the
scape is reduced in some groups of Eucharitidae until it
becomes broader than long and the antenna is no longer
elbowed (e.g., Tricoryna and Stilhula; state 1 ). As dis-
cussed by Gibson (1986), it is difficult to assign an arbi-
trary value to the length:width ratio in order to define the
antenna as geniculate or non-geniculate. A value of 4x
longer than broad is adequate to distinguish the elongate
scape of the majority of Oraseminae and Psilocharitini;
however, reductions in length of the scape in Timioderus,
Indosema, and other Eucharitinae are obvious.

Character 3: Scape of males. Distinct pores (visible on
SEM) on the scape of males are found in Torymidae
(Goodpasture, 1975), Eulophidae (Schauff. 1991), and at
least some Eupelmidae (Gibson, 1990). Males of
Chrysolampus. Euperilampus, and Perilampus (all
Perilampidae) also have small pores on the ventral sur-
face of the scape (state 0; Smulyan, 1936; Darling, 1983a,
1983b, 1986). These pores are clearly visible under high
magnification and may be openings for internal glands
(Darling, 1983b). The pores found in these various groups
indicate that presence may be plesiomorphic. The scape
of males in species of Psilocharis, Neolosbanus, and 1
species of GoUumiella has a ventral patch of small pores
that is visible in slide preparations (Figs. 141-142, 167).
These pores are associated with an apical expansion of
the scape in Neolosbanus and GoUumiella. Examination
of this region using SEM did not reveal external punc-
tures coincident with the pores found on slide prepara-
tions. Scattered minute openings were found in
Neolosbanus townesi, N. anapetus, and N . purpureoven-
tris, but were not found on the scapes of orasemine
species or other Eucharitinae examined. Homology with
pores found in the ingroup and outgroup could not be
confirmed but the states were both given the same code.
Absence of pores from the scape in slide preparations was
considered apomorphic (state 1).

Characters 4 and 5: Number of funicular segments of
males and females. The flagcllomeres between the anel-
lus, if present, and the terminal unfused segment (clava)
are the funicular segments (Fig. 48). The segmentation of
slide-mounted antennae of Chrysolampus and
Chrysomalla show 7 funicular secments with a fused 3-

11

Table 2. Character set used in the analysis of phylogcnetic relationships of Eucharitidae as presented in Figures 2-3. Character states
arc presented in presumed order from plesiomorphic to apomorphic. Values in parentheses follow ing character decriptions are the char-
acter states chosen lor the internode between the outgroup and I:ucharitidae as selected by PAUP, the consistency index, and the reten-
tion index. Differing indices for characters that had different distributions on different tree topologies are separated by a slash.

Antenna

1. Annelliform first flagellomere: 0, present; 1, absent (0, 0.33, 0.87).

2. Lengthof scape: 0, length greater than 3 X width; I, length less than 3 x width (0, 0.50, 0.88).

3. Scape of male: 0, with distinct pores; I, smooth (?, 0.50, 0.86).

4. Number of funicular segments of male (excluding anellus and clava): 7-9 (7, 0.50, 0.88).

5. Number of funicular segments of female: 7-9 (7, 0.33, 0.20).

6. Secondary basal segmentation of funicular segments: 0, absent: 1 , present (0, 0.33, 0.78).

7. Antennal branches of male: 0, absent: 1 . lamelliform; 2, ramose (0, 0.50, 0.60).

Head

8. Facial sculpture: 0, smooth; I. rugulose; 2, reticulate; 3, carinate (0, 0.60, 0.50).

9. Scrobal depression: 0, even depression; I, smooth linear channels; 2, crenulate channels (0, 0.67, 0.50).

10. Malar depression: 0, present; 1, absent (2, 0.50, 0.75).

11. Anteclypeus: 0, distinct; I. indistinct (0. 1.0, 1.0).

12. Apex of clypeus: 0. linear; 1 , lobed (0, 0.50/1 .0, 0.50/1 .0).

13. Mandibles: 0, dentate and broadly curved; I, falcate; 2, reduced (0, 1.0. 1.0).

14. Form of labrum: 0, digits absent, setae marginal; 1, digitate with adoral digits (Perilampinae only); 2, planar and digitate; 3,
reduced in size; 4, complete loss (Darling, 1988a) (2. 1.0, 0).

15. Number of labral digits or marginal setae: 0, more than 5; 1,4 (0, 0.33, 0.73).

16. Posterior genal margin: 0, open; I, closed (Heraty, 1985, 1986) (0, 0.50, 0).

17. Mouthparts: 0, large; I , much reduced (0, 0.33, 0.50).

18. Dorsal occipital margin: 0. rounded: I, carinate (0, 0.25, 0.77).

Mesosoma

19. Prepectus: 0, freely articulating with pronotum; 1, fused with pronotum on different plane; no internal rod: 2, fused with
pronotum on same plane, no internal rod; 3. fused with pronotum and associated with internal prepectal rod {Peiilampus
only) (Heraty, 1989) (0, 1.0, 1.0).

20. Shape of prepectus: 0, narrowed ventrally, reaching tegula and evenly sculptured; 1 , triangular and sculptured, reaching tegu-
la; 2, narrowed ventrally and foveate dorsally; 3, subtriangular. broadly separated from tegula: 4, upper half fmgerlike, nar-
rowly separated from tegula; 5, narrowly separated from tegula and sinuate posteriorly (1, 0.62/0.71. 0.83/0.89).

21. Profile of pronotum: 0, visible in dorsal view; I, hidden (Heraty, 1985; Boucek, 1988) (0, 1.0. 1.0).

22. Scutoscutellar sulcus: 0, diagonal to TSA, dividing axillae; I, transverse and carinate internally, axillae transverse; 2, trans-
verse with prominent internal carina (Heraty, 1989) (0, 1.0, 1.0).

23. Notauli: 0. present; I , absent (0. 0.50. 0.50).

24. Transscutal articulation: 0, present; 1, absent (0, 1.0, 1.0).

25. Mesoscutal ridge (internal): 0, absent; 1 , present (Heraty, 1989) (0, 0.50. 0.89).

26. Mesothoracic spiracle (internal): 0, not enclosed dorsally; I, enclosed by sclerotized cuticle dorsally; 2, broadly enclosed
(Heraty, 1989) (0, 1.0, 1.0).

27. Frenal groove on scutellum: 0, present; 1, absent; 2, as in Perilampinae (0, 0.33/0.29, 0.43/0.29).

28. Propodeal sculpture: 0. evenly sculptured: I. smooth (0. 0.5. 0.89).

29. Femoral groove: 0, rounded and shallow; 1, broadly and evenly impressed; 2, narrow and foveate (0, 1.0. 1.0). Ordered using
character state tree, I *-^ 0 <-^ 2.

30. Lateral axillar surface: 0. large; 1 , reduced (Heraty. 1989) (0, 0.33, 0.33).

31. Scutellar spine morphology: 0, absent; I . short and bifurcating; 2. long and bifurcating, united basally; 3. long spines, not
unitedbasally (0.0.75.0.67).

32. Mesepistemum: 0, ventral margin linear; I, ventral margin broadly rounded and sviollcn anterior to mid coxa; 2, wedge-
shaped anterior to mid coxa (Heraty, 1989) (0. 0.40. 0.82).

Wings

33. Wing setae: 0, large and dense; 1, small or absent (0, 0.33, 0.33).

34. Hind wing vein: 0. well developed; 1, medially indistinct (Heraty, 1985) (0, 0.50. 0.75).

35. Stigmal vein: 0, perpendicular; 1, sharply angled to apex of wing; 2, round (0, 0.40, 0).

36. Postmarginal vein: 0. long; 1, short (0, 0. 14, 0.46).

37. Speculum of forewing: 0, setose (absent); 1 , present ( 1 , 0.20, 0.64).

Metasoma

38. Petiole of female: 0, transverse and not fused ventrally; 1, transverse and fused ventrally; 2, elongate and fused ventrally (2,
1.0, 1.0).

39. Base of petiole: 0, tapered, associated with transverse petiole; 1, truncate basally with dorsal flange; 2, tapered basally, asso-
ciated with elongate petiole; 3, truncate basally, without dorsal flange (0, 1.0, 1.0).

40. Sternal constriction: 0, present; 1 , absent (0, 0.50, 0.92).

41. Ms, of males: 0, no anterior projection; 1 , expanded cup-shaped anterior region projecting forward under petiole (0. 1.0, 1.0).

42. Expansion of valvulae: 0, acicular; 1 , expanded (0, 0.25/0.20, 0.79/0.7 1 ).

43. Lateral apex of first valvula: 0, smooth; 1, with diagonal ridges; 2, 3-4 teeth; 3, several minute teeth (0, 0.50, 0.70).

44. Apex of second valvula: 0, dorsal ridges small or absent; 1, strong dorsal ridges, ridges diagonal and coalescing; 2, lateral
teeth; 3, strong transverse ridges (0, 0.60, 0.80).

45. Hypopygial setae: 0, small or absent; 1 , single row of elongate hairs; 2, dense cluster of apical setae (0, 0.50, 0.82).

46. Gonostylus: 0, separated; 1 , fused (0, 0.50, 0.93).

47. Number of hind tibial spurs (2, 0.50. 0.50).

Immatures

48. Egg shape: 0, stalked; 1 , elliptical (Heraty and Darling, 1 984) ( 1 , 0.50, 0.50).

49. Setal pattern of tergite III: 0. ventral spine present; 1, ventral spine absent (0, 0.50. 0.75).

50. Dorsal setae of first instar: 0. presence on tergites VII, IX, and XI; 1 , loss of setae on tergite XI; 2, loss of setae on VII, IX,
and XI; 3. loss of setae on all but T 1 and II (Heraty and Darling, 1984) (1, 1.0, 0).

51. Tergites I and II of first instar: 0, separated; 1, fused (Heraty and Darling. 1984) (0, 1.0, 1.0).

52. Tergopleural line on first instar: 0, absent; 1, present (Heraty and Darling. 1984) (0, 0.5, 0.86).

53. Caudal cerci on first instar: 0, absent; 1, present (Heraty and Darling, 1984) (1, 0.5, 0).

54. Caudal pad on first instar: 0, absent; 1 , present (Heraty and Darling, 1984) (1,1 .0, 0).

55. Antenna on first instar: 0, present; 1, absent (Heraty and Darling, 1984) (1, 1.0. 0).

56. Cranial setae on first instar: 0, present; 1, absent (Heraty and Darling, 1984) (?, 0.33, 0.33).

57. Postlabium of first instar: 0, not expanded; 1 , expanded ( 1 , 1 .0, 0).

58. Labial plate of first instar: 0. absent; 1, present; 2, secondary (0, 1.0, 0).

59. Third-instar type: 0, hymenopterifomi; 1. pustulate as in Pcrilainpus; 2, pustulate as in Orasema: 3, as in Chalciira: 4, as in
SlilhuliK 5, as in Eiicharis (see Smith, 1912; Clausen. 1940b) (0/1/2/3, 1.0/0.83, 1.0/0.83).

60. Pupa: 0, chalcidiform; 1 , pustulate over petiole (0, 1 .0, 1 .0).

Behavioural

61. Number of eggs laid with each oviposition: 0, 1 egg; 1, 1-10 eggs; 2, more than 10 eggs (1. 1.0. 1.0).

62. Attachment of first instar to host: 0, external; 1, internal (0, 0.5, 0.5).

segmented clava (Darling. 1^)86). The segments of the
clava in Perilampinae appear to be more closely
appressed. again gi\ing a 7-segnienied lunicle. Seven
segments are common in Pteromalidae also, so the 7-seg-
mented tunicle was chosen as the plesioniorphic state for
both outgroup taxa. Within Eucharitidae, the number of
funicular segments is variable and depends on the degree
of fusion in terminal segments and the interpretation of
the anellus. The number of funicular segments of males
(character 4) varied from 7 to 9: this character was treated
as ordered to provide a grouping of species within
Neoloshanus that had 8 or 9 funicular segments.
Unordering this character resulted in no resolution among
species groups of Neoloshanus.

Boucek and Noyes (1987) referred to the absence of
multiporous plate sensilla ("longitudinal sensilla") as a
characteristic of Eucharitidae. Multiporous plate sensilla
were found in almost all of the Eucharitidae examined
(absent in males of the Neolosbamts gemma-group and
Anorasema), although they are reduced in size and
recessed into depressions within the Eucharitinae s.s.

Character 6: Secondary basal segmentation offlagel-
lomeres. The antennae of Psilocharis theocles,
Neolosbamts palgravei, N. townesi, N. purpureoventris,
and Gollumiella show a distinct secondary segmentation
at the base of the funicular segments (Figs. 213-214,
216). which is not unlike the separation between the anel-
lus and F2 (Figs. 211, 215). In Gollumiella, coded as pre-
sent, the segmentation forms a cuplike flange overlapping
the base of the flagellomere (Heraty, 1992). Observation
of this character state is difficult and will require a broad
survey for its distribution using SEM. No secondary seg-
mentation was observed in Oraseminae or other
Eucharitinae, with the exception of Chalcura, which has a
weak basal flange similar to that of Gollumiella.
Secondary segmentation undergoes parallel development
in Psilocharitini, Gollumiella, and Chalcura, although it
may be plesiomorphic for Eucharitinae.

Character H: Facial sculpture. Sculpture of the frons
and lower face is a variable attribute but it is important in
the separation of some lineages of Oraseminae. A com-
pletely smooth face is found in Chrysolampinae and most
Perilampinae and is regarded as the plesiomorphic state
(state 0; Figs. 189, 193-205). Within Eucharitidae, 4
states were recognized with facial sculpture further divid-
ed into rugulose (state 1; Figs. 187-188), reticulate (state
2; Fig. 191), and vertically carinate (state 3). Rugulose
and carinate sculpture patterns were broadly defined to
include several variants, whereas smooth and reticulate
could be more accurately delimited.

Character 9: Scrobal depression. A relatively smooth
or evenly rounded scrobal depression was regarded as
plesiomorphic and was found in the outgroup taxa as well
as most of the Eucharitinae. In Psilocharis and Orasema

glabra, there are 2 smooth channels that terminate in a
small depression at about ().(>-(). Xx the distance between
the toruli and median ocellus (slate 1; Figs. 127. 194—195.
197). State 2, crenulate furrows bordering a raised median
area and reaching the median ocellus, is found only in the
Neolosbanus gemma-group (Figs. 153-154), although
they may be similar to the condition found in Orasema
uichancoi (Fig. 188). The homology of the 2 states is
uncertain.

Character JO: Malar depression. A malar sulcus (visi-
ble as a distinct line) is present in some species of
Chrysolampinae and Perilampinae. Its presence is regard-
ed as plesiomorphic within the Perilampidae based on its
widespread distribution in the Pteromalidae. Absence of a
distinct suture is an autapomorphy of Eucharitidae that
was not coded for in the analysis. However, a linear
depression of the malar space, in a position similar to the
malar sulcus of Perilampidae, occurs in some taxa of
Eucharitidae (Figs. 95, 223). This depression is absent in
Perilampidae (state 0), and its presence in some
Eucharitidae is treated as a derived state (slate 1 ). An apo-
morphic development of the depression into a broad
foveate groove only in some members of the O. uichan-
rw-group was not included in the data matrix (see discus-
sion of phylogeny in revision of Orasema).

Character 11 : Anteclypeus. The anteclypeus (A, Figs.
190, 198), a distinct area along the apical margin of the
clypeus, is found in Perilampidae and most Eucharitidae
(state 0). The margin of the clypeus is rounded and lacks
any distinct area in species of Neolosbanus. Some speci-
mens of A', purpureoventris have the apex of the clypeus
on a different plane so that it appears marginate in some
views (Fig. 210). This was considered as an independent
development, and the clypeus was coded as for other
Neolosbanus. A similar reduction of the anteclypeus
occurs in some species of Austeucharis. Chalcura, and
Schizaspidia although they are probably not homologous
character states.

Character 12: Ape.x of clypeus. In the Neolosbanus
purpureoventris- and N. palgravei-groups, the ante-
clypeus is strongly produced over the base of the mouth-
parts as a semicircular lobe (state 1; Figs. 200-204). The
lobe is less strongly developed in N. townesi (of the pal-
gravei-group) and the A', gressitti-group, and is consid-
ered as absent. It is possible that character 12 is coupled
with character 1 1 and that together they should be treated
as a single apomorphy. Separate coding for characters 1 1
and 12 would be the same as combining them and treating
the character as ordered.

Character 13: Mandibles. The plesiomorphic state is
interpreted as broad, opposable mandibles, which broadly
join or overlap along their apical margin in the same
plane as the ventral margin of the head (state 0).
Apomorphic differences in the mandibles of

14

Chrysolampinae and Perilampinae were disregarded.
Falcate or sickle-shaped mandibles that fold in a plane
perpendicular to the ventral margin of the head (vertical)
are regarded as an autapomorphy of Eucharitidae (state
1). Reductions of the mandibles occur in various genera,
including Pseudometagea and Eucharis (state 2), and the
mandibular configurations of Timioderus and Indosema
are extreme. In Indosema, the mandibles are completely
lost. In Timioderus, the mandibles are reduced and elon-
gate, or in the case of Timioderus peridentatus. ^ill den-
tate, enlarged, and cup-shaped (Figs. 28, 39) as is typical
of the Perilampinae and Pteromalidae. This is treated as a
reversal to the plesiomorphic state.

Characters 14 and 15: Lahrum. A labrum having mar-
ginal digits arranged in the same plane is regarded as an
autapomorphy of Eucharitidae, with a larger number of
digits as the groundplan state for the Chrysolampinae and
Perilampinae (state 0; Darling, 1988a). In Orasema and
the basal Eucharitinae, the labrum is distinctly 4-digitate
(state 1 ) with only minor variations in the number of dig-
its (4-5). Number of digits, presence or absence of digits,
and shape of the labrum are highly variable in
Orasemorpha, Timioderus, and Indosema. In the
Eucharitinae s.s., the labrum is usually 9-16 digitate
(state 0), with a 4-digitate labrum found in some species
of Pseudometagea and Pseudochalcura.

Character 18: Occipital carina. An occipital carina is
found at the junction of the vertex and the occiput behind
the lateral ocelli in most Eucharitidae. Such a carina is not
considered to be homologous with the occipital carina
found in Torymidae. An occipital carina is not found in
the 3 other subfamilies of Eucharitidae, and Perilampidae.
and is rare in Pteromalidae; therefore it is regarded as a
derived character state in the Eucharitidae. In most taxa,
the carina is stable for all members of a genus. However,
a weak carina is found in some individuals of Orasema
koghisiana and O. valgius. In both cases, the carina is
associated with weak striae along the vertex that become
consolidated into a single carina, and this was not coded
as present.

Characters 19 and 26: Fusion of the prepectus:
mesothoracic spiracle. Fusion and shape of the prepectus
can be correlated with changes in internal muscle attach-
ments (Heraty. 1989). Fusion of the prepectus in most
Perilampinae (character 19; state 3) is considered as inde-
pendent from that in Eucharitidae because an internal
apodeme is present, and the mesothoracic spiracle is not
enclosed by cuticle dorsally (character 26; state 0; Heraty,
1989). Fusion of the prepectus is coded differently in
Anorasema and (loUumiella where the prepectus lies on a
different plane from the pronotum (character 19; state 1).
Dorsal enclosure of the mesothoracic spiracle (character
26; state 1 ) was observed in Anorasema. but was difficult
to assess on the smaller (iollumiella and therefore was

coded as missing, although it did appear to be enclosed.
Enclosure of the spiracle and lack of an internal apodeme
suggest that the fusion is homologous with Eucharitinae s.s.

The fusion of the prepectus on the same plane (charac-
ter 19. state 2) in Eucharitinae was considered as the final
step in a transformation series from the fused pronotum in
Gollumiella and Anorasema, which is on a different plane
(state 1); the perilampine type was considered autapomor-
phic and unrelated to states 1 and 2. Coding the prepectal
fusion in Eucharitinae as 2 separate states rather than a
single state allows for more freedom in the placement of
Anorasema and Gollumiella based on other character-
state distributions.

Character 20: Shape of prepectus. The shape of the
prepectus was divided into 5 states. In Chrysolampinae
and Perilampinae. the prepectus extends to the tegula and
is usually narrowed ventrally (state 0; as in Fig. 222) or
broad and triangular (state 1; as in Fig. 217). and was
coded as a multiple state in both outgroup taxa (cf. figures
in Boucek, 1978. 1988; Darling. 1983b. 1986, 1988b).
Character states 0 and 1 are represented in the
Pteromalidae. In Oraseminae, the prepectus is usually tri-
angular (not strongly narrowed ventrally), evenly sculp-
tured, and reaches the tegula (state 1; Figs. 27. 51, 83, 98,
217-218, 220). A similar prepectus is known only for
Pseudometagea (Eucharitinae). In some Oraseminae and
in Neoloshanus and Psilocharis the prepectus is triangular
dorsally and foveate, but strongly narrowed ventrally
(state 2; Figs. 222-225, 227-228). In Stilhula and related
genera, including some species of Schizaspidia, the
prepectus is reduced in length and broadly separated from
the tegula (state 3; cf. Heraty. 1989. figs. 13, 15). In
Chalcura, Austeucharis, and some members of
Schizaspidia , the prepectus is strongly narrowed for most
of its height and dorsally forms a foveate, fingerlike
extension to the tegula (state 4; cf. Boucek, 1988. figs.
945-946). Lastly, the prepectus of Kapala and related
taxa is narrowly separated from the tegula and is sinuate
along the posterior margin (state 5; cf. Heraty. 1989,
fig. 3).

Character 25: Mesoscutal ridge. An internal mesoscu-
tal ridge appears to be an autapomorphy of the
Eucharitinae s.s. (state 1; Heraty. 1989). A mesoscutal
ridge is distinct in Indosema. but absent from other
Oraseminae. Psilocharitini. and Anorasema (state 0).
Individuals of Gollumiella were too small to allow for
resolution of the states of this character, but the mesoscu-
tum appeared to lack the ridge, and was coded as state 0.

Character 2H: Propodeal sculpture. The plesiomor-
phic state for Perilampidae and most Eucharitinae is to
have the propodcum evenly sculptured. In some Orasema
and all Psilocharitini. the propodeal disc is smooth except
for a median carina or alveolate furrow (stale 2). In ln)lli
groups, this state is associated uilii an e\cnl\ rounded

15

and glabrous callus. The propodcum is llai and smooth in
sonic Kapiihi. but lliis uas nol considered as honioiojious
to (he sniooili propodeiini described above. Stale 1 was
interpreted as being derived independently in Orasema
and PsilcKharitini.

Character 29: Femoral i>r<><)ve. An evenly rounded or
indistinct femoral groove was interpreted as the ple-
siomorphic state (state 0). In Psilocharis, the femoral
groove is broadly and evenly impressed, and is associated
with a foveate sternauiar area anteriorly (state I; Figs.
222-225). In Neoloshanus, the femoral groove is narrow
and foveate. with or without an associated foveate ster-
nauiar area (state 2; Figs. 226-234). The character was
ordered using a character-state tree so that there was a
one-step difference from the ancestral state and a two-
step difference between the derived states. There was no
reason to assume that one state was transitional to the
other, and ordering was used to eliminate additional
hypotheses for the Psilocharitini lineage (e.g.. gemma-
group as ancestor to Psilocharis).

Character 32: Mesepisterniim. The plesiomorphic
state for the mesepisternum is a straight ventral margin
(in lateral view) with no expansion anterior to the mid
coxa (Pteromalinae and Chrysolampinae. state 0; Figs.
219, 221-228). The ventral margin of the mesepisternum
is strongly expanded in most species of Orasema,
Orasemorpha. Timioderus, and Indosema (state 1; Figs.
27. 51. 217-218, 220). A similar expansion of the
mesepisternum occurs in the Stilbula and related taxa (cf.
Heraty, 1989, figs. 15. 17). In Perilampinae. Chalcura.
Schizaspidia, Kapala, Isomer a la, and Austeucharis, the
mesepisternum extends as a strong wedge anterior to the
mid coxae (state 2; cf. Heraty, 1989, fig. 13).

Character 35: Stigmal vein. The shape of the stigmal
vein is differentiated into 3 states, listed in Table 2. In
most Eucharitidae, the stigmal vein is usually quadrate to
more than 3x longer than broad, associated with a well-
developed marginal vein, and perpendicular to the
forewing margin (state 0). The angle of the stigmal vein
to the anterior margin is important in separating the
Neoloshanus purpureoventris-gronp (state 1; Fig. 169)
from other groups of Neoloshanus (state 0; Figs.
162-165). Timioderus and the outgroup taxa were coded
as having character state I although homology of these
occurrences is questionable. A circular stigmal vein asso-
ciated with a reduced marginal vein is a synapomorphy of
the Oheza clade (state 2).

Character 36: Postmarginal vein. The length of the
postmarginal vein relative to the marginal vein was treat-
ed as a binary variable. A long postmarginal vein was
defined as being more than 0.5x longer than the marginal
vein, and often reached to the extreme apex of the wing.

Character 38: Petiole of female {Mt i). Three states are
recognized based on shape and degree of ventral fusion of

the petiole: state 0, petiole short, broader than long, and
not fused ventrally, with ventral margins broadly separat-
ed (Figs. 27, 35); state 1, petiole short but lused ventrally
(Fig. 256); state 2, petiole longer than broad, cylindrical,
and fused ventrally. Within the Eucharitidae. states 0 and
I are found only in Timioderus. Indosema. and
Orasemorpha. Chrysolampinae has all 3 states and
Perilampinae has both state I and state 2. The petiole is
not fused ventrally in Chrysomalla (Chrysolampinae)
(personal observation) and in Khicnocoelia
(Pteromalidae) (Heydon. 1989). Heydon suggested that
lack of fusion of the petiole is plesiomorphic. but at the
same time he placed Rhicnocoelia as a derived member of
the Sphegigasterini. Multiple character states found in the
outgroup make it difficult to assess polarity within the
Eucharitidae. Treating this character as ordered and with
the ancestral state fixed as 0 had no effect on the results
of the analysis. The character was treated as unordered.
State 2 was chosen as the plesiomorphic state for
Eucharitidae.

Character 39: Base of petiole. The shape of the base
of the petiole was differentiated into 4 states. In the ple-
siomorphic state (state 0). the base of the petiole tapers to
the condyle of the .short petiole, and is associated with a
short petiole as found in Perilampidae, and some
Oraseminae. Alternate states occur in the Perilampidae
and include an elongate petiole with a strong basal flange
in Chrysolampus. and a transverse petiole with a strong
dorsal flange in some Perilampus. These states were dis-
regarded as they are unique derivations in each group. In
all Eucharitinae, excluding Psilocharis. the base is gradu-
ally narrowed and inserts into the propodeal foramen, the
condyle is small and indistinct, and the petiole is elongate
(state 2). In most Orasema the base of the petiole is trun-
cate basally and abruptly narrowed to the basal condyle of
the petiole, and the basal margin has a distinct basal
flange (state 1 ). The petiole is abrupt but lacks a dorsal
margin in Psilocharis (state 3). State 0 is plesiomorphic
for Eucharitidae. Because state 2, which is associated
with an elongate petiole, is closely linked with character
38, it would be more apomorphic.

Character 40: Sternal constriction. A medial constric-
tion of the second metasomal stemite, Ms,, is interpreted
as plesiomorphic (state 0) in the Eucharitidae. and its
absence as apomorphic (state 1). The constriction origi-
nates at the anterodorsal margin of Ms, at the point of
articulation with the petiole, and curves posteriorly, delin-
eating a semicircular or subtriangular anterior region
(Figs. 27, 35, 51. 54, 61, 74, 78. 126. 138, 256. 258). The
anterior region may be strongly developed and shelflike
with a broad crenulate constriction (Fig. 256), smooth and
broadly rounded with a narrow smooth constriction (Fig.
258), or a short band with a broad crenulate furrow as in
Chrysolampus ohlongiscutella Girault (Chrysolampinae).

16

Different configurations were not distinguished in this
analysis. Presence was estabhshed by a distinct constric-
tion. The shallow indentation found in the Neolosbanus
gemma- and purpureoventris-gvoups (Fig. 273), or condi-
tions in which the anterior region is differentiated only by
sculpture (Fig. 172) were coded as absent (state 1).

Internally, the constriction is associated with a contin-
uous band of muscles which converge on a median ven-
tral ridge within the petiole. Interpretation of homology
with muscles presented in Snodgrass (1956) for the hon-
eybee metasoma was not possible. Each of the gastral
sternites is connected by a fine network of antecostal
muscles that abut on either side of the antecosta of each
segment. However, the antecostal muscles between seg-
ments Ms, and Ms, originate on the anterior dorsolateral
margin of Ms, and insert broadly along the antecosta of
Ms,, suggesting that the antecostal ridge has been lost or
reduced to the dorsal margin of Ms,. The muscles attach-
ing to the constriction insert posterior to, and underneath
(laterally), the antecostal muscles, arguing against the
homology of these 2 muscle groups. The attachment and
shape of the muscle group attaching to the constriction
suggest that it may act as a levitator of the stemite with
respect to the petiole. The same muscle group is present
in a similar position in eucharitine taxa where the con-
striction is absent.

The constriction and distinct anterior region are promi-
nent in all members of the Perilampinae. Within the
Chrysolampinae, the character has multiple states:
Chrysomalla has a broad anterior region and smooth con-
striction (state 0); Chrysolampus and Brachyelutus have a
narrow anterior region with crenulate furrow (state 0);
and some species of Chrysolampus have a sculptured
anterior region lacking a constriction (state 1). A promi-
nent constriction on Ms, was also found in Toiymus
(Bucher, 1948) and Podagrion (both Torymidae), Dipara,
and Lelaps (Pteromalidae; cf. Yoshimoto. 1977. figs.
1-2). as well as in some other ptcromalids and scattered
members of Eulophidae. A broad anterior region and nar-
row constriction was also observed in Philomides rhode-
siensis Risbec, but was absent from other members of the
Philomidinae. The muscles and their homology need to be
studied on a broader scale within the Chalcidoidea to
determine their value in classification. Only presence or
absence of the constriction is treated here.

Charade/ 4J : Ms^ of males. In Timioderus. Indosema,
and Orasemorpha, the anterior region of the second meta-
somal stemite projects cephalad beneath the petiole (Figs.
35, 54, 75. 78). The anterior region is expanded and cup-
shaped, and when contracted, fits securely against the
ventral surface of the petiole. A similar expansion of Ms,
is found in both sexes of the Pseudometagea schwarzii-
group (cf. Hcraty. 1985. figs. 50-51). Pseudometagea
sehwarzii Ashniead also shows an indistinct constriction

of the anterior region and a folding of the anterior margin
similar to that of Orasemorpha didentata (Fig. 78). The
phylogenetic position of Pseudometagea and general dif-
ferences in morphology of the sternite suggest that the
projection in Pseudometagea is not homologous with that
found in Orasemorpha. Timioderus, and Indosema.
Inclusion of this autapomorphy as a multiple state for
Pseudometagea did not affect the analysis, but it was not
included in the final data matrix. A forward projection of
the stemite was not found in the outgroup taxa and was
therefore regarded as apomorphic.

Character 42: Expansion of valvulae. The ovipositor
exhibits various shapes from being acicular or needlelike
(Figs. 264-265, 271-272), having a strong subapical
expansion (Figs. 262, 266-267), or being evenly expand-
ed along its entire length (Figs. 268-270). An acicular
ovipositor is found in the outgroup taxa and other
Chalcidoidea, and is regarded as plesiomorphic. Only 2
states are recognized, acicular (state 0) and expanded
(state 1). because it is difficult to partition differences
between subapically expanded and evenly expanded.
Curvature of the ovipositor was proposed as a character
system but was abandoned because of problems in parti-
tioning states within the expanded ovipositor types. In
almost all species of Orasema, the ovipositor shows a
marked forward curvature that is so pronounced in some
species that the second valvula becomes ventral to the
first (Figs. 93, 262). The curvature is likely associated
with the habit of ovipositing parallel to the leaf surface as
shown for Orasema (Fig. 4). The ovipositor is slightly
curved and elongate in Timioderus and Orasemorpha.
Indosema has a straight ovipositor that is articulated in a
unique manner, and is flipped forward between the coxae
in some preserved specimens. In Neolosbanus, the
ovipositor is expanded but straight and. in the case of A'.
palgravei, associated with a slightly different oviposition
behaviour (Figs. 18-19). Of all the species observed in
Orasema, O. communis is the only species that has a
straight ovipositor with diagonal ridges at the apex of the
first valvula. Anorasema and a few species of
Schizaspidia possess a straight or curved ovipositor,
respectively, which suggests 2 different modes of ovipo-
sition.

Character 43: Lateral ape.x of first valvula. Four char-
acter states are recognized for structures found along the
lateral apex of the first valvula: state 0. processes absent
(Figs. 265, 270. 272); state 1. diagonal ridges (Fig. 267):
state 2, 3 to 4 lateral teeth (Figs. 259, 263): and state 3.
row of 6 to 10 minute teeth (Fig. 261).

Character 44: Apex of second valvula. Strong trans-
verse ridges (state 1: Figs. 29, 31, 99, 267. 269) or lateral
teeth (state 2: Figs. 52, 128b. 270) are sometimes found
on the second valvula. These ridges arc absent from
Anorasema and Chalcura moniana. Minute dorsal ridges

17

on an acicular ovipiisitor are dilTiciili lo discern and
ridges were ofien observed \^itll SLM thai were not visi-
ble with light microscopy. The presence ol minute ridges
(Figs. 265. 272) was coded the same as when completely
absent, and treated as the plesiomorphic state. Presence ol"
lateral teeth was often distinct (Fig. 270). or subject to
interpretation based on the prominence of the medial
ridges (Fig. 261, coded as teeth present).

Character 45: Hypopyiiial setae. Three distinct setal
patterns are recognized at the apex of the hypopygium
(Ms^): state 0. small or minute setae subapically (Figs. 27,
51, 61. 74. 113. 126. 172, 262): state 1, single row of
elongate hairs along posterior margin (Figs. 149, 264):
state 2, cluster of elongate setae around apex of hypopy-
gium. Absence or presence of minute setae (character
state 0) was regarded as plesiomorphic. Character state 1
is unique within the Chalcidoidea and supports a relation-
ship between the Psilocliaris and Gollunilella +
Anorasema. However. Psilocharis aeiili>ma lacks the
setae but definitely belongs to this genus. Conflicts with
other characters render the character homoplastic.

Character 46: Gonostylus. The plesiomorphic condi-
tion is interpreted as a complete separation of the gonos-
tylus from the second valvifer by a diagonal suture (Fig.
257); absence of the suture is considered apomorphic.

Character 48: Egg shape. Heraty and Darling (1984)
used the stalked egg as an autapomorphy for the
Eucharitidae. Egg shape can be determined from ovarian
eggs dissected from the gaster. Species that deposit a
stalked egg possess a dumbbell-shaped ovarian egg which
is narrower at one end, and those that deposit cylindrical
eggs (Perilampinae) have a cylindrical ovarian egg.
Critical-point-dried specimens offered an opportunity to
survey the ovarian eggs, often with very little damage to
the gaster. Some ovarian eggs were recovered from repre-
sentatives of genera that were air-dried, if the gaster was
already split open or crushed. Cylindrical eggs (.state 0)
are found in both Tiinioderus and Indosema whereas their
apparent sister group, Orasemorpha, has distinctly dumb-
bell-shaped ovarian eggs. Chrysolampus sysimhri has also
been shown to possess stalked eggs (Darling and Miller,
1991). Ovarian eggs of Chrysolampus sp. (from West
Australia) and Chrysomalla hesperis are tapered and
fusiform. Eggs of Eucharitidae are more distinctly stalked
and are probably independently derived: however,
Chrysolampinae were coded as possessing a stalked egg
in the analysis (state 1 ).

Characters 49-58: Characters of first-instar larvae.
The characters of the planidial larva were treated by
Heraty and Darling (1984); a reevaluation of some states
for Oraseminae is presented here. Differences from states
coded for Oraseminae in Heraty and Darling (1984)
include the following: a tergopleural line is absent in
Orasema, Oheza, Stilhula, and Pseudochalcura (character

49, discussed further in biology section): and character
states presented for Neohjshanus laeviceps were verified
in N. palgravei (same as Eucharitinae .s..v.). A hatchet-
shaped labial plate (character 55, state I ) is found in all
Eucharitinae. A similar labial plate occurs in only I unde-
scribed species oi' Orasema (Heraty, 1990, fig. 18b: state
2): coding this as the same character state increased the
number of possible tree topologies only within
Eucharitini.

Character 59: Third-instar larvae. The third-insiar lar-
vae have characteristic shapes within major lineages of
Eucharitidae (see Clausen. I94()b). The larva of Orasema
is unusual in that it has enlarged pustules (state 2: Fig. 1 1)
that have been identified in diverse species groups {stri-
atosoma- . assectator- . and z//r/?c/ //ro/'-groups).
Categorizing larvae within the Eucharitinae is somewhat
arbitrary: however, the third instar of N. palgravei was
coded the same as for Chalcura (state 3) based on the
enlargement of the mesothorax behind the head (Fig. 24).
Perilampus has a pustulate third-instar larva but the
enlargements are bandlike over the entire dorsum (Smith,
1917); its larval shape was therefore coded as a separate
character state (state 1) from Orasema.

Character 60: Pupa. The presence of pustules along
the dorsal margin of the petiole (state 1; Fig. 12) is known
only in Orasema. As for the larval pustules, this state is
known for a wide variety of Orasema.

Character 61 : Number of eggs deposited. Three char-
acter states are recognized: 0, 1 egg; 1, 3-10 eggs; and 2,
more than 10 eggs. The distinction is clear in most cases.
All members of Orasema. Neoloshauus laeviceps, and N.
palgravei deposit single eggs in punctures. Members of
the Eucharitinae s.s. deposit large numbers of eggs
(60-10 000) in a single oviposition and usually in cavities
within plant tissue. In Gollumiella antennata. a circle of
100 eggs is deposited around a thrips egg; this was treated
as character state 2 although it is different from anything
known for the Eucharitinae (some Eucharitinae scatter
eggs over the leaf surface but not in association with an
intermediate host). Oviposition habits in the outgroup
laxa arc poorly known and suggest that either single (as in
Perilampus chrysopae) or few eggs (as in Perilampus ful-
vicorni.s) may be the ancestral state (Clancy, 1946; Laing
and Heraty, 1981). Chrysolampus deposits from 1 to 12
eggs into plant tissue (Darling and Miller, 1991): because
of this variability, Chrysolampinae were coded as
unknown.

Character 62: Attachment of first instar. External
attachment to the host is known for Perilampidae and
almost all Eucharitinae, and is regarded as plesiomorphic
(state 0). Internal attachment during the early stages of
the first instar followed by external development on the
pupa (state 1) is known only for Orasema and
Pseudonu'tagea.

18

PARSIMONY ANALYSIS

Character state distributions were analysed using the
Phylogenetic Analysis Using Parsimony program (PAUP
Version 3. On; Swofford. 1990) using various addition
sequences and holding different numbers of initial trees.
OTUs consisted of groups of species, and a large number
of taxa had multiple states (i.e., 0 and 1) for certain char-
acters (Appendix 1 ). Of the 62 character states in the data
matrix, 34 of 47 characters of adults, 2 of 1 1 characters of
immatures, and 1 of 2 behavioural characters had multiple
states for at least 1 OTU. The following results are based
on having multistate characters treated as uncertain (steps
not added at terminal nodes), and having character-state
transformations analysed using the delayed transforma-
tion algorithm (DELTRAN). The DELTRAN algorithm
makes the fewest assumptions of character state change;
derived states are therefore not postulated for sister
groups in which a character is unknown (i.e., larval char-
acters 59 and 60 are synapomorphies of Orasema but are
unknown in other genera of Oraseminae). DELTRAN
also delays changes of states for multistate taxa to the ter-
minal OTU, so fewer models or assumptions of character
state change are proposed at interior nodes.

Analyses were carried out using 2 subsets of the data
matrix presented in Table 2 and Appendix 1. These were
chosen to differentiate between characters of adults and
morphology of immatures plus behaviour. All characters
were treated as unordered except for characters 4 and 29,
which were ordered to provide additional resolution with-
in the Psilocharitini (see discussion for each character).
The ordering of binary characters has no effect on tree
topology or number of steps (always a one-step difference
between states). Unordered multistate characters have no
a priori assumption of transformation (one-step differ-
ence from all states), whereas ordering such characters
requires an ad hoc hypothesis of transformation (i.e.. 0 — >
1 -^ 2 requires 2 steps between states 0 and 2).
Unordered multistate characters support only those taxa
that have the same character state and not taxa that share
a different character state; therefore, the order of transfor-
mation is determined largely by other characters. This can
introduce a source of instability into the data set. but I
believe that unordered characters arc the preferred first
step in an analysis. Ordering of the number of antennal
segments (character 4) is a logical progression (i.e.. 7 <-^
8 <-^ 9 segments). Ordering of character 29 by a character
state tree was done to remove additional hypotheses of a
transition series between the 2 derived states. For both
characters, ordering had minimal effects on the analysis,
as discussed below.

In the first data matrix, characters of adults plus egg
shape (characters l^K. Table 2) were analysed (charac-
ters with least amount of missing data), and a strict con-
sensus solution lor the resulting IS trees is shown in

Figure 1 (Tree length = 156: CI = 0.51; RI = 0.79). In the
second data matrix, characters of adults and immatures,
as well as behavioural information (characters 1-62,
Table 2). were analysed, and a strict consensus solution
for the resulting 20 trees is shown in Figure 2 (Tree
length = 183; CI = 0.53; RI = 0.79). Ordering character
29 did not affect tree length. Ordering character 4 added
one extra step for the transition in Eucharitinae from 7 to
9 funicular segments. No shorter trees or trees of different
topology were found using these different approaches.

The consensus solution from matrix 2 (Fig. 2) had a
higher resolution of relationships within Eucharitinae by
including larval and behavioural characters than by using
characters of adults alone (matrix 1, Fig. 1). The group of
taxa classified as Oraseminae was resolved in both data
sets. Larval stages are unknown in Gollumiello and
Anorasema, and the increased resolution of this group
with the Eucharitini is due to the inclusion of character
61, which refers to the increased number of eggs deposit-
ed by Gollumiella. Because relationships proposed for the
complete data set (matrix 2) provided the strongest evi-
dence of group relationships, and are based on the mini-
mum number of a priori hypotheses concerning character
polarity, character state distributions were analysed only
for the results of this analysis.

Of the 62 characters, 15 had a unit consistency index
of 1.0 and were informative (unit retention index = 1.0)
on all trees (characters 11, 13. 19. 21-22. 24, 26, 29,
38-39, 41, 51, 58, 60-61; Table 2). Eight characters were
treated as autapomorphies (unit retention index = 0) for
either terminal taxa or Eucharitidae on all trees (14. 16.
35. 50. 53-55, 57; Table 2), and were therefore not infor-
mative for determining relationships within Eucharitidae.
Twenty-nine characters had a CI of less than 1 .0, but had
a uniform model of character state change on all trees.
Characters with alternate models of character state change
on different tree topologies are marked by narrow bars on
Figure 2. Characters 12, 20, 27. 42. and 59 can differ by
one step on different tree topologies and had 2 different
CI values that were tree dependent (Table 2).

DISCUSSION OF PHYLOGENY

The phylogeny presented in Figure 2 represents the most
parsimonious solution for the distribution of character
states within the Eucharitidae using the simplest assump-
tions of character state ordering. A character analysis
using only characters of adults does not resolve basal
relationships among Eucharitinae (Fig. 1). The additional
information provided by larval and behavioural characters
is necessary to give increased resolution (Fig. 2). Ant-
host data were not inchidocl in Ihc analysis but sublanii-
lies of ant hosts show a strong correlation w ith the pin -
logeny presented here (sec Table 4). I recognize 3 groups

of Eucharitidac based on the character analysis as pre-
sented in Figures I and 2: the Oraseniinae is recognized
as a monoph) letic group that includes Indosenw.
Orasema, Orasemorpha, and Timioderus; the
Psilocharitini includes the genera Ncoloshonus and
Psilachmis; and the Eucharitini includes the genera
Anorasema, GoUumiella, and the Eucharitinae s.s. (after
Graham, 1969: Riek. 1970: and Heraty. 1985), following
the classification of Eucharitinae proposed by Boucek
(1988). A formal classification of 2 tribes within
Eucharitinae is proposed here to distinguish members of
the subfamily with an independent or fused prepectus.

Alternate hypotheses of relationships were examined
by changing the topology of basal taxa and observing
changes in the number of steps and relative indices. Each
alternate hypothesis resulted in longer trees and slightly
lower CI and RI values. Placement of either the
Timioderus + Indosema lineage or Orasemorpha as
ancestral to other Eucharitidae fixes the expanded and
strongly ridged ovipositor as a groundplan state and
requires at least 2 reversals to an acicular ovipositor.
Placing Psilochahs as ancestral, with Oraseminae and
Neoloshanus as monophyletic, increased the number of
steps to 187. Members of Psilocharis are phenetically
very similar to some Chrysolampinae, but otherwise there
is no support for the hypothesis that they represent the
basal lineage of Eucharitidae. Perhaps the most interest-
ing change was the placement of Anorasema as sister
group to Psilocharitini + {GoUumiella + Eucharitinae
5.S.). This resulted in a tree length of 184 which is only
one step longer than the most parsimonious tree.
Members of Anorasema are phenetically very similar to
species in the Orasema uichancoi-group and both share
an increased number of male funicular segments, elongate
and pilose forewing, and smooth face. All of the alternate
hypotheses above involve changes in taxa where larval
stages are unknown. I believe that the most parsimonious
solution presented in Figure 2 is the most accurate por-
trayal of relationships among major lineages of
Eucharitidae based on the information available, and
forms a sound basis for derivation of a new classification.

The following discussion is restricted to generic level
relationships based on the hypothesis presented in Figure
2. Relationships within genera are discussed within the
body of each revision.

Eucharitidae

The monophyly of the Eucharitidae is supported by 5
aulapomorphies: mandibles falcate (character 13), labrum
with marginal digits arranged in the same plane (14),
labral digits initially reduced in number to 4 or 5 (15),
and pronotum not visible from above (21). The mandibu-
lar and labral character states are lost or modified in at
least a few Eucharitidae (i.e., Timioderus, Indosema, and

P.sciulometaf^ea). An additional autapomorphy for
Eucharitidae not coded in the analysis is the absence of a
malar sulcus or suture, which is found in both states in the
oulgroup and is present in many Pteromalidae. The
reduced number of labral digits is interpreted in the analy-
sis as a groundplan character for Eucharitidae. with a
4-digitate labrum found in the majority of Orasema,
Psilocharitini, GoUumiella + Anorasema. and
Pseudometasiea within the Eucharitini. However, the
majority of species within the Eucharitini possess a multi-
digitate labrum (8-16 digits). Seven funicular segments
for females (5) and a setose speculum (37) are indicated
as groundplan states for Eucharitidae (Table 2), but both
character states probably are plesiomorphic for
Eucharitidae -t- Perilampidae.

Character states of eggs and immature stages provide
the strongest support for monophyly of the family. Heraty
and Darling (1984) proposed stalked eggs (48) as an
autapomorphy of Eucharitidae. A single character of the
first-instar larva that supports the monophyly of
Eucharitidae is loss of dorsal setae on tergites VII and IX
(58). Larval characters 53-58 were previously shown to
support monophyly of the Perilampinae and Eucharitidae
(Heraty and Darling, 1984). This analysis suggests that
presence of cranial setae (56) (found in some Orasema)
and absence of a labial plate (58) are groundplan states of
Eucharitidae. Within Orasema, a labial plate is known
only in Orasema tolteca Mann, which suggests that it
may be plesiomorphic for Eucharitidae (Heraty, 1990).
Heraty and Darling (1984) also proposed that loss of a
single spiracle in the first instar (present only in
Perilampinae and Philomidinae) was a synapomorphy of
Eucharitidae. In addition, ant parasitism supports mono-
phyly of Eucharitidae s.s.

Oraseminae

The Oraseminae is a monophyletic group that includes
Orasema, Orasemorpha, Indosema, and Timioderus.
Monophyly is supported by 5 synapomorphies: scape of
males without pores (3), mesepistemum broadly rounded
(32), ovipositor expanded (42). first valvula with lateral
teeth (43), and second valvula with lateral teeth (44).
Additional characters supporting this clade are the anteri-
or curvature of the ovipositor and perhaps the habit of
depositing eggs parallel to the leaf surface (Fig. 3), a
habit correlated with ovipositor shape. Absence of pores
from the scape of males may be plesiomorphic for
Eucharitidae if the fine pores are independently derived in
Psilocharitini and GoUumiella. The presence of serrate
and lamellate (versus cylindrical) flagellar segments (7)
in Timioderus and the presence of an anellus (1) in
GoUumiella. Anorasema. and Psilocharitini negates the
use of these character states for diagnosing the subfamily
(sensu Heraty, 1985). Immature stages and behaviour

20

may offer 4 additional ciiaracter states to support the
monophyly of Oraseminae: loss of a ventral spine on ter-
gite III (49), larva pustulate (59), pupa pustulate over
petiole (60), and internal parasitism of the host in the first
instar (62). However, the immature stages of
Orasemorpha, Indosema, and Timioderus are unknown
and these character states can be attributed only to
Orasema. The Myrmicinae are the ant hosts (60) for
Orasemorpha and the main hosts for Orasema (Table 3;
see also Table 4); they are predicted to be the hosts for
Timioderus and Indosema.

If larval characters are excluded as possible synapo-
morphies of Oraseminae, then the only support for mono-
phyly of Orasema is that the base of the petiole is trun-
cate and that it has a dorsal flange (39). The monophyly
of the Orasemorpha clade is supported by the presence of
a short, transverse petiole in females (38) and an anterior
projection of Ms^ in males (41). The unfused petiole of
Timioderus is either a reversal or a parallel development
with some Chrysolampinae; however, the basic structure
of the petiole is the same in all 3 genera within that clade,
and is unique for Eucharitidae. These results support both
cylindrical eggs and a ventrally separated petiole as
derived states.

Eucharitinae

The limits of the Eucharitinae are expanded to include
Neoloshanus and Psilocharis. Previous definitions of the
Eucharitinae were limited to the Eucharitinae s.s., which
was based on complete fusion of the prepectus (19),
absence of an anellus ( 1 ), variously modified antennal fla-
gellomeres (7), and usually an acicular ovipositor (42)
(Burks, 1979; Heraty, 1985). Boucek (1988) included
Gollumiella (as Loshanus) and Anorasema in the
Eucharitinae and showed that both had an anellifomi Fl
(1) and 13-segmented antenna. By including
Psilocharitini with the Eucharitinae s.s., monophyly of the
subfamily is supported by the presence of an occipital
carina (18), loss of a constriction from the first gastral
sternite (37), elongate and tapered petiole basally (39),
and loss of the constriction from Ms, (40). The occipital
carina is lost in some Eucharitinae, and the base of the
petiole changes in Psilocharis. A weak basal constriction
within the Eucharitinae is known only for Anorasema.
Characters of the first-instar larva that support monophyly
of the Psilocharitini + Eucharitini are fusion of tergites I
and II (51), presence of a distinct tergopleural line (52),
absence of cranial setae (56). and presence of a labial
plate (58). Character 52 undergoes a reversal in the
Oheza-cVdAc, and the labial plate may be a groundplan
slate for Eucharitidae. The Chakura type of third-instar
larva (59) is treated as a synapomorphy of Eucharitinae
based on its similar occurrence in both Chahura and
Neoloshanus, but it undergoes major changes in the rest

of the Eucharitinae. Based on a shared ponerine ant host
for Neoloshanus and other Eucharitinae. this host subfam-
ily could also be treated as a synapomorphy at this level.

Monophyly of Psilocharitini is supported by the fol-
lowing: base of flagellomeres with secondary segmenta-
tion (6), genal depression distinct (10), prepectus nar-
rowed ventrally and foveate (20), propodeum smooth and
polished (28), postmarginal vein elongate (36), speculum
present (37), and callus polished and slightly swollen.
Character 10 is lost in derived species oi Neoloshanus; 36
and 37 undergo reversals in single species of
Neoloshanus and Psilocharis, respectively, and also show
homoplasy within Orasema. Larval characters are
unknown for Psilocharis, but if similar to Neoloshanus,
then they would be plesiomorphic for Eucharitinae and
offer no supporting characters at this level. One possible
larval synapomorphy is ventral fusion of the cranial
extensions (VFM, Fig. 13), which is discussed in the fol-
lowing section on biology. Within Eucharitinae,
Psilocharitini is distinguished by the independent prepec-
tus and the presence of an anellus in almost all species;
however, these are plesiomorphic features at this level.

Monophyly of Psilocharis is supported by the follow-
ing: scrobal depression with smooth channels (9), femoral
groove broadly impressed (29), and petiole truncate baso-
laterally (39). The latter 2 characters are unique attributes
of Psilocharis. The petiole is always elongate and strong-
ly ribbed and the lateral margins at the base can be flared
laterally. Scrobal channels are also found in Orasema
glahra and are lost in some species of Psilocharis. The
genus Neoloshanus is supported by absence of a distinct
anteclypeus (11), presence of a narrow and sinuate
femoral groove (29), and fusion of the gonostylus to the
second valvifer (46). Characters 1 1 and 29 are unique
attributes of Neoloshanus whereas character 46 is shared
with Eucharitini.

Monophyly of the Eucharitini is supported by the fol-
lowing: absence of pores from the scape of males (3),
dorsal enclosure of the mesothoracic spiracle (26), and
fusion of the gonostylus to the second valvifer (46).
Again, if pores on the male scape are derived states for
the Psilocharitini and 1 species of Gollumiella, then
absence would be a plesiomorphic character state. Fusion
of the prepectus (19) is treated as 2 character states that
support the different lineages within Eucharitini. If these
groups are indeed monophyletic, then tusion of the
prepectus would be a synapomorphy of the Eucharitini
(Heraty, 1992). There are no larval characters supporting
this clade other than an increase in the number of eggs
laid (61). Other members of the Eucharitini usually
deposit their eggs in clusters of 60 to 10 000 eggs, and
only Schizaspidia cnuvnnaia is known to insert only 3 to 4
eggs into oviposition punctures, as do Oraseminae and
Psilocharitini. Single or few eggs deposited in punctures

21

Table 3. List of genera and species of Old World Orascminac and Psilocharitini with known host associations

Taxa

Country Ant Host*

Plant Host (Family)**

Immature

Stages

Known

Reference

Orasemorpha
O. eiihotes
O. myrmicae
O. thdentaia
O. xeniades

Australia
Australia
Australia
Australia

Plieidole sp.
Pheidole sp.
Pheidole proxima
Pheidole
tasmaniensis

Boucek. 1988
PS [BMNH]
Brues, 1934
PS [BMNH]

Orasema

0 . fraudulenta

Africa

Pheidole
megacephala

0. uichancoi

Philippines

Celtis (Fagaceae)
Leucaena (Leguminoseae)

0. sp.

Solomon

Pheidole sp. (W)

(nr rugulosa)

Islands

0. assectator

India

Pheidole sp.

Ceanothus (Theaceae)

0. initiator

India

Ceanothus

0. valgius

Australia

Pheidole sp. (W)

Neolosbanus

N. gemma

Australia

Hypoponera sp. (S)

N. palgravei

Australia

Hypoponera sp. (S)

FUndersia (Rutaceae)

third, pupa Reichensperger, 1913

egg, planidium Ishii, 1932

pupa PS [USNM]

egg to pupa Das. 1963; Kerrich, 1963

Kerrich. 1963
Girault, 1913b

N. laeviceps

Malaysia
Sri Lanka

pupa

PS [JMH]

FUndersia (Rutaceae)

egg to pupa

PS [JMH. TAMU]

Tetrasynandra (Monimaceae)

PS [JMH]

Macaranga (Euphorbiaceae)

PS [JMH]

Lygodium (Schizaeaceae)

PS [JMH]

Rutaceae

egg, planidium

PS [JMH]

Schizaeaceae

PS [JMH]

Artocarpus (Moraceae)

planidium

Clausen, 1940b, 1940c

* Ant determinations by S (= S. Shattuck, ANIC) and W (= E. O. Wilson, MCZ); PS = Present Study.
** Eggs were deposited into leaves for all known plant associations.

may also occur in Anorasema and Chalcura montana,
which also have an expanded ovipositor.

Monophyly of Anorasema and Gollumiella was dis-
cussed in detail by Heraty (1992). Monophyly of the
Eucharitini s.s. (excluding Anorasema and Gollumiella) is
supported by the following: loss of anellus (1), increased
number of labral digits (usually 9 to 16; character 15),
fusion of the prepectus in same plane (19). transverse
axilla (22), presence of internal mesoscutal ridge (25),
and wedge-shaped mesepisternum (32). Character 32
undergoes considerable change within this clade. No
characters of immatures support this clade. The majority
of species deposit their eggs in large numbers into spaces
within preformed plant cavities (flower bracts, under bud
scales, etc.; see Clausen, 1940a, 1940b, 1940c), which
represents a major shift in ovipositional strategies within
the family. Relationships of genera within this clade, as
presented in Figures 1 and 2, are tentative and represent
only an initial hypothesis of relationships. However, the
relationships proposed for Eucharitinae are congruent
with ant hosts, even though ant hosts were not included
within the character matrix used to build the tree.

CONCLUSIONS ON PHYLOGENY

Characters of adults are highly variable within
Eucharitidae and new information based on the taxa
described here creates problems in defining Eucharitidae
at both the family and the subfamily level. More conserv-
ative characters such as larval morphology and behav-
ioural information provided additional resolution neces-
sary to examine relationships among Eucharitidae. The
Oraseminae has been redefined and forms a monophyletic
group containing many of the species of Oraseminae as
defined by Boucek (1988). Two groups are supported
within the Oraseminae: the Orasemorpha clade and
Orasema. The limits of the Eucharitinae are extended to
include the genera Neoloshanus and Psilocharis within
the tribe Psilocharitini. All Eucharitinae with a fused
prepectus are included in the Eucharitini. Larval and
behavioural characters are congruent with the cladogram
constructed using characters of the adults alone, and offer
more unique and unreversed (non-homoplastic) characters
to support the proposed relationships.

Biology

Information on the first-instar larva, other immature
stages in the ant nest, oviposition habits, and ant host is
available for 18 of the 40 known genera of Eucharitidae.
In some groups such as Orasema. Stilhula, and
Schizaspidia, there is considerable information on every
life stage and often for more than 1 species, allowing for
some generalizations on life history. Morphology of the
immature stages, and in particular the sclerotized planidi-
um, is conservative at the generic level and above, and
can be useful in positing relationships at higher taxonom-
ic levels (Heraty and Darling, 1984). Along with informa-
tion on ant hosts and oviposition strategies, inclusion of
such characters within a phylogenetic analysis can serve
to test, supplement, and refine hypotheses derived from
studies of adult morphology. In this section, biologies of
the known members of Oraseminae and Eucharitinae are
treated on a comparative basis. Trends in the evolution of
morphological characters and some behavioural traits
were considered in the earlier section on phylogcny.

All Eucharitidae .v.,v. are parasites of ants. Parasitism is
indirect in thai oviposition occurs in plant tissue away
from the host (Clausen, 194()b. I94()c, 1941). Eggs may
be laid singly or in clusters of up to several thousand
(Clausen. 1940b, 1941; Johnson ct al.. 1986). Upon
hatching, the first-instar larvae, termed planidia. actively
seek out adult ants or Ihrips ( Thysanoptcra) lor transport

to the ant nest (Clausen, 1940b. 1941; Wilson and
Cooley, 1972; Johnson et al., 1986). The planidium trans-
fers to an ant larva where it remains as a first instar until
the ant pupates (Clausen, 1940b, 1941). In Orasema, the
planidium burrows just under the larval cuticle and later
becomes external when the ant pupates (Wheeler. 1907).
In most Eucharitinae, the planidium is always external
(Clausen, 1941c). Pupation and emergence take place
within the ant nest. Adults leaving the nest are ready to
mate and lay eggs.

The function and morphology of the ovipositor suggest
a major dichotomy in the Eucharitidae. Five genera of
Oraseminae and Psilocharitini, plus a few species of
Schizaspidia (Eucharitinae). have an expanded ovipositor.
The structure of this ovipositor is unique within the
Chalcidoidea. and in Orasema and Schizaspidia the
ovipositor is used as a tool to hollow out a chamber in
leaf tissue where 1. or rarely up to 4. eggs are deposited
(Clausen. 1940b; Johnson et al.. 1986). This is in sharp
contrast to the acicular ovipositor generally found in the
Eucharitinae (Heraty. 1985). The acicular ovipositor is
used to pierce plant tissue and deposit a mass of eggs into
pre-existing cavities between bud scales or in undevel-
oped flower buds. Some eucharitines simply scatter eggs
over the leaf surface and Oheza is known to deposit eggs
into the tissue of fruits (Heraty and Barber. 19^)0).

23

Table 3 is a summary of life-history information for
those species of Oraseminae (Old World) and
PsiliKharitini for which the ant host or immature stages
are known. Additional information on habits ol New
World Oiiiscma is found in Heraty (1990) and will be
formally treated in a future publication. Within the limit-
ed list of plant associations, 21 species of Oraseminae are
associated uith 31 plant genera distributed in 22 families
of plants (Heraty. 1990). In the Old World, plant hosts of
Oraseminae are known only for Orasema, and they are
Fagaceae, Leguminoseae, and Theaceae (Table 3).
Psilocharitini have a broad host range and species of
Neoloshamis have been collected on at least 7 families of
plants (Table 3). Ant hosts are more restricted with
records of 8 genera of ants in 4 subfamilies for both
Oraseminae and Psilocharitini (Tables 3 and 4). The fol-
lowing sections present new and previously published
information on the biology of 3 genera of Oraseminae and
Neoloshanus, and provides a review of the biology of the
Eucharitini (Eucharitinae s.s.). Orasema and Neoloshanus
are considered as representative of their respective
groups.

BIOLOGY AND IMMATURE STAGES
OF ORASEMORPHA

Four species of Orasemorpha have been reared from the
ant genus Pheidole (Table 3). The pupa of O. tridentata
was described as having the margins of the abdominal ter-
gites "raised and carinate, the edge of the second tergite
projecting as a distinct, sharp tooth on each side not far
from the median line" (Brues, 1934:205). These struc-
tures may be similar to those described for Orasema, but
no mention was made of pustules along the dorsal margin
of the petiole, a feature known only for Orasema. Pupae
were not recovered along with the type material of O. tri-
dentata (= wheeleri) which is located at the MCZ.

BIOLOGY AND IMMATURE STAGES
OF ORASEMA

The biology and immature stages of the genus Orasema
were described in detail for several species (Wheeler,
1907; Clausen, 1940c, 1941; Parker, 1942; Wheeler and
Wheeler, 1937; Das, 1963; Heraty and Darling, 1984;
Johnson et al., 1986). There is a remarkable consistency
of both oviposition habits and morphology of immature
stages in Orasema. A generalized biology is presented
below. Specific references to differences among species
groups are only briefly discussed.

Oviposition Behaviour

Parker (1942) and Das (1963) described details of adult
oviposition behaviour for Orasema. The ovipositor is
inserted forward into the leaf, and eggs are deposited par-
allel to and just under the surface of the plant tissue (Fig.
4). Females of most species have been observed to
deposit single eggs into punctures. The punctures are
either scattered over the plant surface or placed in short,
straight rows into the stems, the underside of leaves, or
the involucral bracts of various plants (Fig. 3; Parker,
1942; Das, 1963; Johnson et al., 1986). Orasema uichan-
coi (Ishii) was reported to lay eggs in short alternating
double rows on the undersurface of leaves (Ishii, 1932).
In most cases, plant tissue surrounding the length of the
oviposition puncture became brown. Plant tissue was
observed to become scarified around the oviposition
punctures of O. sp. nr aenea on leaves of Muhlenbeckia
(Parker, 1942).

Choice of plants in which species of Orasema oviposit
does not appear to be restricted. Females of Orasema are
known to oviposit in leaves of tea (Theaceae), mango
(Anacardiaceae), and oak (Fagaceae); involucral bracts of
composite flower heads; or even into young banana fin-
gers (Museaceae) (Johnson et al., 1986). In Orasema
viridis Ashmead, females oviposit in undeveloped flower
buds of Haplopappus (Compositae) as well as
Sphaeralcea (Malvaceae) (Johnson et al., 1986). Orasema
tolteca Mann and O. sp. nr aenea have been observed to
oviposit on 4 or 5 different families of plants (Heraty,
1990). Within a species, Orasema is generally consistent
in its choice of plant structure for oviposition. All of the
Old World species are known only to oviposit directly
into leaves (Table 3).

Egg (Fig. 4)

Stalked eggs as typical for Eucharitidae; white with
smooth chorion and short anterior stalk, apical thickening
of stalk indistinct (Parker. 1942; Heraty and Darling,
1984; Johnson et al., 1986). The egg stalk. is usually bent
over the side of the egg within the oviposition chamber
(Fig. 4), not erect as observed for Neoloshanus (Fig. 18).
Mature eggs have a heavily sclerotized (darkened) first-
instar larva occupying almost the entire egg body, with
the head oriented toward the stalk.

First Instar (Figs. 5-9)

The planidium of Orasema differs from those of other
Eucharitidae in that tcrgitcs I and II are separated dorsal-
ly, the mid-ventral seta on tergite 111 is absent, and the
pleural region of tergite IX is separated into a leaflike
ventral plate. A general discussion of characters in rela-
tion to Eucharitinae and the outgroup family Perilampidae
was presented in Heraty and Darling (1984). Heraty and
Darling (1984) and Johnson et al. (1986) reported the

24

absence of a labial plate from Orasema as a loss from the
groundplan state of the Eucharitidae (present in all
Eucharitinae). A weakly sclerotized labial plate was
observed in 1 undescribed species of Orasema from
Mexico, which may offer support for this hypothesis.
Heraty and Darling (1984) suggested that the weak lines
of desclerotization extending from the base of the setae
(Fig. 6) may be a reduced tergopleural line. In Orasema
tolteca, a faint area of desclerotization is apparent along
the tergites in a position homologous to the tergopleural
line in Eucharitinae (Fig. 15) and is distinct from the lines
of weakness associated with the setal bases. An additional
feature found in all species of Orasema examined is the
presence of a broad, ventral extension of the cranium,
which remains separated medially (VFM, Figs. 13-14).

The planidium of Orasema uichancoi, originally
described and illustrated by Ishii (1932). was reexamined
and 3 character states were shown to be incorrectly
described or illustrated: tergites I and II are separated dor-
sally (not fused), labial plate is not apparent (Ishii showed
it as present), and the ventral region of tergite IX is sepa-
rated into a leaflike plate (not shown by Ishii).

The means by which planidia gain access to the ant
nest is unknown, but an intermediate host association
with thrips has been shown for Orasema sp. on
Frankliniella occidentalis (Pergande) (western flower
thrips) (Wilson and Cooley, 1972; specimens examined);
Orasema sp. on Microcephalothrips ahdominalis and
Frankliniella sp. (Beshear, 1974); O. coloradensis Gahan
on Sericothrips sp.; O. viridis Ashmead on immature
thrips (Johnson et al., 1986); and O. assectator on
Empoasca flavescens (F.) (tea leafhopper) or Scirtothrips
dorsalis Hood (tea thrips) (Das, 1963). I found additional
associations with thrips for 5 other species of Orasema.
Planidia remain either lightly attached to the surface of
the thrips abdomen (Fig. 5) or buiTow into the thrips and
feed, without causing distention of the body segments of
the planidia (Wilson and Cooley, 1972; Johnson et al.,
1986). Planidia of Orasema are often found attached to an
immature thrips.

One hypothesis to explain this phoretic behaviour is
that ants capture thrips laden with planidia and carry them
to the nest as food for the brood. Attempts to prove this
direct transfer have not been successful, but under labora-
tory conditions, a worker of Pheidole crassicornis Emery
was observed in the laboratory to pick up a thrips, with
attached planidia. and carry it to the brood pile. Clausen
(1940b) dismissed the possibility of the thrips as an inter-
mediate step to the ant host because no ants were known
to "tend" thrips. but he neglected to consider the possibil-
ity of predation by the ants. There are reliable records of
Wasmannia aiiropuni lata (Roger) preying upon the
cocoa thrips in Trinidad (Ananthakrishnan. 1984). of
Azieca chartifox as a control tor thrips in Brazil

(Ananthakrishnan, 1984), and of Pheidole megacephala
(Fabr.) as a significant predator of Liothrips in Hawaii
(Reimer, 1988). Both Wasmannia and Pheidole are
known hosts of Orasema.

The habits of the early larval stages of Orasema within
the ant nest were described by Wheeler (1907). Wheeler
and Wheeler (1937), and Das (1963). I obtained addition-
al information for Orasema sp. on Pheidole dentata Mayr
and Solenopsis .xyloni x geminata, and for O. .xanthopus
(Cameron) on Solenopsis sp. (? invicta from Matto
Grosso, Brazil; material collected by D. Wojcik and D.
Jouvenaz, Gainesville, Florida, United States; vouchers in
ROM). The habits of the first instar were the same in all
of the species. Unfed planidia were found attached to the
dorsal surface of the second-instar ant larva and burrowed
under the cuticle of the third instar (Figs. 7-9). On
Pheidole dentata, planidia burrowed just behind the
cephalic region (Fig. 8), whereas locations were more
variable for O. .xanthopus. Planidia extracted from the
host larva show an initial distention of the ventral and
unsclerotized regions of the body. Larvae complete a con-
siderable amount of feeding within the host and become
greatly distended (Fig. 8). Upon pupation of the host, lar-
vae become external (Fig. 9) and move to the ventral
region of the thorax of the host, between the deformed
pupal legs (Fig. 10). First-instar larvae show extreme dis-
tention in the final stages with the tergites strongly sepa-
rated.

Second Instar

This instar is more typically hymenopterifonn, white, and
very weakly sclerotized with a single pair of mesotho-
racic spiracles (propneustic). The head region is weakly
delineated, with a small pair of pincerlike mandibles. This
stage is virtually identical to other second-instar larvae
described within the Eucharitidae. The second instar
remains attached to the host in the ventral region of the
host thorax.

Third Instar (Figs. 10-11)

This instar is white and membranous with 2 thoracic and
6 to 7 abdominal spiracles evident. 9 enlarged dorsolater-
al tubercles, a smaller series of lateral tubercles continu-
ing onto the prothoracic segment, a pair of medially
divided tubercles lateral to the oral region, the ventral sur-
face of abdominal segments finely ridged, and the oral
region lacking mandibles and consisting of fine striae
convergent on midline. The structure of the third-instar
larva undergoes a series of changes from the early third
instar (Fig. 10) to the prepupal stage largely based on the
definition of tubercles. Prepupal stages are more promi-
nently tuberculatc. and precursors of the antcnnal seg-
ments are evident along ventrolateral margins.

Third-instar larvae complete development on the host,

25

and then deiach themselves and arc free within the brood
pile. The shrivelled host remains, termed phthisergates
(alter Wheeler l^>()7. 1910). stay within the ant colony.

Pupa (Fiji. 12)

The pupa has 3 enlarged tubercles along the dorsal mar-
gin of petiolar region, the gaster has raised transverse
bands, and lateral and subventral tubercles are often pre-
sent on abdominal ridges. The gaster of males is smaller
than that of females. The structure of the pupa was seen
to be consistent across broad groups within Orasema
based on data from O. fraudulcnta {srriatosoma-group:
Reichensperger. 1913), Orasema assectator (asscctator-
group), and Orasema sp. nr rugulosa (uichancoi-group)
(Table 3). The structure of the pupae of these species sup-
ports the enlarged tubercles over the petiole as a unique
character of Orasema. Diagnostic characters for species
of Orasema are found in the shape and prominence of
tubercles on the gastral ridges.

Pupation and emergence of adults take place within
the ant nest. Adults of Pheidole and Solenopsis were
observed tending and carrying the adults and pupae of
Orasema within the nest. Workers were observed trans-
porting pupae by the dorsal tubercles over the petiole.
Reichensperger (1913) suggested that the tubercles of lar-
vae and pupae might be secretory and act in ant appease-
ment. Vander Meer, Jouvenaz, and Wojcik (1989) found
that pupae and adults of Orasema sp. collected in the nest
of Solenopsis shared the same cuticular hydrocarbon pro-
file of the host brood. The authors proposed that Orasema
may be utilizing colony odour or producing a mimicking
compound, but they downplay the latter hypothesis. More
work needs to be carried out, possibly with a focus on the
tubercles of the third-instar larva and the pupa of
Orasema.

BIOLOGY AND IMMATURE STAGES
OF NEOLOSBANUS

Habits for Neoloshanus are based largely on data collect-
ed by me from 2 populations of A', palgravei observed in
the undergrowth of rainforests in north Queensland,
Australia, and Selangor, Malaysia. The within-nest larval
stages were collected from Hypoponera nesting in rotten
wood at the Australian location. Supplemental informa-
tion was provided by published accounts for N. laeviceps
(Clausen, 1940b, 194()c, 1941), a pupa of A', gemma dis-
sected from a cocoon of Hypoponera. and museum
records of plant hosts.

Oviposition Behaviour

Adults of N. palgravei deposited eggs singly into the
underside of broad-leaf plants belonging to the plant fam-
ilies Rutaceae, Monimiaceae, Euphorbiaccae, and

Lygocilnm (Schizaeaceae). Eggs were deposited at
approximately a 4.5-degree angle to the leaf surface, with
the stalk of the egg erect and slightly protruding from the
opening (Fig. 18). The plant tissue showed no reaction to
freshly deposited eggs (Fig. 18), but became scarified and
swollen around the opening as eggs matured (Fig. 19).
Oviposition punctures were either scattered over the
underside of the leaf (both locations) or deposited alter-
nately in short double rows of about 20 eggs (Malaysian
only). The density of oviposition was extremely high at
both sites and the entire underside of new growth was sat-
urated with punctures. Plants recovered from the wounds,
and older leaves showed decreasing levels of scarifica-
tion. Neoloshanus laeviceps deposited eggs singly into
leaves of Artocarpus (Moraceae) (Clausen. 1940b).
Clausen illustrated a puncture of A', laeviceps with 3 eggs,
but he stated in the text that eggs were deposited singly.
Neoloshanus townesi was collected from leaves of
Castanopsis (Fagaceae) and bamboo (Poaceae) in Papua
New Guinea (Table 3). There are no oviposition records
for species in the N. gemma- or N . purpureoventrls-
groups, but N. gemma was collected from flowers and I
collected adults of A^. purpureoventrls on broad-leaf
understorey plants in Taiwan and Thailand.

Egg (Figs. 17-19)

Undeveloped eggs of A^. palgravei are white with a
smooth chorion. Mean length of egg body (Australian
population) is 0.14 mm (SD = 0.01. n = 6); mean length
of anterior stalk is 0.07 mm (SD = 0.01). The anterior
stalk has a thickened apical extension with a mean length
of 0.07 mm (SD = 0.01). Mature eggs have a lightly scle-
rotized first-instar larva occupying almost the entire egg
body, with the larval head oriented towards the stalk (Fig.
17). Eggs of N. purpureoventrls are the same, and those
of N. laeviceps are similar but lack an apical extension
(Clausen, 1940b). These eggs are similar to those of other
Eucharitidae (Clausen, 1940c: Heraty and Darling, 1984).

First Instar (Figs. 13-14, 20)

Planidia of A', palgravei are similar to those described for
Eucharitinae (Heraty and Darling. 1984) and possess the
following characteristic states: distinct labial plate: dorsal
fusion of tergites I and II: distinct tergopleural line sepa-
rating pleural region on tergites Il-VllI: and 2 ventral
setae on tergite III. Tergite IX shows a partial invagina-
tion of the ventrolateral region of the tergite that is similar
to, but less pronounced than, that in Orasema. Unique
character states for Neoloshanus include a relatively
broad ventral margin of the cranium, which is fused
medially (Figs. 13-14). and tergites lightly sclerotized
and pale brown in colour. The terminal segments of the
planidium of A', laeviceps (illustrated in Clausen, 1940c)
are similar to the Malaysian A', palgravei but have the ter-

26

gopleural line continuing on tergite IX.

Planidia of N. palgravei collected in Malaysia showed
strong differences from the Australian larvae in the fol-
lowing states: cranium dorsally with raised median area
(Fig. 14); seta present at lateral fusion of tergites I and II;
ventral margins of tergites IV-IX strongly sculpted along
posterior margin; caudal cerci short; body length signifi-
cantly shorter (Malaysian, 0.12 ± 0.01 [SD] mm, n = 10;
Australian, 0.14 ± 0.01 mm, n = 10; Student's t, p <
0.01). The Australian population possesses a larger seta
with tuberculate base on tergite VI (Fig. 13), which is
unknown for other Eucharitidae. Additional collections of
larvae may reveal correlations with the minor differences
in adults of N. palgravei such that separate species may
be warranted.

First-instar larvae of A', palgravei move by a looping
motion or can jump for a distance of about 10 mm as
described for other Eucharitidae by Clausen (1940c). No
intermediate host associations were observed on the
plants, and host ants were observed only on the ground.
The high density of eggs and the jumping habits of the
larvae suggest that they "rain" down upon the forest litter.
Within the nest of Hypoponera several partially fed first-
instar larvae were found attached on either side of the
head capsule of mature host larvae (Fig. 20), and on 1
larva within the host cocoon.

Second Instar (Figs. 21-23)

This instar is more typically hymenopteriform, white, and
very weakly sclerotized; the spiracles and mouthparts are
not discernible (Fig. 21). The first-instar exuvium remains
attached to ventral surface. Several second-instar larvae
( 1 per host) were found attached to mature prepupae of
Hypoponera within the host cocoon (Fig. 22). Only 1 sec-
ond instar was attached to a deformed pupa with the
pupal legs exposed (Fig. 23), as is more typical for other
known Eucharitidae.

Third Instar (Fig. 24)

This instar is white and poorly sclerotized, with spiracles
and body segmentation not apparent, entire dorsal surface
minutely tuberculate, and exuvium of first instar not
attached. The shape of the third instar is similar to those
of Chalcura, Schizaspidia, and Stilhiila (Clausen, 1923,
1928, 194()c), and oi Pseudochakitra (Heraty and Barber,
1990).

Pupa (Fig. 25)

The pupa of Neoloshaims palgravei is typically chalci-
doidlike except for raised ridges along the metasomai ter-
gites, as in other Eucharitidae. The cocoon of
Hypoponera was open at the caudal end, and remains of
the host were not found with the pupa. A single female
pupa of A', gemma was collected in 1 of the colonies of

Hypoponera, which was also parasitized by N. palgravei
(Fig. 26). The mature pupa was similar to N. palgravei,
except for characters that distinguish the adults of the 2
species (especially the clypeus and profile of the mesoso-
ma), and to pupae of Eucharitinae.

BIOLOGY AND IMMATURE STAGES OF
EUCHARITINI

The oviposition habits of several species were reviewed
by Clausen (1940a, 1940b, 1940c, 1941) and the ant-host
records were reviewed by Wheeler and Wheeler (1937)
and Johnson (1988). Significantly less information has
been published on the immature stages within the ant
colony, and what little there is, is based on the following
species: Austeucharis fasciiventris (Brues) (Brues, 1919),
Chalcura deprivata (Walker) (Clausen, 1940c), Eucharis
esakii Ishii (= scutellaris Gahan) (Clausen. 1940c),
Kapala terminalis (Clausen, 1940c), Pseudochalcura gih-
hosa (Provancher) (Wheeler, 1907; Heraty and Barber,
1990), Pseudometagea schwarzii (Ashmead) (Ayre,
1962), Schizaspidia convergens (Walker) (Clausen.
1940c), Stilhula cyniformis Rossi (Parker, 1937), and
Stilhula tenuicornis (Ashmead) (Clausen, 1923).

Oviposition Behaviour

Adults of Eucharitinae usually deposit large numbers of
eggs into preformed cavities in plant tissue (Clausen.
1940a, 1940b, 1940c. 1941; Heraty and Darling, 1984). In
a few cases, eggs are scattered on the undersurface of a
leaf (e.g., Kapala terminalis; Clausen, 1940b), are placed
in an erect position on a leaf surface in association with
thrips eggs (Gollumiella antennata (Gahan); Clausen,
1940b, 1941), or are deposited into fruit {Obeza floridana
(Ashmead); Heraty and Barber, 1990). In Schizaspidia
antennata Gahan and Schizaspidia nasua (Walker) (=
Kapala foveatella Girault), 1 to 4 eggs are deposited into
oviposition punctures in the leaf tissue (Clausen. 1940b;
Ishii, 1932). The majority of Eucharitinae, including the
43 described species of Schizaspidia, have an acicular
ovipositor, which is used for penetrating plant tissue. In
the above 2 species of Schizaspidia. the ovipositor is sub-
apically expanded and strongly ridged, as in Orasema. In
Anorasema and Chalcura montana (Girault). the oviposi-
tor is expanded along its length, but smooth.

Egg

The eggs are stalked (as is typical for Eucharitidae) and
white and have a smooth chorion; the anterior stalk some-
times has terminal thickening (Heraty and Darling, 1984).

First Instar (Figs. 15-16)

The larva of Austeucharis impte.ui (Walker) (new infor-
mation on larva removed from body of holotype; Fig. \5)

27

is tspical tor nu>st species in (he sublamilN as described
in Heraiy and Darling (1984). Relevant character stales
are discussed under Neoloshanus. The planidia of Oheza
floridana (Fig. 16). Siilhula. and Psciidocluilciira are
atypical for the subfamily; they lack the tergopleural line
and ha\e only 1 ventral seta on lergite 111. These features
appear to be characteristic of a group of genera within the
Eucharitinae (Heraty and Barber. 1990).

Generally, the planidia of Eucharitinae gain entrance
to the nest by various means of phoretic attachment to
worker ants, independent of any intermediate insect carri-
er (Clausen. 1923. 1940b. 1941; Ayre. 1962; Heraty and
Barber. 1990). One case is known involving an obligatory
thrips association. Clausen (1940a. 1940b, 1941)
described females of Gollumiella anrennata (Gahan)
depositing their eggs "vertically, regularly spaced and in
numbers of up to 100, in the immediate vicinity of a
freshly deposited thrips egg." Oviposition could be
induced only in the presence of the thrips egg. The eggs
all hatch simultaneously and the planidia attach them-
selves to the young thrips. Clausen was not successful in
determining the relationship between the thrips and an ant
host for G. antennata.

Upon entering the host colony, the planidium remains
as an external parasite of the host larvae in Chalcura
deprivata (?)*, Eucharis scutellaris (?). Kapala termi-
nalis, Pseudochalcura gihbosa, Schizaspidia convergens
(?), Stilhula cyniformis, and Stilbula tenuicornis
(Clausen, 1923, 1941; Parker, 1937; Heraty and Barber,
1990). Pseudometagea schwarzii is the only species of
Eucharitinae known to burrow into the larval cuticle,
where it overwinters with its host (Ayre, 1962; verified in
my collections from Windsor, Ontario, Canada).

Second Instar

This is similar to the second instar larvae described for
Orasema (Clausen, 1940c, Heraty and Barber, 1990).

Third In.star

This instar is typically hymenopteriform, lacking pro-
nounced ridges or protuberances (Clausen, 1923, 1940c;
Heraty and Barber, 1990).

Pupa

The pupa is typically chalcidoidlike. In Austeucharis,
Chalcura, Schizaspidia. Stilhuku and Kapala, the pupae
can have marked vesicular swellings on the head and
mesosoma. but pronounced ridges or protuberances over
the petiolar region are lacking (Wheeler. 1907; Clausen,
1940c; Ayre, 1962; Heraty and Barber, 1990).

* Question marks in parentheses refer to my difficulty in inter-
preting Clausen's comments regarding actual observations on
the larvae of these genera.

HOST-ANT RKLATIONSHIPS
OF KLCHARITIDAK

There is little information that can be gained for assessing
host relationships within Eucharitidae by looking at taxa
that are considered to be closely related to Eucharitidae.
Chrysolampinae are primary parasiioids of Curculionidae
and Nitidulidae (Darling. 1986). Perilampinae parasitize a
wide variety of hosts including Curculionidae
(Platypodinae) by Monacon Waterston (Boucek. 1978),
Anobiidae by Steffanolampus Peck (Boucek. 1978),
Eumeninae (Vespidae) by Kromheinius Boucek (Darling,
1988b). Ichneumonidae by Euperilampus Walker
(Darling. 1983a, 1983b), and Chrysopidae. Tachinidae,
and Hymenoptera (Diprionidae and Ichneumonoidea) by
Perilampiis Latreille (Darling. 1983a). Echthrodape and
Philomides are both parasitoids of bees. Only
Eucharitidae s.s. are parasitoids of ants. Chrysolampinae
exhibit a similar method of egg deposition to New World
Orasema by depositing eggs into seed pods, at random
with respect to presence or absence of the host (Darling
and Miller, 1991). but otherwise there are few similarities
with other taxa on methods of gaining access to the host.

Information on the ant hosts of Eucharitidae is summa-
rized in Table 4. Some genera (Orasema and Eucharis)
show a very broad host range, with species found on sev-
eral different genera, or even on different subfamilies of
ants. In other genera (Austeucharis, Chalcura, and
Tricoryna), the hosts are restricted to 1 genus of ants.
Orasema have been reared most often from myrmicine
ants belonging to the genus Pheidole Westwood. with
scattered records in the New World from Solenopsis
Westwood, Wasmaimia Roger, and Tetramorium Mayr
(Wheeler and Wheeler, 1937; Boucek. 1988). Two
notable exceptions are Orasema coloradensis Ashmead,
which was found associated with Formica (Formicinae)
(Johnson et al.. 1986), and a museum record of Orasema
rapo Walker, which was reared from the army ant Eciton
quadriglume (Haliday) (Ecitoninae) (Heraty, 1990). In
the Old World, Orasema and Orasemorpha are known
only from Pheidole.

In the Psilocharitini. 2 species of Neoloshanus have
been reared from the ponerine ant genus Hypoponera.
Hosts of Eucharitini are generally restricted to the
Ponerinae and Formicinae (Table 4). with a few notable
exceptions. The Australian genus Austeucharis has been
reared from Myrmecia Fabricius (Myrmeciinae) (Brues.
1919). Eucharis adscendens Fabricius and Psilogastrellus
punctatus (Foerster) were reared from Messor harharus
Linnaeus (Ponerinae) in Europe (Fahringer and Tolg,
1912); other members of both genera, which are sister
taxa, were reared from Formica Linnc and Cataglyphis
Foerster (Formicinae). Species belonging to 2 closely
related genera of eucharitines, Kapala and Galearia
Brulle, ha\c been associated with nests of Pogonomyrmex

28

Table 4. Ant hosts of the Eucharitidae. Genera of Eucharitidae are listed in a hierarchy based on the cladogram
presented in Figure 2. Psilogastielliis was not included in the cladistic analysis, but is the sister group of
Euchahs. Rearings from host-ant genera presented in parentheses do not belong to the same ant subfamily as the
most common records (presented in last column). Ant genera in which subfamilies are not listed are Eciton
(Ecitoninae), Formica (Formicinae), and Messur (Ponerinae). Numbers in parentheses refer to literature listed
below the table.

ORASEMINAE

Orasema

Orasemorpha

Pheichle {4-5. 11-13, \6).Solenopsis (\3-\6),

TetniDioriiim ( 16), Wasmannia (13), {Formica, 10; Eciton, 16) Myrmicinae

Plieiclole {\3, \6) Myrmicinae

EUCHARITINAE
Psilocharitini

Neoloshanus
Eucharitini

Chalciira

Austeucharis

Schizaspidia

Kapala

Isomerala

Tricoryna

Propsilogaster

Pseudometagea

Euchavis

Psilogastrellus

Oheza

Stilhida

Stilbuloida

Pseudochalcura

Hypoponera {\6) Ponerinae

Odoiitomachus (3, 13), Rhylidoponera (16), {IMyrmecia, 3) Ponerinae

Myrmecia {3. 13) Myrmeciinae

Odoiitomachus {3. 13, \6). Gnamptogenys {16} Ponerinae

Odo/itomachus {4, 13. \6). Pachycondyla {13) Ponerinae

Ectatomma (13) Ponerinae

Rhytidoponera {\3, 16) Ponerinae

Rhytidoponera (16) Ponerinae

Lasius (1,7) Formicinae

Formica (2, 4, 13), Cataglyphis (13), {Messor, 13) Formicinae

Cataglyphis (2), {Messor, 2, 13) Formicinae?

CamponotHs (6) Formicinae

Camponotus (13), Polyrachis (13) Formicinae

Camponotus {3, 13), Calomyrme.x {\3) Formicinae

Camponotus {H, 13) Formicinae

1. Ayre (1962); 2. Boucek (1956); 3. Boucek(1988); 4. Clau.sen (1941); 5. Das (1963); 6. Davis and Jouvenaz
(1990); 7. Heraty (1985); 8. Heraty (1986); 9. Heraty and Barber (1990); 10. Johnson et al. (1986); 1 1. Kerrich
(1963); 12. Van Pelt (1950); 13. Wheeler and Wheeler ( 1937); 14. Williams and Whitcomb (1973); 15. Wojcik
(1988); 16. Unpublished museum records.

Note: Wheeler and Wheeler (1937) is a summary of all rearing information prior to that year and should be
examined for more detailed information.

Mayr (M>rmiciiiaci (Whoolcr. 1907; Gemiiinaiii. 1933).
Neither Kupalu nor CJalcaria have been taken from within
a colony of Potionomyrme.x. and I regard some of the
associations listed above for Euiharis. Kapala. and
Galcaria as doubtful and in need of verification.

In order to evaluate coevolution between hosts and
parasites, the phylogenetic relationships of each group
can be compared for congruence (Kistner, 1979; Mitter
and Brooks, 1983). Brown (1954) proposed 2 main lin-
eages of ants; the poneroid lineage, which included the
Dorylinae. Ponerinae, and Mymiicinae (this last subfami-
ly was considered as derived from the poneroid tribe
Ectatommatini); and the myrmecoid lineage, which
included the Myrmeciinae as sister group to the
Aneuritinae and Formicinae (Wilson, Carpenter, and
Brown, 1967; Wilson, 1971). Taylor (1978) proposed a
slightly different phylogeny with the Myrmeciinae as sis-
ter group to the poneroid lineage. Baroni-Urbani (1989)
proposed the Myrmeciinae -(- Ponerinae as sister group to
the Myrmicinae + Pseudomyrmecinae, although he
strongly questioned the relationships between these 4 sub-
families. Holldobler and Wilson (1990) adopted a similar
scheme but included the army ants as sister group to
Myrmicinae -i- Ponerinae, and placed Formicinae as the
most basal (extant) subfamily of ants. More recent studies
have suggested that the Myrmicinae (together with
Pseudomyrmecinae and Myrmeciinae) either form the sis-
ter group of the Ponerinae, which shares derived character
states with army ants (Bolton, 1990; Ward, 1990;
Shattuck, 1992) or belong to a group including
Myrmeciinae and Formicinae (Baroni-Urbani, Bolton,
and Ward, 1992). If these lineages are compared with the
cladogram produced for eucharitid genera (Fig. 2), dis-
tinctive host correlations are apparent although they do
not necessarily correlate with any of the suggested evolu-
tionary schemes proposed for ants (i.e., Formicinae are
considered as a basal ant lineage, but are hosts to more
highly derived Eucharitinae).

Without a shared host for the Oraseminae or
Eucharitinae (ignoring the single-host records from
Formica and Eciton, and disputed records for
Eucharitinae), it is impossible to determine that either
Myrmicinae or Ponerinae could be the ancestral hosts for
Eucharitidae. The phylogenetic relationships of
Eucharitinae suggest that the Ponerinae are the ancestral
hosts for the Eucharitinae. The genus Austeucharis is
unique in its parasitism of Myrmeciinae. Tricoryna and
Pr()f7siloi>asfer are both parasites of Rliytidoponcra Mayr,
and under some tree topologies are considered as sister
groups. The shift to hosts in the Formicinae appears to be
a derived trait within Eucharitidae. The genera Obeza.
Stilhiila, Stilhiiloida. and Pscudochalcura arc a mono-
phylelic group within Eucharitinae, and Camponotus

species are the hosts for almost all of the species in these
genera (Table 4). PsciuJonwiaiica and Eiuharis +
Psilo^astrelliis are placed within a monophyletic group
that includes the O/xTtz-clade, and all are parasites of
Formicinae. Within the Eucharitinae. these 3 taxa are
probably more distantly related and may represent sepa-
rate colonizations of Formicinae. The Formicinae are dis-
tantly related to the Myrmicinae and Ponerinae, and a
shift in host to this subfamily would be regarded as a col-
onizing host adaptation.

CONCLUSIONS ON BIOLOGY AND
IMMATURE STAGES

Almost no homoplasy occurs for characters based on
morphology of immature stages and the biological traits
of the genera under study (Fig. 2). The 2 subfamilies are
strongly separated by behaviour, morphology of imma-
ture stages, and ant host. In the phylogenetic analysis it
was proposed that the expanded ovipositor, and hence
oviposition into chambers formed in leaf tissue, arose at
least four times in Eucharitidae. The inclusion of
Neoloshanus within the Eucharitinae is supported by bio-
logical traits and features of the immature stages which
are unlikely to be convergent (i.e., fusion of tergites I and
II). This would suggest that the phylogeny is robust and
can be used to predict possible hosts and behavioural
traits within the Eucharitidae.

Strategies of Eucharitidae for gaining entry to the nest
of the ant host are extremely variable (see Clausen,
1940a, 1940c. 1941). Dispersal mechanisms of planidia
through placement of eggs by adults or by activity of the
larva determine the likelihood of success on different host
ants. Development of new dispersal mechanisms by
Eucharitidae probably forms the basis of host shifts to dif-
ferent genera of Formicidae. Host shifts by eucharitids
within groups of closely related species or genera of host
ants would be expected, because related ant genera are
likely to have similar foraging strategies. In the New
World, species of Orasema have developed associations
with Solcnopsis. Tetramorinm, and Wasmannia, which
are probably all ants that are predators of thrips, or would
at least have a similar foraging pattern (size of prey,
search area, etc.) to the proposed ancestral host ant.
Pheidole. Major shifts in the group of ants that are para-
sitized by Eucharitidae may be associated with changes in
dispersal of planidia, through either first-instar behaviour
or oviposition habits of adult eucharitids. Each change in
oviposition behaviour or dispersal of larvae must accom-
modate the behaviour of the ant host, and adaptation to a
new host may result in the rapid evolution of a new group
within Eucharitidae.

30

Revision of Oraseminae and Psilocharitini (Eucharitinae)

In his revision of the Australasian Chalcidoidea, Boucek
(1988) provided revisionary notes and a generic key for
the Oraseminae, which provided a new foundation for
study of the Southeast Asian fauna. New information on
species from the Old World tropics has led to a classifica-
tion that removes from the genus Orasema (sensu

Boucek, 1988), species that are now assigned to 2 new
genera and are here placed in the Eucharitinae. In the fol-
lowing revision, descriptions and keys are provided for
the 6 genera and 56 species that are found in the Old
World tropical regions. Boldface type refers to prominent
character states useful for recognition.

Key to Genera of Oraseminae and Psilocharitini

2(1)

Prepectus distinctly separate from pronotum

(Figs. 217-234); anellus usually present;

hypopygium usually bare or with minute subapi-
cal setae, but if hypopygium with transverse
brush of long setae (Figs. 149, 264), then
propodeal disc smooth and shining laterally
(Psilocharis); antenna usually simple or at most
with blunt dorsal projections (Figs. 47^9); apex
of scutellum always rounded, never with apical
projections 2

Prepectus fused to pronotum, sometimes with
shallow furrow along line of fusion; anellus
usually absent, but if present, then face smooth,
hypopygium often with transverse brush of long
setae, and propodeal disc evenly areolate
{Gollumiella and Anorasema); antenna of male
often with elongate branches; apex of scutellum
rounded, emarginate, or with elongate projec-
tions

Eucharitini (Eucharitinae, see Boucek, 1988)

First gastral sternite (Ms^) with transverse
furrow delimiting small crescentic anterior
region (Figs. 74. 78, 113, 126, 256, 258, 260,
262); dorsal occipital margin usually round-
ed, rarely with carina; face usually with raised
coriaceous, rugose, or reticulate sculpture, rarely
smooth, but not pitted (Figs. 187-192); hypopy-
gium without long hairs, either minute or bare;
ovipositor expanded and ridged, usually curved

ventrally toward head (Fig. 262)

Oraseminae; 3

First gastral sternite (Ms^) evenly rounded
and not constricted (Figs. 268. 271), in some
cases with vague medial constriction (Fig. 273);
dorsal occipital margin with distinct carina;

lace smooth or pitted (Figs. 193-209); hypopy-
gium with or without tranvcrse row of long
hairs; ovipositor expanded or acicular, and usu-

ally straight (Figs. 264-272), but weakly curved

in some N. palgravei

Psilocharitini (Eucharitinae); 6

3 (2) Transscutal articulation (between mesoscutum
and axilla) obliterated (Figs. 36, 50); mandible
usually reduced, maxilla and labium minute or
absent, or maxilla lobate and without palpi 4

— Transscutal articulation complete (Figs.
235-246); mandible falcate, maxilla and labium
usually large, palpi 1- to 3-segmented (Figs.
189-190) ^..5

4(3) Body with metallic blue or green colours;
head and mesosoma strongly sculptured and
either coriaceous or rugulose; body length at
least 3.5 mm (Fig. 27); anellus present;
mandible truncate or spatulate, maxilla small;
Africa Timioderus Waterston, p. 32

— Body black or brown; head and mesosoma
almost smooth; length about 2 mm (Fig. 51);
anellus absent; mandibles absent, maxilla

enlarged and lobate; India

Indosema Husain and Agarwal, p. 38

5(3) Base of petiole gradually narrowed to large
knoblike condyle, and lacking dorsal flange
(Figs. 58. 67); petiole of female transverse

(Figs. 58, 67), hence gaster sessile or nearly so
(Figs. 6 lb, 256); petiole of male about twice as
long as broad, but then distal third o\ half sup-
ported ventrally by swollen anterior expansion
of first gastral sternite (anterior margin of Ms,

advanced; Fig. 78); Australia

Orasemorpha Boucek, p. 40

— Base of petiole truncated, lateral and dorsal
margins sharp and abruptly narrowed lo small
condyle (dorsal view. Figs. 94. 105. 120. 132).
and usually v^ith a sharp dorsal flange (rarely

31

6(2)

rviliKoil. d Fit:. 121 »: both sexes with petiole
usuullv more than twice us lon^ as broad
(rarely as long as broad in some females); in
both sexes anterior margin of first gastral stcrnite
(Ms,) meeting apex of petiole (Figs. 1 13, 126);

circumlropical and Nearctic

Orasema Cameron, p. 54

Base of petiole abruptly narrowed laterally
(truncated) (dorsal view. Figs. 144-145), rarely
with dorsal flange {P. paciftca): clypeal margin
straight with well-defined and narrow ante-
clypeus (Figs. 194-198); anteclypeus with mar-
ginal row of fine setae extending ventrally over
labrum (AS, Fig. 198); femoral groove broad
and evenly impressed (Figs. 223-225), often

reticulate: hypopygium usually with long brush
ol hairs (Figs. 149, 264), rarely with only few
long setae; ovipositor acicular (Fig. 264): Old
World tropics Psilocharis gen. nov., p. 81

Base of petiole gradually narrowed laterally
and dorsally (dorsal vicu. Fig. 159); clypeal
margin strongly lobate or slightly rounded,

anteclypeus absent (smoothly rounded to apex)
(Figs. 199-210); anteclypeus rarely with row of
fine setae (gcnmui-%xo\x\))\ femoral groove nar-
row and foveate (Figs. 227-234). rarely absent
(Fig. 226); ovipositor acicular or expanded;
hypopygium bare or with few small sublateral

setae, never elongate; mostly Indo-Pacific

Neolosbanus gen. nov., p. 93

Revision of the Old World Oraseminae
Timioderus Waterston

Timioderus Waterston, 1916:413. Type species:
Timioderus refringens Waterston; by original designa-
tion.
Originally assigned to the Perilampidae (Waterston,
1916), Timioderus was correctly placed in the
Oraseminae by Boucek (1988). The mandibles of
Timioderus are not falcate as they are for the rest of the
Eucharitidae, and may be well developed and toothed (as
for most Chalcidoidea), truncate, or long and chisel-
shaped. Timioderus is regarded as the sister group of
Indosema. with which it shares the absence of a transscu-
tal articulation, reduced mandibles, and cylindrical egg
without a stalk. Timioderus is similar to Orasemorpha in
having the scape and petiole short, the gastral terga sculp-
tured, the wing setae generally small, and the face and
mesosoma strongly sculptured. The flagellum of
Timioderus is cylindrical, serrate, or ramose, but the basal
anellus is retained in all species. The first gastral stemite
(Ms-,) of males is modified as an anteriorly protruding cup
that holds the apex of the petiole, and the point of articu-
lation with the gaster is markedly posterior to the anterior
margin of the first stcrnite. This mode of articulation is
also found in males oi Indosema and Orasemorplui.

GENERIC DESCRIPTION

Head transverse to subtriangular, 1.0-1.3x as broad as
mesosoma; median ocellus anterior to lateral ocellus, lat-
eral ocellus almost touching occiput (Fig. 34). Face, ver-
tex, and gena sculptured: scrobal depression shallow and
poorly defined, reaching median ocellus; ocellar-ocular

groove present or absent; occiput granulate to finely stri-
ate in circular pattern; occipital carina absent. Malar
depression absent; hypostoma small and not separated
from gena by hypostomal carina. Clypeus transverse,
epistomal sulcus lacking, apex only slightly rounded, not
extended over labrum; anteclypeus narrow, distinct from
postclypeus, and bare. Labrum 4- or 5- digitate or flaplike
with apical setae. Mandibular shape variable, usually
reduced; maxilla and labium strongly reduced, extending
only half distance between foramen and clypeal margin,
stipes forming 2 elongate lobes, labium globose or slight-
ly urn-shaped, palpi not visible. Antenna 10- to 12-seg-
mented; scape short and stout, 2.1-3.3x as long as broad,
usually reaching median ocellus: pedicel short; anellus
present and glabrous: funicle 6- to 8-segmented, segments
cylindrical, serrate, or ramose, without basal secondary
segmentation but with scattered MPS; basal flagellomeres
less than 2. Ox as long as broad, temiinal 3 to 4 segments
fused into distinct clava.

Mesosoma with dorsum evenly sculptured, coriaceous,
or rugulose. Notauli as broad shallow depressions anteri-
orly or absent. TSA obliterated dorsally but suture present
laterally below dorsal margin of lateral axillar surface
(Fig. 36). SSS broadly curved, meeting apparent posterior
margin of mesoscutum. Scutcllum with frenal area sepa-
rated dorsally by weak sculpture, broadly rounded; axillu-
lar sulcus absent. Metanotum extended laterally as small
smooth flange overlapping base of propodeum and partly
covering propodcal spiracle, the latter close to dorsal mar-
gin of propodeum; metanotum strongly excavated.

32

Propodeal disc relatively flat, foramen strongly arched;
callus pronounced, bare, or setose, separated from
mesepimeron by vertical furrow defining anterior callar
region; postspiracular furrow and metepimeral sulcus
marked by deep groove; ventral margin of propodeum
above hind coxa even and strongly ridged, without lateral
processes. Mesopleuron finely sculptured; mesepimeron
evenly swollen and lacking transepimeral sulcus; femoral
groove narrow and deeply impressed, without strong
sculpture; stemaular area of mesepistemum evenly round-
ed (no distinct sulcus or foveae). Prepectus reaching tegu-
la as broad triangular lobe, gradually narrowed ventrally.
Coxae and femora finely sculptured; number of tibial
spurs diagnostic for species; hind tarsi 0.5x length of
tibia, basitarsus short.

Wing. Veins of fore and hind wings broad and well
defined. Forewing 2.3-2.5x as long as broad, subtruncate
at apex (Figs. 43-44); disc with dense short hairs or
microtrichiae, marginal fringe minute; basal area bare;
speculum absent (pilose); costal cell pilose ventrally; sub-
marginal vein with sparse minute setae dorsally; marginal
vein 0.2x as long as wing and as densely pilose as rest of
wing; stigmal vein about 2x as long as wide, angled 45
degrees relative to anterior margin of wing; postmarginal
vein narrow and short, about equal in length to stigmal
vein.

Metasoma with petiole less than 0.7x length of hind
coxa in both sexes. Petiole of female broader than long,
strongly pinched dorsally so as to form small upper and
large lower lobe, narrowed basally to knoblike condyle
(Fig. 37); petiole not fused ventrally, ventral margins
broadly separated. Petiole of male longer than broad,
glabrous and slightly expanded at apex, not fused ventral-
ly, often held in vertical aspect on material examined.
Gastral terga finely rugulose; gaster of female as long as
head and mesosoma; gaster of male slightly longer than
hind femur; Mt^ of female less than 0.6x length of gaster;
Ms^ with broad, smooth basal constriction, anterior region

of female small and crescent-shaped (Fig. 27) and anteri-
or region of male a forward-protruding, cuplike structure
extending under petiole (Fig. 35). Hypopygium with
patch of minute setae on each side of midline. MSj, of
male rounded and setose. Cercus with several short setae
of equal length. Ovipositor sheath broad, not reaching
cercus, gonostylus distinctly separated basally and setose.
Ovipositor subapically expanded, slightly curved anteri-
orly (Figs. 29, 31); first valvula with single strong subapi-
cal ridge followed by lateral line of several small teeth;
second valvula broad with several small lateral teeth, and
smooth medially. Genitalia of males with parameres elon-
gate and bearing several long setae, digitus large and
disc-shaped with several stout marginal spines; aedeagus
subtruncate.

PHYLOGENETIC RELATIONSHIPS

The 5 known species are partitioned into 3 groups. T.
peridentatus is regarded as basal based on a quadrate
head, large mandibles, cylindrical flagellum, and digitate
labrum. The coriaceous or reticulate sculpture of the
mesosoma and occiput are regarded as synapomorphies
for T. coronula and T. ramosus; rugose sculpture, as
found in T. peridentatus and some Orasemorpha, and a
circularly carinate occiput are regarded as plesiomorphic.
A relationship between T. refringens and T. acuminatus is
not justified by synapomorphies. Tentative relationships
proposed among species of Timioderus are {peridentatus
+ ({coronula + ramosus) + acuminatus + refringens)).

BIOLOGY

Unknown. The form of the ovipositor suggests that eggs
are deposited in plant tissue into chambers fomied by the
ovipositor. Ovarian eggs are cylindrical and rounded at
the ends with no evidence of an apical projection.

DISTRIBUTION

Africa (Fig. 275).

Key to Species of Timioderus

Mandible large, each tridentate and slightly cup-
shaped along inner margin, apex overlapping
base of opposing mandible (Figs. 28, 39); head
subtriangular, l.4x as broad as high; funicular

segments cylindrical and moderately setose

T. peridentatus sp. nov., p. 34

Mandible small and narrow with apex acumi-
nate, truncate, or with 2 blunt teeth (Figs. 32, 40,
42); head usually strongly transverse. l.5-1.7x
as broad as high (Figs. 30. 32-33); funicular seg-
ments variable in shape 2

2(1) Head and dorsum of mesosoma rugulose-areo-
late (Figs. 27, 36); occiput circularly carinate;
callus with several small hairs dorsally (Fig. 27);
hind tibia with 1 or 2 apical spurs 3

— Head coriaceous or granulate (Fig. 33), dorsum
of mesosoma coriaceous; occiput granulate or
reticulate, at most with fine strigae; callus bare;
hind tibia usually with 0 or 1 apical spurs (but 9
of T. coronula with 2) 4

3(2) Funicular segments in both sexes cylindrical.

33

4(2)

segments clearly separated (Fig. 45); mesoscu-
tum with narrow medial longitudinal band of
bright blue; mandible with 2 blunt teeth (Fig. 32)
T. refringens Waterston, p. 35

Funicular segments of female serrate, segments
of male lamellate (Figs. 48^9); mesoscutum at
most with narrow longitudinal band of greenish
blue bordered by broad reddish bands; mandible

acuminate apically (Fig. 42)

T. acuminatus sp. nov., p. 36

Funicular segments of both sexes cylindrical or
slightly serrate, apex of each segment with brush

of fine setae that reaches following segment,
with setae strongly adpressed (Fig. 38); calcar
short and peglike; hind tibia with 1 spur (rarely
2) T. coronula sp. nov., p. 37

Funicular segments of female serrate, segments
of male strongly lamellate; setae in both sexes
adpressed or semi-erect in both sexes but seg-
ments clearly separated at apex (Figs. 46-47);
calcar absent; hind tibia with spur reduced or
absent T. ramosus sp. nov., p. 37

Timioderus peridentatus sp. nov.

Figs. 28, 36-37, 39

TYPE MATERIAL

Holotype, 9 , "Mossel Bay,/ Cape Province./ May, 1921."

"S. Africa./ R. E. Turner./ Brit. Mus./ 1921-248."

"HOLOTYPE/ Timioderus/ peridentatus/ Heraty."

Deposited in BMNH.

Paratypes: South Africa: Cape Province: Ceres, ii.l921

(9), xii.1924 (d), R. E. Turner (all BMNH); Oudtshoom,

X.1951(19,SAMC).

DIAGNOSIS

Recognized by having the mandible well developed and
tridentate (Figs. 28, 39), labrum digitate (Fig. 39), head
subtriangular, scape narrow, flagellomeres cylindrical,
calcar prominent, and mesosoma rugulose-areolate.

FEMALE

Length, 3.4 mm. Head and mesosoma dark metallic green
with faint reddish and blue reflections, anterior half of
mesoscutum dark violet medially; propodeum and vari-
able patches on mesepimeron sometimes dark metallic
blue; petiole, gaster, and coxae dark brown with bluish-
green lustre; antennal flagellum, mandible, and femora
dark brown; scape, pedicel, anellus, apex of femora, and
rest of legs yellowish brown. Wings hyaline, venation
clear.

Head subtriangular, 1.4-1.5x as broad as high; occiput
deeply emarginate and slightly invaginated; median ocel-
lus separated by slightly more than own diameter from
occiput; LOL 1.2-1.4x OOL. Face relatively flat, rugose;
scrobal depression broadly impressed above toruli, with
irregular striae extending in a circular pattern around
toruli, scrobal depression dorsally rugose; vertex broad,
rugose, and strongly depressed lateral to lateral ocelli,
shallow ocellar-ocular groove between posterior ocellus

and eye margin; dorsal occipital margin rounded; occiput
circularly carinate. Eyes separated by 1.9-2. Ox their
height. Malar space 0.8-1. Ox height of eye. Clypeal
region rugulose, lateral margin weakly impressed, tentori-
al pit deep; supraclypeal region glabrate or rugulose
medially. Labrum distinctly 5- or 8-digitate with pro-
nounced stiff seta at apex of each digit (Fig. 39); digits
short and placed irregularly around margin. Mandible
well developed, with tip overlapping base of opposing
mandible; each mandible tridentate, apical tooth elongate
and acuminate, basal teeth subequal; mandibles twisted
basally and folded over each other when closed, inner
surface cup-shaped; short row of long setae along outer
ventral margin of each mandible; maxilla and labium
strongly reduced. Antenna 11 -segmented; scape relatively
narrow, 3.0-3.3x as long as broad, ventral surface dis-
tinctly flattened below pedicel; pedicel globose; anellus
present; flagellum l.l-1.4x height of head; funicle 7-seg-
mented, segments cylindrical, surface scabriculous with
sparse, semi-erect setae basally and more densely setose
apically; F2 2.0-2.5x as long as wide, following seg-
ments subequal in length; clava as long as last 2 funicular
segments.

Mesosoma with dorsum rugulose-areolate, sculpture
fine and closely spaced anteriorly but becoming coarser
and more rugose toward apex of scutellum. Mesoscutum
with notauli vaguely impressed. Scutellum with frenal
line represented by narrow glabrous band laterally, nearly
obliterated dorsally, sculpture of frenal area similar to
scutellum, frenum rounded in profile and slightly exceed-
ing metanotum. Metanotum with lateral excavations
smooth and with weak crenulae. Propodeal disc evenly
rugose with septa widely spaced and irregular; callus with
posterior region swollen, globular, and rugose with sever-
al fine hairs dorsally. Mesepimeron smooth to weakly
rugose ventrally, swollen medially, and not divided by
transepimeral sulcus; femoral groove broadly and deeply

34

impressed, slightly crenulate along anterior margin;
mesepistemum rugulose-areolate to scabriculous laterally.
Prepectus and pronotum evenly rugulose. Proepistemum
smooth to weakly reticulate. Coxae weakly coriaceous,
mostly smooth; femora robust and expanded medially,
imbricate with dense short setae; tibiae with dense
adpressed setae; calcar long; hind tibia with 2 large spurs.
Forewing 2.2-2.3x as long as broad.

Metasoma with petiole (Fig. 37) and gaster typical for
genus; terga sparsely setose. Ms, without basal constric-
tion. Ovipositor typical for genus.

MALE

Agrees with female except for the following: head, meso-
soma, and gaster dark blue; antenna dark brown; scape,
coxa and femora with violet lustre; calcar large; hind tibia
with 2 small spurs; petiole with lateral flange.

VARIATION

The paratype females differ from the holotype by having
the mesepimeron entirely rugose (versus smooth), and
proepistemum weakly reticulate (versus smooth).

DISCUSSION

The well-developed and slightly twisted mandibles are
similar to those of Perilampidae and Chrysolampinae
(Pteromalidae). However, the apical tooth is elongate and
more typical of the shape found in most Eucharitidae. The
labrum is digitate (Fig. 39) and similar to that of other
Oraseminae except that it has 5 rather than the more typi-
cal 4 digits. The labrum is reduced in almost all other
Timioderus (Figs. 41^2); its presence in this species is
regarded as plesiomorphic.

DISTRIBUTION

South Africa (N, Fig. 275). Sympatric with T.
acuminatus.

ETYMOLOGY

Combination of the words Perilampidae and Latin dcnta-
tus. meaning toothed; referring to the similarity with non-
falcate mandibles possessed by Perilampidae.

Timioderus refringens Waterston

Figs. 32,41,43,45

Timioderus refriiij^cns Waterston. 1916:413-417.

Malawi: Monkey Bay [BMNH, examined].
Orasema viridicyanea Risbec, 1958:150-152. Zimbabwe:

Penkridgc [SAMC, examined]. New synonymy.

TYPE MATERIAL

Lectotype of Timioderus refringens (here designated), 9 ,
"Type.'" "NYASALAND./ LAKE NYASA,/ MONKEY
BAY./ 1:VI:1915/DR. W. A. LAMBORN." "MB/ 6.1.5."
"J. WATERSTON DET./ Timioderus/ refringens Wtst,/
9." "B.M. TYPE/ HYM./ 5-374." Specimen complete,
head buried in glue.

Paralectotypes: labelled as above with yellow-bordered
co-type labels (2 9 9). Holotype of Orasema
viridicyanea, 9 , "Orasema/ viridicyanea Risbec/ Type/
Penkridge, S. Rhodesia/ 16.1.1928/ R. H. R. Stevenson."
"SAMA 30C7." Adult parts mounted under 2 coverslips
on slide; under first coverslip, head and mesosoma with
right foreleg and mid legs attached, metasoma. ovipositor,
hind leg, and wings; under second coverslip, foreleg and
3 fragments of flagellum.

NOTES ON SYNONYMY

The only noticeable differences between the holotype of
O. viridicyanea and that of T. refringens is in the posses-
sion of a lighter (almost uniformly yellowish brown)
scape and a weak posterior flange on the prepectus, dif-
ferences which I do not consider of value in separating
these specimens as different species.

DIAGNOSIS

This species was well described and illustrated by
Waterston (1916). It is distinguished from other species
of Timioderus by the following: dorsum of mesosoma
rugose-areolate with narrow medial band of bright blue
coloration, otherwise dark green with red and blue reflec-
tions; head transverse (a, Fig. 32); occiput circularly stri-
ate; mandibles narrow and elongate with 2 short apical
teeth (b. Fig. 32); funicular segments cylindrical in both
sexes with short semi-erect setae, apical setae not bridg-
ing segments, and segments well separated (Fig. 45); cal-
car well developed; hind tibia with 1 or 2 elongate spurs;
and gastral terga densely setose.

VARIATION

There is some variation in the degree of bluish coloration
of the head and mesosoma, which is often more extensive
in males; however, the narrow medial band on the mesos-
cutum is consistent.

DISTRIBUTION

Kenya, Malawi, Mozambique. South Africa, Tanzania.
Uganda, and Zimbabwe (R, Fig. 275).

MATERIAL EXAMINED

Malawi: Cholo |no date] (\ 6. BMNF4). Mozambique;
Rikatla |no date] (19. BMNH). Kenya: Kongolai. Pokot.
July (Id, BMNH). South Aerica: Cape Province:
Transkei. Port St. .Iohn[s] [Pondoland). April to May and

35

Ivhiiiar> (159 9. 23d cT, BMNH): Transvaal: Fontaine
nr Piviona; McH)kctsi; Pretoria; September and Januarx to
February (39 9. Id, BMNH. SAMC. USNM); Natal:
InchanLia: Weenen: Dceeniber and February (49 9.
3d d. BMNH). Tanzania: Tanga. September (Id.
BMNH). Uganda: Serere, August (Id, BMNH).
Zimbabwe: Matopo Hills. April (Id. BMNH).

Timioderus acuminatus sp. nov.

Figs. 27.29.42,48^9

TYPE MATERIAL

Holotype, 9, "S. Africa/ R. E. Turner./ Brit. Mus./ 1925-
210." "Cape Province./ Ceres./ April. 1923." "HOLO-
TYPE/ Timioderus/ acuminatus/ Heraty." Deposited in
BMNH.

Paratypes: South Africa: Cape Province: Ceres, i.l921
(Id), iv.l923 (19), iv.l925 (2dd), R. E. Turner (all
BMNH); Cape Peninsular. 25.xii.1971, H. and M.
Townes (19 AEI); Transkei: Umtata, 18.ii-18.iii.l92[?],
[no collector] (Id. BMNH).

DIAGNOSI.S

Recognized by having head and mesosoma rugulose-are-
olate. flagellum of female serrate (Fig. 48). flagellum of
male lamellate (Fig. 49), mandibles acuminate (Fig. 42),
calcar small, hind tibia with 1 or 2 small spurs, and head
subtriangular. This species is morphologically similar to
T. pcridentatiis but differs in the structure of the antennal
flagellum, mandibles, and stouter scape.

FEMALE

Length, 3.4 mm. Head and mesosoma dark metallic green
with strong red and blue reflections, mesoscutum with 2
broad submedian bands of reddish colour bordering a nar-
row medial band of greenish blue; propodcum, lower half
of mesosoma, and coxae dark metallic blue; gaster dark
brown with greenish red reflections; femora dark brown
with faint green reflections; antenna dark brown; apex of
femora and rest of legs yellowish brown. Wings hyaline,
venation clear yellowish brown.

Head subtriangular, 1.5-1.6x as broad as high; occiput
deeply emarginate. not invaginated; median ocellus sepa-
rated by 1.5x own diameter from occiput; LOL 1.0-1.3x
OOL. Face broadly impressed around toruli, strongly
rugulose; scrobal depression broadly impressed above
toruli, with finer coriaceous or rugulose sculpture; vertex
rugulose. ocellar-ocular groove absent; dorsal occipital
margin acute; occiput circularly carinate. Eyes .separated
by 2.()-2.2x height of eye. Malar space 0.8-().9x height of
eye. Clypeal region rugulose or glabrate. lateral margins
indistinct. Labrum rectangular and broader than long with
single digits on each ventrolateral corner, each digit

slightly longer than broad and with long terminal seta.
Mandibles narrow and elongate, broadly overlapping at
tips, apex acuminate and without teeth. Antenna 11 -seg-
mented; scape 2.1x as long as broad, ventral surface flat
and glabrate: pedicel subglobose; anellus present; flagel-
lum 1.2x head height; funicle 7-segmented, F2 cylindri-
cal, following segments slightly serrate (Fig. 46), surface
strongly scabriculous. with moderately dense adpressed
setae, apical segments more densely setose; F2 l.lx as
long as broad, following segments subequal in length;
clava as long as preceding 2 funicular segments.

Mesosoma with dorsum uniformly rugulose-areolate
(may be more finely rugulose anteriorly). Mesoscutum
without notauli. Scutellum with posterior margin of frenal
groove weakly carinate. frenal area finely rugulose,
frenum rounded in profile and exceeding metanotum.
Metanotum with lateral excavations carinate. Propodeum
rugulose-areolate; callus with posterior region strongly
swollen and rugulose, glabrate laterally bare or with patch
of fine hairs dorsally. Mesopleuron rugulose-areolate lat-
erally; femoral groove narrow and deeply impressed,
glabrous medially and along anterior margin. Prepectus
and pronotum rugulose-alveolate. Proepistemum, coxae,
and femora weakly coriaceous or reticulate; calcar and
hind tibial spurs small (hind tibiae of holotype with 2 and
1 spurs, both hind tibiae of paratype with 1 spur).
Forewing 2.4x as long as broad.

Metasoma with petiole and gaster typical for genus,
petiole and Ms, hidden; terga rugulose and sparsely
setose. Ovipositor as in Fig. 29.

MALE

Length, 2.5 mm. Scape brown with iridescent reflections
and mesoscutum more evenly reddish in colour. Antenna
with basal funicular segments lamellate; F2 slightly high-
er than broad, apex cup-shaped, and outer, medial edge
emarginate; following flagellar segments subequal in
length and width. F8 seirate; clava with 2 incompletely
fused segments; funicular segments with dense, semi-
erect setae. Petiole 0.7x length of hind coxa, glabrous.
Gaster slightly longer than hind femur. Genitalia typical
for genus; aedeagus broad and rounded apically.

VARIATION

One paratype female (AEI) differs in having the head
more distinctly transverse (1.6x) but is otherwise similar.
It is much darker in colour with the dorsum of the meso-
soma black with reddish reflections and the medial band
of green very reduced, the lower face with strong reddish
reflections, the scape dark metallic green, and the antenna
black. Other structures are also darker in colour and there
is almost no blue coloration. In addition, the scrobal
depression is rugulose (rather than coriaceous), the
clypeus is glabrate. and the mesopleuron is reticulate-

36

rugulose. Otherwise, this individual is similar to the other

specimens.

BIOLOGY

Ovarian eggs dissected from the gaster were cylindrical,

rounded apically, and with a small apical nib at one end.

DISTRIBUTION

South Africa (Cape Province) (A. Fig. 275).

ETYMOLOGY

From Latin acuminatus, meaning pointed; referring to the
mandibles.

Timioderus coronula sp. nov.

Figs. 35, 38

TYPE MATERIAL

Holotype, 6 . "SOUTH AFRICA/ Grahamstown/ A.
Watsham: 12: 73." "R.IOO." "Timioderus." "HOLO-
TYPE/ Timioderus/ coronula Heraty." Deposited in
BMNH.

Paratype: Nigeria: Zaria, Dumbi Wood, 30.iv.l972. J. C.
Daining(l9,BMNH).

DIAGNOSIS

Recognized by having flagellar segments of both sexes
cylindrical or slightly serrate, each segment with a crown
of dense elongate setae that reach the following segment,
head sculpture granulate, and mesoscutum coriaceous to
finely reticulate.

11 -segmented; scape 2.3x as long as broad, ventral sur-
face slightly emarginate and glabrous; pedicel subglo-
bose; anellus present; flagellum 1 .6x height of head; funi-
cle 7-segmented, each segment slightly serrate, surface
granulate, with short, dense adpressed setae, apex with
crown of dense adpressed setae that reach base of follow-
ing segment (segments appearing fused) (Fig. 38); F2
1.2x as long as broad, following segments subequal in
length; clava elongate, slightly shorter than apical 3 funic-
ular segments.

Mesosoma with dorsum strongly coriaceous to finely
reticulate. Mesoscutum with notauli broadly impressed
anteriorly. Scutellum with frenal line narrow and weakly
impressed, sculpture of frenal area coriaceous, frenum
vertical in profile and only slightly exceeding metanotum.
Metanotum with lateral excavations coriaceous.
Propodeum evenly coriaceous; callus with posterior
region strongly swollen coriaceous and lacking setae.
Mesepimeron coriaceous to reticulate, swollen medially,
and lacking transepimeral sulcus; femoral groove deeply
and narrowly impressed, glabrous along anterior margin;
mesepisternum finely reticulate to coriaceous laterally.
Prepectus and pronotum evenly coriaceous.
Proepisternum finely reticulate. Coxae coriaceous and
bare; femora umbilicate with minute adpressed setae dor-
sally; tibiae with dense, short, adpressed setae; calcar
small; hind tibia with single small spur. Forewing 2.4x as
long as broad.

Metasoma with petiole glabrous, 0.5x length of hind
coxa. Gastral terga rugulose and bare. Ms, broadly con-
stricted and glabrous. Genitalia typical for genus.

MALE

Length, 2.5 mm. Dark metallic green with strong bluish
reflections; mesoscutum with submedian band of reddish
colour bordering narrow medial band of greenish blue;
head mostly metallic blue; antenna, petiole, and base of
gaster dark brown; apex of femora and rest of legs yel-
lowish brown. Wings hyaline, venation light brown.

Head transverse, 1 .7x as broad as high; occiput broad-
ly emarginate, not invaginatcd; median ocellus separated
by slightly more than its own diameter from occiput: LOL
i.lx OOL. Face broadly impressed around toruli, sculp-
ture granulate (very finely reticulate); scrobal depression
shallow, lateral margins obscure, glabrous above toruli;
vertex broad, evenly granulate, ocellar-ocular groove nar-
row and well defined; dorsal occipital margin rounded;
occiput granulate with very weak circular strigac. Eyes
separated by 2.4x their height. Malar space l.Ox height of
eye. Lateral margin of supraclypeal area broadly
impressed and glabrous. Labrum hidden. Mandible nar-
row and elongate, tip broadly overlapping opposing
mandible, each with 2 poorly defined asymmetrical teeth
apically; maxilla and labium greatly reduced. Antenna

FEMALE

Agrees with male except for following: mesoscutum
mostly reddish; scape, pedicel, and legs yellowish brown,
anellus light brown: face mostly rugulose, scrobal depres-
sion coriaceous to granulate: lateral margin of clypeus
hardly impressed; clava with 2 fused segments; notauli
not impressed; hind tibia with 2 well-developed spurs.

DISTRIBUTION

Nigeria and South Africa (Cape Province) (C, Fig. 275).

ETYMOLOGY

From Latin coronula; referring to the apical crown of
dense setae on the funicular segments.

Timioderus ramosus sp. nov.

Figs. 30-3 1 , 33-34, 40, 46-47

TYPE MATERIAL

Holotype. 9. "Aliwal North,/ Cape Province./ 4350 ft.
[1325 ml/ 1 13.1.1923." "S. Africa/ R. E. Turner./ Brii.

37

Mus./ 1^^23-70." "HOLOTYPE/ Timiocloriis/ ramosus
Herat) ." Card mounted, right mid leg mounted separate-
ly, tarsi broken on left fore and mid legs. Deposited in
BMNH.

Paralspes: SoiTH Ai-rica: Cape Province: Aliwal North.
l32.Sm. I-I3.i.l923. R. E. Turner (19, W6 6, BMNH);
Aliwal North, xii. 1922. R. E. Turner (1 d, BMNH).

DIAGNO.SIS

Recognized by having the head and mesosoma coriaceous
(Fig. 33), antenna of female weakly serrate (Fig. 46),
antenna of male strongly lobate (Fig. 47). mandible spatu-
late. calcar lacking, and hind tibia with 1 small spur or none.

FEMALE

Length. 2.9-3.4 mm. Dark metallic green with red and
blue reflections; mesosoma patterned with variable patch-
es of reddish coloration, darker ventrally with more bluish
coloration; gaster, coxae, and femora dark brown with
strong greenish reflections; antenna dark brown; apex of
femora and rest of legs yellowish brown.

Head transverse, 1.5-1.6x as broad as high; occiput
broadly emarginate. not invaginated; median ocellus sep-
arated by 1.5x own diameter from occiput; LOL l.l-1.2x
OOL. Face broadly depressed around toruli, coriaceous;
scrobal depression broad and shallow, with lateral margin
obscure, glabrous just above toruli; vertex evenly coria-
ceous, not impressed lateral to ocelli, the ocellar-ocular
groove shallow; dorsal occipital margin rounded; occiput
reticulate. Eyes separated by 2.1x their height. Malar
space 0.8x height of eye. Clypeus coriaceous to weakly
rugulose. lateral margin weakly impressed, with tentorial
pits deep; supraclypeal region glabrate. Mandibles nar-
row, tips slightly crossing when closed, apex chisel-
shaped; maxilla and labium extremely reduced. Antenna
10- or 1 1 -segmented; scape 2.5x as long as broad, flat
ventrally; pedicel globular; anellus present; flagellum
1.2x height of head; funicle 6- or 7-segmented, the seg-
ments cylindrical to slightly serrate, surface strongly
reticulate with short adpressed setae; F2 1.3x as long as
broad, following segments subequal in length. F8 partial-

ly fused to clava. clava as long as F6-F8.

Mesosoma with dorsum mostly coriaceous; scutellum
weakly striate-rugose, broadly impressed and smooth
anterior to frenum. Mesoscutum with notauli obliterated.
Scutellum with frcnal line narrow and weakly impressed,
frenal area coriaceous, frenum rounded in profile and
only slightly exceeding metanotum. Metanotum with lat-
eral excavations carinate. coriaceous medially.
Propodeum evenly coriaceous; callus with posterior
region strongly swollen, coriaceous, and lacking setae.
Mesepimeron evenly coriaceous, swollen, and not divided
by transepimeral sulcus; femoral groove narrow and
glabrous; mesepisternum coriaceous laterally. Prepectus
and pronotum evenly coriaceous. Lateral lobes of
proepisternum coriaceous, medial plate rugulose. Coxae
and femora weakly coriaceous to smooth, coxae bare,
femora with moderately dense minute setae; hind tibia
with dense covering of minute setae; calcar absent; hind
tibia with 1 small spur or none (absent on holotype).
Forewing 2.3-2.5x as long as broad.

Metasoma with petiole and gaster typical for genus,
gastral terga rugulose and bare; Ms^ glabrous with weak
basal constriction. Ovipositor typical for genus.

MALE

Length, 2.5-2.9 mm. Colour as for female or more com-
pletely bluish in coloration including dorsum and gaster.
Antenna 11 -segmented; basal funicular segments strongly
lamellate, with dense semi-erect short setae; F2 1.6x
higher than broad, apex of F2 cup-shaped, following seg-
ments subequal in length and size of lamella, F8 serrate;
clava as long as F7 plus F8. Petiole 0.5x length of hind
coxa, glabrous. Gaster slightly longer than hind femur.
Genitalia typical for genus.

DLSTRIBUTION

South Africa (O. Fig. 275).

ETYMOLOGY

From Latin ramosus, meaning branching; referring to the
projections of the male flagellomeres.

Indosema Husain and Agarwal

Indosema Husain and Agarwal, 1983:103-104. Type
species: Indosema Indica Husain and Agarwal; by
original designation.

Indosema is regarded as the sister group of Timioderus
based on the reduced labium, obliterated transscutal artic-
ulation, transverse petiole of both sexes, and cylindrical

ovarian eggs. Indosema also shares several character
states with both Timioderus and Orasemorpha, including
having the scape stout, ocelli arranged in an equilateral
triangle, occipital carina lacking, stigma robust,
mesepimeron ventrally expanded, and first gastral stemite
of male (Ms,) expanded and cuplike under the petiole.
Female Indosema have the first castral sternite constrict-

38

ed, the ovipositor subapically expanded, and the clypeus
transverse apically. Unique features of Indosema within
Oraseminae are the head and mesosoma brown and
smooth or lightly reticulate, frenal area indistinct,
forewing with microtrichiae, and Mt, less than 0.3x
length of gaster. Indosema also has the ability to pivot the
ovipositor forward between the coxae so that it may be
directed cephalad and parallel to the body during oviposi-
tion. The mobility of the ovipositor coincides with a
strong medial notch in the hypopygium to accept the
ovipositor.

GENERIC DESCRIPTION

Head transverse, l.lx as broad as mesosoma; median
ocellus anterior to lateral ocellus, lateral ocellus almost
touching occiput. Face including vertex and gena weakly
reticulate; scrobal depression shallow, not reaching medi-
an ocellus; ocellar-ocular groove absent; occiput weakly
coriaceous; occipital carina absent. Malar depression
absent; hypostoma small, not separated from gena by
hypostomal carina. Clypeus transverse, epistomal sulcus
lacking, apex slightly rounded or linear, not extended
over labrum; anteclypeus narrow and bare, distinct from
postclypeus. Labrum reduced (not discernible).
Mandibles absent; maxilla enlarged, lobate, and sclero-
tized (resembling mandibles), palpi and labium absent.
Antenna 10-segmented; scape short and stout, 1.5-2. Ox as
long as broad, not reaching median ocellus; pedicel as
long as broad; anellus absent; funicle 7-segmented, seg-
ments cylindrical and without basal secondary segmenta-
tion but with dense MPS; basal flagellomeres less than
1.5x as long as broad, terminal 3 segments fused into
indistinct clava.

Mesosoma with dorsum smooth or finely reticulate.
Mesoscutum with notauli weakly impressed. TSA obliter-
ated dorsally, with suture present laterally to dorsal mar-
gin of lateral axillar surface. SSS only vaguely impressed
(Fig. 50). Scutcllum with frenal area and scutellum finely
reticulate, separated by transverse glabrous band repre-
senting frenal line; axillular sulcus lacking. Metanotum
extended laterally as small flange overlapping base of
propodeum, not covering spiracle which is close to dorsal
margin of propodeum. Propodeal disc rounded, with fine
reticulate sculpture and faint central furrow; callus pro-
nounced and setose, with anterior vertical furrow (of
Timioderus and Orasemorpha) indistinct; postspiracular
furrow and metepimeral sulcus clearly impressed; ventral
margin of propodeum above hind coxa evenly rounded
and without lateral processes. Mesopleuron finely reticu-
late laterally including femoral groove; mesepimeron
evenly swollen and lacking transepimeral sulcus; femoral
groove broadly and deeply impressed; sternaular area of
mesepisternum evenly rounded (no distinct sulcus or
fovcae). Prepectus reaching tegula as broad triangular

lobe, gradually narrowed ventrally. Coxae and femora
lightly sculptured; tibiae with spurs 1-1-1/2; hind tarsus
0.7x as long as tibia.

Wing. Veins of fore and hind wings broad and well
defined. Forewing 2.4x as long as broad, subtruncate at
apex; disc appearing glabrous but pilose with minute
setae, the marginal fringe absent; basal area bare; specu-
lum absent (pilose); costal cell bare; wing veins lacking
any apparent setae; marginal vein 0.3x as long as wing;
stigmal vein barely distinguishable from marginal vein;
postmarginal vein narrow and about twice as long as
apparent length of stigmal vein.

Metasoma with petiole in both sexes smooth and trans-
verse, less than 0.5x length of hind coxa, narrowed basal-
ly, and fused ventrally. Gastral terga smooth, basal terga
with sparse minute setae, following terga with transverse
band of fine setae; gaster of both male and female as long
as head and mesosoma (Fig. 54); Mt, in both sexes less
than 0.6x length of gaster; Ms, with narrow basal con-
striction, the anterior region of female crenulate, that of
male projecting forward and forming cuplike projection
under petiole. Hypopygium bare. Ms^^ of male narrowly
rounded and setose. Cercus with few elongate setae.
Ovipositor sheath broad, reaching cercus, gonostylus sep-
arated and sparsely setose. Ovipositor broadly expanded
along entire length, only slightly expanded subapically.
and straight (Fig. 51); first valvula with weak subapical
ridge followed by lateral line of 10 minute teeth; second
valvula with several strong lateral teeth, smooth medially.
Genitalia of male with parameres short and bearing few
short setae, digitus disc-shaped and bearing several stout
marginal spines; aedeagus narrow and subacute.

DISTRIBUTION

Northern India (I, Fig. 275).

Indosema indica Husain and Agarual

Figs. 50-54

Indosema indica Husain and Agarwal. 1983:104-106.
India: Uttar Pradesh: Aligarh [AMUA, not examined].

DIAGNOSIS

The single species known for this genus can be recog-
nized based on the generic features. The female was well
described by Husain and Agarwal (1983). The male is
similar to the female except for having the gaster nanow
and elongate, and Ms, constricted but onl) slightly pro-
jecting below the base of the petiole. The male is appar-
ently being described by Husain (Dr. S. I. Farooqi, lARI.
New Delhi, India, pcrs. comm.) and will not be further
discussed here.

39

wmnHA

C\>lk'cicd on leaves of Ricinus communis L. (castor bean)
(Husain and Agarual. 1^83). This plani has large
extrafloral nectaries which may have attracted aduh /.
indica for feeding. My own observations and Boucek
(pers. comni.. 1990) suggest that /. indica is associated
with short grasses. Boucek (1988) speculated that it may
use a short leguminous plant, possibly related to Lotus, as
a plant host tor oviposition. The form of the ovipositor
suggests that eggs are deposited in plant tissue within
chambers formed by the ovipositor. Ovarian eggs are
cylindrical and rounded at the ends with no evidence of
an apical projection.

DISTKIIU HON

Northern India (I. Fig. 275).

M ATKRIAI. EXAMINED

India: Delhi, 14.iv.42. H. U. Khan, on Brinjal (I 9,
BMNH); New Delhi, lARI area, 10,26-28. x.79, Boueek
(19, 96 6, BMNH): lARI. 26.X.79, C. S. Roy (49 9,
lARI): lARI farm. Il.vii.83, B. Lai (19. lARI): lARI,
9-lO.vii 90, J. Heraty (79 9, Id, JMH. TAMU); Ullar
Pradesh: Aligarh. 8.x. 1980. T. Husain (1 9 paratype,
BMNH); Aligarh. 1 1.x. 79, vii.8(). M. Hayat {26 6 ,
BMNH).

Orasemorpha Boucek

Eucharomorpha Girault, 1913b [December]: 94-95. Type
species: Eucharomorpha viridis Girault, by original
designation. Preoccupied by Eucharomorpha Girault.
1913a [September]: 157. Type species: Eucharomor-
pha worcesteri Girault, by subsequent designation
(Gahan and Fagan, 1923:58). Latter genus correctly
treated as junior synonym of Orasema Cameron
(Boucek. 1988:519).

Orasemorpha Boucek, 1988:518-519. Replacement name
and redescription.

The taxonomic history of this genus was provided by
Boucek (1988). A redescription of the genus is presented
here for comparison with other orasemine taxa.
Orasemorpha shares some character states with
Timioderus, including a stout scape, a vertical furrow
dividing the callus, a strongly arched propodeal foramen,
sculptured gaster (some species), and the first gastral ster-
nite extended under the petiole in males. Orasemorpha is
distinguished from Timioderus and Indosema by the fal-
cate mandibles, the digitate labrum, and a complete
transscutal articulation. The strongly sculptured face,
transverse head, short petiole, and first stemite of males
distinguish this genus from Orasema.

GENERIC DESCRIPTION

Head subtriangular (Figs. 55. 66. 73. 76, 187).
1.5-1.6X as broad as high. 1.2-1.6x as broad as mcsoso-
ma; median ocellus anterior to lateral ocellus, lateral ocel-
lus close to occipital margin (Figs. 57, 69, 77). Face, ver-
tex, and gena strongly sculptured: scrobal depression
shallow and poorly defined, partially including median
ocellus; ocellar-ocular groove weakly defined, sometimes
marked by weak striae; occiput with elongate reticulate
sculpture, occipital carina absent. Malar depression

absent; hypostoma well developed, separated from gena
by hypostomal carina. Clypeus transverse, apical margin
linear, not extended over labrum. with epistomal sulcus
shallow; anteclypeus narrow and bare or with minute
setae, distinct from postclypeus. Labrum 4- to 7-digitate.
Mandibles falcate. 2/2, 3/2, or 3/3 dentate; maxilla and
labium reduced, palpi short and broad, not segmented.
Antenna 10- (questionable) to 13-segmented; scape short
and stout but reaching median ocellus, cylindrical, and
slightly flattened just below pedicel; pedicel short and
globose; anellus usually present and glabrous; funicle
usually 8-segmented, rarely 7- or 9-segmented, segments
cylindrical, without basal secondary segmentation, and
with scattered MPS; basal funicular segments less than
2. Ox as long as broad, last 2 to 3 segments fused into
clava.

Mesosoma with dorsum evenly sculptured, finely or
coarsely rugose. Notauli deeply impressed along entire
length and narrowly crenulate. angled to midline but well
separated at TSA. TSA present. SSS angled and meeting
TSA at midline. Scutellum with frenal area weak rugose,
semicircular in dorsal view, usually with weak median
depression; axillular sulcus lacking. Metanotum extended
laterally as smooth flange overlapping base of propodeum
and partly covering spiracle which is close to dorsal mar-
gin of propodeum. Propodeal disc broadly rounded and
sculptured, foramen strongly arched; callus swollen and
glabrous, separated from mesepimeron by vertical furrow
defining anterior callar region; postspiracular furrow shal-
low; ventral margin of propodeum above hind coxa even
and strongly ridged, without lateral processes. Upper
mesepimeron swollen and glabrous or weakly rugulose,
lower mesepimeron sculptured or smooth, transepimeral
sulcus impressed but not prt)mincnt: femoral groove
broadly impressed; sternaular area of mesepisternum

40

evenly rounded (no distinct sulcus or foveae), mesepister-
num swollen ventrally between fore and mid coxae.
Prepectus reaching tegula as broad triangular lobe, only
slightly narrowed ventrally. Coxae and femora weakly
sculptured; tibiae with spurs 1-1-2; hind tarsi 0.6x as long
as tibia.

Wing. Veins of fore and hind wings broad and well
defined. Forewing 2.3-2.4x as long as broad, subtruncate
to broadly rounded apically; disc densely pilose, with
marginal fringe present or absent; basal area, speculum,
and costal cell with pilosity diagnostic for species; sub-
marginal vein with sparse dorsal setae; marginal vein
0.3x as long as forewing and densely pilose; stigmal vein
broad, sessile to club-shaped; postmarginal vein long and
narrow, 0.3-0.6x as long as marginal vein.

Metasoma with petiole of female transverse, in dorsal
view posterior half 3-5x as broad as long, apical margin
broadly emarginate dorsally; anterior half strongly
pinched dorsally (Figs. 58, 67), narrow upper region
glabrous, longer than broad and extending upward into
propodeal foramen; petiole gradually narrowed basally to
knoblike condyle, and fused ventrally. Petiole of male as
in female but less than twice as long as broad, cylindrical
but broader apically, and shorter than hind coxa. Gastral
terga sculptured or smooth, gaster of female as long as
head and mesosoma, gaster of male slightly longer than
hind femur; Mt^ of female less than 0.6x as long as
gaster; Ms^ constricted by broad transverse crenulate or
striate furrow, anterior region of female small and cres-
cent-shaped, anterior region of male protruding forward
as cuplike structure extending under petiole (Fig. 78).
Hypopygium with patch of fine setae on each side of mid-
line or bare. MSj^ of male rounded and setose. Cercus with
several setae of equal length. Ovipositor sheath broad, not
reaching cercus, gonostylus distinctly separated basally
and setose (Fig. 257). Ovipositor subapically expanded,
only slightly curved anteriorly; first valvula with subapi-
cal ridge followed by lateral line of 3 to 4 sharp teeth;
second valvula broad with several strong lateral teeth,
smooth medially. Genitalia of male with parameres elon-
gate and bearing few long setae; digitus disc-shaped, with
several short marginal spines; aedeagus broad, subacumi-
nate at apex.

PHYL()(JENETIC RP:LATI0NSHIP.S

Within OrascDiorpha, 2 groups of species can be recog-
nized. The differences between these groups are minor
and I have not recognized them as formal species groups.
The first group includes 4 species. O. xcniades. O. myrmi-
cac, O. yoc'thcl. and C). tridcntata. These species all have
a relatively smooth, black mescsoma and glabrate gaster.
The antennal flagcllum of O. xcniadcs and O. nidcntara

is strongly narrowed basally. which I consider as a
derived state compared to the rest of the Eucharitidae.
The states attributed to the antenna of O. myrmicae, espe-
cially a reduced number of funicular segments, are ques-
tionable and presently unknown for O. goethei. The loss
of a marginal fringe from the forewing is a derived state
that separates O. goethei, O. myrmicae, and O. tridentafa
from O. xeniades. The second group of species includes 5
species, O. didentata, O. erihotes, O. pytallus, O.
sparsepilosa, and O. varidentata. These species have the
head and mesoscutum strongly rugose, the gastral terga
densely setose, and the eyes almost always setose (bare in
males of O. erihotes). In O. sparsepilosa and O. pyttalus,
the setae of the head and mesosoma are long and dense.
Orasemorpha varidentata, O. didentata, and O. erihotes
are similar morphologically and exhibit only small differ-
ences in setation and shape of the mesosoma.

Within Orasemorpha, of the 2 groups that are evident
based on gastral sculpture and setation, it is difficult to
polarize character states based on the outgroup. The nar-
rowed antennal flagellum of the first group (including O.
xeniades) could be derived (but possibly not shared by all
members). If Timioderus were used as the outgroup, then
the rough sculpture, setose gaster, and broad antennal fla-
gellum would be plesiomorphic. However, using either
Indosema or some species groups of Orasema could
reverse all but the antennal characters. Setose eyes are not
found elsewhere in the Oraseminae and are found only in
some species of Gollumiella and genera in the basal
Eucharitinae. Tentative relationships proposed among
species of Orasemorpha are ((myrmicae + goethei + fri-
dentata + xeniades) + ((jjyttalus + sparsepilosa) + {diden-
tata + erihotes + varidentata))).

BIOLOGY

Orasemorpha didentata, O. erihotes, O. myrmicae. and
O. tridentata have been reared from 2 different species of
Pheidole. Wheeler obtained pupae of. O. tridentata from
a colony of Pheidole proximo Mayr (Brues, 1934). the
pupae of which would have been invaluable for compari-
son with Orasema. Unfortunately these specimens could
not be located at the MCZ. The form of the ovipositor
suggests that eggs are deposited in plant tissue into cham-
bers formed by the ovipositor. Ovarian eggs are smooth
and stalked as is typical of most Eucharitidae.

DISTRIBUTION

Australia and Tasmania (Fig. 275). Few records for
species in this genus exist. Only Orasemorpha erihotes is
found in western Australia; the remaining species arc
eastern. Only 2 species, O. xeniades and O. erihotes. are
known from Tasmania.

41

Key to Species of Orasemorpha

1 Mesosciiium wiih niidlobc reticulate to finely
rugulose; scutellum lightly sculptured, often bare
medially (Fig. 57); gastral terga at most with
very few setae 2

— Mesoscutum with midlobe strongly rugose (Figs.
63. 69, 77), or //rugulose or scabriculous then
scutellum densely sculptured (Fig. 70); scutel-
lum sometimes smooth medially and often with
longitudinal median depression; gastral terga
with dense covering of subdecumbent or
adpressed setae (Figs. 256-257) 5

2(1) Forewings without marginal fringe 3

— Forewings with complete marginal fringe (Fig.
60) O. xeniades (Walker), p. 43

3 (2) Forewing without speculum, evenly pilose; mid-

lobe of mesoscutum and scutellum rugulose;
gastral terga weakly coriaceous; mandibles 3/2
dentate O. myrmicae (Girault), p. 44

— Forewing with speculum; midlobe of mesoscu-
tum smooth to weakly rugulose, scutellum
smooth to coriaceous; gastral terga smooth;
mandibles 3/3 dentate (unknown for some
species) 4

4 (3) Gaster weakly coriaceous; midlobe of mesoscu-

tum weakly rugulose; head and mesosoma of
both sexes dark green; mandibles 3/3 dentate
(Fig. 56) O. tridentata (Girault), p. 45

— Gaster glabrous; midlobe of mesoscutum mostly
bare; head and mesosoma black in female, dark
blue in male. (Dentition unknown.)

O. goethei (Girault), p. 46

5(1) Head (including eyes) and mesosoma with mod-
erate to dense covering of long, erect setae (Figs.
66,69) 6

— Head and mesosoma bare or with short subde-

cumbent setae (Figs. 70, 73, 77, 217); eye setae,
if present, short and sparse (Figs. 76, 187) 7

6 (5) Marginal fringe of forewing complete; forewing

without speculum, the basal area bare but with
narrow band of setae extending along impression
of cubital vein to base of forewing, and costal

cell densely pilose; wing disc densely pilose

O. pyttalus (Walker), p. 47

— Marginal fringe of forewing restricted to pos-
teroapical margin; forewing with speculum and
basal area bare, and costal cell with few minute
setae apically; wing disc with setae sparse and

fine, hardly visible (Fig. 68)

O. sparsepilosa sp. nov.. p. 48

7 (6) Femora yellowish brown; coxae smooth and

completely pilose; mesosoma black with strong
iridescent reflections, dorsum coarsely rugose,
interstices broadly spaced (Fig. 63); gastral terga

moderately pilose

O. varidentata (Girault), p. 49

— Femora dark brown, at least over basal third;
coxae smooth to rugulose, dorsal surface bare;
mesosoma dark green or bluish and sometimes
with faint reddish reflections, the dorsum rugose
to scabriculous, interstices closely spaced (as in
Fig. 187); gastral terga densely pilose 8

8 (7) Hind coxa almost completely smooth; femora

and tibia of hind leg with long dense semi-erect
setae (Fig. 78); eye of female with sparse short
setae (Fig. 76), eye of male with minute setae....
O. didentata (Girault). p. 50

— Hind coxa rugulose to scabriculous; femora and
tibia of hind leg with short dense subdecumbent
setae; eye of female with minute setae or bare,
eye of male bare (Fig. 73)

O. enfto/es (Walker), p. 51

42

Orasemorpha xeniades (Walker)

Figs. 55,57,59-61

Eucharis xeniades Walker, 1839:14-15. Australia: New

South Wales [BMNH. examined].
Orasemorpha xeniades; Boucek, 1988:519.

TYPE MATERIAL

Lectotype (designated Boucek, 1988), 6 . "1441.""
"Sydney." "LECTO-/ TYPE." "Type." "Psilogaster/ xeni-
ades/ Walker." "B. M./ TYPE/ HYM./ 5.620." "d
Orasema/ xeniades (Walk.) LT/ det. Z. Boucek, 1986."
Remounted 6 on white card, flagella. part of left
forewing, and left hind legs missing. Paralectotypes, 2
6 S labelled "Sydney"" and "N.S.W.."" otherwise as
above.

DIAGNOSIS

Recognized by the following: marginal fringe distinct,
speculum present, and gaster almost entirely bare and
smooth. This species is closely related to O. goethei. O.
myrmicae, and O. tridentata but can be differentiated by
the presence of a distinct marginal fringe on the forewing
(Fig. 60).

FEMALE

Length, 2.4-2.8 mm. Body black with strong bluish-green
reflections; antenna, mandible, femora, and apical tar-
someres dark brown; apex of femora and rest of legs yel-
lowish brown (fore and mid tibiae may be dark brown
medially). Wings hyaline, venation pale brown.

Head subtriangular. eyes protuberant and inner mar-
gins only slightly diverging; occiput broadly emarginate;
temples large and broadly rounded behind eyes (Fig. 59);
LOL 0.9-1. Ox OOL. Face broadly rounded, completely
shallow reticulate or finely rugulose; scrobal depression
narrow and broadly rounded with finer sculpture, partially
including median ocellus. Eyes bare, separated by
2.3-2.4X their height. Malar space 1.0-l.2x height of eye.
Clypeal region weakly reticulate, lateral margins of
clypeus at tentorial pits deeply impressed; supraclypeal
area slightly bulging. Labrum 4-digitate. Mandibles 3/2
dentate. Antenna 12-segmented; anellus small; flagellum
1.3x height of head; funicular segments with dense short
adpressed setae; F2 1.8-2.2x as long as broad. I.l-I.3x
F3. following segments subcqual in length, equal in
width; clava obconical and as long as preceding 2 seg-
ments.

Mesosoma with midlobe of mesoscutum reticulate and
lateral lobe strongly swollen and glabrale; axilla swollen
and glabrous; scutellum glabrous dorsally. and reticulate
laterally and basally (Fig. 57). Notauli sharply impressed
and narrowly crenulatc. SSS broadly impressetl with few
transverse carinae. Scutellum l.5-l.6x as long as broad:

frenal line a broad glabrous band dorsally. frenal area
rugulose. broadly impressed medially. Propodeum evenly
rounded and coriaceous to reticulate with few weak longi-
tudinal carinae; callus and metepimeron swollen and
glabrous, callus with weak callar nib. Upper mesepimeron
swollen and glabrous, lower mesepimeron flat and
glabrous, transepimeral sulcus foveolate; mesepistemum
finely reticulate, smooth ventrally. Prepectus broadly tri-
angular, reticulate with pronounced posterior flange.
Pronotum rugose-reticulate. Proepisternum coriaceous.
Fore and mid coxae and femora coriaceous, hind coxa
nearly glabrous; hind leg with tibia and apex of femur
densely short-setose. Forewing 2.3-2.5x as long as broad;
basal area and speculum bare; speculum closed basally by
broad band of setae; costal cell with narrow band of api-
cal setae; disc densely short pilose, distinct marginal
fringe around apical and posteroapical margins; stigmal
vein 1.7-3.3X as long as broad, narrowed basally.

Metasoma with petiole glabrous and 0.3-0.4x as long
as broad. Gastral terga very weakly coriaceous with few
fine adpressed setae, almost glabrous; Ms^ with constric-
tion deeply impressed and crenulate, anterior region
glabrous and semicircular, anterior margin extending for-
ward under petiole. Hypopygium with 1 to 2 minute sub-
lateral setae at apex. (Ovipositor hidden.)

MALE

Length, 1.7-2.0 mm. Body dark brown to black, head and
mesosoma sometimes with faint blue or green reflections;
antenna black, scape and pedicel with metallic blue
reflections; femora and tibiae mostly black, apices of
femora and tibiae, and tarsi brown. Wings weakly infus-
cate.

Head shape as in female; LOL 0.9-1.2x OOL.
Sculpture reticulate to scabriculous. Antenna 12- or 13-
segmented; anellus small; llagellum 1.7-2. Ox height of
head; funicular segments with dense subdecumbent setae,
F2 1.8-2.4X as long as broad, I.2-1.5x F3; clava 1- or 2-
segmented with varying degrees of fusion.

Mesosoma more slender than female; scutellum and
posterior region of axilla weakly reticulate: scutellum
I.6-1.8X as long as broad; frenal area coriaceous, frenal
line as shallow dorsal groove. Propodeum with or without
median carina. Forewing 2.1-2.4x as long as broad;
speculum sparsely pilose (Fig. 60).

Metasoma with petiole 2. Ox as long as broad. ().9-|.2x
as long as hind coxa. ().8-l.2x as long as propodeum and
weakly reticulate dorsally. often with lew weak striae.
Gastral terga glabrous: Ms, strongly constricted, anterior
region glabrous. MSj^ broadly rounded and setose.
Genitalia typical for genus; parameres narrow and elon-
gate, sharp median process, digitus with 3 lo 4 niargmal
spines; aedeagus broad and acuminate at tip.

43

Bioi ()(;v

Reared from PhciJoU' lasnianiensis.

DISTRIKl HON

Southeastern Australia itKiudiiig Tasmania (X, Fig. 275).

MATERIAL EXAMINED

Al'Stralia: New South Wales: Cabramatta. Sydney;
Nerriga; Tooloom Scrub; Sawpit Ck. Mt Kosc[iusko| N.
P.; December to February (39 9, Id, BMNH); South
Australia; Ml Lofty. Adelaide, January (Id, BMNH);
Tasmania; Gladstone, 1 km SSE; Herrick, 1 km E by N
(41.06S 147. 53E); Marrawah, 4 km SW (40.57S
144.40E); The Lea (42.56S 147. 19E); The Lea, 6 km S
Hobart; Huon Riv.. Lea. ex Pheidole tasmaniensis (Id
and partial remains of 5 others mounted with headless
ant, USNM); Wayatinch, 3 km NE by E; April and
December to February (29 9, 22dd, ANIC); Victoria;
Lake Mtn. E of Melbourne; Kinglake N. P. nr Melbourne;
January to February (7 9 9 , 22d d , BMNH).

Orasemorpha myrmicae (Girault) comb. nov.

Fig. 62

Epimetagea myrmicae Girault, 1936;3 [324]. Australia;

Victoria [QMB. examined]. Hedqvist, 1978:243 (list).

Dahms, 1984:842 (notes on type material).
Chalcura myrmicae — Bou&k. 1988:529 (combination).

TYPE MATERIAL

Lectotype (designated as holotype by Dahms, 1984), 9,
"Belgrave V./ F. E. Wilson. Jan 1922." "HOLOTYPE/ T.
9262/ E. C. D. 1984." "Eucharomorpha myrmicae Gir."
[GH] '-LECTOTYPE/ Orasemorpha/ myrmicae (Grit)/
Det. Heraty, '90." Head removed, body mounted with
minor of Pheidole. Head and antenna mounted on slide
labelled '^Tricoryna ectatommae Gir., Paratype.
Eucharomorpha myrmicae Gir. Ty. 9 ." The head is
mounted under a separate coverslip closest to the label
and is correct for Orasemorpha. The 9-segmented anten-
na of T. ectatommae is mounted under the outer coverslip
fragment along with a 10-segmented antenna attributed to
E. myrmicae. A separate 10-segmented antenna, identical
to the other, is under a different coverslip fragment. The
antennae are atypical for Orasemorpha and it is question-
able whether they belong to E. myrmicae. The original
description does not specify the number of specimens
examined. The slide mount is labelled as a paratype.
which indicates that the headless body may not have been
a unique specimen, and thus should not be referred to as a
holotype; the point-mounted body is therefore designated
here as lectotype.

i)ia(;n().sis

Similar to O. tridentata and O. goethei based on the
almost total lack of setae on the gastral terga and no mar-
ginal fringe on the forewing. This species is distinguished
from O. tridentata by the following; gastral terga weakly
coriaceous, mesosoma dorsum (midlobe and scutellum)
rugose-areolate, speculum absent, and mandibles with 3/2
dentition. If the antennae are properly assigned to the lec-
totype then it uniquely possesses a 10-segmented antenna
that lacks an anellus (Fig. 62). This last feature is dubious
and should not be used as a limitation for assigning indi-
viduals to this species.

FEMALE

Length, 1.75 mm (mesosoma and gaster). Mesosoma,
petiole and coxae black with reddish reflections; gaster
and basal two-thirds of femora dark brown with reddish
reflections; apex of femora, tibiae and basal tarsomeres
yellowish brown; antennal flagellum dark brown, scape
lighter brown, pedicel intermediate. Wings hyaline, vena-
tion pale brown.

Head (limited information based on poor slide mount)
with labrum 6-digitate and with 1-4-1 formula. Mandibles
3/2 dentate. Antenna 10-segmented; anellus absent; funic-
ular segments with short dense setae, F2 2. Ix as long as
broad. 1.4x as long as F3, following segments equal in
length and width; clava obconical and longer than preced-
ing 2 segments, composed of 2 partially fused segments.

Mesosoma with midlobe of mesoscutum and scutellum
close-packed rugose-areolate with interstices shallow and
rounded, scutellum smooth just anterior to frenal groove;
lateral lobe polished and .strongly swollen; axilla smooth
dorsally and weakly striate posteriorly, strongly angled to
meet scutellum. Notauli narrow and deeply impressed,
finely crenulate. SSS weakly crenulate. Scutellum round-
ed dorsally; frenal groove a narrow glabrous band dorsal-
ly, frenal area weakly rugose and slightly depressed medi-
ally. Propodeum evenly rounded and weakly rugose,
impressed dorsally and above base of coxae; callus and
metepimeron glabrous, callus with strong medial groove
delimiting anterior region. Mesepimeron slightly swollen
and glabrous, transepimeral sulcus weak; mesepisternum
evenly reticulate laterally, smooth ventrally. Prepectus tri-
angular, reticulate vvith smooth dorsal and posterior
tlanges. Pronotum broadly rounded laterally and lightly
rugose. Proepistemum weakly coriaceous. Coxae weakly
coriaceous; femora scabriculous apically on lateral sur-
face; hind leg with tibia and apex of femur densely short-
setose. Forewing 3. Ox as long as broad; basal area bare;
speculum absent; costal cell with sparse medial band of
fine setae; disc moderately pilose with short fine setae, no
marginal fringe; stigmal vein broad and perpendicular to
wing margin.

Metasoma with petiole glabrous and broader than

44

long. Gastral terga weakly coriaceous and bare; Ms., with
constriction broad and finely crenulate, anterior region
cup-shaped and projecting forward under petiole.
(Ovipositor and sheaths hidden.)

MALE

Unknown.

BIOLOGY

Taken from nest of Pheidole sp. (Girault, 1936).

DISCUSSION

No additional material has been assigned to this species.
See discussion of variation for Orasemorpha tridentata
for similar material.

DISTRIBUTION

Southeastern Australia (M, Fig. 275).

Orasemorpha tridentata (Girault)

Figs. 56, 58

Eucharomorpha tridentata Girault, 1915:230. Australia:

Queensland [QMB, examined]. Dahms, 1986:597

(notes on type material).
Eucharomorpha wheeleri Brues, 1934:203-205.

Australia: New South Wales [MCZ, examined]. New

synonymy.
Orasemorpha tridentata — Boucek, 1988:519.
Orasemorpha wheeleri — Boucek, 1988:519.

TYPE MATERIAL

Lectotype of E. tridentata (designated as holotype by
Dahms, 1984), 9, "HOLOTYPE/ Hy. 3287/ E. C. D.
1985." "Eucharomorpha/ tridentata Girault/ Type." [GH|
"LECTOTYPE/ Orasemorpha/ tridentata (Grit)/ Det.
Heraty, '90." Smashed body on point, no head.
Additional material described by Dahms (1986) was not
discussed in the original description. The slide labelled
"Eucharomorpha tridentata Gir., 9 type" (covered over
by label that docs not state type status) contains the head,
parts of antennae, and 1 forewing. All of the wings are
present on the lectotype indicating that the parts on the
slide were taken from a different specimen. See Dahms
(1986) for additional information on type material.

Lectotype of E. wheeleri (here designated). 9 ,
"Wentworth Falls/ N. S. W. 2800'|854 m]/ Dec. 22. \31."
"Harv. Austr. Exp./ W. M. Wheeler." "M.C.Z./ Type/
1962 [red label]." "Type scries/ Eucharomorpha/
wheeleri/ Brues." "LECTOTYPE/ Orasemorpha/ wheeleri
(Brucs)/ Dct. Heraty "90." Complete specimen, minuten
mounted. Paralcclotypcs, 39 9 with same data. Brucs
(1934) referred to "type and numerous paralypes" but

only the material mentioned above was located at the
MCZ.

DIAGNOSIS

Recognized by the following: marginal fringe of the
forewing lacking, speculum present, gaster almost entire-
ly bare and smooth, and mesosomal dorsum almost
entirely smooth or very weakly coriaceous. This species
is closely related to Orasemorpha goethei. O. myrmicae,
and O. xeniades. It may be distinguished from O. goethei
and O. myrmicae by the presence of a speculum, dorsal
sculpture, and broader forewing, and from O. xeniades by
the presence of a marginal fringe, larger LOL/OOL ratio
(1-1.4) of female, and generally narrower scutellum.
Additionally, it can be separated from O. myrmicae and
O. xeniades by the tridentate mandible (Fig. 56).
However, this last state is unknown for O. goethei.

FEMALE

Length, 1.7-2.4 mm. Head, mesosoma, and gaster,
including petiole and coxae, black with strong greenish
reflections; scape to anellus light brown, pedicel darker
dorsally. flagellum dark brown; mandible dark brown;
femora mostly dark brown to black, apex and rest of legs
dark yellowish brown, apical tarsomere brown. Wings
weakly infuscate, venation pale brown.

Head subtriangular, occiput broadly emarginate; tem-
ples large and abrupt behind eyes in dorsal view; LOL
1-1.4X OOL. Face broadly rounded, completely shallow
reticulate; scrobal depression narrow and broadly rounded
with finer reticulate sculpture, partially including median
ocellus. Eyes bare and protuberant, their inner margins
slightly diverging; separated by 2.3-2.4x their height.
Malar space 1.0-1.2x height of eye. Clypeal region weak-
ly reticulate to scabriculous, lateral margins broadly
impressed; supraclypeal area bulging. Labrum 4-digitate,
digits short and stout. Mandibles 3/3 dentate. Antenna 12-
segmented; anellus small; flagellum I.3x height of head:
funicular segments with dense short adpressed setae
(setae not raised above surface, segments appearing bare),
F2 1.8-2.2X as long as broad, l.l-1.4x F3, following seg-
ments subequal in length and of increasing width to apex;
clava obconical and as long as preceding 2 segments.

Mesosoma with midlobe of scutellum finely reticulate
medially with weakly rounded interstices, the lateral mar-
gins reticulate to smooth; lateral lobe glabrous and
strongly swollen, flattened along anterior dorsal surface;
axilla swollen and glabrous (not narrow as described by
Girault), with posterior margin sharply declivous to level
of scutellum; scutellum weakly reticulate to smooth,
sculpture sometimes appearing weakly striate anterolater-
ally. Notauli sharply and deeply impressed, narrowly
crenulate. SSS weakly crenulate. Scutellum rounded dor-
sally, I.5-1.9X as long as broad; frenal line a narrow

45

grcHive. frenal area snux)ih to coriaceous, vaguely emar-
ginate medially. Propodeuni evenl_\ rounded and weakly
reiiculate lo smooth; callus and metepimeron swollen and
glabrous, separated by sharp metepimeral sulcus; callus
v,nh wcAk callar nib. Upper mescpimeron swollen and
glabrate. lower mescpimeron Hat and weakly reticulate.
with transepimeral sulcus weak; mesepisternum evenly
reticulate, smooth ventrally. Prepeclus broadly triangular,
reticulate with pronounced posterior flange. Pronotum
reticulate. Proepisternum reticulate to smooth posteriorly.
Coxae smooth to weakly reticulate; femora weakly imbri-
cate: hind leg with apex of femur and tibia moderately
pilose with short adpressed setae (hardly visible).
Forewing 2.4-2.6x as long as broad; basal area and
speculum bare, speculum may be closed proximally by
sparse setae; costal cell pilose posteriorly; disc densely
short pilose, marginal fringe absent; stigmal vein narrow.
1.8-2.5X as long as broad and almost perpendicular to
wing margin.

Metasoma with petiole broader than long, finely retic-
ulate or smooth anteriorly (Fig. 58). Gastral terga very
weakly coriaceous with sparse, fine, adpressed setae,
almost glabrous; Ms^ with deeply impressed crenulate
constriction, anterior region glabrous and semicircular
with sharp posterior margin, extending forward under
petiole. Hypopygium with few minute sublateral setae at
apex. Ovipositor typical for genus; first valvula with 3
small teeth along lateral line; second valvula with several
strong lateral teeth.

MALE

Unknown.

VARIATION

There is little morphological variation among the material
examined. The degree of sculpture on the dorsum of the
mesosoma varies from almost complete on the midlobe of
the mesoscutum to almost absent from the entire scutel-
lum, and the dorsal apical region of the scutellum is
always bare.

BIOLOGY

Reared from Pheidole proximo Mayr (Brues, 1934).
Collections of this species have been taken from a dry
sclerophyll Eucalyptus forest and along a riverbank near
Gordonvale.

DISTRIBLTION

Eastern Australia (T. Fig. 275).

MATERIAL EXAMINED

Alistral.ia: A.C.T.: Black Mtn. January (19, ANIC);
New South Wales: Nerriga, 5 km NE. 600 m. dry sclero-
phyll Eucalyptus forest, January to February (19. AEI);

Queensland: Brisbane. 22 km NW: Graham Range,
Babinda; Mt Glorious N. P., 630 m; Mt Nebo; .Mulgrave
Riv.. Bruce Highway. '/- km S Gordonvale [along river];
October and December to April (219 9. AEI, ANIC.
CNC. QMB.TAMU).

Orasemorpha goethei (Girault)

Eucharomorpha ^octhci Girault, 1934:2 (307]. Victoria,
Australia (MUV. examined]. Dahms. 1984:654 (notes
on type material).

Orasemorpha goethei — Boucek. 1 988:5 1 9.

TYPE MATERIAL

Holotype, 9, "Melbourne. V./ F. E. Wilson/ 3.XII.27."
"1452[?]/ Type 9 ." "Eucharomorpha/ 9 goethei Girault/
Type." "F. E. Wilson/ Collection." "ENT - 578."
Complete mesosoma and gaster mounted on point. The
slide with the head should be at the Queensland Museum
but in its place is a note that the slide is "with Mound."
Dahms (1984) states that the type on the slide was
remounted at the BMNH.

DIAGNOSIS

Similar to O. tridentata and O. myrmicae based on the
following: gastral terga almost bare, and marginal fringe
of forewing absent. The species is distinguished by hav-
ing the mesosoma dorsum (midlobe and scutellum) and
gastral terga almost completely smooth, and speculum
present.

FEMALE

Length unknown. Mesosoma. petiole and coxae dark
brown to black, mesosoma with very faint greenish
reflections; gaster and basal two-thirds of femora dark
brown; apex of femora, tibiae and basal tarsomeres yel-
lowish brown; antennal flagellum dark brown, scape
lighter brown, pedicel intermediate. Wings hyaline, vena-
tion pale brown.

Head, missing from only known female.

Mesosoma with midlobe of mesoscutum smooth with
weak superficial sculpture anteriorly and scutellum
glabrous; lateral lobe polished and strongly swollen; axil-
la smooth dorsally. broad, and rounded. Notauli narrow
and deeply impressed, finely crenulate. SSS weakly
crenulate. Scutellum rounded dorsally. on lower plane
than axilla; frenal line narrow dorsally and very finely
crenulate. frenal area weakly rugose and broadly rounded.
Propodeuni evenly rounded, finely and weakly reticulate
medially, slightly impressed above base of coxae; callus
and metepimeron glabrous, with small callar nib.
Mescpimeron smooth, transepimeral sulcus distinct;
femoral groove weakly reticulate: mesepisternum
glabrous. Prepectus triangular, smooth ventrally to reticu-

46

late dorsally, with glabrous dorsal and posterior flanges.
Pronotum broadly rounded laterally and coriaceous.
Proepisternum coriaceous. Coxae weakly coriaceous;
femora smooth to lightly scabriculous apically on lateral
surface; hind leg with tibia and femora densely short-
setose. Forewing 3. Ox as long as broad; basal area bare;
speculum present; costal cell bare; disc moderately pilose
with short fine setae, no marginal fringe; stigmal vein
narrow and perpendicular to wing margin.

Metasoma with petiole 0.2-0.3x as long as broad,
glabrous. Gastral terga almost completely smooth, with
only faint surface sculpture; Ms^ with constriction broad
and weakly crenulate, anterior region smooth.
Hypopygium bare. Ovipositor subapically expanded; first
valvula with subapical ridge followed by 4 sharp lateral
teeth; second valvula with 7 to 8 lateral teeth.
(Gonostylus hidden).

MALE

Length, 1.7-1.9 mm. Black, lower face and mesosoma
laterally with faint greenish or violaceous reflections;
antenna dark brown to black; femora, tibiae, and apical
tarsomeres dark brown, apices of femora and tibiae, and
basal tarsomeres yellowish brown. Wings hyaline.

Head subtriangular, eye only slightly protuberant.
Malar space l.l-1.3x height of eye. Labrum 4- or 5-digi-
tate. Antenna 12-segmented; anellus present; flagellum
1.6-1.8X height of head; funicular segments with dense
subdecumbent setae, F2 1.8-2. Ox as long as broad,
1.2-1.4XF3.

Mesosoma more slender than female; scutellum com-
pletely smooth to very weakly sculptured, 1.8-2.2x as
long as broad; frenal line indistinct dorsally, frenal area
smooth to reticulate. Forewing 2.2x as long as broad;
speculum sparsely pilose; costal cell with single row of
small setae.

Metasoma with petiole 0.9-1. Ix as long as hind coxa,
0.7-1. Ox as long as propodeum, reticulate dorsally.
Gastral terga glabrous; Ms, strongly constricted, anterior
region scabriculous or smooth, cup-shaped and anterior
margin extending forward under petiole. Ms^^ broadly
rounded and setose. (Genitalia withdrawn and not dis-
.sected.)

DISCUSSION

A female in the BMNH (remnants only, labelled 1440c)
probably belongs here as it has the costal cell bare, scutel-
lum slightly indented medially, and the gastral terga
entirely smooth. A female from Tasmania (Dodges Ferry.
29.xii.l979. J. C. Cardale, ANIC) appears to be closely
related to O. goethei but differs in having the scutellum
broader, with side lobe of mesoscutum swollen and not
flattened above, and the basal flagellar segments longer
and narrower than described above. The males described

for this species exhibit a strong sexual dimorphism, which
makes it difficult to associate specimens with females of
the 3 species that lack marginal setae; the male specimens
also could be attributed to either O. tridentata or O. myr-
micae. The males are excluded from O. myvmicae by the
smooth or weakly sculptured scutellum and are consid-
ered to belong to O. goethei only because of their south-
em distribution in Australia. Males of the more northern
O. tridentata are presently unknown. The variation in
sculpture of the anterior region of Ms, is between 2 males
from Shepparton and the male from Toolomb Plateau,
which has the scabriculous sculpture.

DISTRIBUTION

Southeastern Australia (G, Fig. 275).

MATERIAL EXAMINED

Australia: New South Wales: Tooloom Plateau, via
Urbanville, 600-700 m, 8. i. 1977, I. D. Naumann, sweep-
ing low vegetation, subtropical rainforest (Id, UQIC);
Victoria: Shepparton, 15.xii.l974, 1. D. Naumann, sweep-
ing grass by creek, dry sclerophyll forest {2 6 6 UQIC).

Orasemorpha pyttalus (Walker)

Figs. 64, 66-67

Eucharis pyttalus Walker, 1846:21, 87-88. South

Australia [BMNH, examined].
Orasemorpha pyttalus — Boucek, 1988:5 19.

TYPE MATERIAL

Holotype, 9 , "Holo-/ type." "Psilogaster/ pyttalus/
Walker." "B. M. TYPE/ HYM./ 5-615." "9
Orasemorpha/ pyttalus (Walker)/ det. Z. Boucek, 1986."
Petiole and gaster missing.

DIAGNOSIS

Recognized by: entire body covered by dense erect setae,
head triangular (Fig. 66), basal antennal segments yellow-
ish brown, and forewing densely pilose with a prominent
marginal fringe.

HOLOTYPE FEMALE

Length, ca. 2.8 mm. Head and mesosoma dark metallic
green with scattered regions of dark reddish coloration on
clypeus, median line on mesoscutum. axilla, along frenal
line, and most of propodeum; coxae dark metallic green;
flagellum dark brown; scape to anellus yellowish brown;
legs yellowish brown, base of femora slightly darker.
Wings hyaline, venation pale brown.

Head triangular; occiput broadly emarginatc; temples
broadly rounded and densely short erect setose; LOL l.lx
OOL. Face relatively flat and strongh rugose, lower face

47

and vertex covered by dense erect setae (Fig. 66). frons
lateral to scrobal depression sparsely setose; scrobal
depression broadl> impressed and rugose. Eyes slightly
protuberant, covered by dense erect setae, and separated
by 2.3x their height. Malar space l.2x height of eye.
Clypeal region lightly sculptured and dense setose, lateral
margin of clypeus deeply impressed; supraclypeal area
slightly bulging. Labrum 4-digitate. Mandibles 3/2 den-
tate. Antenna 12-segmented (Fig. 64); scape densely
pilose; anellus present; flagellum l.4x height of head;
funicular segments scabriculous with dense short setae
along entire length. F2 l.9x as long as broad. I.5x F2,
following segments subequal in length; clava obconical,
as long as preceding 2 segments.

Mesosoma with dorsum rugose to rugulose and dense-
ly erect pilose; lateral lobe of mesoscutum lightly sculp-
tured and shining; axilla swollen and smooth; scutellum
weak rugulose. Notauli narrowly impressed. SSS narrow
and crenulate. Scutellum as long as broad; frenal line nar-
row and glabrous, frenal area rugulose. Propodeum rugu-
lose with several irregular carinae converging on base of
petiole; callus glabrate and swollen, with small callar nib.
Upper mesepimeron glabrous and swollen, lower
mesepimeron light rugulose, with transepimeral sulcus
distinct; mesepisternum light rugulose. Prepectus triangu-
lar, rugulose with weak dorsal and posterior flanges.
Pronotum evenly rugulose. Proepistemum lightly sculp-
tured and sparsely setose. Fore and mid coxae almost
smooth and densely setose ventrally, hind coxa weak
rugulose to smooth and setose ventrally; femora smooth
with dense semi-erect setae. Forewing 2.4x as long as
broad; basal area bare with band of setae extending along
cubital vein to base; speculum absent; costal cell pilose;
disc densely pilose, marginal fringe distinct; stigmal vein
l.lx as long as broad, constricted at base.

Metasoma missing, probably similar to O. erihotes.

ly related species that has the wings completely and dense-
ly pilose and the scape entirely black and metallic.

DISTRIBLTION

Eastern Australia (P, Fig. 275).

MATKRIAL EXAMINED

AU.STRAIJA: New South Wales: Sams Rd. Bulahedelah
S.F. (32.24S 152. I4E). 8.xii.l986. M. M. Stevens and F.
E. Frindle (29 9. ANIC); Queensland: Mt Glorious.
17. xi. [no collector] (19. AEI); Stanthorpe. 15. i. 1983. Z.
Boucek(l9.BMNH).

Orasemorpha sparsepilosa sp. nov.

Figs. 65. 68-69

TYPE MATERIAL

Holotype, 9. "NSW: Moree/ 1.78. G. Brown." "9
Orasemorpha/ sparsepilosa sp. n./ det. Z. Boucek. 1986."
"HOLOTYPE/ Orasemorpha/ sparsepilosa/ Heraty."
Deposited in BMNH.

Paratypes: Australia: New South Wales: same data as
holotype (2 9 9 , 2d d . BMNH).

ETYMOLOGY

From sparse and pilose, referring to the pilosity of the
body.

DIAGNOSIS

Recognized by: body covered by moderately dense, fine,
erect setae (Fig. 69). head subtriangular with broadly
rounded gena, basal antennal segments light yellow to
nearly white, forewing with fine setae (hardly visible),
and marginal fringe restricted to the postero-apical mar-
gin of wing (Fig. 68).

MALE

Unknown.

VARIATION

One specimen from Queensland (Mt Glorious. AEI) is
tentatively assigned to this species. This specimen is close
to the type except for the following: lacks any of the red-
dish coloration; LOL less than OOL; scutellum with lon-
gitudinal rugae and entire basal area completely (but
sparsely) setose; petiole transverse and weakly rugose lat-
erally with dense erect setae; gastral terga smooth with
dense erect setae; Ms^ with strong crenulate constriction,
and rest of gaster typical. The 2 specimens from New
South Wales are similar to the type but have a dark scape
and pedicel, wings completely and densely setose to base,
and mesepimeron glabrous. Another male (Australia:
New South Wales: Oxford, in BMNH) represents a close-

FEMALE

Length, 3.8 mm. Head dark blue, lower face with patches
of violet or reddish colour, vertex more greenish; mesoso-
ma dark blue-green with strong reddish coloration on
mesoscutum and axilla; dorsomedial regions of lateral
lobe, axilla, and apex of scutellum with stronger bluish
colour laterally; petiole black with reddish reflections;
gaster dark brown with greenish reflections; coxae mostly
dark brown with greenish rencclioiis, apices yellowish
brown; scape to anellus light yellow to white, rest of fla-
gellum brown; mandible light brown; legs beyond coxae
light yellowish brown to yellow. Wings hyaline, venation
clear yellowish brown.

Head subtriangular; occiput broadly emarginate; tem-
ple broadly rounded with short erect setae; LOL 0.8x
OOL. Face strongly rugose to rugulose. lower face and
vertex evenly covered by moderately dense erect setae;

48

scrobal depression shallow and broadly impressed with
irregular striae arching over toruli. Eyes slightly protuber-
ant, with short erect setae and separated by 2.5-3.3x their
height. Malar space 1.2-1.6x height of eye. Clypeus near-
ly smooth with sparse erect setae, lateral margin deeply
impressed to tentorial pit; supraclypeal area finely rugu-
lose and swollen. Labrum 7-segmented. digits with 2-3-2
formula. (Mandibles hidden.) Antenna 12-segmented
(Fig. 65); anellus present; flagellum 1.3-1.4x height of
head; funicular segments with dense subdecumbent setae,
F2 1.6-1.9X as long as broad, 1.4-1.5x F3, following
segments subequal in length, decreasing slightly in width
to apex; clava elongate, almost as long as preceding 2
segments.

Mesosoma with dorsum weakly rugulose and with
moderately dense covering of short erect setae (Fig. 69);
lateral lobe of mesoscutum strongly swollen, weak rugu-
lose laterally and smooth dorsally; axilla swollen and
glabrate, rugulose laterally; scutellum almost entirely
smooth dorsally with weak irregular longitudinal rugae
basally (Fig. 69). Notauli deeply and broadly impressed,
narrowly crenulate. SSS crenulate. Scutellum as long as
broad; frenal line narrow and glabrous, frenal area rugu-
lose. Propodeum rugulose with several irregular carinae
diverging from propodeal midline, ventrally with several
carina converging to base of petiole; callus smooth or
weakly sculptured and bare, swollen, callar nib weak or
absent; metepimeron separated by deep metepimeral sul-
cus. Upper mesepimeron swollen and glabrous, lower
mesepimeron light rugulose, distinct transepimeral sul-
cus; mesepisternum finely reticulate. Prepectus broadly
triangular, rugulose with strong dorsal and posterior
flanges. Pronotum reticulate-rugose. Proepistemum retic-
ulate and bare. Coxae smooth with sparse semi-erect setae
ventrally; femora and tibiae smooth with dense semi-erect
setae. Forewing 2.1-2.3x as long as broad; basal third of
wing, including basal area and speculum, bare; costal cell
with few minute setae apically; disc with short fine setae
(difficult to discern), marginal fringe restricted to extreme
posterior apical margin and along marginal vein (Fig. 68);
stigmal vein 1.4x as long as broad with strongly narrowed
base, angled about 45 degrees to dorsal margin.

Meiasoma with petiole transverse, glabrous medially
and finely sculptured with short erect setae laterally.
Gastral terga weakly coriaceous with moderately dense
covering of fine semi-erect setae; Ms^ with constriction
broad and weakly crenulate. Hypopygium with few short
sublateral setae at apex. Ovipositor typical for genus; first
valvula with 3 sharp teeth forming lateral line; second
valvula with 9 to 10 lateral teeth.

MALE

Length, 3.0-3.6 mm. As in female except: entire coxae
dark; scape, pedicel, and basal third of femora brown.

Antennal flagellum 1.7x height of head, slightly tapering
from base to apex. Metasoma with petiole 2-2. 3x as broad
as long, 0.5-0. 6x as long as hind coxa, twice as long as
propodeum; weakly rugulose dorsally with few erect
setae. Ms., with constriction smooth, anterior region
strongly swollen and glabrous. Ms^, broadly rounded and
setose. Genitalia with parameres short and stout.

DISCUSSION

A female from Queensland is similar to the type material
except as follows: funicle with 7 segments; scape yellow-
ish brown, pedicel and anellus brown; flagellum black;
mandibles 3/3 dentate; mesepisternum rugulose; wing
venation pale brown, wing setae slightly darker and more
visible; head and mesosoma darker green, and legs
orange-brown. The reduction in number of funicular seg-
ments may support the reduced number of antennal seg-
ments reported for O. myrmicae (see discussion for that
species).

DISTRIBUTION

New South Wales and Queensland, Australia (S, Fig.

275).

MATERIAL EXAMINED

Australia: Queensland: Rockhampton. 26-27.xi.l967. J.
and M. Sedlacek (19, BPBM).

Orasemorpha varidentata (Girault)

Fig. 63

Eucharomorpha varidentata Girault, 1936:3 [324].

Australia: Tasmania [QMB, examined]. Dahms,

1986:622 (notes on type material).
Orasemorpha varidentata — Boucek, 1988:519.

TYPE MATERIAL

Holotype. 9, "G. H. Hardy/ Lindisfarne/ 17/ 1/ 1914/
43." "HOLOTYPE/ T. 10049/ E. C. D. 1985."'
"Eucharomorpha/ varidentata Girault/ Type" [GHj.
Mesosoma with hind wing and mid leg on point, fore and
hind leg mounted separately on point; head, antenna, and
basal half of forewing mounted separately. See Dahms
(1986) for details of type material.

DIACJNOSIS

Recognized by having the mesosoma broad and strongly
rugose (Fig. 63) with strong iridescent reflections, basal
flagellar segments yellowish brown, prepectus narrow,
coxae completely setose, basal area of forewing bare,
speculum absent, and first valvula with 3 sharp icoih
along the lateral line.

49

HOl.OIVI'KKKMALK

l.ciigili unknown. Mesosoma black \\ ith strong iridescent
(reddish green and violet) reflections; coxae and gaster
dark brown with reddish rellections; scape to anellus yel-
lowish brown, rest of nagelluni dark brown; legs yellow-
ish brown, femora slightly darker basally. Wings hyaline,
venation light brown.

Head (squashed on slide) with labruni 4-digitate.
mandibles 3/2 dentate, antenna 12-segmented. anellus
present; funicular segments densely short setose, F2 1 .6x
as long as broad. l.Ix F3, the following segments sube-
qual in length; clava slightly longer than preceding 2 seg-
ments.

Mesosoma with dorsum coarsely rugose, interstices
shallow and rounded; axilla and lateral lobe of mesoscu-
tum glabrous and strongly swollen, axilla strongly angled
posteriorly to meet scutellum. Notauli deeply and narrow-
ly impressed, strongly crenulate. SSS strongly crenulate.
Scutellum broad and rounded, 1.3x as broad as long; fre-
nal line broad and glabrous dorsally, frenal area rugose,
depressed medially. Propodeum evenly rounded, coarsely
rugose with smooth septa and moderately pilose covering
of short fine setae (angled dorsally); callus glabrous and
swollen, metepimeron and anterior callar area weakly
rugulose, callus with weak callar nib. Upper mesepimeron
smooth and swollen, lower mesepimeron flat and weakly
rugose, transepimeral sulcus weakly impressed; metepis-
temum finely reticulate to rugose ventrolaterally, smooth
ventrally. Prepectus triangular and rugose-areolate, nar-
row and 0.4x as long as high, with strong posterior
flange. Pronotum rugose-areolate. Proepisternum rugu-
lose. Coxae smooth and completely setose (pilosity of
hind leg not visible on specimen). Forewing with basal
area bare; speculum absent (pilose); costal cell pilose; rest
of wing densely pilose; stigmal vein as long as broad, nar-
rowed basally.

Metasoma with petiole transverse, rugulose dorsally.
Gastral terga smooth to coriaceous with moderately dense
covering of fine adpressed setae; Ms^ with constriction
broad and finely crenulate, anterior region triangular.
Hypopygium bare. Ovipositor typical for genus; first
valvula with strong subapical ridge and 3 sharp lateral
teeth, second valvula with 6 blunt lateral teeth.

MALE

Unknown.

DISTRIBUTION

Tasmania, Australia (V, Fig. 275).

Orasemorpha didentata ((lirault)

l-igs. 74-79

EiH haromorpha didentata Girault. 1940:325-326.

Australia: A.C.T. [ANIC, examined). Dahms,

1983:221-223 (notes on type material).
Orasemorpha bidentata — Bouoek, 1988:519. Misspelling.

TYPE MATERIAL

Lectotypc (here designated), 9 . "F.C.T. Aust./
BlundelTs/ 21.1.1931/ L. F. Graham." "Eucharomorpha
viridis [viridis pencilled out]/ 9 27.4.31/ L. F. Graham
del."' "Eucharomorpha/ PARATYPE/ didentata, Gir. [blue
label]." "LECTOTYPE/ Orasemorpha/ didentata (Grit)/
Det. Heraty '90." Left antenna missing beyond pedicel.
Paralectotypes, 5 9 9. 2c5 d labelled as in lectotype. Two
specimens bear pink labels with "HOLOTYPE" and
"ALLOTYPE." The specimen labelled "holotype" bears
Girault's handwritten determination label, but was not
chosen because of its missing head. No holotype was des-
ignated in the original publication. See Dahms (1983) for
additional notes on the type material.

DIAGNOSIS

Recognized by: midlobe of mesocutum rugose and dor-
sum with sparse to dense covering of short subdecumbent
or adpressed setae (Fig. 77), disc of forewing, including
speculum, pilose with sparse setae continuing to base
(Fig. 79). and gastral terga smooth with dense semi-erect
setae. Differs from O. erihotes by having sparse short
setae on the eye (Fig. 76). hind coxa almost smooth, and
hind femur and tibia with dense, elongate, and semi-erect
setae (Fig. 78).

FEMALE

Length, 2.3-2.8 mm. Head and mesosoma dark green
with areas of reddish reflections prominent on clypeal
region, ocellar-ocular area, midline of mesosculum, axil-
la, frenal line, and propodeum; gaster and coxae dark
brown with faint greenish reflections; scape to anellus
yellowish brown, rest of flagellum dark brown; mandible
dark brown; legs yellowish brown, femora darker brown
in basal half. Wings hyaline, venation pale brown.

Head triangular, gena straight; occiput broadly emar-
ginate; temples broadly rounded with short subdecumbent
setae; LOL 1.0-1.3x OOL. Face strongly rugose, with
subdecumbent pilosity; scrobal depression shallow and
broadly impressed, strongly rugose. Eye slightly protrud-
ing, with sparse short erect setae, as long as facial setae,
eyes separated by 2.4-2.9x their height. Malar space
1.1-1.3X height of eye. Clypeal region weakly sculptured,
lateral margin of clypeus strongly impressed; supra-
clypeal area swollen. Labrum 4- to 7-digitate. digits long.
Mandibles 2/2, 3/2, or 3/3 dentate. Antenna 12-segment-

50

ed; scape densely setose dorsally; anellus present; flagel-
lum 1.4-1.8X height of head; funicular segments
scabriculous with dense semi-erect setae; F2 1.8x as long
as broad, l.l-1.3x F3, following segments subequal in
length, gradually narrowing to clava; clava elongate, as
long as preceding 2 segments.

Mesosoma with dorsum weak rugose, interstices shal-
low and broadly rounded, with short subdecumbent setae;
lateral lobe and axilla swollen and glabrate. Notauli mod-
erately impressed and narrowly crenulate. SSS crenulate.
Scutellum subquadrate to elongate, 1.0-1.6x as long as
broad with elongate median depression; frenal line broad
and glabrous dorsally, frenal area weak rugose.
Propodeum strongly rugose with weak verrucose surface;
callus swollen and glabrous with small callar nib;
metepimeron smooth. Upper mesepimeron glabrous,
lower mesepimeron weak reticulate-rugose, transepimeral
sulcus distinct; mesepistemum weak reticulate-rugose and
smooth ventrally. Prepectus triangular, rugose with dorsal
and posterior verrucose flange. Pronotum and proepister-
num rugulose. Coxa smooth with sparse semi-erect setae;
hind femur and tibia smooth with dense, elongate, semi-
erect setae (Fig. 78). Forewing 2.3-2.5x as long as broad;
basal area sparsely setose along impression of cubital
vein, otherwise bare; speculum lacking (pilose); costal
cell pilose; disc densely pilose with complete marginal
fringe; stigmal vein 1.2-2.7x as long as broad, with or
without basal narrowing.

Metasoma with petiole transverse, 0.3-0.5x as long as
broad, posterior region relatively long and weakly reticu-
late, anterior region as long as broad and glabrous.
Gastral terga smooth with dense subdecumbent to semi-
erect setae, basal terga glabrous medially; Ms^ with con-
striction broad and crenulate, anterior region semicircular.
Hypopygium bare. Ovipositor elongate, only slightly
curved forward; first valvula with weak subapical ridge
and 3 lateral teeth; second valvula with 7 sharp lateral
teeth.

MALE

Length, 2.6-2.9 mm. Body dark brown to black, head and
mesosoma with faint blue-green reflections, prominent on
lower face, scutellum. and ventral areas of mesosoma;
coxae, gaster, and femora dark brown; gaster with faint
reddish reflections; antenna dark brown; mandible light to
dark brown; apex of femora and rest of legs light brown.
LOL ().9-1.2x OOL. Eyes with sparse minute setae, sepa-
rated by 2.3-2.5X their height. Malar space l.()-1.2x
height of eye. Antenna 12-scgmented; flagellum l.4-l.8x
height of head; F2 1.6-1.2x as long as broad, 1.0-1.2x
F3. Midlobe of mesoscutum more finely rugulose; scutel-
lum smot)th apically, without medial depression, 1.4-1.7x
as long as broad. Forewing 2.2-2.3x as long as broad.
Metasoma with petiole as long as broad, 0.6-0. 8x as long

as hind coxa, 0.5-0.7x as long as propodeum; finely retic-
ulate dorsally, glabrous ventrally. Ms^ with constriction
weakly striate, anterior region strongly swollen and
glabrous. Ms^ broadly rounded and setose. Genitalia hid-
den.

VARIATION

Specimens are generally very similar. One female from
Queensland (North Pine River) has only minute eye setae
and distinct blue coloration of the head and mesosoma,
but otherwise agrees with the other material. The series of
males from Ainslie are larger and have a more distinct
green coloration, but 1 small male is almost identical in
colour and size to the paralectotype males.

BIOLOGY

The only known plant association from the collection
records is from the foliage of Protea sp. (Proteaceae).

DISTRIBUTION

Eastern Australia (D, Fig. 275).

MATERIAL EXAMINED

Australia: A.C.T.: Ainslie; Canberra, ex swimming
pool; Canberra, Black Mtn; January to February (599,
7(5 6 , AEI, ANIC, BMNH); Queensland: Lever's
Plateau, via Rathdowney; Nth Pine R[d]; Stanthorpe;
Brisbane; December to January and March to April
(49 9. BMNH, UQIC); South Australia; Adelaide;
Kangaroo Ls.; Kersbrook, ex Protea sp. foliage;
December to January (799, ANIC, UQIC).

Orasemorpha eribotes (Walker)

Figs. 70-71, 73, 187, 217, 256-257

Eucharis eribotes Walker, 1839:13-14. Australia:
Tasmania and New South Wales [BMNH. examined].

Eucharomorpha duhia Girault, 1913b:95. Australia:
Tasmania |SAMA, examined]. Girault. 1915:230
(redescription). Dahms, 1983:233 (notes on type mate-
rial). New synonymy.

Eucharomorpha fuscipes Girault, I913b:95. Australia:
Tasmania |SAMA, examined). Girault. 1915:229
(redescription). Dahms, 1984:642 (notes on type mate-
rial). New synonymy.

Eucharomorpha rlridis Girault, I913b:95. Australia:
Tasmania (SAMA, examined). Girault. 1^)15:229
(redescription). Girault. 1929:331 (new specimen and
additional notes). Dahms, 1986:640-641 (notes on
type material). New synonymy

Psiloi^asteroides eribotes — Girault, 191 3b:94.

Eucharomorpha parti^iUihra Girault. 1940:324. Australia:
Victoria [QMB. examined]. Dahms, 1986:386 (notes

51

on type material). New synonymy.

Epinu-ia\ica ciihotcs — Heclqv ist. 1 97X:243.
Oruscmorpha — Boufek. 1988:519. All above species
names were transferred to this genus.

TYPE MATKRI AL

Lectotype of Eiuharis erihotes (designated Boudek,
1988). 6 , "1440b." "LECTO-/ TYPE." "Co-/ type."
"Psilogaster/ Eribotes/ Walker." "B. M. TYPE/ HYM./
5.619." "(?d) Orasemorpha/ eribotes (Walk.)/ LECTO-
TYPE/ det. Z. Boucek. 1986." Head and thorax with
wings and 1 foreleg mounted on card, antennae missing.
Paralectotype, 6 (headless), same data except "1440a."
The lectotype is from Hobart. Tasmania, and the paralec-
totype is from Sydney, New South Wales. An additional
female labelled as "1440c" (Boucek. 1988) belongs to O.
^oethei and is treated under that species.

Holotype of Eucharomorpha duhia, 6 "Swansea/ Tas.
Lea." "Eucharomorpha/ viridis, fuscipes/ and dubia 6
types [GH]." "T. 1283-4-5./ Eucharomorpha/ viridis Gir/
fuscipes Gir./ dubia Gir ["No. 3" underneath name]/
Tasmania/ see Note Book and Slide" [GH]. One female
and 3 males mounted on card and numbered from 1 to 4,
the holotype of E. duhia is specimen number 3; right
antenna in glue, left antenna missing, left hind wing
intact. See Dahms (1983) for additional information on
type material. The locality on the label is for E. viridis
(Dahms, 1983); the published locality for E. dubia was
Hobart, Tasmania (Girault, 1913a).

Lectotype of Eucharomorpha fuscipes (here designat-
ed), 6 , specimen number 4 from same card listed above;
left antenna and hind tarsi missing. My label "LECTO-
TYPE/ Orasemorpha/ fuscipes (Grit)/ Det. Heraty '90"
added to specimen. Paralectotype, 6, specimen number
2; antenna removed, left hind legs and left wings
removed, covered in mold. Published locality was Hobart,
Tasmania (Girault, 1913a).

Holotype of Eucharomorpha viridis, 9 , specimen
number 1 from same card listed above; left hind wing,
hind legs, and left coxa missing, gaster and petiole
mounted separately with ovipositor exposed. Published
locality was Swansea, Tasmania (Girault, 1913a).

Lectotype of Eucharomorpha partiglahra (here desig-
nated), 9 "Swan Hill, Vic./ 4.1.31/ F. E. Wilson" "SYN-
TYPE/ T.9423/ E. C. D. 1985" "Eucharomorpha 9/ par-
tiglahra Gir. [with red "Paratype" written over label]"
[GH]. "LECTOTYPE/ Orasemorpha/ partiglabra (Gir.)/
Det. Heraty '90"; tip of left antenna broken off. no mid
legs. Paralectotypes, 2 9 9, QMB type numbers T.9421
and T.9422. See Dahms (1986) for additional notes on
type material.

NOTK.S ON SYNONYMY

BouCek (1988) suggested that a synonymy might be pro-
posed for "'dideiilaia. partii>lahra, pyltalus. varidentala
and viridis. if not more." Riek (unpublished notes. ANIC)
proposed a synonymy of viridis, fuscipes, duhia + vari-
dentata; partif^lahra + didentata; and pyttalus + viridis.
There has been considerable confusion over the species
limits in this group, in my opinion, O. varidentala. O.
pyttalus, and O. dideniata are valid species for reasons
discussed in their respective diagnoses. The holotype of
O. duhia has a 1 2-segmented antenna, not 1 1 -segmented
as described by Girault (1913a), and is virtually identical
to that of O. fuscipes. Differences between O. viridis and
O. fuscipes or O. duhia are sexual and based on darker
coloration of the scape and body colour, and smaller
mesosoma. Specimens from Tasmania are more similar to
the lectotype of O. erihotes and are generally darker in
colour, have the midlobe of the mesoscutum. scutellum,
and coxae scabriculous, and the eyes completely bare.
Specimens of O. erihotes collected from New South
Wales and Victoria (including the paralectotype) have
weaker rugulose to smooth sculpture. Differences
between O. erihotes and O. viridis in head shape, pres-
ence or absence of minute eye setae, setation of the wing
base, and coloration are not consistent over the range of
specimens.

Orasemorpha partiglahra appears to be a larger, dark
reddish form of O. erihotes with a verrucose-foveate
prepectus, tridentate mandibles. 7-digitate labrum, and
finely sculptured petiole. Within the paralectotype series
of O. partiglahra, the prepectus is smoothly foveate, the
head and mesosoma are distinctly green in colour, and the
bases of the femora are slightly darkened. In additional
material that could be partially allocated to O.
partiglabra, the labrum was 4- or 5 -digitate. The lecto-
type of O. erihotes has a small intermediate tooth on the
left mandible giving it a partial 3/3 dentition (Fig. 73).
The mandibles and labral digits of the closely related O.
didentata are highly variable and range from 2/2 to 3/3
and from 4 to 7 labral digits.

DIAGNOSIS

Recognized by: midlobe of mesoscutum strongly rugose
to scabriculous (Fig. 70). mesosomal dorsum of female
with sparse to dense covering of short subdecumbent or
adpressed setae (Fig. 217), disc of forewing including
area of speculum pilose, and gastral terga weak to strong
coriaceous with moderately dense covering of fine subde-
cumbent or adpressed setae (Figs. 256-257). Differs from
O. didentata by having the eyes bare (Fig. 73) or some-
times with minute setae in females (Fig. 187), and hind
femur and tibia with dense, short, subdecumbent setae.

52

FEMALE

Length, 2.7-3.5 mm. Head and mesosoma dark green to
black with strong metallic reflections, but sometimes
bluish or reddish in colour; coxae dark brown with metal-
lic reflections; gaster black with green or red reflections;
scape to anellus yellow to light brown, rest of flagellum
dark brown to black; basal half of femora weakly infus-
cate to dark brown. Wings slightly infuscate, venation
pale brown.

Head subtriangular; occiput shallowly emarginate;
temple broadly rounded with short subdecumbent setae;
LOL 0.8-1.3X OOL. Face strongly rugose with sparse
short subdecumbent setae; scrobal depression broad and
evenly impressed, rugulose, often with irregular curved
carinae arching over toruli. Eye slightly protruding (Fig.
187), bare or with minute setae, eyes separated by
2.2-2.5X their height. Malar space 1 .0-1 . Ix height of eye.
Clypeus weakly rugose and moderately setose, lateral
margins deeply impressed at tentorial pits; supraclypeal
area strongly swollen medially. Labrum 4- to 7-digitate,
digits long. Mandibles 3/2 or 3/3 dentate. Antenna 12-
segmented; scape densely setose dorsally; anellus present;
flagellum 1.3-1.5x height of head; funicular segments
scabriculous with dense semi-erect setae, F2 1.6-2. Ox as
long as broad, l.l-1.4x F3. following segments subequal
in length, equal in breadth to clava; clava obconical,
about equal in length to preceding 2 segments.

Mesosoma with midlobe of mesoscutum evenly rugu-
lose to scabriculous, scutellum often smooth medially and
often with weak median depression, entire dorsum cov-
ered by short subdecumbent setae; lateral lobe of mesos-
cutum and axilla swollen and only lightly sculptured,
smooth dorsally. Notauli deeply impressed and weakly
crenulate. Scutellum 1.4-1.7x as long as broad; frenal
line broad and glabrous dorsally, frenal area rugulose,
weakly impressed medially. Propodeum broadly rounded,
coarsely rugose-striate laterally, and lightly sculptured
medially, with striae converging to midline; callus
swollen and glabrous; metepimeron rugulose. Upper
mesepimeron weak rugulose to glabrous, lower
mesepimeron reticulate to scabriculous, transepinieral
sulcus distinct; mesepisternum reticulate to scabriculous
and smooth ventrally. Prepectus broadly triangular and
rugose to rugulose, posterior flange finely sculptured.
Pronotum and proepisternum weakly rugulose. Coxae
bare dorsally, weakly rugulose to scabriculous; hind
femur and tibia finely sculptured and covered by dense,
short, subdecumbent setae. Forewing 2.3-2.5x as long as
broad; basal area bare, at most with narrow band of setae
along impression of cubital vein; speculum lightly to
completely pilose; disc, including costal cell, dense pilose,
with complete marginal fringe including along marginal
vein; stigmal vein 1.4-I.Sx as long as broad, narrowed
basally and club-shaped, slightly angled posteriorly.

Metasoma with petiole transverse, with posterior
region several times broader than long and glabrous to
weak rugulose, and anterior region narrow, as long as
broad, and glabrous. Gastral terga weakly coriaceous to
smooth, with dense adpressed setae, basal terga glabrous
medially; Ms^ with constriction broad and crenulate, ante-
rior region semicircular. Hypopygium with small clusters of
short sublateral setae. Ovipositor elongate, only slightly
curved forward; first valvula with weak subapical ridge and
3 to 4 lateral teeth; second valvula with 8 sharp lateral teeth.

MALE

Length, 2.1-3. 1 mm. Body dark brown to black, head and
mesosoma with metallic reflections of green, red, or blue
(variable); gaster sometimes with faint reddish reflec-
tions; mandible dark brown; antenna dark brown to black,
scape brown to black with metallic blue or green reflec-
tions; femora dark brown with metallic reflections except
extreme apex, rest of legs dark yellowish brown, tibiae
often darker medially. Wings hyaline or very weakly
infuscate, venation pale brown.

Head shape and sculpture as in female; LOL 0.8-1.2x
OOL. Eyes completely bare (Fig. 73), separated by
2.0-2. 5x their height. Malar space 0.8-1.3x height of eye.
Antennal scape stouter than female, only reaching 0.8x
distance to median ocellus; flagellum 1.7-2. Ox height of
head; apex of F2 and F3 broadest, following segments
tapering to apex.

Mesosoma with dorsal sculpture fine rugulose to longi-
tudinally rugulose and appearing weakly striate; scutel-
lum 1.0-1. Ix as broad as long excluding frenal area (Fig.
70). Forewing 2.1-2.4x as long as broad, basal area bare.

Metasoma with petiole slightly longer than broad,
0.6-0.8X as long as hind coxae, 0.5-0.6x as long as
propodeum; finely reticulate dorsally, glabrous ventrally,
sculpture separated laterally by fine groove. Ms, with
constriction striate, anterior region strongly swollen and
glabrous. Ms^^ broadly rounded and setose. Genitalia typi-
cal for genus; paramere narrow and elongate with several
long terminal setae, median process sharp, digitus with
small marginal spines; aedeagus broad and acuminate at tip.

VARIATION

Colour variation ranges from green with patches of red-
dish iridescence on the head and mesosoma (holotype of
O. viridis) through a very dark olive green coloration, and
in some cases, a very strong bluish coloration. Much of
this variation was observed in single collection series
although some was restricted to certain isolated indi\ idu-
als. Males are darker in colour but show similar colour
patterns and variation. The basal area of the wing is bare
in all of the specimens from eastern Australia and
Tasmania, and range from bare to completely pilose in
the individuals from Western Australia.

53

B1()I.()(JV

Reared from PlwicloU' sp. (Boucek, 1988). The only plant
association for this species is "sweeping long grass, dry
sclerophyll forest" in New South Wales.

DISTRIBl TION

Eastern, southwestern Australia, and Tasmania (E, Fig.
275).

MATERIAL EXAMINED

Al'Straiia: A.C.T.: Picadilly Circus (35.22S 148.48E),
1240 m, FIT; Canberra, February, March (39 9, 36 6,
ANIC); New South Wales: Pilliga scrub, via
Coonabarabran, sweeping long grass, dry sclerophyll for-

est: McGarr's Ck. Sydney: Tooma Reserve, 20 km NNE,
Kosciusko N. P.: December to January (3 9 9, 2d d.
ANIC, BMNH. UQIC); Queensland: Kuranda;
Brookficld. nr Brisbane: Daintrec Riv.: Mt Tibrogargan:
December (109 9, BMNH): South Australia: Adelaide,
November (29 9, TAMU); Tasmania: Harford, 5 km E
by S: Dodges Ferry, December to January (7 9 9, 4d,
ANIC); Victoria: Lake Mtn NE of Melbourne: Kinglake
N. P. nr Melbourne: Mitta Mitta Ck. 25 km NNW Omeo;
January to February (3 9 9 , 1 d . ANIC, BMNH); Western
Australia: Glen Forest: Mundaring Weir: Serpentine Riv.,
in nest of Pheidole sp.; October to January (8 9 9 [2 9 9
mounted on cards with ant host], BMNH, WAM).

Orasema Cameron

Orasema Cameron, 1884:105. Type species: Orasema
stramineipes Cameron; by monotypy.

Semora Cameron, 1909:432-433. Type species: Semora
xanthopus Cameron: by monotypy. Not Semora
Peckham, 1892; synonymy by Kerrich (1963).

Eucharomorpha Girault, 1913a:62-63. Type species:
Eucharomorpha worcesteri Girault; designated by
Gahan and Fagan, 1923. Synonymy by Boucek
(1988:520). Not Eucharomorpha [= Orasemorpha]
Girauh, 1913b:95.

Loshanus Ishii, 1932:210. Type species: Loshaniis
uichancoi Ishii; by monotypy. Watanabe, 1958:26
(redescription of genus and new species). Hedqvist,
1978:229 (redescription and new species). Boucek
(1988:521) incorrectly placed Loshanus as senior syn-
onym of Gollumiella Hedqvist. Current synonymy by
Heraty (1992:586) with transfer of L. uichancoi to
Orasema.

Parasemora Gemignani, 1933:192. Type species:
Parasemora freychei Gemignani [location of type
unknown]; by monotypy. New synonymy.

Semorata Strand. 1942:393. Replacement name for
Semora Cameron.

Semorella Ghesquiere, 1946:368. Unnecessary replace-
ment name for Semora Cameron.

Orasema is the most speciose of the orasemine genera,
with possibly more than 100 species throughout the
world. It is comprised of several distinct species groups
that create problems in defining the limits of the genus.
Boucek (1988) suggested that Parasemora may be a
junior synonym of Orasema. Although the type appears
to be lost, this genus is treated here as a junior synonym
based on the description and habitus drawing provided by
Gemignani (1933). 1 have seen no other material from

South America similar to the specimens described by
Gemignani that would suggest another genus in the
Oraseminae. The type species for Loshanus. L. uichancoi
Ishii. 1932. was transferred to Orasema based on features
of the wings, antenna and larva (Heraty, 1992). Boucek
(1988) included O. delicatula (Walker) as part of the
Indo-Pacific fauna (labelled "Australia (?)"). I examined
the type and it belongs to a group of species from South
and Central America that includes O. festiva (Fabr.).
Orasema ranomafanae Risbec (1952) is transferred here
to Stilhula ranomafanae new combination, although it is
an aberrant species that lacks apical scutellar spines as
found in other species of Stilhula.

Orasema differs from other Oraseminae by the follow-
ing: scape always elongate, maxilla and labium large,
propodeal foramen broadly emarginate. petiole cylindri-
cal with truncate base and dorsal flange (prominent in
most species), and gastral terga smooth and polished with
few or no setae. Members of Orasema also have the fol-
lowing: petiole usually longer than broad in both sexes,
rarely as short as 0.8x as long as broad in some females;
first gastral sternite (Ms,) constricted and marked by a
transverse furrow; anterior region of Ms., attached to the
apex of the petiole in both sexes; clypeal margin truncate
with a distinct bare anteclypeus; and ovipositor expanded,
ridged and curved forward in profile (straight only in O.
communis).

(lENERIC DP:S('RIPn()N

Head sublriaiiguiar. 1.2-1.5x as broad as mesosoma;
median ocellus anterior to lateral ocelli (Figs. 92,
129-130); lateral ocellus close to or broadly separated
from occiput. Face smooth or reticulate: scrobal depres-
sion usually shallow and poorly defined, partially includ-
ing median ocellus: ocellar-ocular groove present or

54

absent; occiput smooth or aciculate, occipital carina
rarely present, its dorsal margin usually broadly rounded.
Malar depression absent, present, or forming narrow
foveate channel; hypostoma well developed and separated
from gena by hypostomal carina. Clypeus subquadrate, its
apical margin truncate, epistomal sulcus lacking; ante-
clypeus narrow and bare, distinct from postclypeus.
Labrum usually 4-digitate, rarely multi-digitate, digits
and apical setae always long. Mandibles falcate, 2/3 (all
Old World species) or rarely 3/3 dentate; maxilla and
labium large, labial and maxillary palpi variable in shape,
usually 3-segmented. Antenna 11- to 13-segmented;
scape variable in length, usually narrow and cylindrical;
pedicel as long as broad; anellus present and smooth;
funicle 6- to 9-segmented, segments cylindrical, without
basal secondary segmentation but with scattered MPS;
basal funicular segments usually less than 2. Ox as long as
broad; last 2 to 3 flagellomeres usually fused into indis-
tinct clava.

Mesosoma with sculpture of dorsum variable, usually
reticulate or rugose-areolate medially. Mesoscutum with
notauli usually narrow and deeply impressed along entire
length. TSA complete. SSS deeply impressed, angled for-
ward to midline. Scutellum with frenal area sculptured
and usually separated dorsally by foveate groove; axillu-
lar sulcus present or absent. Metanotum extended lateral-
ly as a smooth flange over base of propodeum and partly
covering propodeal spiracle which is close to dorsal mar-
gin of propodeum; metanotum weakly excavated.
Propodeal disc variable in shape and sculpture, rarely
smooth; callus prominent and broadly rounded, bare or
setose, and lacking vertical furrow; postspiracular furrow
and metepimeral sulcus deeply impressed; ventral margin
of propodeum above hind coxa even and strongly ridged,
without lateral processes. Mesopleuron usually sculp-
tured, upper mesepimeron usually swollen and
transepimeral sulcus present; femoral groove lacking or
broadly impressed and shallow; mesepisternum usually
lacking sternaular area (no distinct sulcus or foveae).
Prepectus broadly triangular and reaching tegula, rarely
narrowed ventrally. Coxae and femora usually sculptured;
tibiae with spurs 1-1-2; hind tarsus 0.7-0.8x as long as
hind tibia.

Wing veins of fore and hind wings well defined.
Forewing 1.9-3.1x as long as broad, acute to broadly
rounded at apex; disc pilose, marginal fringe present (in
all Old World species) or absent; basal area bare or
setose, speculum and costal cell variable in pilosity; sub-
marginal vein with short setae dorsally; marginal vein
().2-().3x as long as wing, and pilose; stigmal vein sessile
or elongate, perpendicular to wing margin; postmarginal
vein short or long.

Mctasowd with petiole cylindrical and fused ventrally.
base abruptly narrowed (truncate) to small, knoblikc

condyle (Figs. 120-121, 132), and usually with prominent
dorsal flange. Petiole of female usually l-2x as long as
broad, rarely as short as 0.8x as long as broad. Petiole of
male longer than in female, usually 2-3x longer than hind
coxa. Gastral terga smooth with scattered short setae,
gaster of female as long as head and mesosoma, gaster of
male slightly longer than hind femur; Mt^ of female vari-
able in relative size; Ms-, constricted by narrow crenulate
or smooth furrow, anterior crescent-shaped region similar
in both sexes. Hypopygium bare or with few minute setae
laterally. Ms^ of male narrowly rounded and setose. Cerci
with variable number of setae of different lengths.
Ovipositor sheath broad, reaching or exceeding cercus,
gonostylus separated and sparsely setose. Ovipositor sub-
apically expanded, usually strongly curved cephalad; first
valvula with strong subapical ridge, followed by 3-10
sharp teeth along lateral line, rarely with diagonal ridges
in place of teeth {O. communis); second valvula narrow or
broad, and with several lateral teeth or complete dorsal
ridges. Genitalia of male with well-developed parameres,
digitus disclike and bearing several stout marginal spines;
aedeagus subacute.

PHYLOGENETIC RELATIONSHIPS

Orasema as described here is a diverse genus, with the
Old World species divided into 6 groups. The relation-
ships between the major groups is shown in Figures 1-2;
the same tree topology was obtained in all analyses. In the
analysis of higher level relationships, the uichancoi-group
was divided into communis and uichancoi subgroups
(treated in this section as only the uichancoi-group). The
Old World species groups are morphologically diverse
and future studies may show a need for further division
into more genera, but a conservative approach is taken
here and only I genus is recognized. The New World
species groups are distinctive and cannot be placed into
any of the Old World species groups. However, most of
the New World species can be aligned with either the
assectafor-group or the uichancoi-group.

The uichancoi-group includes 7 species, Orasema
houccki. O. communis, O. ishii. O. promecca . O.
ru^uh)sa. O. scyri^ii, and O. uichancoi. Monophyly of
this group is based on an increase in the number of funic-
ular segments to 8 or 9 in males (4) and to 8 in females
(5). Additional features found in both the Malagasy and
Indo-Pacific species (but not all species) include a weak
malar depression (10), presence of strong transverse
ridges on the second valvula (44, state 3), and a distinct
occllar-ocular groove dorsally between the lateral ocellus
and eye margin. Association of the Malagasy and Indo-
Pacific species is also based on phenetic similarity with
species having elongate bodies, and long and narrow .
pilose wings. O. communis and O. sc\rii;i arc inlorproled
as sister taxa based on general similarity but no characters

55

distinctl) support the nionophx l\ o\ this group. O. scyrii^i
shares character stales with the hulo-Pacific species in the
iilcluiiu()i-g.roup such as: dorsal occipital margin carinate.
malar depression weak, and second valvuia with trans-
verse ridges. O. communis is a unique species in this
group and has a niuUi-digitate labrum, distinct ocellar-
ocular groove, weak malar depression, and lacks an
occipital carina. The monophyly of the Indo-Pacific
species in the uicliancoi-gioup is justified by the presence
of a small terminal projection from the apex of the male
clava (Figs. 88-89. 95) that is not found elsewhere in the
family. O. uichancoi is regarded as the sister taxon to the
remaining hido-Pacific species by having the basal area
of the forewing bare versus pilose, although it may show
closer affinities to O. ishii and O. promecea based on the
presence of a weak occipital carina and weakly sculptured
or smooth face (plesiomorphic with respect to O. seyrigi).
O. rugulosa and O. houceki form a monophyletic group
based on a foveate malar depression, strongly sculptured
face, distinct ocellar-ocular groove, elongate forewings
(2.7-2. 9x as long as broad), and lack of an occipital cari-
na. Tentatively, relationships within the uichancoi group
are postulated as ({communis 1+ seyrigi) -(- {uichancoi 1+
{ishii + promecea -H {rugulosa + bouceki)))) (Fig. 276).

The striatosoma-group includes 2 species, O. fraudu-
lenta and O. striatosoma. These species are monophyletic
but are difficult to place with any other species groups of
Orasema. The parsimony analysis (Figs. 1-2) places this
group as sister taxon to the assectator-gxoup based on
having reticulate facial sculpture (8), short postmarginal
vein (36; long in O. fraudulenta), and lateral apex of the
second valvuia with several minute teeth (43). Internal
attachment of the first instar (62) is inferred but this is
probably a characteristic of Orasema. Intuitively, I
believe this group is probably more distantly related with-
in Orasema, possibly as a sister taxon to species with a
distinct speculum (37).

The assectator-gxoxxp is comprised of 3 species, O.
assectator, O. initiator and O. nigra, that form a mono-
phyletic group based on having a completely reticulate
face, callar nib, 7 to 10 minute teeth on the first valvuia,
completely reticulate mesosoma dorsum, and a short post-
marginal vein. The assectator-group is placed together
with O. glabra based on the short postmarginal vein (36),
and minute teeth on the second valvuia (43), and in part
on general morphological similarity. Orasema glabra is
unique in its possession of a completely smooth face,
scrobal depression with parallel channels (9) (similar
facial characters to Psilocharis), evenly rugulose
propodeum, and setose callus.

The koghisiana-gmup is monobasic. This species lacks
the broad triangular prepcctus (20) and rounded mesepis-
ternum (32) of the other groups (multistates in the uicluin-
(w-group). and is postulated to have secondarily gained a

larger number of labral digits (15). and reduced wing
setae (33). This species shares a glabrous propodeum (28)
with the \Y//,i,'///.v-group. but this character state is also
found in C'hrysolampinae and basal Eucharitinae and may
be plesiomorphic.

The valgius-gxoup includes 2 species. O. valgius and
O. synempora, that form a monophyletic group based on
having a mid-ventral sulcus on the mid coxa. 3 to 4 lateral
teeth on the first valvuia. laterally glabrous propodeal
disc, and an elongate postmarginal vein. There is a possi-
bility of closer affinities between the valgius- and assec-
lator-gToup^ based on having the face at least partially
reticulate, the mesepisternum broadly rounded ventrally,
and the callus glabrous or with only a few minute setae.

Tentative relationships proposed among the above
complex of species, excluding the uichancoi-group. are
{{koghisiana + {synempora + valgius)) + {glabra +
{{assectator + initiator -i- nigra) +1 {striatosoma + fraud-
ulenta)))) (Fig. 276).

BIOLOGY AND IMMATURE STAGES

The biology and immature stages of Orasema are well
known for several species (see earlier section). They have
been reared as parasites of Pheidole, Solenopsis,
Wasmannia, Tetramorium, Formica, and Eciton. The
only known hosts in the Old World tropics belong to
Pheidole. The genus Pheidole is shared by widely diver-
gent species groups in Orasema, as well as Orasemorpha,
and I postulate that Pheidole is the ancestral host for both
genera. The use of an intermediate host (thysanopteran or
homopteran) has been documented for both New and Old
World species, and may be common for the genus.

Plant hosts have been recorded for several species, and
females always oviposit into chambers hollowed within
the plant tissue by the ovipositor. The plant structure cho-
sen for oviposition is extremely varied in the Neotropical
and Nearctic regions. In the Old World, oviposition into
leaf surfaces is the only known strategy (in O. uichancoi,
O. assectator, and O. initiator). Choosing other struc-
tures, such as involucral bracts, flower stems or fruit
structures, may be a derived behaviour of New World
species. The ovipositor is very conservative in structure
within the genus and similar methods of egg deposition
may be assumed for other species. Orasema communis is
the only known species with a straight ovipositor (versus
ovipositor curved cephalad) and with lateral ridges (ver-
sus teeth) on the first valvuia (Fig. 82), similar to some
Neolosbanus. This may indicate a slightly different
oviposition strategy for what is considered to be a very
basal member of Orasema, but does not affect the general
prediction, for all species, of oviposition into chambers
that are hollowed out in plant tissue by the ovipositor.

The first-instar larvae are plesiomorphic for
Eucharitidae. The later instar larvae are more distinctive

56

and I consider the presence of prominent pustules as a
derived feature of Orasema (and possibly Oraseminae).
Larvae are unknown for O. koghisiana and the valgius-
group. However, their phylogenetic placement between
divergent species-groups of Orasema suggests that larvae
of these groups would possess all of the typical character
states. Adults of the koghisiana- and va/g///5-groups share
some of the character states of Psilocharitini (smooth
propodeum and callus), which I regard as convergent.

DISTRIBUTION

The distribution and proposed phylogeny of Old Word
species of Orasema is presented in Figure 276. Orasema
is distributed throughout the Nearctic, Neotropical,
Ethiopian, Indo-Pacific, and Australian regions.
Collections of Orasema in the Old World tropics are
sparse compared to the New World. The ulchancoi-group
is distributed only in Madagascar and the Indo-Pacific
region (Fig. 276), but may be closely related to a group of
species found in Central and South America. The Indo-
Pacific species of the uichancoi-gmup are considered to
be monophyletic, and form the sister group to O. commu-
nis and O. seyrigi in Madagascar. Within the uichancoi-
group, O. rugulosa, and O. houceki form a derived group
found only in Papua New Guinea. Members of the stri-
atosoma-group are represented by only a few collections
in eastern Africa (Fig. 276). Orasema valgius and O.
synempora are found only along the eastern coast of
Australia, with O. synempora confined to the Papuan sub-
region. Orasema koghisiana is found only on New
Caledonia and the New Hebrides in the Polynesian subre-
gion. Orasema glabra is restricted to southern Africa.
The assectator-gxonp has a widespread distribution in
southern Africa and the Indo-Chinese subregion (Fig.
276), and may form a sister group to similar species that
are widespread in the Nearctic and Neotropical regions.

The occurrence of distinct species-groups with distant
affinities throughout the tropical regions suggests that
Orasema may have initially diverged during the late
Cretaceous or early Eocene as part of the breakup of
Gondwanaland. Only the uichancoi-group has been suc-
cessful in the Oriental region with the most derived
species of this group found in the Papuan subregion.
Relationships within this group to the Madagascar species
may represent either a relictual distribution or collecting
anomaly within continental Africa. Within the assectator-
group, O. assectator and O. initiator may be derived from
an Ethiopian ancestor shared with O. nigra; although
except for affinities with O. glabra and possibly the stri-
atosoma-group, there is no morphological evidence with-
in the group to support this hypothesis. The assectator-
group is restricted to the Indo-Chinese and mainland
Western Malayan subregions of the Indo-Pacific. The val-
gius- and koghisiana-groups are distinct from other
species in the Oriental region and have their closest rela-
tionships to the Ethiopian species groups. The geographic
isolation of the valgius- and koghisiana-groups from
other closely related species groups not found in the Indo-
Pacific region suggests a very old relationship and sup-
ports the notion of an Ethiopian-Australian connection as
proposed by the phylogenetic hypotheses.

Among New World Orasema, there are at least 7 dis-
tinct species groups. Of the species found in the Old
World tropics, the uichancoi-group and assectator-group
both have affinities to species groups in the New World.
These relationships will need to be incorporated into a
more complete biogeographic hypothesis. I am presently
revising the New World species and a more thorough
treatment of the biogeography of this genus will be pre-
sented in that work.

Key to Old World Species of Orasema

Antenna 12- or 13-segmented, funicle 8- or 9-
segmented (Figs. 84-85, 87-89, 95): propodeal
disc evenly sculptured (Figs. 218, 235);
forewing without speculum (pilose) and wing
slightly infuscate (frontispiece. Figs. 86, 96-97)
uichancoi-group. 2

Antenna 1 1 -segmented, funicle 7-segmented
(Figs. 98-100, 108-109. 12.3-125): propodeal
disc evenly sculptured or glabratc laterally (Figs.
247-248): forewing with or without speculum
and wing hyaline (Figs. 104. 110. 117-119. 122)
8

2(1) Labrum with 8 to 10 digits, digits long and cov-
ering mouthparts (Fig. 80): frenal area abrupt
and rugose: axilla usually strongly rugose.

smooth in some females: Madagascar

O. communis Risbec. p. 59

— Labrum with 4 digits: frenal area rounded in pro-
file and smooth or rugose: axilla smooth and
shining or at most with very weak surface sculp-
ture (Figs. 83. 92. 235) 3

3 (2) Occipilal carina raised lo form crcsl bciiiiul ocel-

57

li. distinct in frontal view (Figs. 81. 83): posteri-
or margin of frenal line raised into a distinctive
carina along dorsal margin of frenal area (Fig.
83); Madagascar O. seyrigi Risbec. p. 61

— Occipital carina absent or hardly raised (Figs.
90. 92. 188); frenal area lacking strong ridge or
carina dorsally (Figs. 9 1-92. 218) 4

4 (3) Forewing with basal area bare except for sparse
band of hairs along impression of cubital vein;
stigmal vein with large uncus and appearing Y-
or T-shaped (Fig. 97); antenna 12-segmented in
both sexes (Fig. 87); face weakly rugulose to

almost smooth (Fig. 188); Philippines

O. uichancoi (ls\\\\). p. 62

— Forewing completely and densely pilose (fron-
tispiece. Fig. 96) (sometimes sparsely setose in
small patch just under submarginal vein); stig-
mal vein elongate, usually constricted basally
but without large uncus (if uncus present then
face glabrous); antenna 12- or 13-segmented
(Figs. 88-89, 95); face strongly rugose-areolate
or smooth and polished 5

5(4) Antenna of male 13-segmented (Fig. 88);
forewing 2.4x as long as broad, basal area pilose
but sparsely setose just below submarginal vein;
axillula longitudinally carinate; face and vertex
glabrous; lateral lobe of mesoscutum and axilla
strongly swollen and smooth; female unknown;
Taiwan O. ishii sp. nov., p. 64

— Antenna of male 12-segmented (Fig. 95);
forewing 2.5-2.9x as long as broad, basal area
completely pilose (Fig. 96); axillula smooth or
rugose, otherwise head and mesosomal sculpture
variable 6

6(5) Propodeum sharply angled in profile (fron-
tispiece), disc with complete median ridge; ocel-
lar-ocular groove sharply impressed and either
smooth or areolate; female unknown; New
Guinea O. bouceki sp. nov., p. 65

— Propodeum slightly rounded in profile (Fig. 91),
disc evenly sculptured without complete median
ridge; ocellar-ocular groove shallow or absent
(Fig. 92) 7

7 (6) Face smooth with cheeks lightly pitted; occipital
margin carinate; occiput glabrous; eye margined
by smooth groove; malar depression shallow and
smooth (Fig. 95): female unknown: New Guinea
O. promecea .sp. nov., p. 66

— Face finely and deeply rugulose-areolate (Fig.
90): occipital margin rounded; occiput finely
carinate; eye margined by reticulate groove;
malar depression foveate: New Guinea (New

Britain. New Ireland)

O. rugulosa sp. nov.. p. 67

8(1) Eye small and protuberant; head triangular with
deep pit just below median ocellus at dorsal mar-
gin of scrobal depression (Fig. 102): forewing
lacking speculum (pilose): marginal vein thick-
ened along entire length (Fig. 104); lateral lobe
of mesoscutum sculptured and similar to mid-
lobe (Figs. 98. 101) striatosoma-group. 9

— Eye large; head transverse or subtriangular (Figs.
112, 114, 127. 189, 191-192). without pit below
median ocellus: speculum present: marginal vein
thin (Figs. 110, 117-119, 122); lateral lobe of
mesoscutum polished or reticulate 10

9 (8) Scutellum finely striate, rounded laterally and

axillula not distinguishable from dorsum (Fig.
101); forewing completely pilose except along
impression of cubital vein (Fig. 104); Southern
Africa O. striatosoma sp. nov., p. 68

— Scutellum finely reticulate, abruptly margined
laterally and axillula longitudinally carinate,
axillular sulcus indistinct; forewing pilose, basal

area bare; Northwestern Africa and Yemen

O.fraudulenta (Reichensperger). p. 70

10 (8) Face, including vertex, completely smooth and

polished (Figs. 127. 189-190) 11

— Face completely or partially reticulate (Figs.
112, 191-192) 12

11(10) Labrum 6- to 8-digitate (Fig. 190); propodeal
disc smooth laterally with median areolate band
(Fig. 247); antenna with relatively few MPS

(Fig. 108); New Caledonia

O. koghisiana sp. nov.. p. 71

— Labrum 4-digitate (Fig. 192); propodeal disc
evenly rugulose without median carina or fur-
row; antenna with numerous MPS: Africa

O. glabra sp. nov., p. 72

12(10) Face completely reticulate (Figs. 112. 114.
191-192); propodeal disc evenly reticulate,
rarely weakly sculptured, without differentiated
median band of sculpture; lower mesepimeron
reticulate (Fig. 220); petiole of female 0.8-1. 5x
as long as hind coxa, petiole of male 1.7-2.3x as

58

long as hind coxa; first valvula with 7 to 10 lat-
eral teeth (Fig. 1 16); mid coxa without mid-ven-
tral sulcus; Africa and Indo-China

assectafor-group, 13

— Face smooth to reticulate but usually glabrate
below lower eye margin; propodeal disc smooth
laterally with broad median band of strong
sculpture (Fig. 248); lower mesepimeron
glabrous (Fig. 221); petiole of female 1.2-1.7x
length of hind coxa, petiole of male 2. 3-3. Ox as
long as broad; first valvula with 3 to 4 lateral
teeth (Figs. 262-263); mid coxa with mid-ven-
tral sulcus; Australia valgius-group, 15

13 (12) Propodeal disc reticulate; head 1.4x as broad as
high, with eye large (Fig. 112); mesoscutum
broadly rounded in dorsal view (Fig. Ill); hind
coxa completely reticulate; India and Sri Lanka
O. assectator Kerrich. p. 74

— Propodeal disc weakly sculptured; head 1.2-1.3x
as broad as high, with eye of moderate size
(Figs. 1 14. 191); mesoscutum rounded to sharply
angled anteriorly (Fig. 237); hind coxa weakly
coriaceous basally to glabrous apically 14

14(13) Midlobe of mesoscutum with anterolateral mar-
gin broadly rounded; pronotum without promi-

nence; clypeus and supraclypeal area minutely

reticulate (Fig. 1 14); South Africa

O. nigra sp. nov., p. 75

— Midlobe of mesoscutum with anterolateral mar-
gin sharply angled or produced (Fig. 237);
pronotum sometimes with conical prominence
just below spiracle; clypeus and supraclypeal
area glabrous (Fig. 191); Indo-Chinese and

Eastern Malayan subregions

O. initiator Kerrich, p. 76

15(12) Mesosoma with lateral lobe and axilla smooth
and polished (Fig. 129), mesosoma robust and
elongate (1.3x as long as high); frenal area semi-
circular in dorsal view; forewing with dense,

minute setae; Queensland

O. synempora sp. nov., p. 77

— Mesosoma with lateral lobe coriaceous to rugose
(Fig. 238) and axilla weakly carinate with sur-
face imbricate to reticulate, mesosoma slender
and subquadrate in profile (1.2x as long as high)
(Fig. 221); frenal area abrupt and hardly visible
in dorsal view (Fig. 238); forewing with moder-
ately dense, elongate setae (Fig. 122); eastern
and western Australia

O. valgius (Walker), p. 79

Orasema uic hancoi-gr oup

This group of 7 species is restricted in distribution to
Taiwan, Philippines, New Guinea and Madagascar.
Additional species (not described here; see discussions
under various descriptions) are recognized from Papua
New Guinea.

(JROUP DESCRIPTION

Head broadly subtriangular; face smooth to rugose, frons
often swollen, some species with distinct ocellar-ocular
sulcus, malar depression absent or narrow and foveate.
Antenna 12-segmented and funicle 8-segmented in
females, antenna 12- or 13-segmented and funicle 8- or 9-
segmentcd in males. Mesoscutum and scutcllum rugose to
rugose-areolate, with lateral lobe of mesoscutum and axil-
la swollen and glabrate; lower mesepiineron and callus
variable in sculpture, the callus with 7 or more elongate
hairs. Forewing infuscatc, completely pilose or basal area
bare, speculum absent; disc pilose and marginal fringe
present; stigmal vein variable in shape, postmarginal vein
elongate, more than ().5x as long as marginal vein. Petiole
variable. Ovipositor usually subapically expanded and

slightly curved forward; other features variable.

Orasema communis Risbec

Figs. 80, 82, 84-85

Orasema communis Risbec, 1952:412—414. Madagascar
[MNHP, examined].

TYPE MATERIAL

Lectotype (here designated), cJ , "MADAGASCAR/
BEKILY/ REG SUD DE LTLE.'' "MUSEUM PARIS/
1.37/ A. SEYRIG." "TYPE." "Orasema/ communis/
Risbec." "LECTOTYPE/ Orasema/ communis Risbec/
Det. J. Heraty '90." Left flagellum and right foreleg miss-
ing. Risbec did not designate a holotype in the original
description for Orasema communis. The lectotype male is
based on the handurilten determination label and type
card on the first specimen of the series. Only 1 2 of the 1 7
males mentioned in the original description were exam-
ined. Females of Orasenui communis were labelled as
^'Orasema lyekilensis Risbec." All 1 1 females arc listed in

59

the original description and the name is probably an early
manuscript name of Risbec. There are some discrepancies
with the original label information in Risbec (1952). and
the paraiectotype data is relisted here.
Paralectotypes (examined and here designated):
Madagascar: Bekily. xii.I936 (29 9 , 36 6), i.l937
(19. 2(?d). ii.l937 (19. \6), iii.1937 (19). iv.l937
(19), iv.l938 (\6), ii.l939 (19), i.l94() (Id), ii.l940
(l9);Taolanaro [Fort Dauphin], xii.l936(39 9), v.1937
(36 6): all collected by A. Seyrig (MNHP).

DIAGNOSIS

This species has a distinctive habitus and can be recog-
nized by: body size large, head and mesosoma black,
labrum 8- to 10-digitate with digits narrow and elongate
(Fig. 80). face relatively smooth and flat, F2 3.5-5.0x as
long as broad (Figs. 84-85), lateral lobe and axilla strong-
ly swollen, scutellum rugose with a distinct frenal area,
wing with only extreme basal area bare, and postmarginal
vein elongate. This species differs from O. seyiigi by hav-
ing the ocellar-ocular channel distinct, occipital carina
lacking, vertex not elevated medially (Fig. 80), dorsum of
mesosoma more strongly areolate, and ovipositor larger
and similar in size to other Oraseminae (Fig. 82).

FEMALE

Length. 4.5-5.8 mm. Head, mesosoma, petiole, and
antennal flagellum black, gaster reddish-brown; scape and
pedicel yellowish brown; fore and midlegs, including
coxae, light yellowish brown; hind coxa light brown ven-
trally to black dorsally; femora mostly brown, otherwise
legs yellowish brown; mandible yellowish brown with
dark brown outline, maxilla and labium pale yellowish
brown. Wings hyaline, venation brown.

Head subtriangular, 1.3-1.4x as broad as high; occiput
transverse; lateral ocellus separated from occiput by own
diameter; LOL 0.7-O.9x OOL. Face relatively flat, frons
may be weakly impressed lateral to toruli, glabrate with
only light punctation in lower half of face; scrobal depres-
sion shallow and broadly impressed, finely and irregular-
ly sculptured medially; vertex evenly rounded with
smooth, well-defined ocellar-ocular channel; vertex and
occiput finely strigate just posterior to ocelli, otherwise
glabrous. Eye not especially prominent, eyes separated by
1.9-2.0X their height. Malar space 0.9-1. Ox height of eye,
malar depression narrow and poorly defined. Clypeus
glabrate with only scattered minute setae, epistomal sul-
cus weakly defined, lateral margin deeply impressed to
tentorial pits, anteclypeus subtruncate; supraclypeal area
swollen medially and poorly defined laterally. Labrum 8-
to II -digitate, digits elongate and narrow, setae bristle-
like. Mandible moderately stout; maxilla and labium
large, palpi elongate and 3-segmented. Antenna 12-seg-
mented (Fig. 84); scape stout and cylindrical, almost

reaching median ocellus; pedicel small and globose; anel-
lus present and small; flagellum 2.0-2. 3x height of head;
funicle 8-segmented, segments finely reticulate with
dense, small setae, no MPS evident: F2 1.0-1.2x as long
as scape, 3. 5-5. Ox as long as broad. F2 1.3-1.5x F3. fol-
lowing segments subequal in length and equal in width;
clava subovate. not differentiated from funicle and shorter
than preceding 2 segments.

Mesosoma with midlobe of mesoscutum rugose-areo-
late and broadly rounded; lateral lobe smooth to very
weakly carinate and strongly swollen: axilla irregularly
rugose-carinate and swollen, posterior margin abruptly
margined at SSS; scutellum broadly rugose-areolate.
Mesoscutum with notauli deeply impressed and irregular-
ly carinate. SSS broadly and deeply impressed, irregularly
carinate. Scutellum as long as broad, broadly separated
from TSA at base, strongly sloped posteriorly to meet fre-
nal line; frenal line crenulate dorsally or with band of
crenulate sculpture on either side, frenal area irregularly
carinate or rugose-areolate and with abrupt posterior mar-
gin; axillular sulcus weak and crenulate. Propodeal disc
broadly rounded and evenly alveolate or rugose-areolate;
postspiracular sulcus deeply impressed and mostly
smooth, lacking prominent carina above hind coxa; callus
slightly swollen, smooth laterally and dorsally with dense
patch of short hairs; metepimeron weakly sculptured,
metepimeral sulcus deep and irregular, continuing dorsal-
ly as deep crenulate groove and separating a narrow ante-
rior region. Upper mesepimeron slightly swollen and
mostly glabrate, lower mesepimeron smooth to rugose,
transepimeral sulcus shallow and irregularly foveate;
femoral groove lacking; mesepisternum rugose-areolate
to scabrous, glabrate ventrally and only slightly swollen
anterior to mid coxa. Prepectus narrow ventrally. upper
triangle deep foveate. Pronotum rugose with sharp medial
furrow. Proepisternum swollen and smooth or very weak-
ly sculptured. Fore coxa elongate and smooth, mid coxa
subglobose and finely carinate. hind coxa subglobose,
1.6x as long as broad, and glabrate; hind femur glabrate,
with short dense setae apically: hind tibia densely short
setose, the setae longer on inner margin; hind tibia with 2
large spurs. Forewing 2.3-2.6x as long as broad, 2.6-3. Ix
as long as mesothorax; basal area and along impression of
cubital vein bare; speculum absent: costal cell broad and
densely pilose; submarginal vein with dense row of dorsal
setae; marginal vein 0.29-0.32x as long as forewing; stig-
mal vein subquadrate to elongate and roughly perpendicu-
lar to anterior margin of forewing. with distinct apical
uncus equal to width of stigma; postmarginal vein long
and reaching apex of forewing.

Metasoma with petiole 1 .5-2. Ox as long as hind coxa,
1.4-1.9X as long as propodeum; petiole slightly increas-
ing in width to apex, smooth ventrally and weakly rugose
dorsally. slightly dorsoventrally compressed, with strong

60

basal flange. Mt, 1.0-1.7x as long as hind femur,
glabrous; Ms., with strong constriction, groove narrow
and weakly crenulate, anterior region semicircular.
Hypopygium with few short hairs ventroapically.
Ovipositor short and straight, thickened along entire
length (Fig. 82); first valvula without subapical ridge or
apical line of teeth and with 3 diagonal ridges apically;
second valvula with 6 strong transverse ridges coalescing
dorsally.

MALE

Length, 4.4-5.8 mm. Colour as in female, gaster dark
brown.

Head as in female. Antenna 12-segmented (Fig. 85);
scape thickened medially; funicle 8-segmented, segments
cylindrical and very slightly broader at apices, covered
with dense elongate setae; F2 4.4-5 .4x as long as broad;
clava constricted medially (2-segmented).

Mesosoma with dorsum more strongly sculptured.
Stigmal vein of forewing 6x as long as broad, sometimes
with distinct apical uncus.

Metasoma with petiole 1.4-2.0x as long as hind coxa,
1.8-2. Ox as long as propodeum, cylindrical, smooth ven-
trally and weakly to strongly rugose dorsally, with small
basal flange. Mt^ 0.7-0.9x as long as hind femur; Ms,
narrowly rounded with dense elongate hairs. Genitalia
with strong median process, paramere relatively short and
stout, digitus with 4 or 5 marginal teeth; aedeagus sub-
acuminate.

VARIATION

There is little variation among the type material except in
the shape of the stigmal vein (short and subquadrate to
elongate with a distinct uncus), and sculpture of the
propodeal disc (close areolate to strongly rugose-areo-
late).

DISTRIBUTION

Madagascar (C, Fig. 276).

MATERIAL EXAMINED

Madagascar: Antananarivo |no date], Sikora (l?sex.
BMNH).

Orasema seyrigi Risbec

Figs. 81,83, 86

Orasema Seyrii^i Risbec. 1952:414-416. Madagascar
[MNHP, examined].

TYPE MATERIAL

Holotypc, 9, "MADACJASCAR/ ROGEZ/ FORET
COTE EST." "MUSEUM PARIS/ 1.37/ A. SEYRIG."

"TYPE." "Orasema/ Seyrigi Risbec." Antenna missing
beyond pedicel. The data and description match the speci-
men examined, which is a female, not a male as originally
stated by Risbec (1952).

DIAGNOSIS

Recognized by a crest behind the ocelli (Fig. 81). strong
carina along dorsal frenal margin (Fig. 83), basal area of
forewing bare, and ovipositor minute in relation to the
overall body size. This species appears most closely relat-
ed to Orasema communis.

FEMALE

Length 6. 1 mm. Head black, mesosoma black with blue
reflections, petiole black, gaster dark brown to black;
scape and pedicel yellowish brown, fore and mid coxae
dark brown at base with remainder light yellowish brown,
hind coxa dark brown to black; femur dark brown medial-
ly, otherwise light yellowish brown; mandible light
brown with dark border, maxilla and labium yellowish
brown. Wings lightly infuscate, venation dark brown.

Head subtriangular, 1.5x as broad as high; occiput
slightly emarginate; lateral ocellus separated from occipi-
tal margin by own radius; LOL 0.8x OOL. Face relatively
flat, frons and cheeks slightly swollen medially, glabrate
with only scattered fine punctation over entire face;
scrobal depression narrow and weakly rugulose, lateral
margin broadly rounded, depression with 2 parallel stri-
gate channels reaching ventral margin of median ocellus,
median area swollen and weakly carinate; vertex strongly
impressed next to lateral ocellus, continuing weakly as
ocellar-ocular depression to eye margin, finely transverse-
carinate between depression and occipital margin, ocellar
triangle rugulose; occipital margin abrupt with strong
carina posterior to ocelli, elevated carina fomiing crest in
frontal view; occiput glabrous medially to very weakly
aciculate ventrally. Eye prominent and bulging in frontal
view; eyes separated by 1.7x their height. Malar space
0.8x height of eye, malar depression broad and poorly
defined. Clypeus glabrate with scattered minute setae,
epistomal sulcus weakly defined, lateral margin strongly
impressed and deepest at tentorial pit, anteclypeus sub-
truncate; supraclypeal area swollen medially and poorly
defined dorsolaterally. Labrum not discernible (may be
reduced or missing). Mandible moderately stout: maxilla
and labium normal for genus, palpi elongate and 3-seg-
mented. Scape narrow and cylindrical, converging from
base to apex (bowed medially) and reaching median ocel-
lus, pedicel small and globose (antenna broken beyond
pedicel).

Mesosoma with midlobe of mesosculuni broadly
rounded and rugulose with fine transverse carinae: lateral
lobe strongly swollen and smooth; axilla swollen, smooth
dorsally and obliquely carinate poslcrolatcrally; scutelluni

61

rugose Id rugose-areolaie. Mesoscutuni v\itli iiDiauli
sharply impressed and narrowly crenulate. SSS deeply
toveaie and U-shaped. Scutellum 1.2x as long as broad,
broadly separated Ironi TSA at base by foveate SSS; tre-
nal line broad dorsally and crenulate forming a sharp
ridge at dorsal margin of frenal area, frenal area abruptly
margined, rugose-areolate dorsally to broadly crenulate
ventrally: axillular sulcus weak and crenulate. Propodeal
disc narrow and strongly rounded, strongly rugose-areo-
late medially to glabrate laterally; postspiracular groove
deeply crenulate. forming small carina above base of hind
coxa; callus smooth with patch of dense, long hairs dor-
sally; metepimeron glabrous, metepimeral sulcus strongly
impressed and irregularly sculptured. Upper mesepimeron
slightly swollen and smooth, lower mesepimeron lightly
sculptured, transepimeral sulcus foveate; femoral groove
absent; mesepisternum finely rugulose-alveolate to
rugose, smooth ventrally and only slightly swollen anteri-
or to mid coxa. Prepectus triangular, only slightly nar-
rowed ventrally, medially coUiculate to rugose, swollen
and smooth along dorsal and posterior margins. Pronotum
irregularly carinate to smooth with broad transverse fur-
row. Proepisternum glabrous. Fore coxa elongate and
smooth, mid coxa globose and finely carinate basally,
hind coxa subglobose and glabrate: hind femur slender
and smooth with dense fine setae dorsally along entire
length; hind tibia slightly expanded to apex, with dense
adpressed setae. Forewing 2.7x as long as broad, 3.0x as
long as mesothorax; basal area and along impression of
cubital vein bare; speculum absent; costal cell broad and
densely pilose; submarginal vein with dense row of dorsal
setae; marginal vein 0.28x as long as forewing; stigmal
vein subquadrate and roughly perpendicular to forewing
margin, with small uncus; postmarginal vein long and
reaching apex of forewing.

Metasoma with petiole 1.6x as long as hind coxa, 1.4x
as long as propodeum; petiole rugose, very slightly
increasing in width to apex, cylindrical with weak basal
flange. Mt., l.lx as long as hind femur, glabrous; Ms^
strongly constricted, groove strongly crenulate, anterior
region circular. Hypopygium bare. Ovipositor extremely
reduced and somewhat threadlike (first valvula with-
drawn and hidden) and shorter than length of hind coxa;
second valvula narrow with 4 strong transverse ridges.
Ovipositor sheath acute, gonostylus differentiated as
setose yellowish brown area, but not separated at base by
suture.

DISTRIBUTION

Madagascar (E, Fig. 276).

Orasema uichancoi (Ishii)

Figs. 87, 94, 97, 188, 218, 235, 258-259

Loshanus uichancoi Ishii, 1932:210. Philippines [NIAS,

wing and antenna examined).
GollumieUa uichancoi — Boucek, 1988:522.
Orasema uichancoi — Heraty, 1992:586.

TYPE MATERIAL

Lectotype (here designated), 9 [?], slide of left fore and
hind wing and left antenna labelled "Psilogaster/ L.
Banos." "Losbanus/ uichancoi/ Ishii. 1932/ LECTO-
TYPE/ det:/ J. Heraty "SH." Associated with unlabelled
glass slide of planidium (see below).

No other mounted specimens are known from the
NIAS collection (personal communication, Kazukiko
Konishi, NIAS, and my examination of miscellaneous
Oraseminae in NIAS collection). A slide collection was
sent to me from the NIAS, which was thought to have
been associated with Ishii 's 1932 paper. The collection
included slides of adult parts and planidia that could all
be associated with the species described by Ishii (1932).
The wings are distinct among all of the specimens
described in that paper and matched Ishii 's illustration of
the wing. The pattern of wing pilosity and peculiar stig-
mal vein are sufficient to identify this species. The num-
ber of specimens included as "types" by Ishii was not
stated but may be numerous as he mentions that adults are
"most common" during the dry season, and he was able to
make biological observations on oviposition.

DIAGNOSIS

Recognized by: wing densely pilose, basal area bare
except for a band of hairs along the impression of the
cubital vein, stigmal vein with long robust uncus longer
than width of the stigmal vein (appearing Y- or T-shaped)
(Fig. 97), and the axilla and lateral lobe swollen and
glabrous (Fig. 235). This species is similar to O. ishii but
can be distinguished by having the antenna 12-segmented
in males, eye large but not strongly protruding, gena
rounded or angled, proepisternum strigate-coriaceous, and
labial palpus with terminal segment 2-3x times longer
than broad.

FEMALE

Length. 2.8^.3 mm. Head, mesosoma, and petiole black,
mesosoma with strong blue or green reflections dorsally
and purple retlections sublaterally; gaster dark brown to
black; flagellum dark brown; pedicel, scape and legs yel-
lowish brown; femora yellow to light brown in proximal
half. Wings slightly infuscate. venation pale brown.

Head subtriangular to subquadrate, 1.2-1.5x as broad
as mesosoma, posterior margin of gena broadly rounded
to sharply angled medially; occiput broadly emarginate;

62

lateral ocellus separated from occiput by slightly less than
own radius; LOL 0.8-1. Ox OOL. Face relatively flat,
weakly rugulose to almost smooth with frons and vertex
weakly coriaceous; scrobal depression broad and shallow-
ly impressed, lightly carinate laterally, sculpture delimit-
ing medial glabrous band extending to median ocellus
(Fig. 188); vertex rugose to coriaceous, strongly
depressed next to lateral ocellus, ocellar-ocular groove
shallow and vaguely impressed; temple narrow and rugu-
lose to weakly strigate; occiput aciculate, dorsal margin
with weak carina, carina extending just beyond lateral
ocellus. Eyes separated by 1.7-2. Ox their height and mar-
gined by narrow sulcus. Malar space 0.6-0.9x height of
eye, malar depression vaguely impressed and weakly
sculptured. Clypeus and supraclypeal area glabrate, epis-
tomal sulcus strongly impressed, territorial pits and lateral
margin of clypeus strongly impressed, anteclypeus slight-
ly rounded; margin of supraclypeal area deeply impressed
ventrally. Each mandible with long apical tooth overlap-
ping base of opposing mandible; maxillary palpus 3-seg-
mented; labial palpus 2-segmented, basal segment short,
terminal segment elongate. Antenna 1 2-segmented; scape
stout and cylindrical, reaching 0.6x distance to median
ocellus; pedicel subconate, as long as broad; anellus pre-
sent and about twice as broad as long; flagellum 1.4-1.6x
height of head; funicle 8-segmented. segments densely
setose, sculpture relatively smooth, with numerous MPS;
F2 0.5x as long as scape, 2.1-2.4x as long as broad, fol-
lowing segments subequal in length, equal in width; clava
ovate, as long as preceding 2 segments.

Mesosoma with midlobe of mesoscutum rugose-areo-
late to weakly longitudinally carinate (pronounced only
on SEM specimen; Fig. 235); lateral lobe and axilla
swollen and glabrate; scutellum rugose. Mesoscutum with
notauli deeply impressed and foveate; midlobe subtrian-
gular and strongly elevated along anterior margins.
Scutellum 1.6x as long as broad; frenal line shallow and
foveate, frenal area semicircular in dorsal view and rugu-
lose; axillula weakly carinate. axillular sulcus obscured
by dorsal sculpture. Propodeal disc broadly rounded and
only slightly curved in profile, rugulose to rugose-areo-
late; postspiracular and metepimeral furrow broadly
impressed and irregularly sculptured; callus swollen,
rugulose dorsally and glabrate laterally, with patch of sev-
eral elongate hairs dorsally; mctcpimeron glabrate. Upper
mcscpimeron swollen and glabrate, lower mcsepimeron
weakly sculptured, transepimeral sulcus shallow and
broadly foveate; femoral groove shallow; mesepistemum
rugose-reticulate, swollen, and glabrate ventrally.
Prepectus triangular and rugose. Pronotum mostly
glabrous. Proepisternum weakly strigate with verrucose
or reticulate surface sculpture. Fore and mid coxae
glabrate, hind coxa rugulose dorsally; femora glabrate
with denser fine setae apically; hind tibia with 2 tibial

spurs. Forewings 2.7x as long as mesothorax. 2.4-2.6x as
long as broad; basal area setose along impression of
cubital vein, always bare just posterior to submarginal
vein; speculum absent; costal cell relatively narrow; mar-
ginal vein 0.24-0.27X as long as forewing; stigmal vein
slightly longer than broad and weakly constricted basally,
apical margin subtruncate, uncus elongate and longer than
width of stigmal vein, stigma and uncus appearing Y- or
T-shaped (Fig. 97); postmarginal vein 0.5x as long as
marginal vein, reaching half distance to apex of wing.

Metasoma with petiole 1.3-2.0x as long as hind coxa,
1.1-2.3X as long as propodeum; petiole rugulose dorsally
and weakly carinate to glabrate ventrally with distinct
basal flange (Fig. 94). Mt^ 1.2x as long as hind femur,
glabrous; Ms^ with constriction shallow and glabrous
(Fig. 258). Ovipositor subapically expanded and strongly
curved anteriorly; first valvula with 4 strong lateral teeth
beyond subapical crest (Fig. 259); second valvula with
several strong transverse ridges apically. Gonostylus
broad.

MALE

Length, 2.5-3.5 mm. Colour as in female. Head as in
female. Eyes separated by 1.7-2. Ox their height. Antenna
1 2-segmented; funicle 8-segmented, segments densely
setose, setae papillate and surface appearing scabriculous,
no MPS. Forewing 2.6x as long as mesothorax. 2.2-2.4x
as long as broad. Petiole 2.9-3.4x as long as hind coxa,
2.4— 3. 9x as long as propodeum. slender and finely rugu-
lose. Mt-, 0.9x as long as hind femur. Ms, strongly
impressed and smooth. Genitalia with basiparamere trun-
cate apically with strong median process, paramere short
and broad, digitus with several marginal teeth; aedeagus
broadly rounded.

VARIATION

Females from Bontoc (Mountain Province) show a large
range of variation. These individuals are all large but the
sculpture of the face ranges from weakly sculptured (2
specimens) to rugulose. the uncus of the stigmal vein is
rarely small (usually elongate), and the shape of the gena
ranges from broadly rounded to sharply angled in the
same series. One specimen from Lipa has the basal area
of the wing almost completely setose.

hi()i,()(;y and immature stac.es

Females deposit their eggs in the lower surface of young
leaves of Ccltis philippincnsis (Ulmaceae) and Leucaena
i^laina (Lcguminoscae) (Ishii. 1932). Eggs are deposited
in leaf punctures arranged in 2 short parallel rows (Ishii.
1932). Reexamination of the planidium indicated it was
similar to Ishii 's (1932) description except for the follow-
ing discrepancies and additional comments: body 12-seg-
mcnled with lerga I and I! distinctly separated dorsally.

63

hatchet-shaped sclerite lacking, tergoplcural line lacking,
and tergite IXa leallikc. No differences between this
planidiiim and those of other Orusema were noted, and
the stalked egg is typical for Orasenia. Ishii (1932)
described the flight period as throughout the year, but
common during the dry season in February.

DISTRIBUTION

Philippines (U, Fig. 276).

MATERIAL EXAMINED

Philippines: Luzon: Lipa Btg., 14. ix. 1952, Townes
Family, "Losbanus uichancoi Ishii, Bait. '58." (29 9,
AEI): Mountain Prov., Abatan, Buguio, 60 km S of
Bontoc, 1800-2000 m, 25. iv. 1964 (599, \6),
27. iv. 1964 (19), 9.V.1964 (19), 12. v. 1964 (Ic?),
22-3 l.V. 1964 (19), H. M. Torrevillas (BPBM); Batan-
gas Prov., Citrus Expt. Sta., 26. xi. 1953, J. L. Gressitt
(19, BPBM); Albay Prov., Mt Mayon, 16 km NW of
Lagaspi, 1200-1800 m, 15. v. 1962, H. M. Torrevillas
(Id. BPBM); Biliran: 1927, C. F. Baku ( 1 <? , USNM).

Orasema ishii sp. nov.

Fig. 88

TYPE MATERIAL

Holotype, d, "C. TAIWAN: Tungpu/ 1200 m. Nantou
Hsien/ 20-22. VI. 1980/ C. C. Chen." "HOLOTYPE/
Orasema/ ishii Heraty, 1990." Left forewing, left hind leg,
and tarsi of right hind leg missing. Deposited in TARI.
Paratype: Taiwan: Hassenzan, 21.vi.l932, J. L. Gressitt
(lc5,CAS).

DIAGNOSIS

Similar to O. uichancoi and O. prnmecea. The male is
distinguished by: forewing completely pilose except for a
sparsely setose region below submarginal vein, face
glabrate, axillula longitudinally carinate, and antenna 13-
segmented (Fig. 88).

MALE

Length, 4.2 mm. Body black, mesosoma and petiole with
faint greenish-blue reflections; pedicel and flagellum dark
brown; scape and legs light yellowish brown. Wings
slightly infuscate, venation brown.

Head triangular, 1.3x as broad as mesosoma, posterior
margin of gcna straight in frontal view; occiput broadly
rounded; lateral ocellus almost touching occiput; LOL
0.9x OOL. Face flat and glabrate; scrobal depression
shallow, glabrate with margin broadly rounded; no ocel-
lar-ocular groove, ocellar triangle rugose; temple narrow
and glabrate; occipital carina weak, extending just beyond
lateral ocellus; occiput vaguely strigate, almost smooth.

Eye protuberant, eyes separated by 1.9x their height, ocu-
lar groove narrow and smooth. Malar space 0.9x height of
eye, malar depression absent. Clypeus glabrous, bulging
medially, epistomal sulcus weakly impressed, tentorial pit
and lateral margin of clypeus deeply impressed, ante-
clypeus almost straight. Maxillary palpus 3-segmented,
labial palpus 1 -segmented and only slightly longer than
broad. Antenna 1 3-segmented; scape stout and cylindri-
cal, just reaching median ocellus; pedicel globose; anellus
present; flagellum 2.6x height of head; funicle 9-seg-
mented, segments with dense short semi-erect papillate
setae, no MPS; F2 0.8x as long as scape. 2.5x as long as
broad, following segments subequal in length, only
slightly decreasing in width toward apex of flagellum;
clava ovate and slightly longer than preceding segment,
apex with weakly segregated, cylindrical projection.

Mesosoma with midlobe of mesoscutum and scutellum
rugose; lateral lobe and axilla strongly bulging and
glabrate. Mesoscutum with midlobe subtriangular, verti-
cal face rounded, and anterior lateral margins abruptly
elevated; notaulus deeply impressed and narrowly crenu-
late. SSS broadly impressed and carinate. Scutellum as
long as broad; frenal line narrow and weakly crenulate,
frenal area rounded in profile and rugose-areolate; axillu-
la longitudinally carinate. axillular sulcus obscure.
Propodeal disc rounded, straight in profile, rugose-areo-
late and with irregular lateral carina along postspiracular
furrow that extends to base of petiole; postspiracular fur-
row broad and deeply impressed with few weak trans-
verse carinae; callus swollen and glabrate with patch of
elongate hairs; metepimeral groove broadly and deeply
impressed, continuing dorsally to posterior margin of
wing base, foveate-carinate dorsally. Upper mesepimeron
swollen and glabrous, lower mesepimeron weakly sculp-
tured; femoral groove and transepimeral sulcus broadly
and shallowly impressed. Mesepistemum rugose laterally
with weak verrucose sculpture, rounded and glabrous
ventrally. Prepectus triangular, weakly verrucose with
irregular strigae. Pronotum glabrous. Coxae and femora
glabrate. Forewing 2.8x as long as mesothorax, 2.4x as
long as broad; completely pilose except for sparsely
setose area just below submarginal vein: marginal vein
0.27x as long as forewing; stigmal vein twice as long as
broad and constricted basally, perpendicular to wing mar-
gin, uncus elongate and projecting posteriorly; postmar-
ginal vein 0.8x as long as marginal vein, almost reaching
apex of wing.

Metasoniu with petiole 2.7x as long as hind coxa, 2.8x
as long as propodeum; petiole linear and cylindrical with
small basal flange, rugulose. Mt, as long as hind femur,
glabrous; Ms, with constriction broad and smooth.
Genitalia with elongate median process, paramere stout
basally, abruptly narrowed apically, digitus with 4 mar-
ginal spines; aedeagus long and subacute.

64

FEMALE

Unknown.

DISTRIBUTION

Taiwan (H, Fig. 276).

ETYMOLOGY

Named in honour of Dr. Tei Ishii for his work on the
Eucharitidae.

Orasema bouceki sp. nov.

Frontispiece

TYPE MATERIAL

Holotype, 6 , "PAPUA: Kokoda./ 1,200 ft. [365 m]
X.1933./ L. E. Cheesman./ B. M. 1934-321." "c.f. T 612/
nigra [boxes]/ NOT /delicatula." "Orasema." "HOLO-
TYPE/ Orasema/ bouceki Heraty." Right forewing miss-
ing. Deposited in BMNH.

DIAGNOSIS

Recognized by the following: forewing completely pilose,
mesosoma elongate with propodeum strongly arched in
profile, face and callus rugulose. Similar species with a
smooth face will key to O. bouceki and are discussed in
the included section on related species.

MALE

Length, 3.6 mm. Head, mesosoma, coxae, and petiole
black, mesosoma with faint bluish tinge, not highly
reflective; gaster brown with Ms, black basally; antennal
flagellum dark brown; scape, pedicel, and anellus yellow-
ish brown; mandibles, apex of coxae, and legs yellowish
white, apex of tibiae and tarsi slightly darker. Wings
infuscate, venation dark brown.

Head subquadrate, 1.3x as broad as high; posterior
margin of gena straight; occiput broadly emarginate; ocel-
li large, lateral ocellus close to occiput; LOL 0.8x OOL.
Face relatively flat, frons slightly bulging just below
median ocellus, face including scrobal depression and
clypeal area rugose; scrobal depression narrow, lateral
margin broadly rounded, distinct vertical carina separat-
ing scrobal depression medially; vertex almost smooth,
strongly impressed next to lateral ocellus, ocellar-ocular
sulcus sharply impressed and foveate; temple narrow and
weakly sculptured; occiput strigate, occipital carina
absent. Eyes separated by 1 .7x their height, margined by
narrow reticulate sulcus. Malar space 0.7x height of eye;
malar depression narrow and deeply foveate. Clypeus
rugulose, epistomal sulcus weak, tentorial pit and lateral
margin deeply impressed, lateral margin of supraclypcal
area shallow, anlcclypeus broadly rounded at apex.
Maxillary palpus 3-scgmented and long; labial palpus 2-

segmented and very short. Antenna 12-segmented; scape
stout and cylindrical, almost reaching median ocellus;
anellus slightly broader than long; flagellum 2. Ox height
of head; funicle 8-segmented, segments with dense, semi-
erect papillate setae, surface appearing scabriculous, no
MPS; F2 2.7x as long as broad, 1.3x F3, as long as scape,
following segments subequal in length and equal in
width; clava tapered to apex, slightly shorter than preced-
ing 2 segments.

Mesosoma with dorsum finely and deeply alveolate-
rugose; lateral lobe of mesoscutum and axilla strongly
bulging, lateral lobe glabrate, axilla narrow and weakly
sculptured. Mesoscutum with midlobe subtriangular.
strongly elevated along anterior margins; notaulus deeply
impressed and transversely carinate. SSS broad and
deeply impressed with large transverse carinae. Scutellum
slightly longer than broad, separated at base from TSA by
deep fovea; frenal area and axillula not distinguishable,
apical margin of scutellum crenulate. Propodeal disc
rounded, broadly curved in profile, alveolate-rugose, with
sharp lateral carina along postspiracular furrow dorsally
and extending to base of petiole; postspiracular furrow
narrow and crenulate; callus areolate and flush with rest
of propodeum, with dense patch of short hairs dorsally;
metepimeral furrow vaguely impressed. Upper
mesepimeron glabrous and hardly swollen, lower
mesepimeron rugose-alveolate; femoral groove and
transepimeral sulcus vaguely impressed; mesepisternum
finely rugose-alveolate to umbilicate laterally, glabrous
and flattened ventrally. Prepectus finely alveolate-rugose
and verrucose. posterior margin glabrous and narrowed
ventrally. Pronotum irregularly foveate. Proepisternum
rugose. Fore and mid coxa glabrate with few weak stri-
gae. hind coxa glabrate with weak dorsolateral sculpture;
femora glabrate, tibiae with minute adpressed setae; hind
tibia with 2 spurs. Forewing 3. Ox as long as mesothorax,
2.4x as long as broad; completely pilose, costal cell nar-
row; marginal vein 0.32x as long as forewing; stigmal
vein slightly longer than broad, perpendicular to wing
margin; postmarginal vein 0.5x as long as marginal vein,
reaching half distance to apex of wing.

Metasoma with petiole 2.7x as long as hind coxa, 3.1x
as long as propodeum; petiole slightly sinuate in profile,
irregularly reticulate-carinate and verrucose, without
basal llange. Mt, I.7x as long as hind femora, glabrate;
Ms^ smooth, constriction only vaguely impressed laterally
(considered absent). Genitalia small, additional features
could not be observed on holotype.

FEMALE

Unknown.

65

RKI.ATKI) SPKCIES

Two specimens from Papua New Guinea represent 2
closely related species that are not described here because
the antenna are incomplete on both. These share the fol-
lowing features with Orasema houceki: propodeum
strongly arched with median carina; callus rugulose. not
swollen, and with patch of short hairs: malar depression
crenulate or weakly sculptured; frenal line obscure; and
similar wings. The female (Sepik Riv., Pagwi,
25.viii.1957, Hardy. BPBM) has the face glabrate and
weakly punctate ventrally. body black with strong purple
and blue reflections, dorsum of mesosoma rugose,
proepistemum smooth, and Ms., with constriction deeply
impressed but smooth. The male (Central Prov.,
12. vi. 1983, J. Ismay, BMNH) has the face polished and
weakly punctate to rugulose, body black with purplish
reflections (blue on lateral lobe, axilla, and apex of scutel-
lum), dorsum rugose, proepistemum rugulose basally, and
Ms^ not constricted. The male petiole is longer (3.2x as
long as hind coxa) than O. rngulosa and Mt, is shorter
than the hind femora.

Three pupae (19, 26 6. Fulakora, Solomon Islands,
W. M. Mann, MCZ) were taken from a nest of Pheidolc
sp. (det. E. O. Wilson). These represent yet another
species that shares the above features but differs by hav-
ing a strong blue-green coloration, femur dark brown,
mesosoma dorsum deep areolate, and frenal line more
prominent. The pupae are typical for other members of
Orasema. This is the only pupal and ant-host record
known for the uichancoi-group.

DISTRIBUTION

New Guinea (B, Fig. 299).

ETYMOLOGY

Named in honour of Dr. Z. Boucek for his monumental
effort on reclassifying and organizing information on the
major groups of Chalcidoidea of the Indo-Pacific region.
His work has opened a new world for chalcidologists
everywhere.

Orasema promecea sp, nov.

Fig. 95

TYPE MATERIAL

Holotype, 6 , "Jimmi V. — Baiyer R./ 1750 m. New
Guinea/ 11.6-25.1979/ J. Sedlacek." "HOLOTYPE/
Orasema/ promecea Heraty." Deposited in AEI.
Paratype: Papua New Guinea: Central Dist., Guary |?|.
1900-2 100 m, X.1968. N. L. H. Krauss (I c^, BPBM).

I)ia(;no.sis

Recognized by having the forewing completely pilose,
mesosoma elongate and rugose-alveolate with propodeum
broadly curved in profile, face glabrous, and petiole long
and slender with a small basal flange. Additionally, the
lateral lobe, axilla, upper mesepimeron, and callus are
swollen and glabrate.

MALE

Length, 3.6-3.9 mm. Head, mesosoma, petiole, and base
of coxae black with faint blue-green reflections on head
and mesosoma, vertex with violet reflections; antennal
flagellum black; scape, pedicel, and anellus yellowish
brown: mandibles, apex of coxae, and rest of legs white
to yellow, tip of apical tarsomere of each leg slightly
darker. Wings lightly infuscate, venation dark brown.

Head subtriangular, 1.3x as broad as high, posterior
margin of gena broadly rounded: occiput broadly round-
ed; lateral ocellus separated from occiput by own radius;
LOL 0.7x OOL. Face broadly rounded and glabrous with
cheek lightly pitted, frons lateral to scrobal depression
and cheek swollen: scrobal depression narrow, margins
rounded and indistinct, rugose just below median ocellus;
vertex between ocelli rugulose, ocellar-ocular groove
shallowly impressed; temple narrow and glabrate: occiput
glabrate, occipital carina prominent, extending just
beyond lateral ocellus. Eyes separated by l.7-l.8x their
height, margined by narrow smooth ocular groove. Malar
space 0.7x height of eye, malar depression shallowly
impressed and glabrate (Fig. 95). Clypeus glabrate,
bulging medially, epistomal sulcus weak, tentorial pit
deep, lateral margin of clypeus and supraclypeal area
shallowly impressed, anteclypeus subtruncate at apex.
Maxillary palpi 3-segmented, basal segment elongate,
labial palpi 2-segmented and short. Antenna 12-segment-
ed (Fig. 95): scape stout and cylindrical, slightly expand-
ed ventrally, reaching median ocellus; pedicel globose,
about 1 .5x as broad as anellus; anellus as broad as long;
flagellum 2.0x height of head: funicle 8-segmented, seg-
ments densely setose, setae papillate, no MPS: F2
2.8-3.8X as long as broad, l.l-1.3x F3, 0.8-0.9x as long
as scape, following segments cylindrical and subequal in
length; clava narrow and cylindrical, acute at apex. 1.5x
preceding segment, with 2 incompletely fused segments.

Mesosoma with dorsum strongly rugose-areolate; lat-
eral lobe of mesoscutum and axilla swollen and glabrous.
Mesoscutum with midlobe subtriangular. strongly elevat-
ed at margin; notauli deeply impressed and transversely
carinate. SSS carinate. Scutellum slightly longer than
broad, subtruncate at apex, separated at base from TSA
by SSS; frenal line nanow and foveate, partially obscured
by surface sculpture, frenal area rugulose, crescent-
shaped in dorsal view; axillula glabrate, axillular sulcus
obscure. Propodeal disc rounded and strongly rugose.

66

broadly curved in profile; postspiracular furrow broadly
foveate; callus glabrous and swollen, dorsally with dense
patch of erect hairs; metepimeral sulcus broadly
impressed and glabrate. Mesepimeron glabrate,
transepimeral sulcus broadly impressed; femoral groove
obscure; mesepisternum areolate-rugose laterally,
glabrous ventrally. Prepectus triangular and rugose to
foveate. Pronotum glabrate with broad foveate dorsal and
medial lines. Proepistemum glabrous. Coxae and femora
glabrate, femora with fine, dense setae dorsoapically;
hind tibia slender with 2 apical spurs. Forewing 3. Ox as
long as mesothorax, 2.7-2.8x as long as broad; complete-
ly pilose; costal cell narrow; marginal vein 0.27-0.33x as
long as forewing; stigmal vein 2. Ox as long as broad;
postmarginal vein 0.7x as long as marginal vein, almost
extending to apex of wing.

Metasoma with petiole 3.4-3.7x as long as hind coxa,
3.5-3.7X as long as propodeum; petiole long and cylindri-
cal with small basal flange, areolate with longitudinal
carinae. Mt, 0.8x as long as hind femur, glabrate; Mt-,
constriction narrowly impressed and smooth. Genitalia
with elongate median process, paramere narrow and
short; aedeagus subtruncate.

FEMALE

Unknown.

VARIATION AND ADDITIONAL SPECIES

The paratype is similar to the holotype but differs as fol-
lows: head completely black dorsally, scutellum with
glabrous area anterior to frenal line, which has prominent
violet reflections; hind femur with outer medial surface
light brown. Two other males (not here described) from
the Papuan subregion are similar but differ in several
aspects; male from Papua New Guinea (Wamena,
10-25. ii. 1960, BPBM) has dark brown femora, dark
scape, reticulate mesosoma dorsum, and broadly alveolate
propodeum; male from Bougainville (Crown Prince
Range, 1 l.vi.l956, BPBM) is similar morphologically but
has a completely rugose face excluding the clypeus, a
deep ocellar-ocular groove, and a strongly impressed
alveolate malar depression as in O. nigulosa.

DISTRIBUTION

Papua New Guinea (P, Fig. 276).

ETYMOLOCJY

From Greek pronwces, meaning elongate; referring to the
profile of the mesosoma.

Orasema rugulosa sp. nov.

Figs. 89-93, 96

TYPE MATERIAL

Holotype, 6, "NEW BRITAIN/ Gazelle Pen./ Bainings;
St./ Paul's 350 m./ Sept. 9, 1955." "J. L. Gressitt/
Collector." "HOLOTYPE/ Orasema/ rugulosa Heraty."
Deposited in BPBM.

Paratypes: Papua New Guinea; New Britain: Gazelle
Pen., Gaulim, 130 m, 28.X.1962, J. Sedlacek (19, 26 S,
BPBM); New Ireland: SW, Gilingil Pl'n. 2 m. 5.vii.l956,
J. L. Gressitt (1 9, BPBM).

DIAGNOSIS

Recognized by the following; forewing elongate and
completely pilose (Fig. 96), mesosoma elongate with
propodeum only slightly curved in profile (Fig. 91), and
face finely rugulose-alveolate (Fig. 90).

FEMALE

Length, 2.4 mm. Head and mesosoma including petiole
and coxae dark blue, sometimes with intense purple
reflections on face and lateral aspects of mesosoma and
coxae; gaster dark brown with weak violet reflections,
antenna, mandibles, and legs dark yellowish brown.
Wings weakly infuscate, venation brown.

Head subquadrate, 1 .5x as broad as mesosoma. poste-
rior margin of gena broadly rounded; occiput shallowly
emarginate; lateral ocellus separated from occiput by own
radius; LOL 1.2x OOL. Face rounded, frons bulging lat-
eral to scrobal depression, frons and cheek rugulose-areo-
late; scrobal depression broad and glabrate with weak
medial sculpture, lateral margin marked only by change
in sculpture; vertex rugulose-alveolate with rounded
interstices (Fig. 90), depressed and glabrous just beyond
lateral ocellus, without ocellar-ocular groove; temples
narrow and rugulose; occiput weakly strigate; occipital
carina absent. Eyes separated by 1.5-1.6x their height,
margined by narrow foveate sulcus. Malar space ().7x
height of eye, malar depression narrow, deep, and
foveate. Clypeus glabrate. tentorial pit and lateral margin
deeply impressed, cpistomal sulcus weak, antcclypcus
subtruncate; supraclypcal area glabrate, lateral margin of
supraclypeal area shallow. Palpi 3-segmented, maxillary
palpi long, labial palpi relatively short. Antenna 12-seg-
mcntcd; scape stout and cylindrical, almost reaching
median ocellus; pedicel globose, slightly broader than
anellus; anellus broader than long; flagellum 1.5x height
of head; funicle 8-scgnicntcd. segments moderately
setose, smooth with numerous MPS; F2 1.8x as long as
broad. 1.2x F3, about half length of scape, following seg-
ments subequal in length and equal in width; clava ovate,
as long as preceding 2 segments.

Mesosoma with dorsum rugosc-arcolate; lateral lobe of

67

mcsoscutuiii and axilla rounded and glabrous.
Mesoscutum with niidlobe triangular, strongly elevated
along anterior lateral margin: notaulus deeply impressed
and broadly foveate. SSS carinate. Scutellum l.7x as long
as broad; frenal line foveate. partially obscured by dorsal
sculpture, frenal area semicircular in dorsal view, rugu-
lose dorsally, strongly foveate ventrally. flat in profile;
axillula mostly smooth, without sulcus. Propodeal disc
rounded and slightly curved in profile, strongly rugose-
alveolate; postspiracular and metepimeral sulcus broadly
foveate and shallow; callus swollen and glabrous, with 5
to 6 long hairs dorsally. Mesepimeron glabrous,
transepimeral sulcus foveate and shallow; femoral groove
obscure; mesepistemum rugose, polished along anterior
margin and ventrally. Prepectus triangular; strongly nar-
rowed in ventral half, triangle of prepectus broadly
impressed. Pronotum glabrous with irregular grooves.
Proepisternum glabrous. Coxae and femora glabrate,
femora with short setae dorsoapically; hind tibia slender
(apex in glue). Forewing 2.9x as long as mesothorax,
2.7x as long as broad; completely pilose; costal cell nar-
row; marginal vein 0.27x as long as forewing; stigmal
vein twice longer than broad, expanded and rounded api-
cally; postmarginal vein 0.4x as long as marginal vein,
reaching half distance to wing apex. Hind wing long and
narrow, completely pilose.

Metasoma with petiole 2.1x as long as hind coxa, 1.8x
as long as propodeum; petiole rugulose and longitudinally
carinate with weak basal flange. Mt^ 1.2x as long as hind
femur, glabrous; Ms., constriction smooth. Hypopygium
bare. Ovipositor subapically expanded and strongly
curved forward; first valvula with 4 or 5 teeth beyond

subapical crest; second valvula with 3 strong, widely
spaced, transverse ridges (Fig. 93). Gonostylus broad and
not reaching cercus.

MALE

Length, 3.0 mm. Colour as in female, sometimes with
reddish reflections on mcsosomai dorsum and gaster.

Head as in female, but with shallow foveate ocellar-
ocular depression; LOL 0.9-1. Ix OOL. Eyes separated
by 1.5-1.6X their height. Malar space 0.6-0.8x height of
eye. Antenna 12-segmented, anellus as long as broad; fla-
gellum 1.5-1.6X height of head; funicle 8-segmented,
segments densely setose, no MPS; F2 1.9X as long as
broad, following segments subequal in length; clava
ovate, l.5x as long as preceding segment.

Mesosoma as in female; callus with several erect hairs;
mesepistemum shallowly rugose-areolate. Hind tibia with
2 spurs. Forewing 2.7x as long as mesothorax. 2.6-2.7x
as long as broad.

Petiole 2.9-3.4X as long as hind coxa. 3.2-3.6x as
long as propodeum, and alveolate with irregular, weak
carina, and weak basal flange. Gaster with Mt., as long as
hind femur; Ms^ constriction smooth. Genitalia with basi-
paramere broad and truncate at apex, paramere very nar-
row and short, (digitus hidden); aedeagus broadly rounded.

DISTRIBUTION

Papua New Guinea: New Britain Is. (R, Fig. 276).

ETYMOLOGY

From Latin ruga, meaning wrinkled; referring to the
sculpture of the face.

Orasema striatosoma-group

Two closely related species, O. striatosoma and O.fraud-
ulenta, are included here. This group is very different
from other groups of Orasema but the shape of the head
is similar to other species of Orasema, including O.
susanae Gemignani, from southern South America, and
to some species of Orasemorpha. Species in the striatoso-
wa-group have a divided callus as in Orasemorpha. but
the petiole is elongate and attached apically to the gaster
(not anteriorly as in Orasemorpha).

mesepimeron reticulate; callus swollen and glabrous.
Forewing hyaline and pilose (Fig. 104); basal area bare or
bare only along impression of cubital vein; speculum
absent; disc pilose and marginal fringe present; marginal
vein thickened along entire length; stigmal vein broad and
angled distally; postmarginal vein narrow and short, less
than twice length of stigmal vein. Petiole weakly truncate
basally with weak dorsal flange.

(JROUP DE.SC'RIPTION

Head triangular with eye small and protuberant (Fig.
102); face finely reticulate with deep, sharply impressed
pit just below median ocellus; ocellar-ocular depression
absent. Antenna II -segmented and funicle 7-segmented
in both sexes. Mesoscutum including lateral lobe of
mesoscutum finely reticulate to coriaceous; lower

Orasema striatosoma sp. nov.

Figs. 98-105

TYPE MATERIAL

Holotype. 9. "Kampala. Uganda/ April. 1965/ Denis F.
Owen." "HOLOTYPE/ Orasema/ striatosoma Heraty,
1990." Deposited in AEL

68

Paratypes: Rwanda: 40 km E of Kigali. 1575 ni,
9.xii.l959, E. S. Ross and R. E. Leech (29 9, CAS).
South Africa: Zululand: Spadeni Kop., 9.V.1926. R. E.
Turner (1 9, BMNH); Melmoth, 24.iv.1926, R. E. Turner
(19, BMNH). Uganda: Kampala, x.l965, D. F. Owen
(IcJ, AEI); Kampala, 26.vi.1938. H. Hargreaves (19,
BMNH); Kawanda, vi.l943, T. H. C. Taylor (19,
BMNH).

DIAGNOSIS

Distinguished from O. fraudulenta by the following:
scutellum finely striate (Fig. 101), and basal area of
forewing pilose except along impression of cubital vein
(Fig. 104).

FEMALE

Length, 2.6-2.8 mm. Head, mesosoma, and petiole dark
bluish green, cheek with strong purple reflections, mesos-
cutum with iridescent green reflections, scutellum with
strong green reflections, mesosoma laterally with weak
green to blue reflections; propodeum bluish laterally,
midline of propodeum and frenal area purple. Caster,
coxae, and femora basally dark brown with faint reddish-
blue reflections; pedicel and flagellum dark brown; scape,
trochanters, apex of femora, and rest of legs yellowish
brown; mandible dark brown. Wings hyaline, venation
pale brown.

Head 1.4— 1.5x as broad as mesosoma; occiput weakly
emarginate; lateral ocellus close to occiput, separated by
less than own radius; LOL 0.7-0. 9x OOL. Face relatively
flat, frons broadly impressed midway between toruli and
lower margin of eye, face and vertex finely reticulate;
scrobal depression narrow and shallowly impressed above
toruli, weakly reticulate, and including median ocellus;
temple minute posterior to eye; occiput glabrous, occipi-
tal carina absent but vertex at occiput sharp. Eyes separat-
ed by 2.8-3. Ix their height. Malar space I.4-1.6x height
of eye, malar depression shallow, obliterated close to eye.
Clypeus and supraclypeal area weakly rugulose, lateral
margin, tentorial pit, and epistomal sulcus very deeply
impressed, anteclypcus slightly rounded apically. Maxilla
and labium saclikc with larger sclerites adprcsscd to
membranous sac, no palpi apparent. Antenna 1 1 -segment-
ed; scape stout and cylindrical, reaching top of median
ocellus; pedicel globular; anellus slightly broader than
long; flagellum 1.5-1.7x height of head; funicular seg-
ments moderately .setose, smooth with numerous MPS; F2
().4x as long as scape, F2 2.4-3.0x as long as broad, fol-
lowing segments subcqual in length, equal in width; clava
elongate and cylindrical, as long as preceding 2 segments.

Mesosoma with mesoscutum. including lateral lobe,
finely reticulate to coriaceous; axilla coriaceous to finely
carinatc posteriorly; scutellum excluding frenal area fine-
ly striate with carinae converging on posterior midline.

Mesoscutum with midlobe subtriangular; notaulus broad-
ly and deeply impressed. SSS finely carinate. Scutellum
as long as broad, posterior margin of axilla declivous, and
scutellum on distinctly lower level than mesoscutum; fre-
nal line narrow, frenal area abrupt and rugulose; axillula
not distinguishable; axillular sulcus absent, axillular area
evenly sculptured laterally. Propodeal disc rounded, with
irregular vertical carinae and verrucose surface sculpture;
callus and metepimeron slightly swollen and glabrous,
callus divided into anterior and posterior region by dorsal
extension of metepimeral sulcus; postspiracular and
metepimeral sulci broadly impressed and coriaceous to
smooth. Upper mesepimeron glabrate, lower
mesepimeron and mesepistemum finely reticulate to cori-
aceous, glabrous ventrally. Prepectus triangular and coria-
ceous, sharply angled to midline from edge of mesoscu-
tum. Proepisternum coriaceous. Coxae glabrate; femora
weakly imbricate to smooth with fine short adpressed
setae apically; hind tibia slender with fine adpressed
setae, narrowed apically. Forewing 3. Ox as long as
mesothorax, 1.9-2.1x as long as broad; bare along
impression of cubital vein; costal cell relatively broad and
pilose; disc with small hairs; marginal vein 0.23-0.26x as
long as forewing and thickened along entire length; stig-
mal vein broad and slightly angled distally, almost twice
longer than broad; postmarginal vein narrow, twice as
long as stigmal vein.

Metasoma with petiole 0.9-1. 3x as long as hind coxa,
0.8-1.5X as long as propodeum; petiole subtriangular in
cross-section, weakly carinate to glabrate dorsally, and
coriaceous ventrally. Mt^ 0.5x as long as hind femur,
glabrate with sparse micropunctuation; Ms, with constric-
tion sharp and smooth. Hypopygium with few short sub-
lateral setae at apex. Ovipositor subapically expanded and
strongly curved; first valvula with weak subapical ridge,
several fine lateral teeth beyond subapical ridge; second
valvula with 9 strong transverse coalescing ridges apical-
ly (Fig. 99). Sheath broad at apex, gonostylus separated at
base.

MALE

Length. 2.7 mm. Colour and sculpture as in female.
Antenna 11-segmented (Fig. 100); anellus as long as
broad; otherwise antenna as in female and lightly sculp-
tured. Petiole 1.8x as long as hind coxa. I.7x as long as
propodeum; petiole cylindrical at base and dorsovcntrally
flattened at apex, weakly coriaceous to smooth with weak
median carina dorsally along most of length, apex of peti-
ole emarginate dorsally and separated from Mt, by semi-
circular membrane. Mt, and Ms, as in female. MSj,
enlarged and broadly rounded with sparse, short setae.
Genitalia large (Fig. 103), basiparamerc broad and trun-
cate apically with sharp median process, paranicre long
and broad, weakly scleroti/.cd; acdeagus broadly rounded.

69

DISTRIBl TION

Eastern and southern Africa (S, Fig. 276).

ETVMOI.OdY

From Latin stria and soma; referring to the striate scutei-
lum.

Orasema fraudulenta (Reichensperger) comb. nov.

Psilogaster frauduh'ntus Reichensperger, 1913:21 1-214,
figs. 1-m, 12. Ethiopia: Harrar [ZFMK, examined].

TYPE MATERIAL

Lectotype (here designated), 9 , "Psilogaster/ fraudulen-
tus/ n. sp./ Type! R." "bei Pheidole/ megaceph./ Fab."
•TYPUS." "Harrar, Abesi./ V/ Kristensen." "Coll.
Reichensperger." "Museum Koenig BONN." Female with
head and antennae enclosed in pupal exuvium and wings
missing, mounted on same card with males and ant host;
female between 2 males. Paralectotypes: 2 S S and 2
pupae of both sexes, mounted on same card as holotype.

DIAGNOSIS

Distinguished from O. sthatosoma by having the scutel-
lum finely reticulate, and basal area of the forewing bare.

FEMALE

Length, 2.9 mm. Dark metallic blue-green with faint iri-
descent reflections; mesosoma with bluish reflections
sublaterally; gaster dark brown with faint purple or green
reflections; antenna pale brown; coxae and most of femo-
ra dark brown with greenish reflections; trochanters, apex
of femora, and rest of legs yellowish brown. Wings hya-
line, venation pale brown.

Head 1.5x as broad as mesosoma; occiput slightly
rounded; lateral ocellus close to occiput, separated by less
than own radius; LOL 0.8x OOL. Face relatively flat,
frons broadly impressed midway between toruli and
lower margin of eye, face and vertex finely reticulate;
scrobal depression narrow and shallowly impressed above
toruli, weakly reticulate, and including median ocellus,
with deep pit below median ocellus; temple narrow and
weakly sculptured; occiput glabrous, occipital carina
weak or absent, vertex at occiput sharp. Eyes separated by
2.6x their height. Malar space 0.7x height of eye, malar
depression absent. Clypeus weakly reticulate, lateral mar-
gin of clypeal area, tentorial pit, and basal groove deeply
impressed, anteclypeus slightly rounded. Maxilla and
labium elongate, no palpi apparent. Antennal segments
not discernible within pupal exuvium; flagellum 1.3x
height of head; flagellomeres densely setose, smooth with
sparse MPS.

Mesosoma with dorsum finely reticulate, including lat-

eral lobe of mesoscutum and axilla. Notauli and SSS nar-
rowly impressed and weakly foveate. Scutellum slightly
longer than broad, posterior margin of axilla rounded, and
scutellum below level of mesoscutum: frenal line a nar-
row glabrous band, not impressed, frenal area finely rugu-
lose; axillular sulcus absent, axillula longitudinally cari-
nate. Propodeal disc rounded, finely reticulate-rugulose;
postspiracular and melepimcral sulcus narrow and shal-
low; callus divided into anterior and posterior regions by
dorsal extension of metepimeral sulcus; callus swollen
and glabrous, metepimeron weakly sculptured. Upper
mesepimeron weakly reticulate, lower mesepimeron and
mesepisternum finely reticulate, glabrous ventrally.
Prepectus and pronotum finely reticulate, prepectus weak-
ly angled to midline below mesoscutum. Proepistemum
finely reticulate, strongly swollen and smooth posteriorly.
Coxae weakly coriaceous to smooth; hind femur imbri-
cate; hind tibia stout along entire length, slightly nar-
rowed apically. Forewing 2.4x as long as mesothorax,
2.0-2. Ix as long as broad; basal area and costal cell bare;
disc moderately pilose with short hairs; marginal vein
0.26x as long as forewing and thickened along entire
length; stigmal vein broad and slightly angled distally,
slightly longer than broad; postmarginal vein elongate
and broad, 0.5x as long as marginal vein.

Metasoma with petiole 0.9x as long as hind coxa. 0.8x
as long as propodeum; petiole cylindrical and slightly
flattened dorsoventrally at apex, aciculate to smooth api-
cally. Mt^ 0.5x as long as hind femur, glabrate with
sparse minute piliferous punctures; Ms^ strongly con-
stricted, furrow glabrous, anterior region circular.
Hypopygium bare except for short hairs at apex.
Ovipositor subapically expanded and curving slightly
cephalad; first valvula with subapical ridge and lateral
line of 6 teeth (second valvula withdrawn). Gonostylus
broad and separated at base.

MALE

Length, 3.0 mm. Generally agrees with female.

Head 1.5x as broad as mesosoma; LOL 0.8-1. Ox
OOL. Eyes separated by 2.7-2.9x their height. Malar
space 1.2-1.4X height of eye. Antenna 1 1-segmented;
scape stout and cylindrical, slightly narrowed at apex,
reaching top of median ocellus; pedicel subconate: anel-
lus slightly broader than long; flagellum 1.8-1.9x height
of head; funicular segments moderately setose and lightly
sculptured; F2 0.6x as long as scape, F2 2.8x as long as
broad; following segments subequal in length, equal in
width; clava elongate and cylindrical, as long as preced-
ing 2 segments.

Mesosoma more slender than in female.

Metasoma with gaster elongate, glabrate; Mt^ l.lx as
long as hind femur. Ms^^ small, rounded, and sparsely
setose. Genitalia relatively small, basiparamere truncate.

70

and apex with sharp median process; paramere long,
broad, and poorly sclerotized with few elongate setae on
outer margin; aedeagus broadly rounded.

of the Pheidole larva. He stated that the larva is similar to
that described for Orasema (Wheeler, 1907) with a series
of nodules along the body segments on each side.

BIOLOGY AND IMMATURE STAGES

The type material was taken from a nest of Pheidole
megacephala Fab. The pupa of this species is similar to
that of other Orasema, and includes 3 nodules along the
dorsal margin of the petiole, and a series of raised ridges
and nodules along the tergal lines of the gaster
(Reichensperger, 1913). Reichensperger refers to the "mit-
tlere Larvenstadium" as attaching to the prostemal region

DISTRIBUTION

Ethiopia and Yemen (F, Fig. 276).

MATERIAL EXAMINED

Ethiopia [Abyssinia]: Harrar (29 9, USNM). Yemen [W
Aden Prot.]: Wadi Dareija, SW of Dhala, ca. 1500 m,
28. ix. 1937, H. Scott and E. B. Britton (Id, \16
[abdomen missing], BMNH).

Orasema koghisiana-group

This is a monobasic group and is peculiar among Old
World Oraseminae in that the face is smooth and relative-
ly flattened, and the labrum has 6 to 8 elongate digits
(usually 4 in other orasemine taxa) (Figs. 189-190). It is
unusual within Orasema by possession of a prepectus that
is narrowed ventrally and with the medial area almost
glabrous (Fig. 219), the stemaular area reticulate, and the
mesepistemum not strongly swollen.

Orasema koghisiana sp. nov.

Figs. 107-108, 1 10, 189-190, 21 1-212, 219, 236, 247

TYPE MATERIAL

Holotype, 9 , "NEW CALEDONIA:/ Mtns des Koghis/
400-600 m, 1.1969." "N. L. H. Krauss/ Collector/ BISH-
OP MUSEUM." "HOLOTYPE/ Orasema/ koghisiana
Heraty." Deposited in BPBM.

Paratypes: New Caledonia: Mtns des Koghis, 400-600
m, i.l969 {lis 6), ii.l973 {26 6), N. L. H. Krauss
(BPBM); same data, 200^00 m, i.l969 (1(5, BPBM);
Yahoue, ii.l966 (Ic5), ii.l978 (lc5), N. L. H. Krauss
(BPBM). New Hebrides: Efate Is.: Vila, 0-200 m,
ii. 1973 (1 9), ii.l973 (Id); i-ii.l977 (1 (5). i. 1976 (2c? d),
iii.1970 (Id), ii. 1970 (Id) xi.l978 (Id, CNC), N. L. H.
Krauss (all but 1, BPBM); Epi Is.: Vaemali, 100-150 m,
6-10.viii.1967, J. and M. Sedlacek (19. wings and anten-
nae on slide no. 9574 BPBM); Lowekcwou, 0-100 m,
31.viii.l979. W. C. Gagne. G. M. Nishidai, and G. A.
Samuelson (19, Id. BPBM); Ambrym Is.: Ranon to Mt
Toyo, 0-500 m, 2.ix.l979. Gagnc. Nishida. and
Samuelson (Id. BPBM): Erromango Is.: Dillon Bay.
()-l()() m. iii.1978. N. L. H. Krauss (Id. BPBM): Macwo
Is.: Sounwari. 15'^ 23S 168° 07E. 0-360 m. 4-5. ix. 1979,
Gagnc. Nishida. and Samuelson (Id. BPBM); Malekoula
Is.: Port Sandwich (Id, BPBM): Sanlo Is.: Matantas.
0-100 m, ll.ix.i979, G. M. Nishida and G. A.

Samuelson (Id, BPBM); Tongariki Is.: 0-300 m,
29.viii.1979, G. M. Nishida (3dd, BPBM); Vanoua
Lava Is.: Sola to Chelva Riv., 0-260 m, 16. ix. 1979,
Nishida and Samuelson (Id, BPBM).

DIAGNOSIS

Recognized by the following: face (Fig. 189), lateral lobe,
and axilla smooth, labrum 6- to 8-digitate, stemaular area
reticulate (Fig. 219). and propodeum glabrous laterally
(Fig. 247).

FEMALE

Length, 2.6-3.1 mm. Head and mesosoma glossy green or
blue-green with varied patches of reddish reflections;
gaster dark brown to black; mesepimeron and propodeum
with blue reflections; coxae dark brown with faint green-
ish reflections; antennal flagellum black: scape and legs
yellowish brown. Wings hyaline, venation dark brown.

Head subtriangular, 1.2-1.4x as broad as mesosoma,
occiput weakly rounded; lateral ocellus separated from
occiput by own radius; LOL 0.7x OOL. Face broadly
rounded, smooth, and polished with scattered fine setae,
cheek lightly pitted. Irons swollen lateral to scrobal
depression; scrobal depression narrow and strongly
impressed, each scrobal channel irregularly foveate (Fig.
189); temple narrow and polished; vertex with weak
depression along ocellar-ocular groove; occiput aciculate.
dorsal margin lightly strigate. not carinate. Eyes separated
by 1.6x their height. Malar space 0.8x height of eye.
malar depression hardly impressed. Clypeus glabrate.
cpistomal sulcus weakly impressed; anteclypeus only
slightly rounded. Labrum 8-digitatc. setae bristlclike.
Mandible moderately slout; palpi 3-segmenlcd. Antenna
1 1-segmented; scape slender, reaching median ikcIIus:
pedicel as long as broad; ancllus present, very small; lla-
gellum I. Ox height of head; funicic 7-scgmented: seg-
ments densely setose, surface smooth, with numerous

71

MPS; F2 l.8x as long as broad, following segments sub-
equal in length, equal in width, each clearly separated
(Figs. 108, 21 1-212); clava acute, not differentiated from
funicle.

Mcsosonui v\ ith midlobe of mesoscutum transversely
carinate anteriorly to rugose posteriorly; lateral lobe and
axilla glabrous and swollen; scutelluni lightly rugulose to
glabrate. Notauli and SSS shallow foveate. Scutelluni
slightly longer than broad, base separated from TSA by
small fovea; frenal line narrowly foveate, frenal area
glabrous, crescentic in dorsal view; axillula mostly
glabrate, axillular sulcus foveate. Propodeal disc glabrous
laterally, narrow median band of alveolate sculpture
forming a band dorsally to postspiracular furrow, ventral
band deeply foveate; callus polished and rounded, sparse-
ly setose dorsally. Mesepimeron glabrate. transepimeral
sulcus weakly foveate; femoral groove broad, weakly car-
inate anteriorly; sternaular area marked by triangular band
of reticulate sculpture. Proepisternum glabrate. Coxae
glabrate, mid coxa weakly strigate; femora glabrate, mid
and hind femur with fine, dense setae apically. Forewing
2.3x as long as mesothorax, 2.5x as long as broad (Fig.
110); basal area and speculum bare; costal cell and disc
pilose; stigmal vein broad, only slightly longer than wide;
postmarginal vein 5. Ox as long as stigmal vein, reaching
half distance to apex of wing.

Metasoma with petiole 1.2x as long as hind coxa, l.lx
as long as propodeum; petiole irregularly carinate, with
small basal flange. Mt, longer than hind femur, glabrate;
Ms., with constriction deeply impressed and smooth.
Hypopygium with few minute setae. Ovipositor expanded
subapically; first valvula with sharp lateral ridge from
apex to broadest subapical width (margin of ventral sur-
face that receives second valvula), and 3 to 4 lateral teeth;
second valvula broad with several sharp teeth laterally.
Gonostylus broad and setose, basal separation not visible
in available material.

MALK

Length. 2.0-2.6 mm. Colour as in female except scape
and hind femur medially dark brown to black w ith strong
blue-green reflections.

Head 1.4-I.5X as broad as mesosoma, cheek sparsely
punctate; ocelli large, LOL 0.7-1. Ox OOL; occipital cari-
na present or absent (obscured with occipital carinae).
Labrum 6- to 8-digitate. Mandible moderately stout; palpi
3-segmented. Antenna 1 1 -segmented; tlagellum 1.3-l.4x
height of head; funicle 7-segmented, MPS dense.

Mesoscutum with midlobe finely alveolate to scabricu-
lous with faint indication of transverse carinae; scutellum
rugose; frenal area abrupt apically. Forewing 2.1-2.2x as
long as broad; stigmal vein l.0-1.5x as long as broad,
almost perpendicular or distinctly angled distally.

Petiole 1.9-2.7X as long as coxa, 1.9-2. 4x as long as
propodeum; irregularly carinate with verrucose surface
sculpture. Mt^ as long as hind femur. Genitalia elongate,
paramere with long subapical seta, basiparamere with
sharp median process; aedeagus broad.

VARIATION

Specimens from New Caledonia and New Hebrides are
considered conspecific although there is some minor vari-
ation. Specimens from the New Hebrides differ in having
the head of females 1.3-1.4x as broad as the mesosoma
(versus 1.2x in New Caledonia), and the mesosoma dor-
sum with stronger sculpture (scutellum rugose and axilla
weakly carinate).

DISTRIBUTION

New Caledonia and Hew Hebrides (K, Fig. 276).

ETYMOLOGY

Named after the type locality. Mount Koghis.

Orasema glabra-group

This monobasic group is recognized by the completely
smooth head (Figs. 126-127), scrobal depression with
parallel channels, occipital carina absent, propodeum
evenly rugose, basal area of the wings and speculum bare.
and first valvula with 9 or 10 minute lateral teeth. Some
features are shared with the O. coloradensis-group of
species found in the New World.

Orasema glabra sp. nov.

Figs. 119. 125-128, 130, 132

TYPK MATERIAL

Holotype, 9, "S. AFRICA; Transvaal/ 15 km E. Klaserie/
18-31. XII. 1985/ H. and A. Howden." "HOLOTYPE/
Orasema/ glabra Heraty." Deposited in CNC.
Paratypes: South Africa: Cape Province: Somerset East
[?], 27-31.i.l93l. R. E. Turner (1 9, BMNH); Transvaal:
Klaserie. 15 km E. Guernsey Farm. 19-3l.xii.1985, W.
Mason (19, CNC); same locality. 19-3 l.xii. 1985. M.
Sanborne, yellow pan trap (19, slide no. 943, CNC).

72

DIAGNOSIS

Recognized by the following: face, lateral lobe, axilla,
posterior half of scutellum, and frenal area glabrous (Figs.
127, 130). Distinguished from species in the valgius- and
assectator-groups by the following: propodeal disc even-
ly rugose and relatively flat, mid coxa without a mid-ven-
tral sulcus, and petiole short (0.9-1. 2x as long as hind
coxa) (Fig. 126).

FEMALE

Length, 2.2-2.5 mm. Head, mesosoma. and petiole black,
head and mesosoma laterally with blue to purple reflec-
tions, mesosomal dorsum with green reflections; gaster
dark brown; pedicel, flagellum, and femora dark brown;
scape, apex of femora, and rest of legs yellowish brown.
Wings hyaline, venation pale yellowish brown.

Head subtriangular l.lx as broad as long (Fig. 127),
1.2-1.3X as broad as mesosoma; occiput transverse; LOL
1.4-1. 9x OOL. Face broadly rounded, smooth and pol-
ished with scattered minute pits; scrobal depression nar-
row, lateral margin broadly rounded medially with 2
impressed vertical channels ending in shallow pit about
0.6x distance to median ocellus (Fig. 127); temple broad
and weakly aciculate; no ocellar-ocular depression;
occiput aciculate, dorsal margin broadly rounded, occipi-
tal carina absent. Eyes separated by 1.4-1.5x their height,
ocular groove narrow and smooth. Malar space 0.6x
height of eye, malar depression broad and shallow.
Clypeus transverse and glabrate, lateral margin and epis-
tomal sulcus shallowly impressed, anteclypeus subtrun-
cate. Labrum 4-digitate. Mandible moderately stout, palpi
3-segmented. Antenna 1 1 -segmented; scape stout and
cylindrical, reaching 0.7x distance to median ocellus;
pedicel 1.7x as long as broad, slightly broader than anel-
lus at apex; anellus small; flagellum 0.9-1. Ox height of
head; funicle 7-segmented, segments densely setose with
numerous MPS; F2 1.4x as long as broad, following seg-
ments subequal in length, and slightly increasing in width;
clava conate, slightly longer than preceding 2 segments.

Mesosoma with midlobe of mesoscutum and scutellum
finely reticulate; lateral lobe and axilla glabrous, lateral
lobe strongly swollen, axilla rounded (Fig. 130).
Mesoscutum with notauli deeply impressed and crenulate.

SSS foveate. Scutellum as long as broad, separated at
base from TSA by narrow foveate SSS; frenal line nar-
rowly and shallowly impressed, frenal area semicircular
and glabrous; axillula glabrate, axillular sulcus foveate
and prominent. Propodeal disc evenly rugose, subtriangu-
lar, and relatively flat; postspiracular and metepimeral
sulci narrowly impressed and without transverse carinae;
callus and metepimeron glabrous, callus with several
short setae. Mesepimeron glabrate, transepimeral sulcus
shallow; femoral groove obscure; mesepisternum reticu-
late laterally, glabrate anteriorly and ventrally. Prepectus
reticulate. Pronotum glabrate with irregular medial line.
Proepisternum smooth. Coxae and femora smooth to
weakly imbricate; hind femur moderately setose dorsally.
Forewing 2.4x as long as mesothorax, 2.4-2.6x as long as
broad; basal area and speculum bare; costal cell relatively
narrow; rest of wing moderately pilose; stigmal vein more
than twice as long as broad, perpendicular to wing mar-
gin; postmarginal vein 0.35-0.4x as long as marginal
vein, reaching less than half distance to apex of wing.

Metasoma with petiole 0.9-1.2x as long as hind coxa,
0.8-1. Ox as long as propodeum; petiole granulate, with
strong basal flange (Fig. 132). Mt^ l.l-1.4x as long as
hind femur, glabrate; Ms, constriction abrupt and rugu-
lose. Ovipositor subapically expanded and gently curved
forward; first valvula strongly excavated ventrally,
notched subapically beyond subapical crest, with strong
lateral ridge from ventral margin of crest to dorsal medial
margin, several fine lateral teeth beyond subapical crest,
apex finely aciculate; second valvula broad and smooth
with several strong lateral teeth (Fig. 128). Gonoslylus
narrow, setose apically, and distinctly separated from sec-
ond valvifer.

MALE

Unknown.

DISTRIBUTION

South Africa (G, Fig. 276).

ETYMOLOGY

Name refers to the glabrous face and mesosomal dorsum.

Orasema assec/a/or-grou p

The differences between species of this group are small
and more material is necessary to verify the stability of
character states described. It is noteworthy that both
Orasema assectator and O. initiator were found oviposit-
ing on leaves of tea, and (). assectator was reared from
Pheidolc sp. nesting under a tea plant. Possibly the pre-
sent Ethiopian/Indo-Chinese distribution of this group

may be a recent event associated with the movement of
tea throughout the region. Similarity of this group to
species of the Orasema hakeri-p\n\\) in the New World is
based on the following: mesosoma and propodeum evenly
reticulate, callus bare, stigmal vein narrow and perpendic-
ular, postmarginal vein short, wings hyaline, and petiole
short.

73

GROUP DKStRIPTION

Face completely reticulate: occipital carina absent; with-
out occllar-ocular depression. Malar depression absent.
Antenna 1 1 -segmented and funicle 7-segmented in both
sexes. Mesosoma completely reticulate dorsall> including
lateral lobe and axilla: axillular sulcus absent and axillula
indistinct from scutellum: callus swollen and glabrous:
lower mesepimeron reticulate. Forewing hyaline: basal
area and speculum bare: disc moderately setose: stigmal
vein narrow and elongate, perpendicular to wing margin
or nearly so: postmarginal vein poorly defined and usual-
ly less than 0.3x as long as marginal vein. Petiole
0.8-1.5X as long as hind coxa in females, 1.7-2.3x as
long as hind coxa in males, with basal flange. Ovipositor
expanded, subapical ridge broadly rounded: first valvula
with lateral line of several minute teeth from apex to
ridge. Gonostylus setose, separated at base from second
valvifer.

Orasema assectator Kerrich

Figs. 111-112

Orasema assectator Kerrich, 1963:367-368. India:
Assam [BMNH, examined].

TYPE MATERIAL

Holotype, 9, "ASSAM/ Tocklai/ 1962/ Ex Ants/ CIE.
18496 [744/5 on left end]." "Type." "Orasema female/
assectator sp. n./ G. J. Kerrich det 1963/ HOLOTYPE."
"B.M. TYPE/ HYM./ 5.2066."

Paratypes: 89 9 (BMNH), 49 9 (USNM), same data, or
with "on tea" instead of "ex ants" and CIE No. 18869.

DIAGNOSIS

Differs from other species in the assectator-gro\x^ by:
head slightly transverse (Fig. 112), supraclypeal area
glabrate [stated as finely reticulate in original descrip-
tion], mesoscutum broadly rounded anteriorly (Fig. Ill),
axillular sulcus indistinct and scutellum rounded laterally,
propodeal disc evenly sculptured and without median
carina, pronotum without lateral prominence below spira-
cle, and coxae completely reticulate. Orasema assectator
is known only from the type locality and this may only be
a small form of O. initiator. However, the differences list-
ed here are distinctive.

FEMALE

Length, 1.5-1.9 mm. Head, mesosoma, and petiole dark
blue: gaster dark brown with faint metallic sheen: femora
brown medially with faint metallic reflections: antenna,
apex of femora and rest of legs yellowish brown, flagel-
lum slightly darker. Wings hyaline, venation clear.

Head slightly transverse with large eyes, 1.3-1.4x as

broad as high. 1.1 1.3x as broad as mesosoma: occiput
broadly rounded: lateral ocellus almost touching occipital
margin: LOL 1.4-1.9x OOL. Face relatively flat, reticu-
late: scrobal depression shallow, lateral margin rounded
and glabrate, weakly sculptured medially: vertex more
finely reticulate than face: temple narrow and coriaceous:
occiput coriaceous, dorsal margin abrupt. Eyes separated
by 1.6-1.8X their height. Malar space 0.7-0.8x height of
eye. Clypeus and supraclypeal area glabrate and slightly
swollen, tentorial pits deep, lateral margin of clypeus
shallowly impressed, frontogenal sulcus parallel and
meeting middle to outer margin of torulus. Labrum 4-dig-
itate. Mandible moderately stout: maxilla and labium
large, maxillary palpus 3-segmented and enlarged, labial
palpus 1 -segmented and elongate. Antenna 11 -segment-
ed: scape cylindrical, reaching 0.7x distance to median
ocellus: pedicel subconical: anellus large: flagellum
1.2-1.4X height of head: funicular segments sparsely
setose basally to densely setose apically, with numerous
MPS: F2 0.5x as long as scape, F2 1.3-1.8x as long as
broad, following segments subequal in length, equal
in width: clava ovate, slightly longer than preceding 2
segments.

Mesosoma with dorsum completely and finely reticu-
late including scutellum and axilla: lateral lobe of mesos-
cutum coriaceous or more finely reticulate than rest of
dorsum. Mesoscutum with midlobe subtriangular with
anterior margin broadly rounded: notauli narrowly and
deeply impressed. SSS foveate. Scutellum slightly longer
than broad, widely separated at base from TSA by single
shallow medial extension of SSS; frenal line forming nar-
row glabrous band dorsally, frenal area finely rugulose to
reticulate; axillula reticulate. Propodeal disc slightly
rounded, reticulate; postspiracular furrow and metepimer-
al sulcus shallow and broadly impressed: callus slightly
swollen, glabrous laterally with small callar nib;
metepimeron reticulate. Upper mesepimeron weakly cori-
aceous, lower mesepimeron and mesepisternum reticu-
late, glabrous anteriorly and ventrally. transepimeral sul-
cus shallow foveate; femoral groove obscure. Prepectus
triangular and reticulate. Pronotum finely reticulate, even
dorsally and without any prominence. Proepisternuni
reticulate. Fore and mid coxa umbilicate. hind coxa retic-
ulate: hind femur weakly coriaceous, broad and slightly
flattened. Forewing 2.5x as long as mesosoma, 2.5-2.6x
as long as broad: basal area and speculum bare, costal cell
with few setae in single medial line: stigmal vein narrow
and twice as long as broad, very slightly angled distally;
postmarginal vein indistinct, about twice length of stig-
mal vein.

Metasoma with petiole 0.6-().8x as long as hind coxa,
0.6-0. 8x as long as propodeum: petiole cylindrical and
1.5x as long as broad, verrucose. Mt, 0.8-1. Ox as long as
hind femur, glabrous: Ms, with constriction sharp lateral-

74

ly. Second valvula of ovipositor broad with more than 10
strong teeth along lateral margin, transverse ridges
extending medially from base of teeth but not complete.

MALE

Unknown.

VARIATION

Two specimens from southern India (Yercaud, 7.vi.l982,
G. J. Spencer, 1 9, CNC) and Sri Lanka (Nugegoda Prov.,
7.vii.l970, P. B. Karunaratne, 1 9, CNC) are similar to
the type material of O. assectator. The Sri Lankan speci-
men is more distinctly rugulose than described for O.
assectator, it has a broader mesosoma and is more similar
to O. initiator or O. nigra. The south Indian specimen has
more complete reticulate sculpture, the scutellum flat-
tened dorsally. axillula glabrous, and the axillular carina
conspicuous as compared to other species in which the
scutellum is broadly rounded laterally. The shape of the
head, antenna, sculptured mesepimeron. and short petiole
are consistent and are considered diagnostic. Both speci-
mens have a body length of 2.8 mm and the forewing is
only 2.2x as long as broad. These specimens are consid-
ered as closer to O. assectator than to O. initiator and
emphasize the small differences between the 2 species.

BIOLOGY

The biology of this species was described by Das (1963)
and Kerrich (1963). Females deposit their eggs into punc-
tures on the underside of tea leaves. Secondary fungal
infections of oviposition punctures resulted in Sewing
Leaf Blight on tea (Das, 1963). Larvae were associated
with immature stages of Empoasca flavescens (Fabr.)
(Hemiptera). which were presumed by the authors to be
the intermediate host necessary to gain access to the ant
nest. Orasema assectator was reared from a species of
Pheidole. Larval and pupal stages were described by Das
(1963) and are typical for Orasema.

DISTRIBUTION

Known only from northeastern India, with questionable
records from southern India and Sri Lanka (A, Fig. 276).

Orasema nigra sp. nov.

Figs. 113-116

TYPE MATERIAL

Holotype, 9. "Roy. Natal Natl. Park/ 1.24.71 So. Africa/
H. and M. Towncs." "HOLOTYPE/ Orasema/ nigra
Heraty." Deposited in AEI.

Paratypes: South Airica: Natal: Royal Natal N. P..
27. i. 1971. H. and M. Townes (29 9. AEI). Uganfm:
Kawanda, 30.iv[71.1963, D. J. Grealhead, ex Leucoptera

sp. [with large handwritten "?"] on coffee arabica {\S ,
BMNH).

DIAGNOSIS

Differs from other species in the assectator-grouTp by:
head subtriangular (1.3x as broad as high; Fig. 1 14). supr-
aclypeal area reticulate, mesoscutum broadly rounded
anteriorly, propodeal disc weakly sculptured with median
carina, pronotum even and without lateral prominence,
and coxa weakly coriaceous basally to glabrate apically.

FEMALE

Length, 2.5 mm. Head and mesosoma dark bluish black,
gaster dark brown; flagellum dark brown; coxae and
femora black; scape, apex of femora and rest of legs yel-
lowish brown. Wings hyaline, venation pale brown.

Head subtriangular (Fig. 1 14), 1.3x as broad as high,
1.2-1.3X as broad as mesosoma; occiput broadly emar-
ginate; lateral ocellus separated from occiput by own
radius; LOL l.Ox OOL. Face broadly rounded, complete-
ly reticulate; scrobal depression narrow, lateral margin
broadly rounded, coriaceous medially; temple relatively
broad and coriaceous; occiput weakly aciculate; vertex
rounded. Eyes separated by 1.9-2. Ox their height. Malar
space 0.8-0.9X height of eye, malar depression broad and
vaguely impressed (Fig. 1 15). Clypeus weakly .sculptured,
supraclypeal area minutely reticulate, tentorial pit strong-
ly impressed, lateral margins of clypeal area shallowly
impressed. Labrum 4-digitate. Mandible moderately
stout; maxilla and labium large, maxillary palpus elongate
and 3-segmented, labial palpus elongate and 2-segment-
ed. Antenna 1 1 -segmented; scape slender and cylindrical,
reaching 0.7x distance to median ocellus; pedicel glo-
bose; anellus glabrous and small; flagellum l.lx height of
head; funicular segments moderately setose with numer-
ous MPS; F2 0.5x as long as scape, 2. Ox as long as broad,
F3 slightly longer than broad, following segments sube-
qual in length and slightly broader; clava ovate, as long as
preceding 3 segments.

Mesosoma quadrate in profile; dorsum finely reticu-
late. Mesoscutum with mid and lateral lobes broadly
rounded anteriorly; notauli and SSS narrow, deeply
impressed, and crenulate. Scutellum slightly longer than
broad, broadly separated from TSA at base by deep fovea,
both scutellum and frenal area rounded in profile; frenal
area vaguely separated by indistinct frenal line; axillula
obliquely carinate, axillular sulcus indistinct. Propodeal
disc broadly rounded, weakly reticulate, median area are-
olate with inegular vertical carina: postspiracular furrow
broad and fovcatc; callus and metcpimeron swollen and
glabrous; metepimeral sulcus narrowly impressed.
Mesepimeron glabrous, transepimeral sulcus foveate;
femoral groove narrow and fovcatc; mcsepistemum finely
reticulate, glabrous niid-venlrally anil anteriorly.

75

Prcpectus triangular, glabralc ventrally and deeply
fovcaie dorsally. Pronoium and proepisiornum coria-
ceous. Coxae coriaceous basally, smooth apically; hind
femur smooth ventrally, coriaceous mid-dorsally. and
sparsely setose. Forewing 2.5x as long as mesothorax.
2.2-2.3X as long as broad; basal area and speculum bare
except few sparse setae; costal cell with sparse medial
band of setae; disc of wing moderately setose; stigmal
vein narrow, 3. Ox as long as broad, almost perpendicular
to wing margin; postmarginal vein 2.0x as long as stigmal
vein.

Metasoma with petiole 1.0-1.5x as long as hind coxa
(Fig. 113), 1.0-1. 5x as long as propodeum; petiole cylin-
drical, weakly rugose-carinate. Mt^ l.lx as long as hind
femur, glabrate; Ms, constriction sharp and weakly crenu-
late laterally. Ovipositor (Figs. 113, 1 16); second valvula
broad and glabrous with several strong teeth along lateral
margins, connected dorsally by weak transverse ridges.

MALE

Length, ca. 1.5 mm (head missing). Colour, sculpture,
and wings as in female. Propodeum glabrate with median
carina. Petiole 2. Ox as long as hind coxa, 2.2x as long as
propodeum. Genitalia difficult to discern on specimen but
apparently typical for genus.

BIOLOGY

Unknown. The association with Leucoptera (Homoptera:
Diaspididae) for 1 specimen is unlikely, and it was proba-
bly collected together with the host plant.

DISTRIBUTION

South Africa (Natal) and Uganda (N, Fig. 276).

Orasema initiator Kerrich

Figs. 117-118.191. 220, 237, 260-26 1

Orasema initiator Kerrich, 1963:368. India: Assam
[BMNH, examined].

TYPE MATERIAL

Hololype, 9, "ASSAM/ 1955/ G. M. Das/ CLE. Coll/
14226 [S.N. 9 on right end, Reg 987/4 on left end]."
"Orasema sp. 9/ G. J. Kerrich del 1955/ HOLOTYPE."
"B.M. TYPE/ HYM./ 5.2067."
Paratypes: same data as holotype |29 9. BMNH].

DIAGNOSLS

Differs from other species in the (/.v.v<'(7(//o/-group by:
head subtriangular (1.2-1.3x as broad as high) (Fig. 191 ).
supraclypeal and clypeal area glabrous and narrowed at
toruli, mandibles slender with long subapical teeth, mid-
lobe of mesoscutum with anterior lateral corners promi-

nent (Fig. 237), propodeal disc weakly sculptured with
median carina, pronoium sometimes with conical promi-
nence just below spiracle (Fig. 237, arrow), and coxae
coriaceous basally to glabrate apically.

FEMALE

Length, 2.1-2.8 mm. Body black; pedicel and flagellum
dark brown; coxae and femora (including trochanters)
black; scape, apex of femora and rest of legs yellowish
brown. Wings hyaline, venation clear.

Head subtriangular. 1.2-1.4x as broad as high,
1.2-1.5X as broad as mesosoma; occiput broadly emar-
ginate; lateral ocellus separated from occiput by less than
own radius; LOL 1.3-1.7x OOL. Face weakly rounded,
reticulate; scrobal depression narrow, lateral margin
rounded and glabrous, weakly coriaceous medially; ver-
tex more finely reticulate posterior to line drawn across
vertex between median and lateral ocelli; temple narrow
and coriaceous; occiput coriaceous, dorsal margin abrupt.
Eyes separated by 1.6-1.8x their height. Malar space
0.6-0.9X height of eye. Clypeus and supraclypeal area
glabrous and swollen, tentorial pit strongly impressed, lat-
eral margin of clypeal area shallowly impressed, fronto-
genal sulcus converging to inner margin of torulus.
Labrum 4-digitate. Mandible slender with elongate sub-
apical teeth (Fig. 191); maxillary palpus 3-segmented and
stout, labial palpus 2-segmented. Antenna 1 1 -segmented;
scape cylindrical, reaching 0.9x distance to median ocel-
lus; pedicel globular; anellus small; flagellum l.l-1.2x
height of head; funicular segments densely setose and
reticulate; F2 0.4x as long as scape, F2 1.4-1.5x as long
as broad, following segments subequal in length, equal in
width; clava ovate, as long as preceding 3 segments.

Mesosoma robust, dorsum finely reticulate, lateral lobe
of mesoscutum and axilla weakly reticulate, scutellum
rugose-reticulate. Mesoscutum with midlobe subtriangu-
lar, anterior lateral comers sharp and protruding; notaulus
narrowly and deeply impressed crenulate. SSS crenulate
and extending to TSA as united medial furrow; frenal line
shallowly impressed. Scutellum as long as broad, widely
separated at base from TSA, both scutellum and frenal
area rounded in profile, frenal area coriaceous; axillula
not distinct from surface sculpture. Propodeal disc broad-
ly rounded, coriaceous with weak median carina; post-
spiracular and metepimeral sulci broad and irregularly
foveate; callus slightly swollen and glabrous, with promi-
nent callar nib; metepimeron glabrous. Upper
mesepimeron glabrous; transepimcral sulcus shallow
foveate; lower mesepimeron and mcsepisicrnum reticu-
late, mesepisternum glabrous anteriorly and ventrally;
femoral groove obscure. Prepectus triangular and finely
reticulate. Pronotum weakly reticulate, sometimes (see
variation) with sharp conical projection below spiracle
(prominent in dorsal view). Proepisternum coriaceous.

76

Coxae coriaceous basally and glabrous apically; femora
glabrate. Forewing 2.4-2.6x as long as mesothorax,
2.4-2. 5x as long as broad (Fig. 118); basal area and
speculum bare; costal cell with sparse medial band of
setae; disc moderately setose; stigmal vein narrow and
longer than broad, very slightly angled distally; postmar-
ginal vein indistinct, 2-3x as long as stigmal vein.

Metasoma with petiole 0.7-l.lx as long as hind coxa,
0.9-1.2X as long as propodeum; petiole rugose to reticu-
late, broad and narrowed at base. Mt^ l.lx as long as hind
femur, glabrous; Ms^ constriction sharp laterally and
smooth. Second valvula of ovipositor broad with several
prominent lateral teeth at apex, connected dorsally by
weak transverse ridges.

MALE

Length 1.7-2.3 mm. Colour as in female except flagellum
light to dark brown; scape black; tibiae dark yellowish
brown to black, hind tibia usually black. LOL l.l-1.5x
OOL. Eyes separated by 1.8-2. Ix their height. Flagellum
1.2-1.4X height of head; funicular segments densely
setose, setae semi-erect and giving flagellum distinct
fuzzy appearance; F2 1.6x as long as broad, following
segments quadrate and beadlike; clava with 2 incomplete-
ly fused segments and strong ventral notch. Mesosoma as
in female. Forewing small (Fig. 117), 2. Ox as long as
mesothorax, 1.9-2. Ix as long as broad; marginal vein
0.27-0.32X as long as forewing. Petiole 1.7-2.8x as long
as hind coxa, 1.7-2.3x as long as propodeum. Genitalia
typical for genus; paramere long with 2 apical setae;
aedeagus broadly rounded.

VARIATION

Little geographical variation exists among males and
females for the fine reticulation of the vertex, densely
setose antennal flagellum of males, or the presence of a
prominent nib on the callus. The conical projection just
anterior to or below the mesothoracic spiracle is weak in
some specimens although usually evident. In the 1 speci-
men from Vietnam the projection is weak but the anterior
lateral comers of the mesoscutum are prominent. In the
specimen from Iriomote Island, the prominence is barely
discernible and the midlobe is rounded, but it otherwise
fits the description.

DISTRIBUTION

Known from the Indo-Chinese subregion and ranging
from India to the southern Ryukyu islands of Japan (I,
Fig. 276).

MATERIAL EXAMINED

Japan: Ryukyu Islands: Iriomote Is.. Shira Lama.
7.xi.l963, H. Hasagawa ( 1 c5 , NIAS). Taiwan:
Praomonszu, 2 km S Keelung, 16.viii.l958, K. S. Lin
{\6. TARI); Nantou Hsien: Lushan, 1000 m,
27-3 l.v. 1960, K. S. Lin and L. Y. Chou (Id, TARI);
Wushe, 1150 m, 7-8. x. 1982, K. C. Chou (19, 36 S ,
TARI); Taichung Hsien: Chiapaotai. 750 m,
14-18. X. 1980, K. S. Lin and C. H. Wang (29 9, Id,
TARI); Kukuan, 730 m, 14-17. x. 1980. K. S. Lin and C.
H. Wang {56 6. TARI); Pingtung Hsien: Kenting Park.
22.iii.1980. K. S. Lin (19, TARI). Vietnam: Di Linh
(Djiring), 1200 m, 22-28. iv. 1960, L. W. Quate (Id,
BPBM).

Orasema va/g/as-group

This group consists of 2 species, O. valgius and O.
synempora, that are closely related based on the follow-
ing: propodeal disc glabrate laterally with a broad median
band of sculpture, mid coxa with a mid-ventral sulcus,
postmarginal vein elongate, first valvula with 3 to 4 later-
al teeth, and the petiole of female generally longer than in
other closely related groups of Orasema (1.2-1.7x as
long as hind coxa versus 0.9-1.2x in O. glabra and
0.8-1.5X in the assectator-gxo\x\)).

lum bare; disc pilose, forewing with marginal fringe; stig-
mal vein narrow and elongate, perpendicular to wing mar-
gin or angled distally; postmarginal vein well defined and
more than 0.4x as long as marginal vein. Petiole 0.9-1. 7x
as long as hind coxa in female, 2. 3-3. Ox as long as hind
coxa in male, the base with dorsal flange. Ovipositor sub-
apically expanded and subapical ridge prominent: first
valvula strongly narrowed distal to subapical ridge, with
3^ minute or large teeth along lateral line. Gonostylus
broad and setose, separated at base from second valvifer.

(JROUP DESCRIPTION

Face at least partially reticulate, often glabrate below
lower margin of eye. Antenna 1 1 -segmented and funicle
7-segmenled in both sexes. Mesosomal dorsum variously
sculptured; lower mesepimcron glabrous; callus rounded
and glabrate. with or without setae; inid coxa with mid-
ventral sulcus. I'orcwing hyaline: basal area and specu-

Orascma synempora .sp. nov.

Figs. 121. 123. 129

TYPE MATERIAL

Holotype, 9. "15.035S I45.09E/ 3 km NE Mt Webb. 1-3
Oct. I9S0 0/ J. C. Cardalc/ ox ethanol." "collected/ al

77

lighi." "AUST. NAT./ INS. COLL." "HOLOTYPR/
Orascma/ synempora Heraty." Deposited in ANIC.
Paratypcs: Australia: Queensland: Ayr, 30 S, 9.ix.l95().
E. F. Riekdd, ANIC); Ayr, 63S. IZ-ix-igL^S, E. F. Riek
(26 6, ANIC): Bowen. \5 W, 24.ix.1950, E. F. Riek
(2cJ<?. ANIC); Herberton, 12-23 km W. via Watsonville.
1.V.1967. D. H. Colless (2d 6 , ANIC); Hope Vale
Mission. 7 km N. 4.x. 1980. J. C. Cardale (Id, ANIC):
Maitland Downs, 3 km S, 15. iv. 1980. G. F. Hevel and
J. A. Fortin (Id, USNM); Mt Webb, 3 km NE,
1-30. X. 1980, J. C. Cardale, at light (3dd, ANIC);
Nomianton. 2-8. iii (3dd, AEI); Rounded Hill, 5 km W
by N. nr Hope Vale Mission, 7.x. 1980, J. C. Cardale
(2d d , ANIC): Ellis Beach, 9.3 km N, 30. iv, 1 .v,
8.V.1990. J. Heraty. on kangaroo grass. (49 9, 24dd,
JMH, ROM, TAMU); Ellis Beach. 9.7 km N. 18.iv.l987,
E. C. Dahms and G. Sames (Id, QMB).

DIAGNOSIS

Distinguished from O. valgius by the following: mesoso-
ma robust, lateral lobes of mesoscutum, axilla, and frenal
area glabrous (Fig. 129), mesepistemum with a narrow
glabrate band ventral and anterior to frenal line, and
wings with minute setae. Additionally, the flagellar seg-
ments are tightly adpressed (Fig. 123), and the callus is
glabrous and evenly swollen with no projection.

FEMALE

Length, 3.1-3.5 mm. Body dark glossy blackish green
with iridescent reflections; pedicel and flagellum black;
scape and legs yellowish brown, mid and hind femora
infuscate near base, hind femur dark brown with faint
metallic lustre in basal half on outer side. Wings hyaline,
venation brown.

Head subtriangular, l.l-1.2x as broad as long, 1.3x as
broad as mesosoma, LOL 0.7-0.9x OOL. Face broadly
rounded and reticulate, with face below torulus mostly
glabrate; scrobal depression narrow and rounded, finely
reticulate medially with narrow sublateral glabrous bands:
temple relatively broad and aciculate; occiput aciculate,
dorsal margin abrupt posterior to ocelli and rounded later-
ally, occipital carina absent. Eyes separated by 1.5-1.6x
their height. Malar space 0.7-0. 8x height of eye, malar
depression broad and vaguely impressed. Clypeal area
glabrate and swollen, epistomal sulcus weak, tentorial pit
deeply impressed, lateral margin of clypeal area shallow,
anteclypeus truncate. Labrum 4-digitate. Mandibles with
apical tooth reaching base of opposing mandible; palpi
relatively long and 3-segmented, second segment short in
both. Antenna 1 1 -segmented; scape stout and reaching
0.7x distance to median ocellus; pedicel subconical; anel-
lus large; flagellum ().9x height of head: funicular seg-
ments apprcssed, with short adpressed setae becoming
denser toward apex (Fig. 123). MPS numerous; F2 0.5x

as long as scape. F2 2. Ox as long as broad, following seg-
ments quadrate and equal in length; clava ovate, indistinct
from funicular segments, as long as preceding 2 segments.

Mesosoma robust and elongate (Fig. 129), 1.3x as long
as high, midlobe of mesoscutum and sublateral areas of
sculellum weakly reticulate; lateral lobe and axilla
swollen and glabrate. Notauli and SSS strongly and nar-
rowly impressed. Scutellum as long as broad, narrowly
separated at base from TSA; frenal line weakly impressed
and crenulate; frenal area glabrate and semicircular in
dorsal view; axillula colliculate, axillular sulcus weakly
impressed but distinct. Propodeal disc rounded, laterally
glabrous to very weakly punctate, medially with broad
reticulate furrow, furrow broader at base and weakly
rugulose: postspiracular furrow shallow and irregularly
foveate; callus rounded and glabrous; metepimeral sulcus
narrow and smooth, extending dorsally as narrow sculp-
tured furrow separating small anterior and large posterior
areas. Upper and lower mesepimeron glabrous,
transepimeral sulcus sharp; femoral groove reticulate with
distinct glabrous band anterior to femoral groove;
mesepistemum mostly glabrous, broad sternaular area
finely reticulate. Prepectus weakly reticulate to smooth.
Pronotum mostly glabrate. Proepistemum weakly collicu-
late along anterior and lateral margins, mostly glabrous.
Coxae and femora mostly glabrous, femora apically with
weak imbricate sculpture and dense minute setae, mid
coxa with longitudinal mid-ventral groove. Forewing
2.0x as long as mesothorax, 2.5x as long as broad; basal
area and speculum bare, costal cell narrow and with only
few scattered and minute setae; disc with dense, minute
setae; marginal vein 0.3x as long as forewing; stigmal
vein narrow and more than twice longer than broad,
slightly angled distally; postmarginal vein elongate. 0.4x
as long as marginal vein.

Metasoma with petiole 1.2-1.3x as long as hind coxa,
1.0-l.lx as long as propodeum; petiole cylindrical and
reticulate, weakly truncate basally and dorsal flange weak
(Fig. 121). Mt-, 1.2x as long as hind femur, glabrous; Ms,
with constriction sharp and smooth. First valvula of
ovipositor with 3 small teeth distal to crest: second valvu-
la broad with 7 strong lateral teeth, ridges strong dorsally
but not meeting along midline. Gonostylus broad, setose,
and separated from second valvifer.

MALE

Length, 2.5-3.0 mm. Colour darker and more bluish than
in female, rarely more reddish, scape dark brown and
sometimes with faint metallic reflections, femora dark
brown except at apex. Eyes separated by 1.7-1.9x their
height. Malar space 0.8-0.9x height of eye. Antenna 11-
segmentcd, flagellar segments closely adpressed; F2 1.5x
F3; clava indistinct from preceding segments. Mesosoma
less robust, sculpture pattern as in female; mesepistemum

78

as in female but band anterior to femoral groove may be
weakly reticulate. Fore wing 2.1x as long as mesosoma,
2.1-2.6X as long as broad. Petiole 2.1-2.6x as long as
hind coxa, 2.1-2.4x as long as propodeum; almost
smooth to reticulate dorsally. Mt, 0.8x as long as hind
femur; Ms^ with constriction sharp. Genitalia typical for
genus, paramere long, broad, and weakly sclerotized, with
2 apical setae, aedeagus broadly rounded.

BIOLOGY

I collected this species in a relatively small patch of
mixed kangaroo grass, Themeda triandra Forskal
(Poaceae), and Cassia mimisoides Linnaeus
(Leguminoceae), surrounded by an open Eucalyptus for-
est. Individuals were collected sporadically, and their host
plant could not be isolated. Females placed on bouquets
of either plant were unresponsive.

DISTRIBUTION

Australia (Queensland) (M, Fig. 276).

ETYMOLOGY

From Greek synemporos, meaning fellow traveller: in
honour of my wife, Laura, and my daughter, Joanne.

Orasema valgius (Walker)

Figs. 122, 124, 131, 192, 221, 238, 248, 262-263

Eucharis valgius Walker, 1839:11. New South Wales,

Australia [BMNH. examined].
Psilogasteroides valgius — Girault, 1913b:94.
Parapsilogaster valgius^GxrowXt, 1915:233.
Epimetagea valgius — Hedqvist, 1978:243.
Orasema valgius — Boucek, 1988:521.
Orasema pheidolophaga Girault, 1913b:96. Australia:

Victoria [SAMA, examined]. Synonymy by Boucek,

1988:521.

TYPE MATERIAL

Lectotype off. valgius (designated by Boucek, 1988), (5,
"LECTO-/ TYPE '" "Type." "N.S.W." "Sydney."
"Psilogaster/ Valgius/ Walker." "B.M. TYPE/ HYM./
5.617." "(5 Orasema/ valgius (Walker)." Flagellum and
hind legs missing, wings and legs in glue.

Lectotype of O. pheidolophaga (here designated), 9,
"Reared from pupae obtained/ in ncsl of Pheidole sp/
Geelong Victoria/ H. W. Davcy." "Orasema phe-/
dolophaga Gir./ 6 9 types." [GH] "I?]. 1286/ Orasema/
pheidolophaga Gir/ Victoria/ also slide." "S.A. Museum
specimen." "LECTOTYPE/ Orasema/ pheidolophaga
Gir./ Del. J. Hcraty '90." Six syntypes of both sexes,
mounted on card with headless minor of Pheidole (sec
Dahms, 1986). Slide: head, antennae, and wings.

'"Orasema phedolophaga [= pheidolophaga] Girault, S ,
9 types" [GH].

DIAGNOSIS

Distinguished from O. synempora by having the body
smaller and more slender, lateral lobe and axilla sculp-
tured (Fig. 238), frenal area carinate, mesepisternum
broadly reticulate (Fig. 221), and fore wing with longer
setae (typical of other species).

FEMALE

Length, 1.7-2.5 mm. Head, mesosoma, and petiole black-
ish-green, sometimes with greenish or iridescent reflec-
tions; gaster dark brown to black with faint greenish
reflections, antennal flagellum dark brown; scape, pedi-
cel, and femora dark brown with faint metallic lustre or
completely yellow; apex of femora and rest of legs yellow
to yellowish brown. Wings hyaline, venation brown.

Head subtriangular, 1.3x as broad as high, 1.4-1.5x as
broad as mesosoma; LOL 0.6-1. Ox OOL. Face relatively
flat, frons reticulate, face below toruli mostly glabrate;
scrobal depression shallow, weakly sculptured; temple
narrow and weakly sculptured; occiput aciculate, dorsal
margin broadly rounded, occipital carina absent. Eyes
separated by 1.6-2. Ox their height. Malar space 0.7-1. Ox
height of eye, malar depression absent. Clypeal area
glabrate, epistomal sulcus absent, tentorial pit deeply
impressed, lateral margin of clypeal area shallow, ante-
clypeus subtruncate; supraclypeal area slightly swollen.
Labrum 4-digitate. Mandible with apical tooth just over-
lapping base of opposing mandible; palpi 3-segmented,
second segment of maxillary palpus 3x as long as broad,
second segment of labial palpus short, as long as broad.
Antenna 11-segmented (Fig. 124); scape cylindrical,
almost reaching median ocellus; pedicel subconical: anel-
lus small; flagellum l.l-1.4x height of head; funicular
segments distinctly separated, densely setose along entire
flagellum, with numerous MPS; F2 0.3-0.5x as long as
scape, F2 1.4-1.7x as long as broad, following segments
slightly subequal in length, slightly broader than F2;
clava ovate, as long as preceding 2 segments.

Mesosoma 1.2x as long as high (Fig. 221); midlobe of
mesoscutum reticulate, scutellum rugose-reticulate with
verrucose surface; lateral lobe variously sculptured (never
glabrous); axilla weakly carinate and imbricate to reticu-
late. Mesoscutum with notauli deeply and broadly
impressed and crenulate. SSS broadly impressed and
crenulate. Scutellum 1.2x as long as broad, separated at
base from TSA by broad fovea (Fig. 238); frenal line
broad and irregularly foveatc, frenal area vertical and
hardly visible in dorsal view, vertically carinate or rarely
almost glabrate; axillula rugose, axillular sulcus obscured
by sculpture. Propodcal disc broadly rounded, glabrous
laterally with broad median bami c^f ruguloso sculpiure.

79

band bri)adcr dorsally: postspiracular furrow strong and
fovcato: callus rounded and glabralo with lew short setae
dorsally; metepimcral sulcus shallow, irregularly
impressed without dorsal extension. Mesepimeron
glabrous, transepimeral sulcus broad and irregularly
impressed; femoral groove obscure; mesepisternum retic-
ulate laterally, glabrous anteriorly and ventrally.
Prepectus weakly coriaceous to smooth. Pronotum mostly
smooth. Proepisternum glabrous. Coxae mostly smooth,
weakly coriaceous dorsally with weak mid-ventral sulcus;
femora glabrate basally and ventrally, femora imbricate
and minutely setose dorsally. Forewing 2.6x as long as
mesothorax, 2.2-2.5x as long as broad (Fig. 122); basal
area and speculum bare; costal cell sparsely setose and
relatively broad; disc with dense, relatively long setae;
marginal vein 0.27-0.29x as long as forewing; stigmal
vein narrow, more than 3x as long as broad and slightly
angled distally; postmarginal vein elongate. 0.6x as long
as marginal vein.

Metasoma with petiole 1.3-l.7x as long as hind coxa,
1.1-1.6X as long as propodeum; petiole cylindrical and
reticulate to rugose. Mt, equal to length of hind femur,
glabrate; Ms^ constriction broad and smooth (Fig. 262).
Ovipositor (Fig. 262); first valvula with lateral line of 2
large and 2 small teeth distal to subapical ridge, inner sur-
face with dense short setae (Fig. 263); second valvula
broad, with several strong lateral teeth, connected by
weak transverse ridges.

ra resulting in what could be called a dark and light
morph ol this species. However, a complete range of
scape colour has been found on specimens with darkened
femora at a single locality. Individuals with completely
yellow femora have not been collected with the darker
colour morphs. Males are always darker in coloration and
may have the scape and femora black. Both £. val^ius
and O. pheidolophui^u conform to the dark morph and
were described with darkened femora.

ADUmONAL .SPECIES

A similar, undescribed. species, which occurs in Australia
(Mt Glorious, sclerophyll forest, 16-20.ii.l961, L. and M.
Gressitt, BPBM), differs by having the face glabrous, lat-
eral lobe of mesoscutum glabrous and swollen, and the
midlobe, scutellum. and mesepisternum with fine reticu-
late sculpture. It shares the smooth propodeum and callus,
elongate petiole, and slender mesosoma that are typical
for O. vali>ii4s. The male and female are in poor condition
and are not here described.

BIOLOGY

Reared from Pheidolc (Girault, 1913b). The collection
records do not offer much information on habitat other
than that individuals were collected in long grass in open
forest, or in wet forest. The association with long grasses
could suggest a similar habitat preference with O. synem-
pura.

MALE

Length, 2.3-2.5 mm. Colour as in female but darker. Eyes
separated by 1.8-2.2x their height. Antenna 11 -segment-
ed; scape reaching median ocellus; flagellum 1.4-1.9x
height of head; F2 1.8x as long as broad. Mesosoma as in
female. Forewing 2.1-2.5x as long as broad; stigmal vein
stout, sharply angled distally. Petiole 2. l-2.6x as long as
hind coxa, 2.4-3.5x as long as propodeum; petiole cylin-
drical, reticulate to smooth ventrally and apically. Gaster
small. 0.8x as long as hind femur. Genitalia typical for
genus, paramere elongate with terminal seta, small medi-
an process, digitus with 5 marginal spines; aedeagus sub-
truncate.

VARIATION

There is little variation in shape of the mesosoma, but
considerable variation in sculpture. The midlobe of meso-
scutum and scutellum are almost always strongly and
evenly sculptured, but the lateral lobe varies from weakly
to strongly sculptured. Facial sculpture varies from
almost completely reticulate to nearly smooth on the face
and vertex, and the petiole may be rugose to reticulate.
The coloration of the scape ranges from dark brown with
a weak metallic lustre to a prominent yellow. This is
sometimes correlated with similar coloration of the femo-

DISTRIBUTION

Known only from eastern Australia (V. Fig. 276). A sin-
gle female of O. valgius was recorded from Kuranda,
north Queensland. The northern record suggests that this
species is sympatric with O. synempora. O. valgius
exhibits a similar pattern of distribution to Psilocharis
theocles and Orasemorpha eribotes, but has not been
recorded from western Australia or Tasmania.

MATERIAL EXAMINED

Australia; A.C.T.: Bendora; Black Mtn. 300 m;
Canberra; December to February (39 9, 1 <5 . AEI,
ANIC); New South Wales: Baleman's Bay, 33-39 km W;
Bondi S.F., via Bombala; Botany Bay; Cabramatta;
Canley Vale; Casula; Corang Riv. via Nerriga; Dorrigo
N. P.. 1000 m; Mt Dromedary. 330 m. light trap; Mt
Elliot. NE Gosford; Gibraltar Range, via Glen Innes, 900
m; Palm Creek Royal N. P.; Pilliga Scrub, via
Coonabarabran; Springwood. Blue Mts. 350 m; Tooloom
Plateau, via Urbanville. 600-700 m; Wiangaree N. P.,
Brindle Ck. 1000 m; October to March (33 9 9,16 S,
ANIC, BMNH. CNC. MCZ. QMB. UQIC. USNM);
Queensland: Adelaide; Bald Mtn area, 380^50 m via
Emu Vale; Fleurian Peninsula. Deep Creek Conservation
Park; Kangaroo Is., Rocky Riv., Flinders Chase N. P.; Mt

80

Byron area, D'Aguilar Range, long grass, open forest;
Cooloola, nr Freshwater Lake, open forest; Stanthorpe;
Kuranda; Stanthorpe, 700 m; nr Wilson's Peak, via
Teviot Gap, 700-800 m; November to January, and
March to April (179 9, 19c? c?, AEI, ANIC. BMNH,
BPBM, JMH, ROM, TAMU, UQIC); Victoria: Cann

Valley Highway, 7 km SW NSW border; Cann Riv.,
80-120 m, sweeping low vegetation, wet sclerophyll for-
est; Club Terrace, 120 m; Grampians, Rose's Gap;
Growler Ck, Lind N. P.; Mt Beauty; February to March,
October, and December (119 9, 18(5(5, ANIC. TAMU.
UQIC).

Revision of Psilocharitini (Eucharitinae)
Psilocharis gen. no v.

Type Species: Eucharis theocles Walker, 1839:1 1-12; by
present designation.

Psilocharis is included within the Eucharitinae based on:
presence of an occipital carina, an acicular ovipositor, and
absence of a constriction on the first gastral stemite. This
genus is distinguished from other Eucharitinae by: anellus
usually present, clypeal margin transverse with row of
fine elongate setae along margin of anteclypeus (Figs.
193-198), malar space with complete longitudinal depres-
sion (Fig. 223), mesepimeron mostly polished with
broadly impressed femoral groove, stigmal vein slightly
broader than marginal vein, base of petiole truncate later-
ally (Figs. 144-145), ovipositor acicular, and ovipositor
sheath narrow and cylindrical (Fig. 264). In most species,
the apex of the hypopygium has a band of elongate hairs,
a character state shared with Gollumiella and Anorasema
of the Eucharitinae. Psilocharis, along with Neoloshanus,
is placed as the sister group of the rest of the Eucharitinae
based on the following combination of plesiomorphic
character states: prepectus not fused to pronotum, anellus
usually present, and mesothoracic spiracle not enclosed
dorsally. The immature stages and ant host are unknown.

GENERIC DESCRIPTION

Head subtriangular, 1.3-1.5x as broad as mesosoma;
median ocellus anterior to lateral ocelli, lateral ocellus
separated from occipital margin by at least its own radius
(Fig. 136). Face, vertex, and gena polished; scrobal
depression shallow and poorly defined laterally, usually
with 2 parallel smooth channels extending part way to
median ocellus (Figs. 194-19.5. 197); occiput finely acic-
ulate, occipital carina present (Figs. 136, 223); ocellar-
ocular groove absent. Malar depression broadly and
deeply impressed along entire length; hypostoma large,
separated from gena by prominent hypostomal carina and
extending mesally over base of mandible. Clypeus slight-
ly broader than long, subcqual to supraclypcal area; apical
margin subtruncate or slightly rounded, epistomal sulcus
weak or absent; anteclypeus distinct and with row of fine

elongate setae along dorsal margin that extends ventrally
over labrum (difficult to observe and sometimes visible in
sublateral view; Fig. 198). Labrum 4-digitate, setae long
and spatulate. Mandibles falcate and 2/3 dentate; maxilla
and labium elongate, maxillary and usually labial palpi 2-
segmented. Antenna 8- to 1 1 -segmented; scape narrow
and elongate, scape of female without pores, scape of
male with minute pores ventrally (visible only in slide
preparations); pedicel short, at most 1.5x as long as
broad; anellus present or absent; funicle 5-to 7-segment-
ed, segments cylindrical or rarely moniliform, with basal
secondary segmentation and scattered multiporous plate
sensilla in both sexes; basal funicular segments less than
2.2x as long as broad, terminal 3 or 4 segments fused into
distinct clava.

Mesosoma with dorsum usually strongly sculptured,
sometimes weakly reticulate to smooth. Mesoscutum with
each notaulus well defined along entire length. TSA com-
plete. SSS broadly curved and angled to midline at TSA.
Scutellum with frenal area separated by foveate or crenu-
late groove; axillular sulcus distinct and foveate.
Metanotum extended laterally as smooth tlange overlap-
ping base of propodeum and partly covering propodeal
spiracle; spiracle close to dorsal margin of propodeum.
Propodeal disc rounded and glabrous laterally (Fig. 249)
with broad medial band of rugose-areolate sculpture,
sculpture broadest dorsally and reaching postspiracular
furrow; callus polished and not strongly swollen, with
few to several long hairs dorsally. and with or without
callar nib; postspiracular furrow and metepimeral sulcus
marked with narrow foveate groove; ventral margin of
propodeum above hind coxa even and strongly ridged,
without lateral processes. Mesopleuron mostly smooth;
mesepimeron slightly swollen, transepimeral sulcus
marked by narrow foveate groove; femoral groove broad-
ly impressed or foveate; sternaular area of mesepisternum
with wedge-shaped foveate groove (SA. Fig. 224).
Prepectus reaching logula as triangular, rovcatc lobe dor-
sail) . strongly narrowed and smooth \entrall) . Coxae and
femora smooth and shining; hind tarsus O.Sx shorter than

81

tibia: fore aiui iiiiil lihiac with I spur, liimi tibia with I or
2 spurs.

Wini>. Veins of fore and hind wing well defined.
Forewing 2.0-2. 5x as long as broad, broadly rounded api-
cally: basal area usually bare; speculum present (bare);
costal cell pilose or bare; disc densely pilose and with dis-
tinct marginal fringe; submarginal vein with row of dorsal
setae; marginal vein ().26-().33x length of forewing and
densely pilose as rest of wing; stigmal vein elongate and
broader than marginal vein; postmarginal vein less than
l.5x length of .stigmal vein (Figs. 146-148).

Metasoma with petiole more than 1 .4x length of hind
coxa; petiole sublriangular in cross-section and fused ven-
trally, lateral margin of petiole base parallel or expanded
and abruptly narrowed to articulation (Figs. 144-145).
base with or without dorsal flange. Gastral terga polished,
gaster of female as long as head and mesosoma, slightly
smaller in male; Mt^ of female less than 0.6x length of
gaster; Ms, smooth, without medial transverse constric-
tion. MSjj of male narrow, rounded apically, and densely
setose. Cercus with few long setae and 1 seta twice length
of others. Hypopygium with row of elongate hairs along
apical margin (Figs. 149, 264), rarely with 2 minute setae
on either side of midline at apex. Ovipositor sheath nar-
row and elongate, sometimes exceeding cercus by more
than its width. Gonostylus indistinctly separated basally,
and setose. Ovipositor acicular; first valvula smooth, sec-
ond valvula with weak dorsal ridges (Fig. 265). Genitalia
of male with paramere well developed and bearing 2 stout
setae, digitus disclike with 3 to 4 stout marginal spines;
aedeagus narrow and subacute (Fig. 151).

ETYMOLOGY

Combination of Greek psilos, meaning smooth or bare,
and eucharis, meaning pleasing; feminine gender.

PHYLOGENETIC RELATIONSHIPS

Monophyly of Psilocharis is supported by several charac-
ter states, including the following: anteclypeus distinct
with marginal setae, petiole with lateral margins parallel
and abruptly narrowed at base (39), scrobal depression
with short parallel channels and apex of each channel
forming smooth circular impression (9), labial palpi usu-
ally 2-,segmented, and femoral groove broadly impressed
(29). A distinct anteclypeus (without marginal setae) and
abrupt base of the petiole are also found in some species
of Orasema, and O. glabra also has scrobal channels.
These character states are treated here as convergent
developments within Orasema.

Most species of Psilocharis possess a row of long
hairs on the hypopygium (45). A similar marginal row of
hairs is found also in Gollumiclla and Anorascma. but
otherwise it would appear to be a unique character state
of Eucharitidae (or Chalcidoidea that I know oO- It is pos-

sible that the row of hairs was present in the groundplan
of Hucharitinae, with multiple losses or modifications
postulated as derived features. I have interpreted this
character state as an independent derivation within
Psilocharis.

Psilocharis acnii>ma is the sister group of the remain-
ing species based on the following: mesoscutum com-
pletely imbricate (apomorphic), proepisternum glabrous
(plesiomorphic), and hypopygium with only 2 small lon-
gitudinally aligned hairs (plesiomorphic). A marginal row
of hairs on the hypopygium is a derived state shared by
the remaining species (unknown for P. monilicera),
which may have been secondarily lost in P. pacifica (pre-
sent in its sister group, P. dahmsi). A completely and
strongly sculptured dorsum, as in Neoloshanus, is ple-
siomorphic. Therefore, a polished lateral lobe and axilla
is a derived state for P. dahmsi, P. joanneae, P. pacifica,
and P. theocles, and the polished axilla and scutellum in
P. aeiiigma is independently derived. A single specimen
from Argentina, which is closest to P. aenigma. has an
evenly sculptured scutellum. The former group, which
includes P. theocles, is further supported by a glabrous
frenum, but again this is homoplastic. A polished scutel-
lum, smooth scrobal depression, and dorsal flange at the
base of the petiole are shared by P. pacifica and P. dahm-
si. A lack of channels from the scrobal depression and
reduction of the callar nib may support a relationship
between P. joanneae and P. pacifica + P. dahmsi, but
both states are found to some degree in P. theocles. There
are no derived states to support a relationship between P.
joanneae and P. theocles.

The species P. afra, P. hypena, P. monilicera. and P.
pentella have prominent channels in the scrobal depres-
sion (weak in P. pentella) with a distinct impression at the
apex of each channel, a prominent callar nib, and a stri-
gate temple. Any of these states could be plesiomorphic
within Psilocharis. The species P. afra and P. hypena are
almost identical, but otherwise have no supporting apo-
morphies. Psilocharis monilicera and P. pentella both
have the basal area of the forewing pilose, the ventral
margin of scape in both sexes carinate, and the eyes
sparsely setose.

Tentatively, relationships among species of
Psilocharis may be hypothesized as {aenigma -f- {(monil-
icera + pentella) + hypena + qfra) -\- (theocles -^joanneae
+ (jjacifica -)- dahmsi))) (Fig. 277).

BI()LO(JY

Plant and ant hosts are unknown. 1 have collected P. theo-
cles in dry open Eucalyptus forests near Brisbane and
Perth, Australia, and P. hypena in rainforest near Kuala
Lumpur, Malaysia. Adults were scattered on plants along
paths, but no ant or plant hosts could be isolated.

82

DISTRIBUTION

Psilocharis is distributed throughout the southern
Ethiopian, Malagasy, and Indo-Pacific regions (Fig. 277).
Psilocharis pacifica is 1 of the few species of Oraseminae
or Eucharitinae found in the Polynesian subregion, and
the only species (other than some Chalcura and
Schizaspidia) known to occur as far east as Fiji.
Psilocharis theocles has a similar distribution to
Orasemorpha erihotes and Orasema valgius along the
eastern and southwestern coast of Australia. A single
specimen (in CAS) of an undescribed species known
from Argentina is morphologically very similar to P.
aenigma. As with the single specimen of Neoloshanus
known from Uruguay, this range extension to the
Neotropical region needs to be verified by at least 1 other
specimen.

Based on the phylogenetic hypotheses discussed
above, it would appear that the Oriental distribution of P.
hypena (Indo-Chinese + Malayan + Philippine subre-
gions) and P. pentella (Malayan) is derived from the

Ethiopian region, and not from the Australian region. The
4 species found in the Australian, Papuan, and Polynesian
(New Caledonia and Fiji) subregions {P. dahmsi, P. joan-
neae, P. pacifica, and P. theocles) appear to form a
monophyletic group that may share a common ancestor
with the Ethiopian species. The morphological differ-
ences between the Oriental and African species and the
Australasian species are distinctive, and suggest an early
vicariant event between Africa and Australia similar to
that found in the Oraseminae. The possible existence of a
species in the Neotropical region, which is most closely
related to the Malagasy species, would add further sup-
port to a hypothesis of an older widespread Gondwanan
distribution. Psilocharis hypena (Oriental) and P. afra
(Ethiopian) exhibit few morphological differences, which
could suggest a more recent faunal exchange between the
Ethiopian and Oriental regions, rather than vicariance
involving a shift of the Indian subcontinent from Africa
during the Palaeogene.

Key to Species of Psilocharis

1 Mesoscutum with midlobe rugose-areolate, later-
al lobe glabrous or strongly sculptured but not
umbilicate: hypopygium with transverse band of
8 to 10 long hairs along apical margin (unknown
for 2 species) 2

— Mesoscutum completely imbricate; hypopygium
with 2 minute hairs on each side of midline at
apex, hairs aligned longitudinally; Madagascar...
P. aenigma sp. nov., p. 84

2(1) Inner ventral margin of scape carinate and scape
cylindrical or strongly expanded, not reaching
median ocellus; female without anellus; basal
area of forewing setose; dorsum of mesosoma
weakly rugose; South Africa 3

— Ventral margin of scape evenly rounded; may be
slightly expanded apically but then scape long
and reaching median ocellus; female with anel-
lus; basal area of forewing bare or at most with
few sparse setae (Figs. 146-148); dorsum of
mesosoma and frenal area either strongly sculp-
tured or glabrous 4

3 (2) Antennal funicle with 6 or 7 segments; ante-
clypcus narrow and straight: frenum glabrous;
southern Africa P. monilicera sp. nov.. p. 8.'>

— Funicle with 5 segments; anteclypcus broad and

rounded; frenum vertically carinate; Sumatra

P. pentella sp. nov.. p. 85

4 (2) Lateral lobe, axilla, and frenum smooth and pol-

ished (Fig. 137); temple smooth or at most very
weakly aciculate 5

— Lateral lobe and axilla completely rugose or car-
inate, frenum vertically carinate (smooth in some
hypena; Figs. 239-240); temple finely strigate
dorsally 8

5 (4) Scutellum completely smooth and polished (Fig.

137); mesoscutum with midlobe scabrous 6

— Scutellum rugose-areolate, at least in basal half;
mesoscutum with midlobe transversely carinate
or rugose-areolate 7

6 (5) Hind tibia and scape yellowish brown; Fiji. New

Hebrides P. pacifica sp. nov., p. 86

— Hind tibia and scape black with blue reflections;
New Guinea P. dahmsi sp. nov.. p. 87

7 (5) Head subcircular in frontal view (Fig. 193); eye

large, malar space 0.4-().5x height of eye; anten-
nal flagellum of females 0.6-0.7x height of head
(Fig. 138). ().8x height of head in males;
Australia P.joanneae sp. nov.. p. 88

83

8(4)

Head siibtriangular in frontal view (Fig. 194);
eye smaller (as in other species), malar space
().6-0.9x height of eye; antennal llagellum of
females 0.8-1. Ox height of head (Fig. 139),
1.0-I.2X height of head in males (Fig. 141);
Australia P. theocles (Walker), p. 89

Lower face below eye strongly narrowed, poste-
rior margin of gena not or only slightly visible in
frontal view (Fig. 195). head width at lower mar-
gin of eye, including posterior genal margin.

1.8-2.4X vertical distance between eye margin

and apex of clypeus; Africa

P. afra sp. nov.. p. 90

Lower face broad, posterior margin of gena
rounded and distinct in trontal view (Figs.
196-197). head width at lower margin of eye
2.5-2.9X vertical distance between eye margin

and apex of clypeus; Oriental region

P. hypena sp. nov., p. 91

Psilocharis aenigma sp. nov.

Fig. 144

TYPE MATERIAL

Holotype, 9 , "Ambohitsitondrona/ XL55 (Vadon)."
"Madagascar/ Ambohitsitondrona/ XL 1955 (Vadon)."
"sp. (Oras.).'" "HOLOTYPE/ Psilocharis/ aenigma
Heraty." Deposited in BMNH.

DIAGNOSIS

Differs from all other species as follows: mesoscutum
completely imbricate (versus rugose or carinate), hypopy-
gium with 2 pairs of sublateral setae, proepisternum
glabrous, and gonostylus several times longer than broad.

FEMALE

Length, 2.4 mm. Body, including coxae, black; antenna,
femora, and rest of legs yellowish brown, pedicel and fla-
gellum slightly darker. Wings hyaline, venation clear pale
brown.

Head subtriangular; occiput shallow emarginate; LOL
1.2x OOL. Face broadly rounded and polished with
sparse, decumbent setae; scrobal depression not reaching
median ocellus, with parallel channels reaching 0.6x dis-
tance to median ocellus, median groove absent; occipital
carina weakly developed, extending just past lateral ocel-
lus; temple strigate. Eyes separated by 1.2x their height.
Malar space 0.5x height of eye. Clypeus sparsely setose,
epistomal sulcus absent. Antenna 1 1 -segmented; scape
reaching 0.8x distance to median ocellus, cylindrical;
pedicel 2.2x as long as broad; anellus stout; flagellum
0.9x height of head; funicle 7-scgmcnted; F2 2. Ox as long
as broad, slightly shorter than pedicel, following seg-
ments subequal in length and slightly broader apically;
clava ovate, slightly longer than preceding 2 segments.

Mesosonni with mesoscutum entirely imbricate, axilla
and scutellum smooth and polished. SSS foveate with
strong transverse carinae. Scutellum l.3x as long as
broad, narrowly separated from TSA basally, apex sub-
truncate; frenal line narrow and weakly foveate. frenal

area glabrous, broadly crescent-shaped in dorsal view;
axillula glabrous, axillular sulcus foveate. Callus with few
elongate hairs dorsally. callar nib absent. Femoral groove
broad and shallowly impressed, finely reticulate; ster-
naular area foveate. Proepisternum glabrous. Coxae
glabrous, imbricate basally; femora polished with sparse
short setae; hind tibia slender with 1 long spur. Forewing
2.4x as long as broad; basal area bare; speculum moder-
ately sized with regular margins; stigmal vein slightly
longer than broad; postmarginal vein 2.4x as long as stig-
mal vein.

Metasoma with petiole l.5x length of hind coxa, 1.5x
length of propodeum; petiole strigate-verrucose dorsally
and carinate laterally; base parallel-sided, without distinct
lateral flange (Fig. 144), dorsal flange absent. Caster typi-
cal. Hypopygium with 2 long hairs on each side of mid-
line, first hair directly anterior to second. Ovipositor acic-
ular. Gonostylus narrow and elongate. 6.3x as long as
broad (other species 3-4x as long as broad).

MALE

Unknown.

DISTRIBUTION

Madagascar (E. Fig. 277).

ETYMOLOCJY

From Latin aenigma, meaning mystery; referring both to
the set of character states and to distribution.

ADDITIONAL SPECIES

A single undescribed female, similar to P. aenigma, is
known from Cordoba, Argentina (2 km S Dean Funes,
8.ii.l951, E. Ross, CAS). It is similar in sculpture of the
mesoscutum and in several other states to P. aenigma, but
is a distinctly different species. The hypopygium is
immersed in glue and the setation of the hypopygium
could not be observed. This would be the only
Neotropical record of this genus, but needs to be verified
by additional collections.

84

Psilocharis monilicera sp. nov.

Fig. 134

TYPE MATERIAL

Holotype, 6 , "S. Afr. Grahamstown/ 5-7.1.86 W. Mason/
500 m (MT)." "HOLOTYPE/ Psilocharis/ monilicera
Heraty." Deposited in CNC.

Paratypes: South Africa: E. Transvaal: [Pondoland] Port
St. John, 5-30.iv.l923 (IcJ), 16-28.iv.l924 (19), R. E.
Turner (both BMNH).

DIAGNOSIS

Recognized by the following: scape only of male
enlarged, both sexes with inner ventral margin of scape
carinate. funicular segments of male moniliform (Fig.
134), basal area of the forewing sparsely setose, scutel-
lum lightly rugulose, and lateral lobe of mesoscutum and
axilla weakly sculptured.

MALE

Length, 1.2-1.6 mm. Head, mesosoma, and petiole dark
blue or bluish green; coxae, femora, and gaster dark
brown; antenna, apex of hind femora, and rest of legs yel-
low. Wings hyaline, venation clear.

Head subcircular; occiput broadly emarginate; LOL
l.lx OOL. Face broadly rounded, polished with sparse
decumbent setae; scrobal depression with parallel chan-
nels reaching just over half distance to median ocellus,
apex of each channel with distinct semicircular depres-
sion, median groove absent; occipital carina extending to
lateral ocellus; temple broad and weakly aciculate. Eye
with minute setae, eyes separated by 1.6-1.8x their
height. Malar space 0.7-0.9x height of eye. Clypeus
glabrate, tentorial pit deeply impressed; anteclypeus very
narrow and straight. Antenna 1 1 -segmented; scape reach-
ing 0.8x distance to median ocellus, enlarged along entire
length and inner ventral margin carinate (Fig. 134), apical
half broadly emarginate ventrally; pedicel as long as
broad; anellus present; flagellum l.l-l.2x height of head;
funicle 7-segmented; F2 cylindrical, 1.5-l.6x as long as
broad, F3-F8 moniliform and as long as broad; clava
ovate, shorter than preceding 2 segments.

Mesosoma with dorsum lightly rugulose; lateral lobe
of mesoscutum and axilla completely sculptured but
sculpture shallow and irregular. SSS broadly impressed
and weakly crcnulatc. Scutcllum 1.6x as long as broad,
subtruncate at apex; frcnal line shallow and foveate. fre-
nal area glabrous, crescent-shaped from above; axillula
glabrate. axillular sulcus narrow. Callus with several
elongate hairs and prominent callar nib. Femoral groove
broad, irregularly reticulate to rugulose in anterior half;
sternaular area narrow and shallowly foveate.
Procpistcrnum weakly carinate. Coxae glabrate; hind
femora weakly imbricate to smooth, with sparse setae;

hind tibia stout with moderately dense fine setae, with 2
spurs, the outer spur small. Forewing 2.0-2.2x as long as
wide; basal area evenly pilose; speculum bare but mar-
gins poorly defined; stigmal vein twice as long as broad;
postmarginal vein slightly longer than stigmal vein.

Metasoma with petiole l.5-1.7x as long as hind coxa,
1.5-1.8X length of propodeum; petiole subtriangular in
cross-section, ribbed dorsally, glabrate to aciculate sublat-
erally, with weak ventral keel; base parallel-sided, with-
out distinct lateral flange, dorsal flange absent. Mt, as
long as petiole. Genitalia typical for genus.

FEMALE

Length, 1.5 mm. Colour basically dark brown with dor-
sum bluish green. Occipital carina weak (difficult to dis-
cern from occipital strigae); temple glabrous. Tentorial pit
not deeply impressed. Antenna 10-segmented; scape
reaching median ocellus, cylindrical and with short carina
on inner ventral margin (surrounding depression under
pedicel); anellus absent; flagellum broadened apically,
F4— F7 enlarged. Mesoscutum with midlobe and anterior
region of scutellum weak rugose-areolate; lateral lobe
smooth; axilla smooth with weak carinae posteriorly.
Forewing with basal area bare and with few scattered
setae; gaster typical; ovipositor, sheath, and hypopygium
hidden.

DISTRIBUTION

Known only from 2 localities in South Africa (Cape
Province; M, Fig. 277).

ETYMOLOGY

Combination of Latin nionile. meaning necklace or bead-
like, and ceros, meaning horned; referring to the rounded
antennal flagellomeres.

Psilocharis pentella sp. nov.

TYPE MATERIAL

Holotype. 9, 'iNDONESIA: Sumatra. Aceh/ Gunung
Lcuser Nat. Pk./ Ketambc Res. Sta. Jan. 1990/ per DC
Darling. IIS 900003." "Malaise trap, primary/ rainforest.
400 m./ Mature forest. Terrace 4/ Light gap. 3°4rN,
97°39'E." "HOLOTYPE/ Psilochari.s/ pentella Heraty."
Deposited in MZB.

Paratypes: Indonesia: Sumatra: Aceh: same data as holo-
type (19, ROM); same locality and collector, 350 m.
ii.I990. young forest. Terrace 3. closed canopy. IIS
90001 1 MT (I 9, MZB); same locality and collector. 350
m. primary rainforest I-30.X.I989, D. C. Darling, 890006
MT(l9.ROM).

85

ni\(;N()sis

Recogni/cd b\ ihc lollowing: inner venlral margin ot
female scape expanded and carinate in the apical third,
anellus absent, funicle with 5 segments, basal area of
forewing pilose, and scutcllum rugulosc-areolate.

Hypopygium with 10 to 11 long hairs along apical mar-
gin. Ovipositor acicular. Gonostylus narrow and elongate.

MALE

Unknown.

FEMALE

Length, 2.0-2.9 mm. Head, mesosoma, and petiole black,
mesosoma with faint bluish reflections; coxa and gaster
dark brown; femora mostly light brown, apex of femora,
rest of legs, and .scape yellowish brown; antennal pedicel
and flagellum brown, basal segments lighter. Wings hya-
line, venation light brown.

Head subcircular; occiput broadly emarginate; LOL
0.9-1. Ox OOL. Face broadly rounded, polished with
dense short adpressed setae; scrobal depression with par-
allel channels shallow and reaching 0.7x distance to
median ocellus, apex of each channel with distinct semi-
circular depression, median groove absent; occipital cari-
na extending beyond lateral ocellus; temple broad and
weakly aciculate. Eye with minute setae, eyes separated
by 1.6-1.7X their height. Malar space 0.7x height of eye.
Clypeus glabrate, lateral margin and tentorial pit hardly
impressed; anteclypeus broad and rounded ventrally.
Antenna 9-segmented; scape reaching median ocellus,
enlarged ventrally and apical ventral margin carinate, api-
cal half broadly emarginate ventrally; pedicel 1.4x as
long as broad; anellus absent; flagellum 0.9x height of
head; funicle 5-segmented; F2 expanded subapically,
1.8-2.2X as long as broad, F3-F6 subequal in length;
clava oblong, nearly as long as preceding 3 segments.

Mesosoma with midlobe of mesoscutum and scutellum
rugulose-areolate; lateral lobe irregularly transverse-stri-
gate, axilla longitudinally strigate. SSS broadly impressed
and crenulate. Scutellum 1.5x as long as broad, subtrun-
cate at apex; frenal line shallow and crenulate, frenal area
with broadly spaced vertical carinae, abrupt and hardly
visible dorsally; axillula longitudinally carinate, axillular
sulcus narrow. Callus with several elongate hairs and
prominent callar nib. Femoral groove broad, irregularly
rugose-areolate in anterior half; sternaular area narrow
and shallowly foveate. Proepisternum and coxae
glabrous. Hind femur smooth, with sparse setae; hind
tibia slender with moderately dense fine setae, with 2
spurs, the outer spur small. Forewing 2.3-2.4x as long as
wide; basal area evenly pilose; speculum bare but mar-
gins poorly defined; stigmal vein 2-3x as long as broad,
strongly angled toward apex; postmarginal vein 0.6-1.6x
as long as stigmal vein.

Metasoma with petiole 1.8-2. Ox as long as hind coxa,
1.8-2. Ox length of propodeum; petiole subtriangular in
cross section, ribbed dorsally and laterally, with weak
ventral keel; base parallel-sided, with distinct lateral
flange and no dorsal flange. Gaster typical for genus.

DISTRIBUTION

From Sumatra in Malayan subregion (N, Fig. 277).

ETYMOLOGY

From Latin penta; referring to number of funicular seg-
ments.

Psilocharis pacifica sp. nov.

Figs. 133, 137

TYPE MATERIAL

Holotype, 9 , "FIJI: Vanua Levu I:/ Mt Dalaikoro,/ sum-
mit area,/ 700-790 m, lO.X. 1979.'" "#233.'" "S. N. Lai, 0.
A. & S. L./ Samuelson, Colls./ BISHOP Museum/ Ace.
#1979.387." "HOLOTYPE/ Psilocharis pacifica Heraty."
Deposited in BPBM.

Paratypes: New Hebrides: Efate Is. (NW), Maat (Mat,
Ambryn Vill.), 3 m, 15.viii.l957, J. L. Gressitt (1$,
BPBM).

DIAGNOSIS

Differences from P. dahmsi: scape completely yellowish
brown, antennal flagellum longer (1.2x head height ver-
sus 0.9x; Fig. 133), hind tibia with 1 spur, and petiole of
female 1.6x length of propodeum (versus 2.1-2.2x).

FEMALE

Length, 1.6 mm. Body dark metallic green or reddish;
antenna including scape yellowish brown; femora black
with strong greenish reflections; scape, pedicel, tibiae,
and tarsi dark yellowish brown. Wings hyaline, venation
clear yellowish brown.

Head subtriangular, eye large, and face elongate (Fig.
133); occiput broadly emarginate; ocelli small, LOL l.Ox
OOL. Face relatively flat and polished with sparse,
decumbent setae; scrobal depression reaching median
ocellus, otherwise obscured and probably as in P. dahmsi
(smooth and without channels); occipital carina weakly
developed, extending to dorsal eye margin; temple nar-
row and glabrate. Eyes separated by 1.3x their height.
Malar space 0.4-0.5x height of eye. Clypeus sparsely
setose, cpistomal sulcus weakly impressed. Antenna 11-
segmcnted (Fig. 133); scape reaching 0.8x distance to
median ocellus, cylindrical; pedicel 1.2x as long as broad;
anellus short; flagellum l.2x height of head; funicle 7-
.segmented; F2 2. Ox as long as broad, slightly longer than
pedicel, following segments subequal in length and

86

slightly broader apically; clava ovate, slightly shorter than
2 preceding segments.

Mesosoma with midlobe of mesoscutum scabrous; lat-
eral lobe and axilla smooth and polished with sparse
setae; scutellum glabrous. SSS deeply foveate with wide-
ly spaced, prominent carinae. Scutellum 1.4x as long as
broad, base well separated from TSA by broad depres-
sion, apex subtruncate; frenal line narrow and weakly
foveate, frenal area glabrous, broadly semicircular in dor-
sal view; axillula glabrate, axillular sulcus broadly
foveate. Callus slightly swollen and glabrous with few
small dorsal setae, callar nib absent. Femoral groove
broadly and shallowly impressed, glabrous; sternaular
area broadly foveate. Proepisternum mostly glabrous,
weakly carinate in anterior half. Coxae glabrate; femora
polished with sparse short setae; hind tibia slender, slight-
ly expanded apically, with 1 long spur. Forewing distort-
ed and width not measurable but appears typical for
genus; basal area bare; speculum small with irregular
margins; stigmal vein 1.5x as long as broad; postmarginal
vein slightly longer than stigmal vein.

Metasoma with petiole 1.7x length of hind coxa, 1.5x
length of propodeum; petiole broad and subtriangular in
cross-section, weakly aciculate with prominent carina on
lateral margin and prominent dorsal flange basally. Caster
typical for genus. Hypopygium obscured, but 1 long api-
cal hair issuing from between folded tergites (both HT
and PT). Ovipositor acicular. Gonostylus narrow and
elongate.

MALE

Unknown.

VARIATION

The mesosoma of the paratype has a more distinct reddish
coloration and the occipital carina does not extend lateral-
ly to the dorsal eye margin.

DISTRIBUTION

P. pacifica is the only species in this genus with an ocean-
ic (Polynesian) distribution, and the only known species
within the Orascminae and basal Eucharitinae to reach the
Fiji Islands (P, Fig. 277).

f,tymol()(;y

From Pacific Ocean; referring to the oceanic distribution
of the species.

Psilocharis dahmsi sp. nov.

typp: matp:riai.

Holotypc. 9. "NEW GUINEA: NETH./ Swart Val.: W./
ridge 1800-2000 m./ Nov. 19, 1958." "PICRA-

NOPTERIS." "J. L. Gressitt/ Collector." "HOLOTYPE/
Psilocharis dahmsi Heraty." Right antenna missing.
Deposited in BPBM.

Paratypes: Papua New Guinea: same data as holotype
(29 9, 1(5, BPBM); Swart Val., Karibaka, 1500 m,
1 l.xi.l958, Picranoft erus [!] (1 c5, BPBM).

DIAGNOSIS

Differences from P. pacifica: scape and tibiae dark brown
with metallic reflections, antennal flagellum shorter (0.9x
height of head versus 1.2x); hind tibia with 2 spurs, and
petiole of female 2.1-2.2x (versus 1.6x) length of
propodeum.

FEMALE

Length, 2.0-2.2 mm. Head and gaster black, mesosoma
with strong bluish reflections; antenna dark brown to
black, scape and tibiae dark brown with metallic reflec-
tions; femora black with strong greenish reflections;
scape, pedicel, and tarsi dark yellowish brown. Wings
hyaline, venation brown.

Head subtriangular, eye large, and face elongate;
occiput narrowly emarginate; ocelli small, LOL l.lx
OOL. Face relatively flat and polished with sparse,
decumbent setae; scrobal depression not reaching median
ocellus, without parallel channels but with weak median
groove; occipital carina weakly developed, extending half
distance to eye margin; temple narrow and glabrate. Eyes
separated by 1.4x their height. Malar space 0.5-0.6x
height of eye. Clypeus sparsely setose, epistomal sulcus
weakly impressed. Antenna 1 1 -segmented; scape reach-
ing 0.8x distance to median ocellus, cylindrical; pedicel
l.Sx as long as broad; anellus stout; flagellum 0.9x height
of head; funicle 7-segmented; F2 twice as long as broad,
slightly shorter than pedicel, following segments subequal
in length and slightly broader apically; clava ovate, slight-
ly longer than preceding 2 segments.

Mesosoma with midlobe of mesoscutum rugulose and
with faint transverse ridges apparent; lateral lobe and
axilla smooth and polished with sparse setae; scutellum
glabrous. SSS deeply foveate with widely spaced, promi-
nent carinae. Scutellum twice as long as broad, base well
separated from TSA by broad depression, apex subtrun-
cate; frenal line narrow and weakly foveate, frenal area
glabrous, broadly semicircular in dorsal view; axillula
glabrous, axillular sulcus broadly foveate. Callus with
few small dorsal setae, callar nib absent. Femoral groove
broadly and shallowly impressed, glabrous; sternaular
area broadly foveate. Proepisternum mostly glabrous,
weakly carinate in anterior half. Coxae glabrous; femora
polished with sparse short setae; hind tibia slender, slight-
ly expanded apically. with 2 long spurs. Forewing
2.4-2.5X as long as broad; basal area setose disially;
speculum small with irregular margins; stigmal vein as

87

long as broad; postmarginal vein l-l.3x as long as stig-
mal vein.

Mctasoma with petiole 1.7-2.1x length of hind coxa.
2.I-2.2X length of propodeum; petiole ribbed, with
prominent dorsal flange basal ly. Gaster typical for genus.
Hypopygium with 4 or 5 long hairs along apical margin.
Ovipositor acicular. Gonostylus narrow and elongate.

MALE

Differs in the following characters: F2 2.4x as long as
broad; petiole 2.4x length of hind coxa, 3.3x length of
propodeum; petiole slightly subtriangular in cross-
section, with 4 strong parallel ribs dorsally; gaster typical
(genitalia withdrawn).

BIOLOGY

Collected from Picranopteris (or Picranoft erus [?]). I
assume this is a plant, although I can find no literature
reference to this generic name.

DISTRIBUTION

Known only from New Guinea within the Papuan subre-
gion (D. Fig. 277).

ETYMOLOGY

Named in honour of E. C. Dahms at the Queensland
Museum in appreciation for his work on sorting and clari-
fying Girault's type material.

Psilocharis joanneae sp. nov.

Figs. 138, 193

TYPE MATERIAL

Holotype, 9, "15.04S 145.07E/ Mt Webb Nat. Pk/ 27-30
Apr. 1981 Q/ I. D. Naumann/ ex ethanol." "collecting/ at
light." '-AUST. NAT./ INS. COLL." "HOLOTYPE/
Psilocharis/ joanneae Heraty." Deposited in ANIC.
Paratypes: Australia: Queensland: same data as holo-
type (6 9 9, 2(? (5, ANIC); Hope Vale Mission, 14 km W
by N, 15.16S I44.59E, 7-l().v.l981, 1. D. Naumann. col-
lected at light (19, ANIC); Mt Webb N. P..
28-3().ix.l980, J. C. Cardalc, collected at light (59 9,
ANIC).

DIAGNOSIS

P. joanneae is closely related to P. theocles, P. dahmsi,
and P. pacifica: all have axilla and lateral lobe nearly
smooth, scutcllum quadrate with reduced sculpture,
frenum glabrous, and occiput transverse or only slightly
emarginate. It can be distinguished from these species in
that the eye is relatively large and the malar space is
reduced (Fig. 193). the antennal flagcllum is short, and
the proepisternum is carinate.

FEMALE

Length. 1.6-1.9 mm. Head, mesosoma. and petiole black,
mesosoma with strong blue-green reflections; gaster and
femora dark brown, apical tergites of gaster with faint
greenish reflections; antenna, apex of femora, and rest of
legs yellowish brown. Wings hyaline, venation pale
brown.

Head subcircular (Fig. 193); occiput broadly emar-
ginate; LOL l.()-1.2x OOL. Face broadly rounded, pol-
ished with sparse decumbent setae; scrobal depression
reaching 0.6x distance to median ocellus, parallel grooves
absent, median groove crenulate; occipital carina weak,
extending just beyond lateral ocellus; temple narrow and
smooth. Eyes large, separated by l.2-1.4x their height.
Malar space 0.4-0. 5x height of eye. Clypeus sparsely
setose, lateral margin weakly impressed, epistomal sulcus
barely discernible. Antenna 10- to 11-segmented (Fig.
138); scape not reaching median ocellus, slightly expand-
ed medially, narrowing apically and flattened on ventral
surface under pedicel; pedicel 2.3x as long as broad; anel-
lus stout; flagellum 0.6-0.7x head height; funicle 6- or 7-
segmented, F8 may be partially to completely fused with
clava; F2 only slightly longer than broad, following seg-
ments subequal in length and slightly increasing in width;
clava ovate, wider than funicular segments and as long as
preceding 3 segments.

Mesosoma with midlobe of mesoscutum rugose-areo-
late, often appearing transversely carinate; lateral lobe
and axilla polished with sparse setae; scutellum rugose
with apex proximal to frenum glabrous. SSS narrow and
deeply impressed, crenulate. Scutellum 1.6x as long as
broad, subtruncate at apex, base narrowly separated from
TSA; frenal line narrow and shallow foveate, frenal area
glabrous, narrow, and crescent-shaped in dorsal view;
axillula weakly carinate. mostly glabrous; axillular sulcus
narrow and shallow foveate with strong lateral carina.
Callus with several short hairs along posterior margin,
callar nib weakly developed or absent. Femoral groove
broadly impressed, reticulate to lightly rugose; sternaular
area reticulate to broadly foveate. Proepisternum carinate.
Coxae glabrate. femora smooth to weakly imbricate dor-
sally; hind tibia slender at base, tapering to twice that
width at apex, with 2 spurs, outer spur minute and hardly
discernible from apical .setae. Forewing 2.2-2.4x as long
as broad; basal area and speculum bare; stigmal vein
twice as long as broad; postmarginal vein 1.5x as long as
stigmal vein.

Metasoma with petiole l.5-2.1x as long as hind coxa.
1.4-2. Ix as long as propodeum; petiole subcylindrical in
cross-section, flnely carinate to ribbed, ventral keel pre-
sent, without dorsal flange basally. Gaster typical for
genus. Hypopygium with 9 or 10 elongate hairs along
apical margin.

88

MALE

Length, 1.7 mm. Colour and sculpture as in female, but
lower half of face with prominent reddish reflections.
Antenna 1 1-segmented; anellus as long as broad (not
disc-shaped as in other species); shape of antenna and fla-
gellomeres as in female; flagellum 0.8x height of head.
Petiole 1.9-2.0X as long as hind coxa, 2.0-2. Ix as long as
propodeum.

DISTRIBUTION

North Queensland (Australia) as part of the Papuan subre-
gion (J, Fig. 277). Only 1 specimen of the other
Australian species, P. theocles, has been collected in
north Queensland.

ETYMOLOGY

Named after my daughter, Joanne.

Psilocharis theocles (Walker) comb. nov.

Figs. 139, 141, 145-146, 151, 194,213,223,264-265

Eucharis theocles Walker, 1839:11-12; Australia:
Sydney [BMNH, examined].

Psilogaster theocles — Girault, 1913b:93 (combination);
Girault, 1915:232.

Orasema gemma Girault, 1934:2; Australia: Queensland
[QMB, examined]. Girault, 1936:3 (erroneously pub-
lished twice). Dahms, 1983:249 (notes on type materi-
al). Synonymized by Boucek, 1988:521.

Orasema theocles — Boucek, 1988:521 (combination).

TYPE MATERIAL

Lectotype of Psilogaster theocles (designated by Boucek,
1988), 9, "Lectotype." "Type." "Sydney." "Psilogaster/
theocles (1839)/ Walker." "B.M. TYPE/ HYM. 5.618."
"d Orasema/ theocles/ (Walk.)/ det. Boucek 1978."

Lectotype of Orasema emma (designated as holotype
by Dahms, 1983, and as lectotype by Boucek, 1988), 9,
see discussion of type material by Dahms (1983). Labels
added to the card-mounted specimen include "HOLO-
TYPE/ T.8819/ E.C.D. 1983" and "Lectotype of emma."
The label "T8819/ ECD 1983" was added to the slide
mount.

dia(;n()sis

This species has the following similarities to P. Joanneae,
P. dahmsi, and P. pacifica: lateral lobe, axilla, and
frenum smooth and polished, scutellum quadrate, and
occiput broadly emarginate. It has the following differ-
ences from these species: head subtriangular, eye relative-
ly smaller and malar space larger (().6-().9x height of eye;
Fig. 194), scutellum rugose, and proepisternum weakly
carinate or smooth.

FEMALE

Length, 1.8-2.2 mm. Head, mesosoma, and petiole dark
brown to black and usually with faint bluish-green or red-
dish reflections; antennal flagellum, coxae, and gaster
dark brown, flagellum sometimes lighter basally; femora
dark brown (except apex) to completely yellowish brown
or yellow; scape and rest of legs yellowish brown. Wings
hyaline to slightly infuscate. venation brown.

Head subtriangular (Fig. 194); occiput only very
weakly emarginate; LOL 0.8-l.lx OOL. Face broadly
rounded, polished, and sparsely setose; scrobal depression
reaching 0.8x distance to median ocellus, parallel grooves
weakly impressed, median groove, if present, short and
smooth; occipital carina weak, extending laterally to dor-
sal margin of eye; temple narrow and glabrate. Eyes sepa-
rated by 1.4-1.6X their height. Malar space 0.6-0. 8x
height of eye. Clypeus sparsely setose; lateral margin
weakly impressed, epistomal sulcus barely discernible.
Antenna 9- to 1 1-segmented (Fig. 139); scape not reach-
ing median ocellus, cylindrical but flattened ventrally
below pedicel; pedicel 2.3x longer than broad; anellus
prominent; flagellum 0.8-1. Ox height of head; funicle 5-
to 7-segmented; F2 1.8x as long as broad and shorter than
pedicel, following segments subequal in length and only
slightly broader apically; clava ovate, as long as preced-
ing 3 segments, clava and F7-F8 sometimes incompletely
fused.

Mesosoma with dorsum rugose-areolate; lateral lobe of
mesoscutum and axilla smooth and polished with sparse
setae; scutellum rugose-areolate with margins weakly
sculptured to glabrate. SSS narrow and deeply impressed,
crenulate. Scutellum 1.5-l.6x as long as broad, subtrun-
cate at apex; frenal groove strongly or weakly impressed,
frenal area glabrous; axillula glabrous, axillar sulcus
deeply impressed and with strong carina. Callus with
patch of dense elongate setae dorsally, callar nib promi-
nent or absent. Femoral groove broad (Fig. 223), weakly
sculptured to glabrate; sternaular area rugose to lightly
reticulate. Proepisternum weakly carinate to glabrate.
Coxae glabrate, femora weakly imbricate dorsally, mostly
smooth; hind tibia slender, only slightly wider at apex,
with 2 spurs. Forewing 2.2-2.4x as long as broad (Fig.
146); basal area with few sparse setae proximal to specu-
lum; stigmal vein 2-3x as long as broad; postmarginal
vein 1.0-I.5X as long as stigmal vein.

Metasoma with petiole 1.5-2. Ix as long as hind coxa,
1.5-2. Ox as long as propodeum; petiole subtriangular in
cross section, ribbed dorsally. aciculate to elongate reticu-
late sublaterally, without dorsal flange basally (Fig. 145).
Gaster typical (Figs. 264-265). Hypopygium with 8 to 10
elongate hairs along apical margin.

89

MALF

Length. 1.9-2.1 iiitii. Colour and sculpture as in female.
but face with prominent greenish reflections. Antenna 1 1 -
segmented (Fig. 141). shape of segments as in female;
scape with prominent pores; flagellum 1.0-1.2x height of
head. Petiole l.7-2.4x as long as hind coxa. 2.0-2.7x
length of propodeum; petiole cylindrical, ribbed dorsally,
aciculate ventrally and without ventral keel. Gaster typi-
cal, as long as petiole. Genitalia typical (Fig. 151).

HABITAT

Collections have been made along rainforest margins on
low vegetation or in dry sclerophyllus Eucalyptus forests.
I have collected this species at 2 locations in Australia:
near Brisbane, adults were collected from short plants in
the middle of a hydro-line cut; near Perth, adults were
collected from short plants along a trail in an open Jarrah
forest. Adults were collected from short, non-flowering,
broad-leaf plants, but no direct association with a particu-
lar species could be made.

DISTRIBUTION

Distributed in the Australian subregion in eastern
Australia. Tasmania, and southwestern Australia (T. Fig.
277). Numerous records have been made for southeastern
Australia and only 1 record from north Queensland
(Yungaburra) on the Atherton Tablelands. The western
population is almost identical to the eastern populations.

MATERIAL EXAMINED

Australia: A. C. T.: Black Mtn; Gibraltar Falls: Wombat
Ck; Woods Reserve, nr Gibraltar Falls; January to March
(6$ 9, 4dc?, ANIC); New South Wales: Brindle Ck;
Canley Vale; Casula; Congo. 8 km SE by E of Moruya;
Dorrigo; Mt Dromedary nr Narooma. 700 m; Gibraltar
Range, rainforest margin; Monga State Forest, dry sclero.
Eucalyptus forest; Rotary Lookout. 19 km NW Milton;
Tooloom Scrub; Tooloom Plateau, via Urbanville,
600-700 m, sweeping low vegetation, rainforest, also at
light; nr Wilson's Peak, via Teviot Gap. 700-800 m.
sweeping low vegetation, rainforest; October to March.
July (399 9, 38(5(5, ANIC. BMNH, CNC. UQAB);
Queensland: Bald Mtn area, via Emu Vale, 1000-1300 m;
Mt Glorious, rainforest, sclerophyll forest; Mt
Tibragargan; Maleny. 7 km SW. 300-450 m. sweeping
low vegetation, rainforest/ araucaria cunninghamii planta-
tion boundary; Yungaburra, Atherton Dist.; December to
March (179 9, 16 6. ANCI. BPBM. BMNH. QMB.
ROM, UQAB); Tasmania: Arve Riv. (43.()9S, 146.48E);
14 km S Bronte Park (42.15S, 146.29E); Bronte Park. 9
km S; Edwards Rd, Hart/. Mtns (43.07S, 146. 47E);
Franklin Riv. (42.13S, 146. OlE); Frodsham's Pass
(42.49S 146.23E): Geeveston, 13 km W; Gladstone. 1 km
SSE (40.58S, 148.0 IE); Nunamara. 10 km ENE (41.22S.

147.24E); Port Arthur; Wilmot. 14 km SW by S (41.30S.
146.()5E); January to March (55 9 9. 31 (5 c5. AEI. ANIC.
BMNH): Victoria: Omeo. 7 km SW by W; Growler Ck.
Lind N. P.; February (29 9, ANIC); Western Australia:
Crowea S. F. nr Pemberton, October to December (3 9 9,
1(5, ANIC).

Psilocharis afra .sp. nov.

Figs. 135-136. 148-150, 195,222

TYPE MATERIAL

Holotype, 9 , "W. Uganda/ Kalinzu Forest/ 5000 ft. 1 1525
m] July 1972/ H. Falke." "HOLOTYPE Psilocharis/ afra
Heraty." Deposited in CNC.

Paratypes: Cameroon [Kamerun]: R. Thaxter (19,
MCZ). Kenya: Malindi. forest near coast, v. 1973, H.
Falke (109 9, slide no. 883. CNC; 1 9. ROM): Nanyuki,
2100 m. 37° 07'E. 0° 02'S. 22.iv.1973, Hans Gonget (19,
ZMC); Nairobi. Ngong Rd forest. 11. xii. 1990-3.1.1991,
B. D. Gill. FIT (1 9, 1(5. JMH). Malawi: Nyika Plateau.
2200 m 12.vii.l972. J. Eccott (3(5(5, CNC). South
Africa: E. Transvaal, Malta forest, 20 km W
Trichardtsdal. 22.xii.1985. M. Sanborne (lc5, CNC);
Kruger N. P.. Satara, 16.xii.l985, M. Sanborne (19,
CNC); [Pondoland] Port St. John, 1 8-3 l.iii. 1924, R. E.
Turner (1(5, BMNH). Uganda: same data as holotype
(109 9, CNC; 29 9, JMH); Busbinyi(?). iii.1939. H. C.
Taylor, T.1085 (4c5 c5 , BMNH); Fernando, Moka,
l.ii.l933. W. H. T. Tams (19. BMNH); Kayonza forest,
Kigezi Dist., v. 1972, H. Falke (29 9, CNC); Jinja,
Mabira forest, 5-14. iv. 1973, H. Falke (19, CNC);
Kawanda. x.l942, T. H. C. Taylor, T.1085 (1 9, BMNH);
Kawanda. I.vii.l942. xi.l942. xii.1943. T. H. C. Taylor,
T.1085 (39 9, lc5, BMNH); Lutotoana [?], iii.1939, T.
H.C.Taylor (1(5, BMNH).

DIAGNOSIS

Differences from other species: lower face elongate with
the posterior margin of the gena hidden in frontal view,
scrobal depression with distinct parallel grooves, occipital
carina pronounced, midlobe and scutellum completely
scabrous to reticulate, and femoral groove of both sexes
partially to completely reticulate.

FEMALE

Length, 1.6-2.0 mm. Black; head, gaster. and mesosoma
laterally with faint greenish reflections, mesosomal dor-
sum with strong greenish reflections (mesosoma all black
in some); antennal flagellum and femora brown; scape,
tibiae, and tarsi yellowish brown. Wings hyaline, vena-
tion brown.

Head subtriangular. eye large, making face appear nar-
row and elongate; occiput emarginate; LOL 1.0-1.8x

90

OOL. Face relatively flat, polished with sparse decum-
bent setae; scrobal depression reaching just over half dis-
tance between toruli and median ocellus, channels dis-
tinct, apex of each channel with distinct semi-circular
depression, smooth median groove present (Fig. 195);
occipital carina sharp, extending to eye margin; temple
broad, obliquely strigate dorsally. Eye with sparse minute
setae, separated by 1.2-1.4x their height. Malar space
0.4-0.6X height of eye. Clypeus bare to moderately
setose; lateral margin sharply impressed, epistomal sulcus
absent. Antenna 11-segmented (Fig. 140); scape almost
reaching median ocellus, cylindrical; pedicel 1.8x as long
as broad; anellus present; flagellum 0.8-1. Ox height of
head; funicle 7-segmented; F2 1 .4-2.0x as long as broad,
following segments subequal in length, flagellum gradu-
ally increasing in width apically; clava acute, as long as
preceding 2 flagellomeres.

Mesosoma with midlobe of mesoscutum and scutellum
scabrous to reticulate (Fig. 222); lateral lobe irregularly,
transverse-carinate; axilla longitudinally carinate. SSS
broadly and deeply impressed with several large trans-
verse ribs. Scutellum twice as long as broad, broadly
rounded apically, narrowly separated from TSA at base;
frenal line narrow and foveate, frenal area vertically cari-
nate, vertical, and hardly visible from above; axillula
reticulate, axillular sulcus narrow. Callus with few long
setae dorsally and with prominent callar nib. Upper
mesepimeron glabrous, lower mesepimeron rugose,
transepimeral sulcus broadly foveate; femoral groove
broadly impressed, anterior half of groove reticulate,
sculpture occurring in irregular patches (Fig. 222); ster-
naular area narrow and shallowly foveate. Proepisternum
smooth to very weakly carinate. Fore and hind coxae
weakly imbricate to smooth with sparse setae, mid coxa
weakly strigate; femora weakly imbricate with moderate-
ly dense setae; hind tibia stout with dense, semi-erect
setae and 2 spurs, outer spur minute and barely dis-
cernible. Forewing 2. l-2.5x as long as broad; basal area
and speculum bare; stigmal vein twice as long as broad;
postmarginal vein as long as stigmal vein (Fig. 148).

Metasoma with petiole 1.3-2.Ix length of hind coxa.
1.4-2. Ox length of propodeum; petiole subtriangular in
cross section, dorsally ribbed, weakly carinate to acicu-
late sublatcrally, without dorsal flange at base. Gaster
typical for genus. Hypopygium with 10 elongate hairs
along apical margin (Fig. 149). Ovipositor acicular, as in
Fig. 150.

MALK

Length. 1..V2.4 mm. Colour and sculpture as in female.
Antenna 10- or 1 1 -segmented (Fig. 135); scape reaching
median ocellus, cylindrical but (laltened ventrally: anellus
minute or absent; Hagcllum 1.6-1.7x height of head; F2
2.0-3.0X as long as broad. Petiole 2.0-2.6x length of hind

coxa, 2. 3-3. Ox length of propodeum; petiole carinate dor-
sobasally, otherwise aciculate, cylindrical. Gaster small,
shorter than petiole. Genitalia typical for genus.

VARIATION

Little variation was observed in most of the specimens
examined including those from Cameroon and South
Africa. The male from Lutotoana (Uganda) has a rugose
face and completely carinate petiole. A rugose face was
found nowhere else in Psilocharis and this specimen was
regarded as aberrant. Males collected from the Transvaal
are similar to the paratypic males but have a slightly
shorter petiole (2. Ox length of hind coxa). The female
from Kruger National Park has very light sculpture on the
lateral lobe of the mesoscutum (similar to P. momlicera).
The females from Kawanda (Uganda) and Malawi have
the head and mesosoma black, hind coxa smooth with
only a few lateral setae, and the femoral groove almost
entirely reticulate.

DISTRIBUTION

Southern Africa (A, Fig. 277); probably sympatric with P.
momlicera.

ETYMOLOGY

From Latin for African; referring to the distribution.

Psilocharis hypena sp. nov.

Figs. 142-143, 147, 196-198. 224-225, 239-240. 249

TYPK MATERIAL

Holotype, 9, "BRITISH N. BORNEO:/ Tenompok/
10-19.11.1959." "T. C. Maa/ Collector/ BISHOP."
"HOLOTYPE/ Psilocharis/ hypena Heraty." Deposited in
BPBM.

Paratypes: Malaysia: Sabah: Laiwan. 14-19.1.1959. T.
C. Maa (lc5, BPBM); Mt Matang, 28.1.1914, G. E.
Bryant (1(5. BMNH); Sensuron, 9-11.1.1959, T. C. Maa
(19, BPBM); Tenompok. 10-19. ii. 1959. T. C. Maa
(15 9 9. 9(5 d, BPBM); Tenompok, 1460 m. Jessclton. 48
km E. 10-19. ii. 1959, T. C. Maa ( 1 c5 , BPBM);
Tenompok, 13.ii. 19.59. T. C. Maa (5 9 9, 2c5c5, BPBM);
Tenompok. 15.ii.l959, T. C. Maa (1 9 , lc5,BPBM);
Sarawak: Bau Dist., Bidi, 90-240 m. 3.ix.l958. T. C.
Maa (19, BPBM).

I)IA(;nosi.s

Differences from other species: lower face broad with the
posterior margin of the gcna rounded and prominent in
frontal view (Figs. 196-197), scrobal depression with
smooth parallel channels, occipital carina weak, midlobe
of mesoscutum and scutellum completely scabrous to
reticulate (Figs. 239-240), and femoral groove polished

in female (sometimes coarsely reticulate anteriorly), par-
tially reticulate in male (Figs. 224-225).

FKMALK

Length, 1.7-2.7 mm. Black, mesosoma with strong green-
ish reflections, gaster with violaceous reflections basally:
antenna and femora dark brown, tibiae and tarsi brown to
yellow ish brown. Wings hyaline, veins brown.

Head subtriangular (Figs. 196-197); occiput broadly
emarginate; LOL 1.0-1.6x OOL. Face broadly rounded,
polished with sparse decumbent setae; scrobal depression
reaching 0.6x distance to median ocellus, parallel chan-
nels weakly impressed, apex of each channel with distinct
semi-circular depression, median groove absent; occipital
carina weak or absent, extending to lateral ocellus; temple
broad, weakly strigate dorsally. Eye with small erect
setae, eyes separated by 1.0-1.3x their height. Malar
space 0.4-0.6X height of eye. Clypeus sparsely setose to
bare except for apical margin of elongate setae, lateral
margin weakly impressed, epistomal sulcus absent or
weakly impressed. Antenna 11 -segmented; scape almost
reaching median ocellus, cylindrical but flattened on ven-
tral surface below pedicel; pedicel 2. Ox as long as broad;
anellus present; flagellum 0.7-0.9x height of head; funi-
cle 7-segmented; F2 1.2-1.6x F3, following segments
subequal in length, slightly increasing in width to apex;
clava conate, as long as preceding 2 segments.

Mesosoma with dorsum rugulose-areolate (Figs.
239-240); lateral lobe and axilla carinate. Scutellum 1.4x
as long as broad, sharply rounded at apex; frenal groove
narrow and foveate, frenal area vertically carinate. verti-
cal, and hardly visible in dorsal view; axillula rugose,
axillular sulcus narrow and not deeply impressed. Callus
with 5 to 13 short hairs, callar nib prominent. Upper and
lower mesepimeron smooth, transepimeral sulcus broadly
foveate; femoral groove broadly impressed and glabrous;
sternaular area broad and deeply foveate anteriorly,
abruptly narrowed posteriorly. Proepisternum carinate.
Coxae glabrate; hind femora weakly imbricate to smooth
with dense short setae; hind tibia narrow basally, broad at
apex, with moderately dense, semi-erect setae and 2
spurs, outer spur small. Forewing 2.2-2.4x as long as
broad; basal area and speculum bare; stigmal vein twice
as long as broad; postmarginal vein shorter than stigmal
vein (Fig. 147).

Metasoma with petiole 1.5-1.8x as long as hind coxa,
1.5-1.9X length of propodeum; petiole subtriangular in
cross section, ribbed dorsally, aciculate to smooth sublat-
erally, without dorsal flange at base. Gaster typical for
genus. Hypopygium with 8 to 10 elongate hairs along api-
cal margin.

MALK

Length. 1.7-2.4 mm. Colour and sculpture as in female.
Ocelli large, LOL 1.0-1.7x OOL. Eyes separated by
1.1-1.3X their height. Face generally more setose than in
female. Antenna 11 -segmented (Figs. 142-143); scape
reaching top of median ocellus, cylindrical, with small
pores ventrally; tlagellum 0.9-l.lx height of head; F2
1.8x as long as broad. Petiole 1.9-2.5x length of hind
coxa, 1.8-2.8X length of propodeum; petiole cylindrical
and strongly carinate, ventral keel not apparent.

HABITAT

I have collected specimens on understorey plants within a
rainforest in Malaysia, and also at 2 localities, both high-
elevation rainforest, in Nantou Hsien province of Taiwan.

VARIATION

Over the range of this species, there is variation in the
length of the antennal flagellum. density of setae on the
callus, presence or absence of setae on the eye, relative
size of the eyes to their separation, and relative length of
the petiole. Individuals from Borneo consistently have the
shortest flagellum and petiole, eye densely setose, callus
with 11 to 13 hairs, and a glabrous frenal area.
Individuals from Taiwan are not included within the type
material because their antennal flagellum is shorter
(0.9-1. Ox head height for females. 1.0-1.2x for males),
petiole is longer (2.0-2.4x length of hind coxae for
females, 1.9-2.4x for males), dorsal sculpture is weak
(Fig. 239), frenal area is glabrous (Figs. 239, 249). and
callus has only 3 to 7 hairs. The Taiwanese population is
distinct, but collections from Japan have forms that con-
form with specimens from either Borneo or Taiwan.
Specimens from the Indo-Chinese, West Malayan, and
Philippine subregions generally agree with the Borneo
(East Malayan) forms but have a longer flagellum and
petiole (within range of the East Malayan specimens) and
various combinations of sculpture and pilosity. The range
of variation suggests that only 1 species should be recog-
nized.

DISTRIBUTION

Widely distributed in the Indo-Chinese and Malayan sub-
regions (H, Fig. 277).

OTHER MATERIAL EXAMINED

India: H.P.I?], Newchowk[?]. on grass; T. Nadu,
Mangarai Forest; September and October (19, \6 .
BMNH, JMH). Indonesia: Sumatra: Aceh, Gunung
Leuser N. P., Ketambe Res. Sta., primary young rainfor-
est, closed canopy: same data but 400 m, mature forest,
light gap; October to November (19, Id, MZB, ROM).
Malaysia: Selangor: 21 km W Gombak, 900 m, rainfor-
est, June (1(5. JMH). Japan: Kyushu: Kamiozoegawa.

92

Fuji, Saga Prefecture. August and October (49 9, 36 6,
KUEC). People's Republic of China: Guangdong: Ding-
Hu Mtns, 60 km W of Guangzhou, June (36 6. BMNH).
Fukien: Chunggan. Bohea Hills. July to August (2$ 9,
TARI); Kiangsi: Kwangche, Chienmen, August (26 6,
TARI). Philippines: Los Banos (1$, USNM). Taiwan:
Nantou Hsien: Meifeng, 2150 m; Tungpu, 1200 m;

October (2\9 9 , 26 6 , TARI); Taichung Hsien:
Chiapaotai, 750 m, October (19,16 TARI).

ETYMOLOGY

From Greek hypene, for mustache; referring to clypeal
setae.

Neolosbanus gen. nov.

Type Species: Eucharis palgravei Girault, 1922:105; by
present designation.

Neolosbanus includes a group of species that were origi-
nally misplaced in Loshanus Ishii by Watanabe (1958).
Watanabe (1958) transferred the species Parapsilogaster
laeviceps Gahan and Psilogaster nishidai Ishii and
Nagasawa to Loshanus, and described an additional
species, Loshanus grcssitti Watanabe. Hedqvist (1978)
described Loshanus petersoni, and placed Eucharis pur-
pureovenths Cameron in the genus Gollumiella Hedqvist.
Boucek (1988) transferred G. purpureoventris to
Orasema Cameron and treated L. laeviceps, L. nishidai,
and L. petersoni as junior synonyms of O. purpureoven-
tris. Gollumiella was treated as a subjective junior syn-
onym of Loshanus Ishii based on the description of the
type species Loshanus uichancoi Ishii, 1932 (Boucek,
1988). Examination of the type material for L. uichancoi
resulted in the transfer of this species to the genus
Orasema, and Gollumiella was given revised status in the
Eucharitinae by Heraty (1992). Orasema purpureoventris
is transferred here to Neolosbanus. Parapsilogaster laevi-
ceps is raised from synonymy with Orasema purpure-
oventris. and is placed in Neolosbanus. Psilogaster
nishidai and Loshanus petersoni are placed here as junior
synonyms of A', palgravei, which is the type species of
Neolosbanus.

Neolosbanus is distinguished from other Eucharitidae
by the following features: sculpture of head smooth or
punctate, dorsal occipital margin carinate, clypeal margin
rounded and without an anteclypeus, palpi 3-segmented.
prepectus not fused to pronotum and fovcate. femoral
groove foveate, petiole gradually narrowed basally, and
first gastral stemite smooth (not medially constricted).

(;enkri( de.scription

Head subtriangular, 1.2-1.7x as broad as mesosoma;
median ocellus anterior to lateral ocelli or arranged in a
line, lateral ocellus close to occipital margin. Face smooth
and polished or with pits and punctures; scrobal depres-
sion smooth and broadly impressed; occiput smooth and

broadly concave in dorsal view, occipital carina distinct;
ocellar-ocular groove absent. Clypeus as high as wide,
apical margin rounded and smooth, broadly rounded, or
strongly produced over base of mouthparts; epistomal sul-
cus present or absent; anteclypeus absent. Malar depres-
sion present or absent; hypostoma separated from gena by
hypostomal carina and extending medially over base of
mandible. Labrum 4-digitate, setae of digits long and
spatulate. Mandibles falcate and 2/3 toothed (Figs. 200.
203-204, 206); maxilla and labium elongate, maxillary
and labial palpi each 3-segmented, palpi long and thin
with only few minute setae. Antenna 10- to 1 3-segment-
ed; scape elongate and usually cylindrical, usually reach-
ing median ocellus; pedicel short, not much longer than
broad; anellus present or absent; funicle 7- or 8-segment-
ed, segments cylindrical with basal secondary segmenta-
tion (Figs. 214—216), and densely setose; basal funicular
segments more than 2.5x as long as broad, rarely shorter,
terminal 2 or 3 segments fused into clava.

Mesoscutum with notauli well defined along entire
length. TSA complete. SSS usually fovcate, slightly
angled to midline, and reaching TSA through medial
depression. Scutellum with frenal area usually separated
by distinct frenal groove, frenal area polished or weakly
sculptured; axillular sulcus weak. Metanotum usually
extended laterally as smooth flange overlapping base of
propodeum and partially covering propodeal spiracles;
spiracles close to dorsal margin of propodeum
(Neolosbanus townesi and A^. laeviceps with weak flange
and spiracle exposed; Fig. 250). Propodeal disc rounded
and smooth laterally, usually with medial band of rugose-
areolate sculpture; callus only slightly swollen and pol-
ished with few to many setae but without callar nib.
Mesopleuron smooth; mesepimeron relatively flat,
transepimeral sulcus usually present (mesoplcural sulci
absent only in A', townesi); femoral groove narrow and
foveate, continuing to ventral margin of mesepistemum;
sternaular area of mesepistemum foveate in some species.
Prepectus reaching tegula as triangular or fingerlike dor-
sal lobe, strongly narrowed ventral ly. dorsal lobe fo\ calc
(rounded in N. townesi). Proepistemum flat and glabrous.

93

Coxae and femora sniooih and shining; tibiae with spurs
I- 1-2; hind tarsus only slightly shorter than tibia.

Win}> veins of fore and hind wings well defined.
Forewing 2.3-2.7x as long as broad, broadly rounded api-
cally; disc densely pilose, marginal fringe short; submar-
ginal vein with short setae dorsally; marginal vein
0.25-0. 36x as long as forewing and pilose; stigmal vein
less than 3x longer than broad, as broad as or narrower
than marginal vein; postmarginal vein short or long.

Mctasoma with petiole of both sexes more than 1.3x as
long as hind coxa, gradually narrowed basally, without
dorsal or lateral flange (Fig. 159). cylindrical and fused
ventrally. Gastral terga smooth and shining, usually bare,
gaster of female as long as head and mesosoma or slightly
shorter, gaster of male slightly longer than hind femur;
Mt^ of female more than 0.9- 1. Ox as long as hind femur;
Ms, smooth and without basal constriction. Hypopygium
bare or with few short setae on either side of midline. Ms„
of male densely setose apically. Cercus with few elongate
setae of equal length. Gonostylus narrow or broad, fused
to second valvifer. Ovipositor needlelike or expanded,
sculpture variable between species. Male with paramere
well developed and bearing several stout setae, digitus
disclike with several stout marginal spines; aedeagus nar-
row or broad.

ETYMOLOGY

Combination of Greek neo, meaning new, and loshanus
from the genus name originally proposed for specimens
collected from Los Banos in the Philippines; gender mas-
culine.

PHYLOGENETIC RELATIONSHIPS

Of the 4 species-groups recognized in Neoloshanus, the
gemma-gxouT^ has 3 species (apoanus, gemma, and
wusheanus), the purpureoventris-group has 3 species
(purpureoventris, storeyi, and watanahei), the palgravei-
group has 6 species {laeviceps, nepalensis, palgravei,
pilosus, taiwanensis, and townesi), and the gressltri-group
has 4 species {anapetus, gressitti, kokiireanus, and vio-
laceus). Unnumbered characters discussed below were
not used in the phylogenetic analysis of higher relation-
ships as they are considered of value only at the species
level.

The gemma- and purpureoventris-group^ both have a
needlelike ovipositor (42) and cylindrical gonostylus or
sheath (46). Both of these states are plesiomorphic for
Neoloshanus and are shared with at least I. but not all. of
the outgroups. The presence of a unique (crenulate)
scrobal depression (9), broad gonostylus, and absence of
MPS in males distinguish the gemma-group from other
species of Neoloshanus. No resolution of species within
the gemma-group was possible. The presence of a
glabrous frenum suggests a close relationship between N.

wusheanus and N. apoanus: however, this character state
is found throughout the outgroup taxa and its polarity is
uncertain.

The remaining species are interpreted as monophyletic
based on an increase in the number of funicular segments
in males (from 7 to S or 9) (character 4; Tabic 2), and a
glabrous occiput (with reversal in the ,i,'/-<',v.s/7//-group).
Three additional synapomorphics include the apex of the
clypeus extended as a strong lobe (12). prepecius nar-
rowed to a fine ridge along the ventral half, and siernaular
area smooth, but these are all proposed to have undergone
reversal in the gressird-group. The instability of these last
characters suggests that the purpureoventris- and pal-
gravei-groupa are sister groups, and that the expanded
ovipositor in the palgravei- and gressitti-groups may have
been independently derived within Neoloshanus. This is
further suggested by morphological differences of the
ovipositor found in the gressitti-group, including absence
of lateral ridges on the first valvula (43). widely separated
lateral teeth on the second valvula (44). and an even
thickening of the ovipositor (versus subapical expansion).

The purpureoventris-group is monophyletic based on
presence of 9 funicular segments in males (4). and stig-
mal vein strongly angled distally (35). The ovipositor is
needlelike (42), which is regarded as plesiomorphic for
Eucharitidae. and the gonostylus is narrow and elongate,
sometimes exceeding the cercus. Within the purpureoven-
tris-group, N. purpureoventris and A^. storeyi may be sis-
ter species based on having a striate frenum. The raised
ocellar triangle of A', storeyi is autapomorphic within
Neoloshanus.

Monophyly of the palgravei-group is based on the fol-
lowing: ovipositor subapically expanded, first valvula
with oblique ridges (43). and second valvula narrow with
the ridges coalescing dorsally (44). all states that are
found in some or all Oraseminae. Except for A', townesi,
other species in this group share a 7-segmented funicle in
females (5). and strong projection of the clypeal margin
(12). Neoloshanus townesi is treated as the derived sister
species to A', laeviceps based on loss of the anellus ( 1 ),
expansion of the scape of males, and an enlarged truncate
MSj^. with autapomorphic reversals to a rounded clypeal
margin (12) and independent derivation of an 8-segment-
ed funicle in females (5). Except for loss of the anellus,
character states of A^. townesi are also shared with species
in the gressitti-group. Of the remaining species in the pal-
gravei-group. N. pilosus and A', taiwanensis share a
densely pilose mesosoma. and together are treated as the
sister group to A', nepalensis and A', palgravei. although
no synapomorphics were found to support this latter
group.

Monophyly of the gressitti-group is supported by the
8-segmcntcd female funicle (5). and second valvula broad
and smooth with strong lateral teeth (44). Monophyly of

94

this group is proposed by a reversal in clypeal shape (12)
and shape of the ventral half of the prepectus (narrow but
not ridgelike) (20). A yellow femur is found only in N.
gressltti and A^. kokureanus and may support a closer rela-
tionship between these species. Monophyly of the
remaining species is based on presence of a reduced
malar depression (10), and complete facial sculpture (8).
The presence of stemaular sculpture and a striate frenum
in these taxa may be plesiomorphic. Males of N. gressitti
are unknown but may have a more strongly sculptured
face, as in A', kokureanus. If the reduction of the ster-
naular sculpture and glabrous frenum are autapomorphic
for N. gressitti, then N. gressitti and N. kokureanus would
form the sister group to the remaining species in this
group. Monophyly of A^. anapetus and A', violaceus is
suggested by the sculptured occiput, expanded scape of
males, and brown femur, which may all be plesiomorphic,
but adults of both sexes also have a strongly sculptured
face and are morphologically very similar. No further
relationships among these species could be assessed.

The relationships among species of Neoloshanus may
be described as {{gemma + wusheanus + apoanus) +
{{watanahei + {purpureoventris + storeyi)) + {{townesi +
{laeviceps + {{pilosus + taiwanensis) + nepalensis + pal-
gravei))) + {{gressitti + kokureanus) + {anapetus + vio-
laceus))))) (Figs. 278-279).

BIOLOGY AND IMMATURE STAGES

The immature stages of N. palgravei are described in the
earlier section on biology of Psilocharitini. I reared both
N. gemma and A', palgravei from the cocoons of
Hypoponera (Formicidae: Ponerinae). There is no ques-
tion that these 2 species belong to well-separated clades
within Neoloshanus and it may be postulated that most
species of Neoloshanus are parasitic on Hypoponera.
Except for one derived group of Eucharitinae (on
Formicinae), other members of this subfamily are almost
all parasitic on Ponerinae. Almost all known hosts for
Oraseminae are Myrmicinae, with a few rare records of
Formicinae and Ecitoninae (Heraty. 1990). No ant host is
known for the proposed sister group, Psilocharis.

Detailed plant-host records for A', palgravei suggest
that eggs are deposited into the leaves of a wide variety of
host plants. Additional plant records are known only for
A', townesi; adults were collected on Castanopsis (same as
for A', palgravei) and bamboo, which suggests similar
habits for members of the palgravei-grou^. The type
material of A', gemma was collected from a flower and
may indicate a different strategy for species with an acic-
ular ovipositor.

First-instar larvae of N. laeviceps and A', palgravei
share derived character states with other Eucharitinae.
The mature larvae of A', palgravei and the pupae of A'.
gemma and A', palgravei are simple and lack any of the

derived character states of Orasema (see Wheeler, 1907;
Heraty, 1990). The mature larvae and pupae are ple-
siomorphic in morphology and hence do not support
inclusion within Eucharitinae; however, absence of any
shared states with Oraseminae argue against this genus"
having evolved from within Orasema from groups that
share a similar propodeal morphology and expansion of
the ovipositor.

Whether the ancestor of a monophyletic group com-
prised of Oraseminae and Eucharitinae had an expanded,
strongly ridged ovipositor is unknown. It is absent
(needlelike) from all of the proposed outgroup taxa and is
therefore assumed to be plesiomorphic for Eucharitidae.
Small differences in the morphology of the first and sec-
ond valvulae, and incongruence with other characters,
suggest that the expanded ovipositor was derived once or
twice within Neoloshanus, and independent from
Oraseminae. The ovipositor is utilized in a similar way to
Oraseminae for depositing eggs into plant tissue in cham-
bers hollowed out by the ovipositor. The method of egg
deposition in Psilocharis and the gemma- and purpure-
oventris-gro\xx>?>, which have a needlelike ovipositor, is
unknown; however, other Eucharitinae with a similar
ovipositor deposit eggs into preformed cavities in plant
tissue or on the leaf surface.

DISTRIBUTION

Sixteen species oi Neoloshanus are distributed throughout
the Indo-Pacific region, excluding the Australian subre-
gion (Figs. 278-280). Extreme disjunctions for this genus
are represented by records of Neoloshanus palgravei in
Algeria and another species (possibly new) in Uruguay
(see description oi N. violaceus). The record of the speci-
men from Uruguay appears to be valid but odd, consider-
ing that it is regarded as a member of a derived group
found only in the Malayan, Papuan, and Polynesian sub-
regions.

Hypoponera are widely distributed around the world in
tropical and subtropical regions in predominantly mesic
habitats (Brown, 1973), and limitations to the distribution
of Neoloshanus cannot be attributed to the distribution of
the host ant. The same may be said in conjunction with
the lack of specificity toward plant hosts. Species of
Neoloshanus are virtually all restricted to the Indo-Pacific
region, and occur only within the Papuan subrcgion of
Australia. One specimen oi N. palgravei, which is identi-
cal to specimens from the Indo-Chinese subregion. is
known from Algeria. This may be an inaccurate data label
or a true extension of the range into northern Africa. If
true, then I would explain this as western dispersal of the
most common Indo-Pacific species.

The disjunct placement of the specimen from
Uruguay, as a derived member close to A', violaieus of
the gressitti-group, is difficult to explain. The most closc-

95

1\ related species is rmiiul in Malaysia and the collection
data appear to be valid (see description of N. vioUucus).
It is possible that this species was recently colonized in
South America by human intercession as part of ship's
ballast or some other accidental means. I find it unlikely
that this is part of a relictual disiributit)n of species in this
genus, but this is a possibility.

There is little resolution of species relationships to
suggest biogeographic hypotheses in the Indo-Pacific
region. Relationships do suggest that species found in the

Papuan and Polynesian subregions are the more highly
derived, and that distributions of extant taxa may have
originated from a mainland Indo-Chinese ancestor. Only
A', pali^ravei is widely distributed throughout the region,
which indicates good dispersal capacity, but regional
variation shows that some isolation is evident. The above
hypotheses are concordant with an evolution and radia-
tion of species since the Miocene and connection of the
Australian plate to the Malaysian Archipelago (Keast,
1981; Axelrod and Raven, 1982; Noonan, 1985).

Key to Species of Neolosbanus

1 Scrobal depression with 2 vertical crenulate

channels extending from torulus to median ocel- 4(1)
lus, median area between channels polished
(Figs. 153-154); funicle of both sexes 7-seg-
mented (Figs. 152, 157-158); MPS absent in
male, present in female; ovipositor needlelike....
gemma-group. 2

— Scrobal depression completely smooth; funicle

of female 7- or 8-segmented, funicle of male 8- —

or 9-segmented (Figs. 168, 171, 180-186); MPS
present in both sexes; ovipositor needlelike or
expanded 4

2(1) Frenal area roughly sculptured and vertical; hind
tibia dark brown to black; eye large (Fig. 153);
Australia N. gemma (Girault), p. 98

5 (4)

— Frenal area glabrous and angled 45-80 degrees

to dorsum (Fig. 152); hind tibia yellowish
brown; eye moderate in size (Fig. 154) 3

3 (2) Postmarginal vein 3.3^x length of stigmal vein
(Fig. 160); sternaular area rugose-alveolate;
antennal flagellomeres closely appressed and —

cylindrical (Fig. 157); funicular segments of
male enlarged basally, F2 broader than pedicel,
following segments tapering to apex; Indo- 6 (5)

Chinese subregion

A^. wusheanus sp. nov., p. 99

— Postmarginal vein 2.8-3.2x length of stigmal —
vein; sternaular sculpture foveate; antennal fla-
gellomeres of male clearly separated and slightly

flared apically (Fig. 158); F2 as broad as pedicel,
following segments equal in width; female 7(4)

unknown; Philippines

N. apoanus sp. nov.. p. 100

Stigmal vein strongly angled relative to anterior
wing margin (Fig. 169); postmarginal vein more
than 3x as long as stigmal vein (Fig. 169);
ovipositor needlelike (Figs. 271-272); gonosty-
lus long and narrow, sometimes exceeding cer-

cus; antenna of male 13-segmented

purpureovcntris-group, 5

Stigmal vein almost perpendicular relative to
anterior wing margin (Figs. 162. 164-165); post-
marginal vein short, at most slightly longer than
stigmal vein; ovipositor expanded and strongly
ridged (Figs. 266-270); gonostylus broad and
tapering to apex, not exceeding cercus; antenna
of male 11- or 12-segmented 7

Head with shallow groove along anterior margin
of ocellar triangle, groove extending from anteri-
or margin of median ocellus to outer margin of
lateral ocellus and continuing around ocellus to
occipital carina (Fig. 179); New Guinea,
Queensland A^. storeyi sp. nov.. p. 101

Head without any indication of groove, vertex
and ocellar triangle evenly rounded 6

Frenum glabrous, at most with single faint medi-
an carina; Philippines. Palau Islands

A^. watanabei sp. nov.. p. 102

Frenum vertically striate; Indo-Chinese.

Malayan. Sulawesi subregions

N . purpureoventris (Walker), p. 103

Face and frenum completely polished (Figs.
201-205); stigmal vein more than 1.5x as long
as broad, postmarginal vein more than 1 .5x as
long as stigmal vein (Figs. 162, 165); ovipositor

96

usually subapically expanded, rarely thickened
along entire length; first valvula with diagonal
lateral ridges (Figs. 267, 269); second valvula

narrow, apical ridges meeting along midline

palgravel-group. 8

— Face smooth, partially punctate, or completely
sculptured (Figs. 176, 206-208), and frenum
glabrous or carinate (Fig. 254); stigmal vein as
long as broad, postmarginal vein less than 1.5x
as long as stigmal vein (Fig. 164); ovipositor
thickened along entire length; first valvula com-
pletely smooth (Fig. 270); second valvula broad,
smooth medially with strong teeth on lateral
margin gressitti-gmup, 13

8 (7) Anellus absent; scape expanded apically, strong-

ly so in male (Figs. 171, 184); frenal groove
absent or weakly impressed dorsally; dorsum of
mesosoma lightly sculptured (Figs. 241-242);
Ms^ of male broader than petiole and truncate
apically 9

— Anellus present (may be small and indistinct but
always visible with transmitted light); scape nar-
row and cylindrical (Figs. 180-182); frenal
groove foveate; dorsum of mesosoma strongly
rugose-areolate to areolate (Figs. 243-244); Ms^
of male narrower than petiole, rounded apically.
10

9 (7) Prepectus swollen medially (Fig. 226); mesoso-

ma almost completely glabrous dorsally and lat-
erally (Figs. 241, 250); funicle of female 8-seg-

mented; New Guinea

N. townesi sp. nov., p. 104

— Prepectus foveate medially (Fig. 227); mesoso-
ma lightly sculptured dorsally (Fig. 242); funicle

of female 7-segmented; Indo-Chinese

N. laeviceps (Gahan), p. 105

10(8) Head, mesosoma, and petiole moderately to
densely pilose (Figs. 202. 229); length of head in
dorsal view short at median ocellus, not exceed-
ing diameter of median ocellus (Fig. 178);
scrobal depression broadly and deeply impressed
11

— Body sparsely setose (Figs. 227-228, 23()-2.'^3),
petiole completely bare; length of head in dorsal
view long at median ocellus, exceeding diameter
of median ocellus by more than l.5x (as in Fig.
179); scrobal depression shallow and indistinct
12

11 (10) Head and mesosoma moderately pilose, eye with
sparse setae; propodeum with median carina;
hind tibia dark brown to black; in male, petiole
2.2-2.9X as long as hind coxa and 2.2-2.7x as
long as propodeum; female unknown; China,
Taiwan N. taiwanensis sp. nov., p. 106

— Head and mesosoma densely pilose, eye with
dense setae (Fig. 202); propodeum smooth, with-
out median carina; hind tibia yellowish brown;
in male, petiole 2.6-3.5x hind coxa and 2.7-3.4x
length of propodeum; Malayan subregion, ?New
Guinea N. pilosus sp. nov., p. 107

12(10) Flagellum of female more than 1.5x height of
head, flagellum of male more than 2.2x height of
head; callus with more than 15 hairs (Figs. 230,
252); hind tibia white or yellowish brown in

both sexes; Nepal

N. nepalensis sp. nov., p. 108

— Flagellum of female less than 1.4x height of
head, flagellum of male less than 2.1x height of
head; callus bare or with few hairs (Figs. 231.
253), if pilose (India. Nepal, and Algeria) then
hairs short, fine, and less than 10 are present;
hind tibia white to dark brown in female, always
dark brown in male; Indo-Pacific region and
?Algeria A^. palgravei (Girault), p. 109

13 (7) Femora yellow; frenum polished (may be rugose

in male); face of female polished with few scat-
tered pits lateral to supraclypeal area (Fig. 176).
face of male punctate to weakly rugose; Ms^^ of
male broad and truncate 14

— Femora dark brown to black except extreme
apex; frenum with widely spaced longitudinal
carinae (Figs. 232-233. 245, 254); face in boili
sexes completely pitted, coriaceous, or rugulose
(Figs. 206-208); MSj^ of male narrow and round-
ed 15

14 (13) Anellus absent; head and mesosoma dark brown

to black; lateral lobe weakly sculptured to
smooth, scutellum partially carinate; male

unknown; E. Caroline Islands (Ponape)

N. gressitti (Watanabe), p. 1 14

— Anellus present (Fig. 177); head and mesosoma
dark blue-green; lateral lobe transversely striate,
scutellum rugose-areolate; New Guinea and

Solomon Islands

N. kokiireanus sp. now. p. 114

97

15(13) Midlobe of nicsoscutum and scutcllum finely
and shallowly rugose-areoialc doisally (Fig.
245); hind tibia yellowish brown; scape of male
expanded apically and with distinct apical flange
on inner side; basal flagellomeres of male com-
pressed, F2 noticeably broader than F3 (Fig.
186); mesosoma dark green; Borneo and
Philippines A', anapetus sp. nov., p. 1 15

— Midlobe of mesoscutum and scutellum scabrous
or deeply rugose-areolate dorsally; hind tibia
yellow or brown; scape of male cylindrical or
expanded medially (not apically); basal flagel-
lomeres of male cylindrical, F2 not noticeably
broader than F3 (Fig. 185); mesosoma dark blue
or black, sometimes with strong violaceous

reflections; New Guinea

N. violaceus sp. nov., p. 1 16

Neolosbanus gemma-group

This group of 3 species has the ovipositor acicular,
scrobal channels crenulate, funicle of both sexes 7-seg-
mented, and the clypeal margin weakly rounded. These
species are uncommon but widespread in the Indo-Pacific
region from Taiwan to northern Australia.

GROUP DESCRIPTION

Head 0.3-0.4x as long as wide; ocelli arranged in tri-
angle, median ocellus not exceeding line drawn across
anterior margin of lateral ocelli. Face smooth with scat-
tered short setae; scrobal depression shallow with 2 paral-
lel crenulate grooves extending from torulus to median
ocellus, surface slightly raised and polished between
grooves; occiput aciculate. Clypeal margin broadly
rounded with line of fine setae along apical margin,
clypeus not strongly produced over base of mouthparts,
epistomal sulcus weak or absent. Malar depression linear
and strongly impressed along entire length. Antenna 11-
segmented; scape narrow and cylindrical in both sexes,
presence of pores on male scape not assessed; anellus pre-
sent; funicle of both sexes 7-segmented; female with
sparse MPS, male without MPS.

Mesosoma with dorsum of mesoscutum transversely
carinate (Fig. 155) or irregularly transverse-areolate
(appearing striate); scutellum rugose-alveolate; frenal line
foveate; frenal area glabrous or carinate, vertical angle to
dorsum variable; axillula glabrous, axillular sulcus weak.
Mesepimeron mostly smooth; femoral groove and
transepimeral sulcus foveate; mesepistemum with wedge-
shaped sternaular sculpture. Prepectus foveate and trian-
gular dorsally, gradually narrowed ventrally, polished
below upper triangular lobe. Forewing 2.4x as long as
broad (Fig. 160); speculum and basal area bare; stigmal
vein narrow, 1.5-2.5x as long as broad; postmarginal vein
more than 2x as long as stigmal vein.

Caster with basal tcrga glabrous. Ovipositor needle-
like; first valvula smooth; second valvula with minute
dorsal ridges connecting medially. Gonostylus elongate
and broad (partially concealed on material available), not
exceeding cercus. MSj^ of male narrow and rounded api-

cally. Genitalia of male with paramere narrow and elon-
gate, bearing 2 stout setae; aedeagus subacute.

Neolosbanus gemma (Girault) comb. nov.

Figs. 26, 153, 159

Orasema gemma Girault, 1932:4 [289]. Queensland,
Australia [QMB, examined]. Dahms, 1984:646 (notes
on type material).

TYPE MATERIAL

Lectotype (designated Boucek. 1988), 9, "ex. flower."
"Kuranda, Q./ March 1921 F.P.D." "LECTOTYPE."
"SYNTYPE/ T.8969/ E.C.D. 1983." "Orasema/ gemma
Gir/ Cotypes." "LECTOTYPE/ 9 Orasema/ gemma Grit./
det. Z. Boucek, 1986."

Paralectotypes: same data as lectotype, type numbers
T.8967-68, T.8970-71, 4 9 9. Additional slide material:
"Orasema/ gemma Gir/ 9 type." "SYNTYPES/
T.8967-T.8971 ECD 1983 [red label]."

DIAGNOSIS

Close to yV. wusheaiuis but differs in that frenum abrupt
and roughly sculptured, femora, trochanters, and hind
tibia uniform brown with reddish reflections. LOL/OOL
ratio large, and antennal segments distinctly separated.

FEMALE

Length, 2.2 mm. Head and mesosoma dark metallic
green, strongest colour on mesosomal dorsum; mesosoma
laterally and coxae with additional faint reddish reflec-
tions; gaster black; antennal flagellum dark brown; femo-
ra, trochanters, and hind tibia uniform brown with reddish
reflections; scape, fore and mid tibiae, and tarsi dark yel-
lowish brown. Wings hyaline, venation clear yellowish
brown.

Head subcircular (Fig. 153); occiput broadly rounded;
median ocellus separated from occiput by own diameter;
LOL 1.2x OOL. Face broadly rounded, smooth or with

98

several weak punctures lateral to torulus, setae moderate-
ly dense and minute; occipital carina extending to dorsal
eye margin; temple narrow and weakly aciculate. Eyes
large, separated by 1.4x their height. Malar space 0.4x
height of eye. Antenna 1 1 -segmented; pedicel slightly
longer than broad; flagellum 1 .Ox height of head; funicu-
lar segments densely setose with scabriculous surface
sculpture; F2 narrow. 2.7x as long as broad, about twice
as long as pedicel, following segments subequal in length,
slightly wider towards apex of flagellum; clava ovate, as
long as preceding 2 segments.

Mesosoma with dorsum deeply rugose-alveolate to
scabrous, septa of midlobe of mesoscutum aligned in
transversely carinate pattern; lateral lobe transversely car-
inate; axilla obliquely carinate. Scutellum slightly longer
than broad; frenal line weak and obscured by dorsal
sculpture, frenal area polished with irregular vertical cari-
nae, nearly vertical in profile. Propodeum smooth with
rugose-areolate median band; callus with several short
setae. Coxae polished with sparse setae; femora polished
with moderately dense short setae; hind tibia with 2 dis-
tinct spurs. Forewings 2.2x as long as broad, 2.7x as long
as mesothorax; stigmal vein perpendicular to wing mar-
gin, 2.5x as long as broad; postmarginal vein 3.2x as long
as stigmal vein.

Metasoma with petiole 1.8x length of hind coxa, 1.6x
length of propodeum; petiole ribbed dorsally and carinate
ventrally (Fig. 159). Ovipositor acicular.

MALE

Unknown.

VARIATION

The lectotype has sparse setiferous punctation similar to
that found in N. gressitti and N. kokurcanus of the gressit-
r/-group. All other specimens have the face completely
smooth.

BIOLOGY

Removed from cocoon of Hypoponera sp. in a colony
also parasitized by N. palgravei. Pupa (Fig. 26). No host
remains or cast skins of earlier instars remained in the
cocoon. 1 collected this species along a path in a rainfor-
est. The type material was collected from an unidentified
flower near this locality.

DISTRIBUTION

Known only from a single locality in north Queensland
(Australia), which falls within the Papuan subregion (G,
Fig. 278).

MATERIAL EXAMINED

Australia: Queensland: Kuranda. I ..S km SE.
16-17. V. 1980, I. D. Naumann and J. C. Cardalc, collected

at light (19, ANIC); Kuranda, 2.8 km N on Black
Mountain Rd, 23.iv.90, J. Heraty, ex cocoon of
Hypoponera sp. (1 9 pupa, JMH).

Neolosbanus wusheanus sp. nov.

Figs. 152, 154-157, 160

TYPE MATERIAL

Holotype, 9, "Wushe, Taiwan/ 1150 m IV-2-83/ Henry
Townes." "HOLOTYPE/ Neolosbanus/ wusheanus
Heraty." Deposited in AEI.

Paratypes: Taiwan: Nantou Hsien: same data as holotype
(Ic?, AEI); Tungpu, 1200 m, 16-20.iv.l984, K. C. Chou
and C. H. Yung (19, TARI).

DIAGNOSIS

Distinguished from N. gemma by the following: frenum
glabrous and extended in dorsal view (Figs. 152, 155),
hind tibia and trochanter yellowish brown, hind femur
black with greenish reflections, LOL/OOL ratio small,
and flagellar segments closely appressed (Fig. 152). The
antennal flagellum of the male is distinct from N.
apoanus.

FEMALE

Length, 2.2-2.4 mm. Head, mesosoma, and petiole dark
metallic green; gaster black with green reflections; anten-
nal flagellum dark brown to black; coxae and femora
black with green reflections; scape, trochanters, apex of
femora, and rest of legs yellow to light brown. Wings
hyaline, venation brown.

Head subcircular (Fig. 154); occiput weakly emar-
ginate; median ocellus separated from occiput by less
than own diameter; LOL 0.7x OOL. Face broadly round-
ed, polished, setae sparse and minute; occipital carina
weak and extending to lateral ocellus; temple narrow and
aciculate dorsally. Eyes large, separated by 1.6-1.7x their
height. Malar space 0.6x height of eye. Antenna 1 1 -seg-
mented; pedicel as long as broad; flagellum 0.9-1. Ox
height of head, funicular segments densely setose with
reticulate surface sculpture; F2 1.8x as long as broad, fol-
lowing segments subequal in length, only slightly wider
towards apex of flagellum; clava ovate, as long as preced-
ing 2 segments.

Mesosoma dorsally with scutellum and posterior half
of midlobe of mesoscutum areolate. midlobe anteriorly
and lateral lobe transversely carinate; axilla obliquely car-
inate (Fig. 155). Scutellum as long as broad: frenal line
narrow and shallow, frenal area crescent-shaped in dorsal
view, broadly rounded in profile and angled about 45
degrees to dorsum, glabrous. Propodeum smooth \\ iih
rugose-areolate median band: callus with several short
setae. Coxae glabrate; femora polished with moderately

99

dense short setae: hind tibia with 1 spur surrounded by
long apical fringe. Forewing 2.4-2.5x as long as broad
(Fig. IdO), twice as long as mesosoma; stigmal vein per-
pendicular to wing margin, twice as long as broad; poste-
rior marginal vein 2.6x as long as stigmal vein.

Mi'disoma with petiole 1.7-1.8x as long as hind coxa,
1.5-1.7X as long as propodeum; petiole ribbed dorsally
and carinate ventrally. Ms, with weak medial constriction
(as in Fig. 273). Ovipositor acicular (Fig. 152).

MALE

Length, 2.4 mm. Colour and sculpture as in female. Ocelli
larger, and wings slightly infuscate. Antenna 1 1 -segment-
ed; scape almost reaching median ocellus; anellus small;
flagellum 2. Ox height of head; F2 stout, 2.3x as long as
broad and as long as scape, following segments subequal
in length and width; clava cylindrical, twice length of pre-
ceding segment. Petiole 2.4x length of hind coxa, 2.4x
length of propodeum. Caster as long as petiole; Ms, with
weak medial constriction. Cenitalia typical for genus,
with strong median process, paramere elongate with long
terminal seta and 3 long setae basally; aedeagus narrow
and weakly sclerotized.

VARIATION

Single specimens from Thailand and Vietnam differ
slightly from the type material and are not included as
part of the species description. The female from Vietnam
has a distinctive reddish coloration, all tibiae are dark
brown with reddish reflections, and the petiole is strongly
ribbed. The female from Thailand is similar to the type
material but differs in the length of the petiole (2.3x
length of hind coxa) and enlarged eyes (malar space 0.4x
height of eye).

DISTRIBUTION

This species, as broadly defined, occupies an Indo-
Chinese distribution (U, Fig. 278).

OTHER MATERIAL EXAMINED

Thailand: Chingdao, 5-1 l.iv.l958, native collector (19,
BPBM). ViiiTNAM: Karyu Danar, 200 m, 1 3-28. ii. 1961,
N. R.Spencer (19, BPBM).

ETYMOLOGY

From Wushe, the type locality.

Neolosbanus apoanus sp. nov.

Figs. 158, 161

TYPE/ Neolosbanus/ apoanus Heraty." Deposited in

MCZ.

Paratypes:PHii.ipPiNEs: Mindanao: Ml Apo. Galog Riv..

6000' 11830 m|, 28. ix. C. F. Clagg (Id, MCZ); Mt Apo.

Tia Ridge. 65()0" [ 1980 m|. ix. C. F. Clagg (1 d, MCZ).

dia(;nosis

Recognized by: flagellar segments separated, elongate,
and slightly flared (Fig. 158). frenum smooth, and legs
honey yellow.

MALE

Length, 2.7-2.9 mm. Black with bluish green reflections,
strongest reflections on mesosoma dorsum; gaster black;
antennal flagellum black; coxae black with greenish
reflections; mid and hind femora brown medially; scape,
trochanters, base, and apex of mid and hind femora, all of
fore femur, tibiae, and tarsi light honey yellow. Wings
hyaline, venation brown.

Head subtriangular; occiput broadly rounded; median
ocellus separated from posterior margin by less than own
radius; LOL l.l-1.4x OOL. Face broadly rounded, pol-
ished; occipital carina sharp, extending almost to eye
margin; temple narrow and smooth. Eyes separated by
L4— L6x their height. Malar space 0.6-0.7x height of eye.
Antenna 11-segmented (Fig. 158); pedicel as long as
broad, only slightly broader than F2 at base; flagellum
2.4— 2. 5x height of head; funicular segments with dense,
semi-erect setae and scabriculous surface sculpture; F2
2.6x as long as broad, slightly longer than scape, follow-
ing segments subequal in length, only slightly decreasing
in width, each funicular segment elongate, cylindrical,
and widely separated between segments; clava cylindri-
cal, 1.5x as long as preceding segment.

Mesosoma with dorsum rugose-areolate; lateral lobe of
mesoscutum transversely carinate; axilla obliquely cari-
nate. Scutellum 1.3x as long as broad, broadly rounded at
apex; frenal line narrowly impressed, frenal area
glabrous, crescentic in dorsal view, angled 70 to 80
degrees to dorsum. Propodeum smooth with rugose-areo-
late median band; callus with several short setae. Coxae
glabrate; hind femur polished with sparse short setae;
hind tibia with 2 spurs. Forewing 2.1-2.3x as long as
broad, 2.2x as long as mesothorax; stigmal vein angled
distally, 1.5x as long as broad (Fig. 161); postmarginal
vein 3x as long as stigmal vein.

Metasonia with petiole 2.2-2.3x as long as hind coxa,
2.1-2.7X as long as propodeum; petiole carinate to ribbed
dorsally, finely carinate to aciculate ventrally. Gaster as
long as petiole. Genitalia not visible in available material.

TYPE MATERIAL FEMALE

Holotype, 6, "Galog Riv./ 6,000 ft. [1830 m] Sept. 12." Unknown.

"Mt Apo, Mindanao/ Phil. Islds., C. S. Clagg." "HOLO-

100

DISTRIBUTION

Known only from a single collection from the Philippines
(A, Fig. 278).

ETYMOLOGY

From Mount Apo, the type locality.

Neolosbanus purpureoventris-group

This group of 3 species is recognized by: ovipositor acic-
ular, scrobes smooth, and stigmal vein sharply angled.
These species are common and widespread in the Indo-
Pacific region including northern Australia.

GROUP DESCRIPTION

Head 0.3-0.4x as long as wide; ocelli arranged in tri-
angle, median ocellus meeting line drawn across anterior
margin of lateral ocelli. Face smooth with scattered short
setae; scrobal depression shallow and completely smooth,
partially including median ocellus; occiput glabrous.
Clypeal margin lobate and bare, strongly produced over
base of mouthparts, epistomal sulcus weak or absent.
Malar depression linear and strongly impressed along
entire length. Antenna 11 -segmented in females, 13-seg-
mented in males; scape narrow and cylindrical in both
sexes, pores present on ventral surface of male scape;
anellus present; funicle 7-segmented in females, 9-seg-
mented in males; both sexes with MPS.

Mesosoma with dorsum scabrous to rugose-areolate
dorsally; frenal line narrow and foveate. frenal area
glabrous or carinate and angled 80 to 90 degrees to dor-
sum; axillula glabrous, axillular sulcus present or absent.
Mesepimeron mostly smooth; femoral groove and
transepimeral sulcus foveate; mesepisternum without
wedge-shaped sternaular sculpture, at most with single
fovea anteriorly. Prepectus foveate and fingerlike dorsal-
ly, reduced to a narrow ridge in ventral half. Hind tibia
with 2 apical spurs. Forewing 2.3-2.5x as long as broad
(Fig. 169); speculum and basal area bare; stigmal vein at
least 3x as long as broad, angled approximately 45
degrees to anterior margin of forewing; posimarginal vein
more than 3x as long as stigmal vein.

Caster with basal terga glabrous. Ovipositor needle-
like; first valvula smooth; second valvula with minute
dorsal ridges connecting medially. Gonostylus elongate
and narrow, sometimes exceeding cercus by more than its
width. Ms^ of male narrow and rounded apically.
Genitalia of male with paramcre narrow and elongate,
bearing 2 stout setae; aedeagus subacute.

Neolosbanus storeyi sp. nov.

Fig. 179

TYPE MATERIAL

Holotype, 9 , ''NEW GUINEA: NE./ Nondugl. 2200-/
2700 m. V-28-1959." "C. D. Michener/ Collector/ BISH-
OP." "HOLOTYPE/ Neolosbanus/ storeyi Heraty."
Deposited in BPBM.

Paratypes: Australia: Queensland: Cape York, Lankelly
Ck, Mcllwraith Range, vi.l932, Darlington (1$, MCZ);
same data, 28. vi. 1932 (Id, MCZ). Indonesia: Seram
[Amboina]: F. Muir (6(5(5, MCZ). Papua New Guinea:
Port Moresby, 20 km SE, 26.1.1985, J. Ismay (19,
BMNH); Riv. Tor (mouth), 4 km E of Hoi Maffen,
4.vii.l959, T. C. Maa (19, BPBM); Bulolo. 900 m.
13.ii-13.iii.l979, J. Sedlacek (15, AEI); Baiyer Riv.,
1100 m, 25.i-6.ii.1979, J. Sedlacek (3d 5. AEI); Lae.
0-100 m, ix.l968. N.L.H. Krauss (15. BPBM); Brown
Riv., 30. ix. 1959, T. C. Maa (15, BPBM); Eliptamin
Valley, 1200-1350 m, l-15.vii. 1959, W. W. Brandt (1 5,
BPBM); Bougainville (S.): Kokures 690 m, 16.vi.l956,
E.J. Ford, Jr. (Id, BPBM).

DIAGNOSIS

Recognized by the following: ocellar triangle slightly
raised above surface of the vertex and anterior margin
with a shallow curving sulcus (Fig. 179), ocellar-ocular
sulcus shallow, mesosomal dorsum scabrous-alveolate,
frenal area carinate, and propodeum with a deep areolate
median channel.

FEMALE

Length, 2.2-3.2 mm. Head and mesosoma black with
blue or purplish-blue retlections, gaster and femora dark
brown to black with reddish reflections; pedicel and Ha-
gellum dark brown; tibiae brown medially; scape, apices
of tibiae, and tarsi honey yellow. Wings slightly infuscate
or hyaline, venation pale brown.

Head subcircular; occiput v\cakl\ emarginale; LOL
1.0-1.4X OOL. Face broadly rounded, polished with
sparse setae; ocellar triangle raised and bordered by shal-
low sulcus (Fig. 179); occipital carina weak, extending to
lateral ocellus. Eyes separatctl by 1.2-1.6x their height.
Malar space 0.4-0.8x height of eye. Anteclypeus lobate.
lateral margin of anteclypeus sinuate, extending well
below ventral marcin of siena. Antenna I l-secmenled:

flagollum ().9-I.2x height of head; funicular segments
with short, dense, semi-erect setae and numerous MPS;
F2 2. 0-3. Ox as long as broad; clava acuminate.

Mesosoma dorsal ly with moderately spaced scabrous-
areolate sculpture; posterior iialf of axilla and SSS longi-
tudinally carinate. Scutellum as long as broad; Irenal area
vertically carinate; axillular sulcus not apparent.
Propodeum with median band of deep areolate sculpture.
Coxae and femora glabrate, hind femur with dense, fine
setae in apical half. Forewing 1.9-2.9x longer than broad.

Metasoma with petiole 1.6-2.5x as long as hind coxa.
1.5-2.1X as long as propodeum; petiole subtriangular.
almost completely smooth dorsally, and carinate to ribbed
sublaterally. Ms, smooth, otherwise gaster typical for
species-group. Gonostylus not extending beyond apex of
metasoma.

MALE

Length, 2.7-2.9 mm. Coloration as female. Antenna 13-
segmented; pores present on ventral surface of scape; fla-
gellum 2.0-2. 3x height of head; F2 3.1-4.0x as long as
broad, 0.9-1. Ix as long as scape. Petiole 2.3-2.6x as long
as hind coxa, 1.9-3.3x as long as propodeum.

DISTRIBUTION

Papuan subregion (S, Fig. 278). This species occurs in
strict sympatry with A^. palgravei in Australia (Cape York
Peninsula), and Papua New Guinea (Bulolo, Baiyer River).

ETYMOLOGY

Named in honour of Ross Storey from the Department of
Plant Industries in Mareeba, Queensland — a truly out-
standing individual.

Neolosbanus watanabei sp. nov.

Loshanus sp.— Watanabe, 1958:28. Hedqvist, 1978:230
(list and key).

TYPE MATERIAL

Holotype, 9, "PHILIPPINES, Negros/ Oriental: Cuemos
de/ Negros 7 km W Valencia/ 700 m. 7-13 JUN 1987/
ROM 873057. DC Darling." "HOLOTYPE/
Neolosbanu.s/ watanabei Heraty." Deposited in ROM.
Paratypes: Caroline Islands: Palau Islands, Koror
Island, 10.ii.l948, H. S. Dykas, limestone ridge (Id,
BPBM). Philippines: Luzon: Mountain Prov., Ifuagao,
Mayoyao, 1200-1500 m, 3.ix.l966, H. M. Torrevillas
(19, BPBM); same locality. 1000-1500 m, 6.vii.l966,
H. M. Torrevillas. light trap (19. BPBM); Negros
Oriental: same data as holotype (Id, ROM); L.
Balinsasayao. 1-7. x. 1959. L. Quate and C. M.
Yoshimoto, bait traps, chicken intestines, (Id, BPBM);
Basilan: Island of Basilan, Baker (Id, MCZ).

I)IA(;n()sis

Recognized by the following: margins of ocellar triangle
smooth and not separated from the vertex, mesosomal
dorsum broadly spaced rugose-alveolate, frenal area
glabrous, and propodeum with a shallow areolate median
channel.

FEMALE

Length, 2.5-3.3 mm. Head and mesosoma dark metallic
green; metasoma, coxae, and femora dark brown to black
with faint reddish reflections; flagellum dark brown,
becoming darker apically; scape and rest of legs honey
yellow. Wings hyaline, venation pale brown.

Head subcircular; occiput weakly emarginate; LOL
1.0-1.2X OOL. Face broadly rounded, polished with
sparse setae; occipital carina weak, extending to lateral
ocellus. Eyes separated by 1.2-1.3x their height. Malar
space 0.6x height of eye. Anteclypeus broadly rounded,
lateral margin of anteclypeus evenly rounded; extending
well beyond ventral margin of gena. Antenna 1 1 -seg-
mented; flagellum 1.2-1.3x height of head; funicular seg-
ments with short, dense, semi-erect setae and numerous
MPS; F2 2.4-2.8X as long as broad; clava acuminate or
rounded.

Mesosoma with dorsum, including lateral lobe of
mesoscutum, with broadly spaced rugose-areolate sculp-
ture; posterior half of axilla and SSS longitudinally cari-
nate. Scutellum as long as broad; frenal area glabrous;
axillular sulcus weakly carinate. Propodeum with median
band of shallow areolate sculpture. Coxae and femora
glabrate, hind femur with dense, fine setae in apical half.
Forewing 2.3-2.4x longer than broad.

Metasoma with petiole 1.7-1.8x as long as hind coxa,
1.5-1.8X as long as propodeum; petiole subtriangular in
cross-section, weakly carinate to smooth dorsally and
sublaterally, weakly ribbed on lateral and ventral comers.
Ms, smooth, otherwise gaster typical for species-group.
Gonostylus sometimes extending beyond apex of metaso-
ma by more than its width (not exceeding width in holo-
type).

MALE

Length, 2.3-2.6 mm. Colour as in female. Eyes separated
by 1.3-1.4X their height. Antenna I3-segmented;
unknown if pores on ventral surface of scape. Flagellum
1.9-2.2X height of head; F2 2.9-4.4x as long as broad.
0.9-1. Ox as long as scape. Forewing 2.3-2.4x as long as
broad. Petiole 2.2-2.5x as long as hind coxa, 2.4-2. 8x as
long as propodeum.

DISTRIBUTION

Philippine subregion and Caroline Islands (W. Fig. 278).

102

ETYMOLOGY

Named for Dr. Watanabe for his works on Micronesian
eucharitids.

Neolosbanus purpureoventris (Cameron) comb. nov.

Figs. 166-169, 209-210, 234, 246, 255, 271-272

Eucharis purpureoventris Cameron, 1909:232. Kuching,

Borneo [BMNH, examined].
Orasema purpureoventris — Boucek, 1988:520.

TYPE MATERIAL

Lectotype (designated by Boucek, 1988), 9, "LECTO-/
TYPE." "Kuching/ Hewitt." "Hy. 6." "P. Cameron Coll./
1914-110." "Eucharis/ purpureoventris/ Cam. Type/
Borneo." "B.M. TYPE/ HYM./ 5.367." "c? Gollumiella/
purpureoventris/ (Cameron)/ see Hedqvist, 1978/ det. Z.
Boucek, 1980." "LECTOTYPE/ 6 Orasema/ purpure-
oventris/ (Cam.)/ det. Z. Boucek, 1986." Point-mounted
male with no head. Housed in BMNH.

mented (Fig. 166); flagellum l.l-1.2x height of head;
funicular segments with short, dense, semi-erect setae and
numerous MPS; F2 2.5-2.7x as long as broad, about 0.5x
length of scape; clava acuminate.

Mesosoma with dorsum, including lateral lobe of
mesoscutum, closely spaced rugose-areolate to scabrous
(Fig. 246); posterior half of axilla longitudinally carinate.
Scutellum as long as broad; frenal area with weak, widely
spaced transverse carinae; axillular sulcus weakly cari-
nate. Propodeum completely smooth or with median band
of foveate sculpture (variable within localities; Fig. 255).
Coxae and femora glabrate, hind femur with dense, fine
setae in apical half. Forewing 2.4-2. 5x as long as broad
(Fig. 169).

Metasoma with petiole 1.5-1.8x as long as hind coxa.
1.4-1.8X as long as propodeum; petiole subtriangular.
smooth dorsally and sublaterally. weakly carinate on lat-
eral and ventral comers. Caster typical of species-group;
MSt smooth. Gonostylus exceeding apex of gaster in
specimens from Malaya and Vietnam, otherwise with-
drawn or shorter than apex of gaster.

NOTES ON SYNONYMY

This species was placed in the genus Orasema by Boucek
(1988) as a senior synonym of Psilogaster laeviceps
Gahan^ Psilogaster nishidai Ishii and Nagasawa, and
Loshanus petersoni Hedqvist. It is distinct from these
other species based on the features outlined in the group
diagnosis. Psilogaster nishidai and L. petersoni are trans-
ferred here to Neolosbanus palgravei as junior synonyms,
and P. laeviceps is given species status in Neolosbanus.

DIAGNOSIS

Recognized by the following: margins of ocellar triangle
smooth and not separated from vertex, dorsal sculpture of
mesosoma closely spaced rugose-alveolate or scabrous,
frenal area carinate, and propodeum completely smooth
or with areolate median channel.

FEMALE

Length, 2.2-^.0 mm. Head black, mesosoma black with
bluish reflections, gaster dark brown to black; coxae and
femora dark brown to black and sometimes with metallic
reflections; antenna testaceous to black; hind tibia honey
yellow to dark brown, tarsi yellowish brown. Wings hya-
line, venation pale brown.

Head subcircular to subtriangular (Fig. 209); occiput
weakly emarginate; LOL l.0-l.2x OOL. Face broadly
rounded, polished with sparse setae; occipital carina
weak, extending halfway between ocellus and eye mar-
gin. Eyes large, separated by 1.2-1.3x their height. Malar
space 0.5-0.8X height of eye. Anteclypeus lobate, lateral
margin of anteclypeus sinuate, extending well below ven-
tral margin of gena (Figs. 209-210). Antenna 1 l-seg-

MALE

Length. 1.9-2.9 mm. Colour as in female. Antenna 13-
segmented (Fig. 168); scape with patch of minute pores
on the ventral surface (Fig. 167); flagellum 1.7-2.3x
height of head; F2 4.3x as long as broad. 0.8-1.2x as long
as scape. Petiole 2.1-2.5x as long as hind coxa, 2.6-2.9x
as long as propodeum. Ms^ sometimes with weak indenta-
tion along line of constriction (as in Fig. 273).

VARIATION

There is considerable variation over the range of this
species due to the amalgamation of 2 or 3 distinct forms.
Extremes of variation are found in single collections from
a broad range of localities. Specimens from Borneo.
Malaya (Malaysia), and Larat conform to the lectotype in
coloration (head and mesosoma dark blue to blue-green,
hind tibia honey yellow), sculpture of the mesosomal dor-
sum (scabrous), and propodeum (strong areolate medial
band). However, all females from Malaya have the
gonostylus exceeding the apex of the gaster. This last fea-
ture occurs sporadically in different collections, and may
be an artifact of preservation. Specimens from southern
Thailand, Vietnam, and Taiwan form a group with similar
dorsal sculpture, but differ in having the head and meso-
soma almost entirely black, hind tibia usually dark brown,
and propodeum completely smooth or with a faint median
carina. A northern group from Nepal, north Thailand.
Laos, and Fukien province (China) include larger individ-
uals that have the head and mesosoma dark green, hind
tibia yellow or dark brown (locality dependent), mesoso-
mal dorsum with closely packed areolate sculpture, and
propodeum with a narrow band of weak areolate sculp-

103

lure. Differences are difficult to quantify and I believe
that onis a single species should be recognized here.

BI()L()(;V

The host-ant and larval stages are unknown. The egg. as
dissected Ironi the abdomen, is stalked, and similar to
those of other Eucharitidae. I collected adults from broad-
leaf undersiory plants in rainforests of Taiwan and
Thailand, but could not determine the host plant. The
overall elevation of the collections ranges from 300 m in
Thailand to 2000 m in Nepal.

DISTRIBUTION

Widespread in the Indo-Chinese, Malayan, and Sulawesi
(Larat) subregions (P, Fig. 278). This species occurs in
strict sympatry (same locality and collection date) with A'.
palgravei (Taiwan: Tungpu; Vietnam: Da Lat), N. laevi-
ceps (Taiwan: Sun Moon Lake, Tungpu), A^. pllosus
(Vietnam: Da Lat), A', townesi (Papua New Guinea:
Bulolo). and A', nepalensis in Nepal.

MA TKRIAI. KXAMINKI)

Indonesia: Tanimbar: Larat, xii (Id. MCZ). Laos:
Vientianne Prov.: Ban Van Hue, April (19. BPBM).
Malaysia: Panang. Cameron Highlands, Tanah Pata,
1700 m, August (Id. BMNH). Ni;pai.: Godavari. 2000 m,
July (19, CNC). Peoples Republic of China: Hainan Is.,
Tien Fong Mtns, May; Fukien: Shuipeikieh. Shaowu;
Kwatun. Chungan: Tachulan, Shaowu: July to September
(13 9 9, 4dd, BMNH, TARl). Taiwan: Nantou Hsien:
Sun Moon Lake, W of Techuache, 900 m, edge of rain-
forest; Tungpu, 1200 m; March to April. June to July,
September to October (29 9. 20d d, JMH. ROM,
TARI). Thailand: Chiangmai Prov.: Doi Suthep,
500-900 m; Wat Suthep; November (79 9, 1 d, BPBM);
Suphanburi Prov.: Khao Yai N. P., elephant-crossing
trail, 300 m, rainforest undergrowth; Khao Yai N. P,
Haew Sawat waterfall, 300 m, rainforest undergrowth;
July ( 1 9 , 8 d d , JMH, ROM). Vietnam: 20 km S Da Lat,
1300 m; Mt Lang. Bian, 1500-2000 m; Fyan, 1200 m; Ap
Hung-Lam, 21 km NW of Di Linh, 1 100 m; May to
October (3 9 9 , 26 d d , BPBM).

Neolosbanus palgravei-group

This group of 6 species is recognized by: face smooth,
clypeal margin well developed, and ovipositor broad with
lateral ridges on the second valvula. Species are common
and widespread in the Indo-Pacific region including
northern Australia. Females deposit eggs into punctures
in the leaves of a wide variety of broad-leaf plants.

GROUP DESCRIPTION

Head 0.2-0.4x as long as wide; ocelli arranged in line,
median ocellus close to occipital margin and exceeding
line drawn across anterior margin of lateral ocelli (Fig.
178); face glabrous; scrobal depression glabrous and shal-
lowly or deeply impressed; occiput glabrous. Clypeal
margin strongly produced as rounded lobe over mouth-
parts (rounded in A', townesi), usually with well-defined
epistomal sulcus. Malar depression absent. Antenna 10-
or 1 1 -segmented, scape cylindrical or expanded apically,
scape of male with pores on ventral surface below pedi-
cel; anellus present or absent; funicle 7- or 8-segmented
in females, 8-segmented in males; both sexes with MPS.

Mesosoma with dorsum rugose-areolate to smooth;
frenal area glabrous, angled 60 to 90 degrees to dorsum of
mesosoma; axillula and axillular sulcus diagnostic.
Mesepimeron glabrale, transepimeral sulcus shallow
foveate (absent in A', townesi); femoral groove foveate or
absent, continuing ventrally between fore and mid coxae
to midline, glabrous between groove and base of coxa
(Figs. 227-231). Prepectus foveate (smooth in A'.

townesi) and narrowly triangular dorsally. reduced to nar-
row ridge in ventral half. Forewing 2.2-2.6x as long as
broad; basal pilosity diagnostic; stigmal vein of forewing
usually narrow and elongate, more than 1.5x as long as
broad; postmarginal vein 1 .5-2.0x as long as stigmal vein.
Gaster with basal terga setose or glabrous. Ovipositor
subapically expanded and straight or only slightly curved;
first valvula with 2 or 3 oblique ridges apically; second
valvula narrow with several strong transverse ridges,
ridges connecting medially. Gonostylus broad, gradually
narrowing to apex, not exceeding cercus. MSg of male
narrow or broad. Genitalia of male with paramere narrow
and elongate, bearing 2 or 3 stout setae; aedeagus suba-
cute to truncate apically.

Neolosbanus townesi sp. nov.

Figs. 170-175. 199, 226, 241, 250

TYPE MATERIAL

Holotype. 9, "Kassam Pass/ 1300 m. N. Guinea/
1.10-23.1979/ J. Sedlacek." "HOLOTYPE/ Neolosbanus/
townesi Heraty." Deposited in AEI.
Paratypes: Papua New Guinea: Bulolo. 900 m,
1 3.ii-l 3.iii. 1 979. J. Sedlacek ( 14d d , AEI); Mt Suckling,
Mau 1, 300-1000 m, 13.vii.l979, J. L. Gressitt. on
Castanopsis (19. BPBM); Morobe Dist.. Wau. 1400 m.
20. xii. 1961, L. W. Quate (19, BPBM); Wau, Mt

104

Mission, 1800 m, 22.iii.1966, J. L. Gressitt, on bamboo
(1 9, slide no. 88-23-19, BPBM); Tapo (= Tapu), 3 km W
of Kainanm, 1 650 m, 22.x. 1959, T. C. Maa (19, BPBM);
Minj, W Highlands, 8-13. ix. 1959. T. C. Maa (19,
BPBM); New Ireland: Ridge above "Camp Bishop," 15
km up Kait Riv., 250-750 m, 14.vii.l956, J. L. Gressitt
(1(5, BPBM).

DIAGNOSIS

Differs from other Neoloshanus by the following: anellus
absent, prepectus smooth and evenly rounded (Fig. 226),
mesosoma with dorsum almost completely smooth (Fig.
241), frenal line only faintly indicated, and Ms., of female
with a transverse band of fine rugulose sculpture.

FEMALE

Length, 2.2-2.7 mm. Dark brown to black; scape and
pedicel light brown; extreme apex of femora, tibiae, and
tarsi white to pale yellowish brown. Wings lightly infus-
cate, venation brown.

Head subtriangular (Fig. 199); occiput broadly and
deeply emarginate; ocelli large, median ocellus separated
from occipital margin by more than its own radius; LOL
1.1-1.4X OOL. Face broadly rounded, cheek sunken, pos-
terior margin of gena abrupt and finely punctate; scrobal
depression shallow and smooth; occiput glabrous, occipi-
tal carina extending just beyond lateral ocellus. Eyes sep-
arated by 1.8-2. Ix their height. Malar space 0.8-1. Ox
height of eye. Clypeus broadly rounded, slightly extend-
ing over base of labrum. Antennae 11 -segmented (Figs.
170-171); pedicel short, slightly broader than F2; scape
reaching median ocellus, expanded apically and broadly
excavated on ventral surface (Fig. 199); anellus absent;
flagellum 1.6-1.9x height of head; funicle 8-segmented,
segments with MPS sparse and small; F2 4.2-5.7x as
long as broad, 1.4-1.7x F3, following segments about
equal in length and width.

Mesosoma with dorsum glabrous, except midlobe of
mesoscutum with weak transverse sculpture; lateral lobe
smooth and swollen (Fig. 241); axilla smooth and round-
ed. SSS deeply impressed and carinate. Scutellum as long
as broad, apex broadly rounded in dorsal view; frenal area
glabrous; axillula glabrous, axillular sulcus absent.
Propodeum completely smooth (Fig. 250); callus with
patch of several short setae. Femoral groove absent.
Prepectus smooth and swollen medially (Fig. 226). Coxae
elongate, hind femur moderately setose, tibiae and tarsi
densely setose. Forewing (Fig. 175).

Metasoma with petiole 1.3-2. Ox as long as hind coxae,
1.6-2. Ox as long as propodeum; petiole glabrous with few
weak longitudinal carinae. Basal mctasomal tergites
glabrous; Ms^ weakly sculptured basally (Fig. 172).
Ovipositor (Fig. 174). diagonal ridges of second valvula
strong and converging medially.

MALE

Length, 2.2-2.6 mm. Colour as in female. Antenna longer
and stouter than in female; scape more strongly expanded.
Petiole 2.0-2.3X as long as hind coxae, 2.5-3.2x as long
as propodeum. Ms^ twice as broad as petiole, apex broad-
ly emarginate. Genitalia enlarged and prominent; aedea-
gus broad and emarginate apically (Fig. 173).

BIOLOGY

Collected on Castanopsis (Fagaceae) and bamboo
(Poaceae).

DISTRIBUTION

New Guinea including New Ireland (T, Fig. 279).

ETYMOLOGY

Named in memory of Dr. Henry Townes.

Neolosbanus laeviceps (Gahan) comb. nov.

Figs. 183-184, 200, 227, 242, 251, 268-269

Psilogaster laeviceps Gahan. 1940:429-430. Peradeniya.

Sri Lanka [USNM, examined].
Parapsilogaster laeviceps — Ghesquiere, 1946:368

(replacement name).
Losbaniis laeviceps — Watanabe, 1958:26; Hedqvist.

1978:230 (Hst and key).
Orasema purpureoventris—sensu Boucek. 1988. in part.

TYPE MATERIAL

Holotype, 9. "Peradinyia/ Ceylon II-'30." "CP Clausen/
Coll." "Clausen no./ 2435." "6336/ 4 m." "Type No./
53550/ USNM." "Parapsilogaster/ laeviceps/ Gahan/ 9
Type" (USNM).

Paratypes: Sri Lanka [Ceylon]: Peradeniya, ii.l930. C. P.
Clausen 2435 (299, no. 53550, USNM); Peradeniya,
25.vii.1913 (1 9, no. 53550, USNM). Additional slide.
"Parapsilogaster/ laeviceps Gah/ Max. palpus and
Antenna from Clausen No 2435." "Para 53550/ Type No.
USNM [red label)."

NOTES ON SYNONYMY

Boucek (1988) incorrectly treated this species as a junior
synonym of A'. (= Orasema) purpnreoventris. The
enlarged ovipositor, expanded scape, and consistently
reduced pattern of sculpture belonging to N. laeviceps
clearly distinguish this species from N. purpnreoventris.

DIAGNOSIS

Recognized by the following: anellus absent (Figs.
183-184), mcsosomal dorsum weakly sculptured, scape
of male expanded apically and grooved ventrally to
receive the pedicel (Fig. 184), and scutellum rounded in

105

lateral profile. Distinuuishod (Voiu N. lowncsi by: margin
ol the clypeus lobaic and cxicnding vcntrally over the
labriim (Fig. 200), funicle of female 7-segmented (versus
8). and prepeetus foveate. The female was thoroughly
described b\ Gahan (1940). The male was previously
unknown and is described here in detail.

M.ALE

Length, 2.2 mm. Dark, brown to black; antenna,
trochanters, and legs beyond apex of femora yellowish
brown. Wings hyaline, venation pale brown.

Head subtriangular. occiput broadly emarginate; medi-
an ocellus separated from occipital margin by less than
own diameter; LOL 0.9x OOL. Face rounded and
glabrale; scrobal depression narrow and strongly
impressed, lateral margin indistinct; occiput glabrous;
occipital carina weak, not reaching dorsal eye margin.
Eyes separated by 2.0x height of eye. Malar space l.Ox
height of eye. Clypeus extended as strong lobe ventrally
over labrum, epistomal sulcus present, tentorial pit deep.
Antenna 11 -segmented; pedicel short, slightly broader
than F2; scape strongly expanded apically, inner margin
strongly produced, ventrally excavated to receive pedicel,
with minute pores on ventral surface below pedicel (Fig.
184); anellus absent; flagellum 1.6x height of head; funic-
ular segments with MPS sparse and small; F2 2.5x as
long as broad. 1.7x F3; following segments subequal in
length and width, base of each segment narrower than
apex; clava 1 -segmented.

Mesosoma with dorsum lightly rugose (Fig. 242); lat-
eral lobe of mesoscutum and axilla smooth and very light-
ly sculptured. SSS carinate. Scutellum longer than broad,
apex narrowly rounded in dorsal view; frenal area
glabrous; axillula weakly carinate. dorsal margin obscure.
Propodeum almost completely glabrous; callus with few.
small setae dorsally (Fig. 251). Femoral groove narrow
and weakly foveate. Prepeetus triangular and foveate
above, strongly narrowed ventrally. Pronotal sulcus sharp
and smooth. Coxae and femora glabrate. hind femora
slightly expanded medially, hind tibia gradually expanded
to apex and slightly flattened. Forewing (Fig. 162).

Mctasoma with petiole 2.2x as long as hind coxa. 2.5x
as long as propodeum; petiole carinate dorsally. Basal
metasomal tergites glabrous. MSj^ broad and subtruncate
at apex. Genitalia large and broad, paramere short and
bearing 3 stout setae; aedeagus broad and subtruncate
(Fig. 163).

2-segmented in I specimen from Ceylon). Mesoscutum
and axilla rugose to weakly carinate; propodeum smooth
or weakly sculptured medially. Petiole weakly or strongly
carinate. Ms, glabrous. Ovipositor with diagonal ridges of
first valvula reduced or absent.

biol()(;y

Clausen (194()b. 1940c. 1941) discussed biology and the
morphology of the first-instar larva of A', laeviceps (see
earlier discussion in biology section). Adults are known
to deposit single eggs into leaves of Artocarpus
(Moraceae) (Clausen, 1940b).

DISTRIBUTION

Broadly distributed in the Indo-Chinese and mainland
Malayan subregions (L. Fig. 279). The most northern
record is from Tsushima Island, which lies between
Korea and Japan and would be included in the Palaearctic
realm according to Gressitt (1956) and Schuh and
Stonedahl (1986). This species is broadly sympatric with
N. pilosus and N. taiwanensis and has been collected in
strict sympatry with Neoloshanus purpureoveiitris in
Taiwan (Sun Moon Lake. Tungpu).

MATERIAL EXAMINED

India: Tamil Nadu: Anamalai Hills. Cinchona. 1150 m;
Kerala: Periyar A. San.; Thadripukudisai; September to
October (149 9, BMNH. CNC. lARl). Japan; Honshu:
Aichi Prefecture, Yakusa, Toyota, PT, July to August
(29 9, CNC); South Ryukyu Islands: Ishigakishima Is.,
Omoto-dake, 100-250 m; Tsushima Island, Izuhara-
Ariakeyama; May and September (299, BPBM. KUEC).
Laos: Namgueng Riv.. Phou Kan Khouei. 300-600 m,
Sayaboury Prov.: Sayaboury; Vientianne Prov.: Ban Van
Hue, February, April, and December (3 9, BPBM). Sri
Lanka: Kandy, 10 km W; Katugastota, 550 m;
Nugegoda; Udugama, Kanneleya Jungle, at light;
February, May. August (6). and November (109 9,
BPBM. BMNH). Taiwan: Yangminshan. Site 2. 1991. H.
Y. Yong. YPT (299. CNC); Nantou Hsien: Sun Moon
Lake: Tianmu. Tungpu. 1200 m; Taipei: Keelung;
Paomenszu, 2 km S Keelung City; Taipei; June to July,
October to November (1369 9, TARl). Vietnam: Pleiku,
20 km N, 650 m. May (19, BPBM).

Neoloshanus taiwanensis sp. nov.

FEMALE

Length, 1.9-2.1 mm. Colour as in male. Tentorial pits
shallow. Antenna lO-segmented and more slender; scape
slightly expanded with inner margin produced as apical
flange; funicle 7-segmcnted; F2 3.0-4.3x as long as
broad, 1. 3-1. 8x F3; clava usually I -segmented (distinctly

TYPE MATERIAL

Holotype, d, "N. TAIWAN: Wufeng/ 400 m., Hsinchu
Hsien/ 14-16. Vll. 1982/ K. C. Chou and C. C. Pan."
"HOLOTYPE/ Neoloshanus/ taiwanensis Heraty."
Deposited in TARI.
Paratypes: Taiwan: Hsinchu Hsien: same data as holo-

106

type (8c? d, TARI); Chienhsi, 5.xi.I981. K. S. Lin (Ic?,
TARI); [in Chinese characters], 9.xii.l961, S. C. Chiu
(1(5, TARI). People's Republic of China: Hunan:
Changteh, Yangshan, 1 1 .x. 1 938, T. C. Maa (\6, TARI).

DIAGNOSIS

Distinguished from N. pilosus by the following: eye with
sparse setae, body moderately pilose with elongate setae,
propodeum with a median carina, petiole of male robust
and as broad as the hind coxa (2.2-2.9x as long as hind
coxa), and hind tibia dark brown.

MALE

Length, 2.5-3.0 mm. Body black; antennal flagellum, all
femora, and hind tibia dark brown to black; scape brown;
fore and mid tibiae and tarsi yellowish brown. Wings
hyaline, venation clear yellowish brown.

Head subtriangular to subcircular; occiput slightly
emarginate; head strongly narrowed medially, dorsal
length not much broader than median ocellus, median
ocellus separated from occiput by less than its own
radius; LOL 1.6-1.7x OOL. Face relatively flat, glabrate
with moderately dense, fine hairs; scrobal depression
broadly impressed; occiput glabrous, occipital carina
prominent, extending laterally to eye margin. Eyes sepa-
rated by 1.6-1.7X height of eye. Malar space 0.7-0.8x
height of eye. Clypeus extended as strong lobe ventrally
over labrum, epistomal sulcus weak. Antenna 12-seg-
mented; pedicel short, as wide as F2; scape narrow, not
reaching median ocellus; anellus present; flagellum
1.7-1.9X height of head; funicle 8-segmented, with few
MPS; F2 3.8x as long as broad; F2 1.2-1.4x F3, follow-
ing segments subequal in length; clava with 2 incomplete-
ly fused segments.

Mesosoma with dorsum rugose-areolate, with moder-
ately den.se covering of short, fine hairs; lateral lobe of
mesoscutum and anterior face of midlobe transversely
carinate. SSS longitudinally carinate. Scutellum 1.4x as
long as broad, apical margin narrowly rounded in dorsal
view; frenal area angled about 45 degrees to dorsum,
crescent-shaped in dorsal view; axillula glabrous to weak-
ly carinate. axillular sulcus weakly fovcatc. Propodeum
smooth with medial carina and sparse erect hairs; callus
with patch of dense hairs. Mesepimeron with few scat-
tered setae; femoral groove broad, foveate to weakly
crenulatc. Pronotal sulcus foveate. Coxae and femora
smooth with moderately dense setae. Forewing 2.3-2.4x
as long as broad; speculum and basal area bare; stigmal
vein more than twice as long as broad, and perpendicular
to wing margin.

Metasoma with petiole 2.2-2.9x as long as hind coxa,
2.2-2.7X as long as propodeum; petiole longitudinally
carinate with sparse erect hairs dcirsally. thick in cross-
section and as broad or broader than hind coxa. Mt, to

Mt^ with semi-erect elongate hairs. MSj^ narrow and
rounded. Genitalia moderately sized, paramere narrow
and elongate; aedeagus subacute.

FEMALE

Unknown, but based on N. pilosus, the sexes are not
expected to differ greatly.

DISTRIBUTION

Taiwan (I, Fig. 279).

ETYMOLOGY

From Taiwan; referring to the type locality.

Neolosbanus pilosus sp. nov.

Figs. 178, 180,202,229,243

TYPE MATERIAL

Holotype, 9, "VIET NAM: Fyan/ 900-1000 m./ 1 l.VII-
9. VIII. '61." "N. R. Spencer/ Collector." "Collection of
BISHOP MUSEUM." "HOLOTYPE/ Neolosbanus/ pilo-
sus Heraty." Right antenna broken. Deposited in BPBM.
Paratypes: Vietnam: same data as holotype (18$ $,
14(5(5, BPBM); Fyan, 1200 m, 1 l.vii-9.viii.l96l, N. R.
Spencer (79 9 , 10(5 5, BPBM); Da Lat, 6 km S.
1400-1500 m, 9.vi-7.vii.l96l. N. R. Spencer (39 9.
BPBM); Da Lat, 20 km S, 1300 m, 12. ix. 1960, J. L.
Gressitt (19. BPBM); Di Linh (Djiring), 1200 m,
22-28.iv.l960, L. W. Quate (59 9, BPBM); Di Linh, 28
km N, 22-28. iv. 1960, S. Quate (15, BPBM); Blao
(Balao), 500 m, 1 4-2 1.x. I960, C. M. Yoshimoto (19,
BPBM).

dia(;n()SIS

Distinguished from A^. taiwanensis by the following: body
with denser and shorter pilosity, eye with very minute
setae, propodeum completely smooth without median
carina, petiole longer and narrower than hind coxa, and
hind tibiae yellowish brown.

FEMALE

Length, 2.6-3.4 mm. Body dark brown to black, mesoso-
ma with faint greenish reflections; antenna, coxae, and
femora dark brown to black; tibiae, tarsi, and apex of
femora yellowish brown. Wings hyaline, venation clear
testaceous.

Head subtriangular; occiput deeply emarginate; head
strongly narrowed medially, dorsal length not much
longer than median ocellus (Fig. 178); median ocellus
separated from occipital margin by less than its own
radius; LOL 0.7-1. 1 x OOL. Face broadly rounded,
smooth with covering of dense, fine setae; scrobal depres-
sion broad and strongly impressed, including median

107

cKelliis: occiput glabratc: occipital carina extending to eye
margin but widely separated from it in dorsal view (Fig.
178). Eyes separated by 1.5-l.7x their height. Malar
space 0.7-0.9X height of eye. Clypeus extended as strong
lobe ventrally over labrum. without epistomal sulcus.
Antenna I I -segmented; pedicel slightly longer than
broad, as wide as F2; scape reaching median ocellus;
anellus present; flagellum l.3x height of head; funicle 7-
segmented. segments with few MPS; F2 4.2x as long as
broad. F2 l.2x F3, following segments subequal in
length; claval segments not discernible.

Mesosoma with midlobe of mesoscutum irregularly
transverse-carinate (Fig. 243); lateral lobe weakly trans-
verse-carinate; axilla carinate, carinae widely spaced;
scutellum with widely spaced and deep rugose-areolate to
scabrous sculpture; entire dorsum with dense, short setae.
Scutellum only slightly longer than broad, apex broadly
rounded in dorsal view; frenal area angled 70 to 80
degrees to dorsum, not visible in dorsal view; axillula
glabrous, axillular sulcus weakly carinate. Propodeum
glabrous and without median sculpture; callus and
metepimeron densely setose. Mesepimeron sparsely
setose posteriorly and ventrally; femoral groove broadly
foveate dorsally and narrowly foveate ventrally.
Prepectus and most of pronotum setose. Pronotal sulcus
smooth. Coxae smooth and densely setose; femora weak-
ly imbricate and densely setose. Forewing 2.3-2.6x as
long as broad; completely pilose on both surfaces (specu-
lum absent); stigmal vein perpendicular or slightly angled
to wing margin; postmarginal vein indistinct but longer
than stigmal vein.

Metasoma with petiole 1 .9-2.6x as long as hind coxa,
1.8-2.1X as long as propodeum; petiole subtriangular in
cross-section and carinate dorsally. smooth sublaterally
with 2 parallel carinae ventrally, and dorsal patch of
dense setae basally. Mt, to Mt^ sparsely setose. Genitalia
moderately sized, paramere narrow and elongate; aedea-
gus subacute.

DISTRIBITION

Vietnam (P. Fig. 279) and likely a similar species on Irian
Jaya.

ADDITIONAL SPECIES

A male from Irian Jaya (Vogelkop: Kebar Val. W. of
Mano Kwari, 550 m, 4-3 1 .i. 1962, Quate, BPBM) is simi-
lar to N. pilosus. but the propodeum is carinate and the
petiole has only 2 parallel carinae dorsally. This probably
represents another closely related species.

Neolosbanus nepalensis sp. nov.

Figs. 203-204, 230, 252

TYPE MATERIAL

Holotype, 9, "NEPAL, Ktmd./ Godavari. 5000* [1525
m]/ 12.viii. [handwritten] 1967/ Mai. Tr.. Can. Exp."
"HOLOTYPE/ Neolosbanus/ nepalensis Heraty."
Deposited in CNC.

Paratypes: Nepal: Kathmandu: Godavari, 1525 m,
23.vii.1967 (19, lOc? d), 25.vii.1967 (Id), 29.vii.1967
{26 6), 4-7. viii. 1967 (19), 8.viii.l967 {\6),
12.viii.l967 (569 9, 96 6), 15.viii.l967 (19, 26 6),
16.viii.i967 {\6), Can. Exp. MT (all CNC); Godavari,
1525 m. Royal Botanical Gardens. July, Can. Nepal Exp.
(29 9. Ic?, CNC); Godavari, 1830 m, l-3.viii.1967, MT,
Can. Exp. {\6, CNC); Godavari 7.2, 5.ix.l972 (49 9,
\6), 10.ix.l972 (79 9, \26 6), Mani and party (all
USNM); Godavari 7.4, 20-2 1. ix. 1972, Mani and party
(69 9,29dd,USNM).

DIAGNOSIS

Distinguished from A', palgravei by the following: stig-
mal vein angled proximally. callus with dense patch of
hairs (Fig. 252), antennal flagellum long in both sexes,
hind tibia of both sexes white, and mesosomal dorsum
with sparse setae (Fig. 230).

MALE

Length, 2.6-3.6 mm. Colour as in female, hind tibia dark-
er brown medially. Antenna 12-segmented (Fig. 180); fla-
gellum 1.7x height of head; funicle 8-segmented; F2 1.3x
F3. Forewings 2.3-2.4x as long as broad. Petiole
2.6-3.5X as long as hind coxa. 2.7-3.4x as long as
propodeum. Mt, to Mt, with semi-erect hairs. Genitalia
typical for group.

etymolo(;y

From Latin pilosus, meaning hairy; referring to the pilo.se
covering of the head and mesosoma.

FEMALE

Length, 2.4-2.9 mm. Mesosoma and petiole dark brown
to black with green reflections; head, gaster, antennal fla-
gellum, coxae, and femora dark brown; scape yellowish to
light brown, apex of femora, tibiae, and tarsi light yellow-
ish brown to white. Wings hyaline, venation pale brown.

Head subtriangular; occiput broadly emarginate; medi-
an ocellus separated from occiput by slightly less than its
own diameter; LOL 0.8-1. Ox OOL. Face broadly round-
ed, glabrate; scrobal depression narrow and weakly
impressed, including median ocellus; occipital carina
extending to dorsal margin of eye. Eyes separated by
1.6-2. Ox height of eye. Malar space 0.7-0.9x height of
eye. Clypeus extended as strong lobe ventrally over
labrum. epistomal sulcus absent. Antenna 1 1 -segmented;

108

pedicel short, slightly broader than F2; scape not reaching
median ocellus; anellus present; flagellum 1.5-1.7x
height of head; funicle 7-segmented, segments with
numerous MPS; F2 3.5-4.2x as long as broad, l.l-1.3x
F3, following segments subequal in length; clava with 1
or 2 segments incompletely fused.

Mesosoma with dorsum rugose-areolate and bare; lat-
eral lobe of mesoscutum and anterior face of midlobe
transversely carinate; posterior margin of axilla longitudi-
nally carinate. Scutellum as long as broad, apical margin
rounded in dorsal view; frenal area angled 90 degrees to
dorsum; axillula glabrous, axillular sulcus carinate.
Propodeum with irregular sculpture, sometimes strongly
or weakly developed as median furrow; callus with patch
of fine elongate hairs (more than 15; Fig. 252). Femoral
groove broad and shallowly foveate. Coxae and femora
glabrate. Forewing 2.3-2.5x as long as broad; basal area
and speculum bare; stigmal vein angled distally about 70
to 80 degrees to wing margin, rarely perpendicular.

Metasoma with petiole 1.9-2.4x as long as hind coxa,
1.7-2. Ox as long as propodeum; petiole longitudinally
carinate with weak ventral keel. Basal metasomal tergites
glabrous.

MALE

Length, 2.2-2.7 mm. Colour as in female. Antenna 12-
segmented; pedicel narrower than F2; anellus present; fla-
gellum 2.2-2.5X length of head; funicle 8-segmented; F2
only slightly longer than F3. Petiole 2.6-3.3x as long as
hind coxa, 2.7-3.4x as long as propodeum. Ms^^ narrow
and rounded. Genitalia moderately sized, paramere nar-
row and elongate; aedeagus subacute.

DISTRIBUTION

Nepal (N, Fig. 279).

ETYMOLOGY

From Nepal; referring to the type locality.

Neolosbanus palgravei (Girault) comb. nov.

Figs. 13-14. 17-25. 165. 181-182. 201. 205. 228. 231,
244, 253, 266-267

Orasema pali^ravci Girault. 1922:105-106. Australia:
Queensland |QMB. examined). Dahms, 1986:376
(notes on type material).

Psilogastcr nishidai Ishii and Nagasawa, 1941:292-294.
Caroline Islands: Elgi. Airai and Palau [not examined,
type not located). Synonymy with Orasenut piiipure-
oventris (Cameron) by Boucek, 1988:520. New syn-
onymy.

Loshanus nishidai — Walanabe, 1958:27-28; Hedqvist,
1978:230 (list and key).

Loshanus petersoni Hedqvist, 1978:229. Philippines:
Palawan, Tagembung [ZMC, examined]. Synonymy
with Orasema purpureoventris by Boucek. 1988:520.
New synonymy.

Orasema purpureoventris — sensu Boucek. 1988. in part.

Orasema indica Snehalatha and Narendran, 1992:355.
India: Kerala, Trichur [USNM, examined]. New syn-
onymy.

TYPE MATERIAL

Lectotype (here designated) of Orasema palgravei, intact
9 , no. T.9405 (number 3 from outer specimen), mounted
with 3 other 9 9 syntypes (no. T.9403-T.9404, T.9406)
on card labelled "Orasema palgravei Gir., c?/ 9 types
[GH].'" Published data: "Greenhills, Cairns, May 31. jun-
gle, many specimens." Additional type material: slide 4
(Dahms, 1986), "Type male, Orasema palgravei Gir."
[GH]. See Dahms (1986) for details. Type material for
Psilogaster nishidai not found. Holotype of Loshanus
petersoni, 9 , "Philippines, Palawan/ Mantalingajan/
Tagembung 1150 meters/ 19 Sept 1961/ Noona Dan
Exped. 61-62." Holotype of Orasema indica, 9 ,
"Holotype" Snehalatha, S. and Narendran, T. C. 1991."
"SL 151."

NOTES ON SYNONYMY

Boucek (1988) treated N. palgravei and N. purpureoven-
tris as different species of Orasema, with O. purpure-
oventris being the senior synonym for Parapsilogaster
laeviceps Gahan, Psilogaster nishidai Ishii and
Nagasawa, and Loshanus petersoni Hedqvist. Type mate-
rial was examined for all of the above species, except for
P. nishidai, which could not be located. However, P.
nishidai was well represented by material from the
Caroline Islands. Neoloshanus palgravei is treated here as
a widespread species in the Indo-Pacific region. Loshanus
petersoni and Orasema indica are junior synonyms of
Neoloshanus palgravei.

Specimens of both P. nishidai and N. purpureoventris
have been collected in the western Caroline Islands.
There is no doubt that P. nishidai belongs to Neoloshanus
based on the subapically expanded and strongly ridged
ovipositor (Ishii and Nagasawa. 1941). Specimens of P.
nishidai from the Caroline Islands match the original
description, and have a sessile stigmal vein, whicii 1 liave
seen from no other comparable species at this locality.
Psilogaster nishidai is similar to what I recognize as the
Papuan race of /V. palgravei and is treated here as a junior
synonym.

DIA(JN()SIS

The morphological limits of this species are broad and
encompass considerable regional variation (see
Variation). This species is recogni/cd by the following:

KW

face elabrato (Fig. 205), occiput broadl\ cniargitiaic. sec-
ond nagelloniere elongale (Figs. 181-1X2). stignial vein
perpendicular (Fig. 165), and femora dark brown. It may
be further distinguished from A', nepalensis by the shorter
flagellum in both sexes, and callus bare or with few short
seiae(Figs. 227, 231,253).

FEMALE

Length, 1.9-2.7 mm. Dark brown to black with faint
bluish reflections; coxae and femora dark brown to black
except extreme apex of femora; antenna yellowish brown
to dark brown; scape, fore and midlegs beyond femur,
and hind tarsi yellowish brown to white; hind tibia white
to dark brown (see Variation). Wings hyaline, venation
light or dark brown.

Head subtriangular; occiput broadly emarginate; medi-
an ocellus separated from occipital margin by less than its
own diameter; LOL about equal to OOL. Face broadly
rounded; scrobal depression shallow and poorly defined;
occipital carina extending laterally to eye margin. Eyes
separated by 1.4-1.7x their height. Malar space 0.6-0.9x
height of eye. Clypeus extended as strong lobe ventrally
over labrum, with or without epistomal sulcus (see
Variation). Antenna 11-segmented (Fig. 180); scape nar-
row, sparsely setose, reaching median ocellus; pedicel
short, narrower or equal in width to flagellum; anellus
present; flagellum 1.3-1.4x height of head; funicle 7-seg-
mented (8-segmented in 2 females from Nepal), segments
with numerous MPS; F2 about 1.2-1.3x F3, following
segments subequal in length; clava with 2 or 3 fused seg-
ments usually discernible.

Mesosoma with dorsum rugose-areolate or areolate
(Fig. 244); lateral lobe of mesoscutum usually transverse-
ly carinate; axilla irregularly carinate. Scutellum slightly
longer than broad, apex rounded in dorsal view; frenal
area angled 90 degrees to dorsum; axillula glabrate. axil-
lular sulcus weakly carinate. Propodeum with median
carina (Fig. 253) or irregular foveate channel; callus bare
or with sparse (less than 10) short setae. Femoral groove
broad and shallowly foveate (Figs. 228, 231). Pronotal
sulcus broad and foveate. Coxae and femora glabrate.
Forewing 2.2-2.6x as long as broad; basal area and
speculum bare; stigmal vein elongate, more than twice as
long as broad, rarely sessile (short and as long as broad),
but always perpendicular to wing margin (Fig. 165).

Metasoma with petiole l.8-2.8x as long as hind coxa,
I.5-2.8X as long as propodeum; petiole longitudinally
carinate, usually linear in profile but sometimes slightly
sinuate. Basal metasomal tergitcs bare. Ovipositor
straight or weakly curved forward (Fig. 267).

MALE

Length, 2.1-2.6 mm. Colour as in female, hind tibia
always dark brown with apex white or yellowish brown.

Antenna 12-segmented (Fig. 181 ); anellus present: flagel-
lum 1.9-2.1X height of head; funicle 8-segmented; F2
1.2x F3, following flagellomeres cylindrical and subequal
in length; clava with 2 fused segments. Forewings
2.2-2.4X as long as broad. Petiole carinate, long, and nar-
row, 2.8-4. 1 X as long as hind coxa, 2.1-3.4x as long as
propodeum (regionally variable). Ms^, narrow and round-
ed. Genitalia moderate sized, paramere narrow and elon-
gate; aedeagus subacute.

BI()L()(;Y(Figs. 17-25)

I observed the habits of individuals from populations in
Queensland and West Malaysia. At both localities,
females deposited eggs on the underside of a wide range
of broad-leaf plants. In Australia, host plants included
Tetrasynandra laxiflora (Monimiaceae), Lygodium retic-
ulatum (Schizaeaceae), Macaranga suhdentata
(Euphorbiaceae), and the most common host plant,
Fliudersia pimenteliana (Rutaceae). Plant hosts were sim-
ilar in Malaysia where eggs were deposited into new
growth of broad-leaf understorey shrubs or saplings,
including ferns. One additional host record was made on
Castanopsis (Fagaceae) in Papua New Guinea, which is
the same plant record as for N. townesi. Females deposit-
ed about 20 eggs in short double rows of alternating
punctures on the underside of a leaf. Oviposition was
extremely dense on some leaves and the entire underside
of each leaf (new growth) was saturated with oviposition
scars. Where numbers of adults were high, females and
males were observed congregating on sunlit vegetation.

Females deposited single eggs into each leaf puncture
formed by the ovipositor. Freshly deposited eggs could be
observed through a small circular hole in the leaf surface,
which was not marked by any alteration of the surround-
ing leaf tissue. Eggs were deposited nearly vertical to the
leaf surface, with the stalk erect and protruding from the
opening. As eggs matured, the leaf tissue formed a brown
cyst around the egg, and the margins of the cyst elevated
around the opening of the oviposition chamber. This was
more pronounced in the Australian plants than in the
Malaysian plants, which showed similar encrustments of
the punctures after the planidia had emerged. The under-
sides of many leaves were almost completely encrusted
by this scar tissue. Older leaves showed that leaf tissue
recovered from the scarring.

First-instar larvae were lightly sclerotized (pale yellow-
ish brown) and barely visible against the surface of the leaf.
Planidia were able to move by either slow inching move-
ments or by jumping 10 to 20 mm. No thrips or immature
Homoptera were observed on the leaves that could be inter-
preted as an intemicdiate host cairier. The host ants were
not found on the vegetation, but were common on the for-
est tloor. I assume that the planidia "rain" down upon the
host below, but did not observe this behaviour.

110

All stages of A^. palgravei were observed within the
host-ant nest in the Australian population. On the larval
ant host, planidia were attached to the lateral posterior
edge of the larval head capsule. Planidia remained exter-
nal and were attached only by the mouthparts. First-instar
larvae were turgid, showing evidence of some feeding,
but never greatly distended. Further development took
place on the host pupa within the cocoon, as is typical for
other Eucharitidae. Each parasite completed development
on a single ant host. Morphology of life stages and com-
parison with other Eucharitidae is presented in the earlier
section on biology.

I collected this species from undergrowth in rainforest
habitats in Australia and Malaysia and from a mixed
dipterocarp forest in Malaysia. Collections have also been
taken from a mixed dipterocarp forest and mangrove
swamp in Papua New Guinea and from a treefem ravine
in the Caroline Islands. Plant-host records suggest that this
species is associated with lush vegetation in a closed forest.

VARIATION AND DISTRIBUTION

Neoloshamis palgravei is widely distributed throughout
the Indo-Pacific region (Fig. 280), with a single specimen
belonging to this species known from Algeria (Boucek,
1988; BMNH, examined). Different patterns of geograph-
ical variation are found in the Indo-Pacific region, which
can be categorized into 4 distinct groups as outlined in
Table 5. Each of these forms may represent different
species but there are no patterns of variation that indicate
clear breaks in the populations. Variation is most evident
in the presence (Fig. 205) or absence (Fig. 201) of the
epistomal sulcus between the clypeus and supraclypeal
area, presence of a median groove or channel on the
propodeum (Fig. 253), white or dark hind tibia in the
female, and relative size of the petiole and hind coxa.
Although regional variants appear distinct, each character
state can usually be found to some degree in another
region. The distribution of the 4 geographical races is
shown in Figure 280. The 4 variants are discussed below
followed by a discussion of intermediate or disjunct dis-
tributions of character states.

Neoloshanus ncpalcnsis can be used to polarize char-
acter states among the races of A', palgravei outlined
below. Using this assumption, the absence of an epistom-
al sulcus, white hind tibia in males and females, and a rel-
atively short female petiole (1.5-2. Ox longer than
propodeum) would place the Malayan race as having a
plesiomorphic form, the Australian form as intermediate,
and the Papuan and Carolincan populations as derived.
Alternatively, it may be that the present distribution is the
result of morphological differentiation from a widespread
ancestor similar to the Australian race.

Malayan (M. Fig. 280). All of the specimens collected
on Borneo, Java, the island ot Mindanao in the

Philippines, and Malaysia (peninsular) can be attributed
to this group based on the lack of a clypeal groove, white
tibia in both the males and females, and a smaller ratio of
the petiole to propodeal length (Table 5). The coxae and
petiole are generally more robust than in the other
regions, but this is difficult to quantify. Unlike the eastern
races, the Malayan race lacks an epistomal sulcus except
for females from Selangor (Malaysia), which have a weak
depression. Tibial colour is intermediate in western
Malaysia, with a faded brown coloration of the hind tibia
in both males and females; however, females collected in
Selangor (Malaysia) are identical to females from
Australia.

Australian (A, Fig. 280). Specimens collected from
Australia (Queensland) and Papua New Guinea are recog-
nized by having a white tibia in females and a dark tibia
in males (with no intermediates). The clypeus has a dis-
tinct epistomal sulcus in all specimens. The propodeum is
carinate in females and grooved in males. The ratio of
petiole to propodeal length (1.8-2.8x) is large in both
sexes. Females collected in Taiwan and the Okinawa
Islands are similar to the Australian forms, but the epis-
tomal sulcus is intermediate as a narrow, transverse
depression. Wing veins of southern forms are all dark
brown but to the north and in Malayan forms the wing
veins are a light yellowish brown colour.

The Malayan and Australian races are difficult to sepa-
rate based only on females. Males differ by having the
hind tibia white in the Malayan race and dark in speci-
mens from Australia and Taiwan. Females from Borneo,
Singapore, and Thailand have a subcircular head and
short petiole (Table 5) and lack an epistomal sulcus
(swollen medially). Specimens attributed to the
Australian form from India, Nepal, and Algeria have a
slightly longer petiole and a subtriangular head and there
is a vague indication of an epistomal sulcus. The
propodeum has a distinct median carina in specimens
from India, Nepal, and Algeria, and a foveate or areolate
groove in specimens from Borneo, Java, Malaya,
Thailand, the Philippines, and Australia. The specimen
from Algeria is the only record of this genus from outside
of the Indo-Pacific region (except for a single South
American specimen), and is virtually identical to the indi-
viduals from India and Nepal. Confusion in the separation
of females of the Australian and Malayan races empha-
sizes the lack of distinct boundaries between races.

Papuan (P, Fig. 280). This race occurs throughout
New Guinea and extends north to the Philippines.
Females and males have the hind tibia dark, the cpislomal
sulcus usually distinct (may be absent or intermediate),
and the wing veins usually dark. The propodeal line may
be distinctly carinate, irregularly carinate. or areolate. The
petiole is usually very Ihin and elongate in males, and the
hind coxae are small in proportion to the nicsosoma.

Ill

Table 5. Major geographic variants of Neoloshanus pal}>ra\ei (Girault) with associated character states.
Moasuroinonts arc the ratio ol" the petiole length lo ihe hind coxal length (PTL/CXI.) and to the propodeal
length (PTl./F'PL); the mean value is given first, followed by the number of specimens measured, in paren-
theses, then the minimum-to-maximum range.

Character

Sex

Malayan

Australian

Papuan

Carolinean

Clypeal groove

Absent

Present

Present

Present

Medial propodeal

Foveate channel;

9 carinate;

Channel

Channel with

channel or line

both sexes may

channel in 6

or groove

parallel

have parallel or

or may be

carinae

single carinae

obliterated

Tibial colour

9

light

light

dark

dark

6

most light

dark

dark

dark

Stigmal vein

elongate

elongate

elongate

sessile

PTL/CXL

9

2.47 (9)

2.30(11)

2.48 (9)

2.14(10)

2.08-3.04

1.89-2.50

1.87-2.64

1.65-2.41

6

3.04 (5)

3.12(10)

3.18(8)

2.79-3.36

2.98-3.46

2.76-4.11

PTL/PPL

9

1.78(9)

2.22(11)

2.21 (9)

2.23(10)

1.47-2.01

1.82-2.79

1.87-2.64

1.92-2.54

6

2.59 (6)

3.17(10)

2.97 (8)

2.14-3.3

62.91-3.39

2.60-3.34

Carolinean (C, Fig. 280). This form can be recognized
by a rugose-areolate mesosoma with a vague median
depression on the scutellum, and small petiole-to-hind-
coxa ratio. The propodeum has a median areolate channel
which is sometimes bordered by weak carinae. The hind
tibia is dark in both males and females, and the stigma is
short and visible only as a swelling on the marginal vein.
This form was previously treated as Neoloshanus
nishidai. The variation of character states overlaps with
similar slates found in the Papuan forms, and the syn-
onymy proposed by Boucek (1988) appears justified. This
may just be an isolated island population of the Papuan
form.

MATERIAL EXAMINED

[Australian form) ALC.iiRiA |?|: Biskra (BMNH).
Australia: Queensland: Dunk Is.; Gillies Highway, 3.3
km W of Little Mulgravc; Hope Vallc Mission, 14 km W
by N (15-16S 144.5E); Hope Valle Mission. 14 km W by

N; Iron Range, Cape York Peninsula; Kuranda, 3, 2.8,
and 5.4 km N on Black Mtn Rd, Kuranda Range, rainfor-
est; Kuranda 200 m, 1000 m; Kuranda Range S. F.;
Kuranda, 1.5 km SE; Lankelly Ck, Mcllwraith Range. C.
York; Leo Creek Rd, 500 m, Mcllwraith Range, 20 km
NE Coen; Moses Ck, 4 km W by E Mt Finnigan (15.47S
145.17E); Mossman Gorge; Mt Tozer. 9 km ENE (12.43S
143.17E); Rocky Scrub. Mcllwraith Range, C. York; Tam
O'Shanter S. F., Mission Beach, N Rd, 50 m; Upper
Mulgrave Riv., via Gordonvale; Shipton's Flat (15.47S
145. 14E); all tnonths except September (1369 9,
163d c? , ANIC. BMNH, BPBM, CNC, JMH, MCZ,
QMB, ROM, TAMU). India: Karnataka: Mudigere,
October (19. BMNH); Kerala: Periyar, October (59 9,
BMNH). Japan: Ryukyu Islands: Okinawa Is.. Yona;
Amami-Oshima Is.: Hatsuno; Shinmura; Tojyomura;
Tokuno Shima Is., Mikyo, 200 m; Macaranga [?], J. L.
Gressitt; July. October to November (5 9 9, 5dd,
BPBM, KUEC). Nt-PAL: Lothar. nr Birganj. 150 m;

112

Pokhara, 910 m; September (399, BPBM. CNC). Papua
New Guinea: Amok; Baiyer Riv., 1 100 m; Brown Riv., 5
m; Bulolo, 900 m; Port Moresby, 20 km NE; October,
December to March (79 9, 1(5, AEI, BPBM, BMNH).
Taiwan: Nantou Hsien: Tungpu, 1200 m; Wushe, 1250
m; Taitung Hsien: Hung t'ou Hsu [Lanyu], May, August
to October (4 9 9, TARI).

[Malayan form] Indonesia: Java: Pekalongan;
Pelaboean, Ratoe, no dates (39 9, 16, MCZ, USNM);
West Kalimantan: Gunung Palung N. P.. 100-400 m,
1°15'S 110°5'E, primary rainforest, closed canopy MT
(29 9, MZB, ROM). Malaysia: Brunei: Labi, 200 m,
mixed dipterocarp forest, August to October (29 9,
BMNH). Selangor: Gombak, 26.4 km W, Univ. Malaysia
F.S.C.; Kuala Lumpur, Univ. Malaya, Rimba lima rain-
forest trail, 100 m; June, October, December (69 9, 26,
JMH. TAMU); Sarawak: Forest Camp, 19 km N
Kalabakan; Sandong, Kampong Tapuh, 300-400 m;
Sarikei Dist., Rejang Delta; Semongok; SW of Tapuh;
Tawan District. Kalabakan, primary forest; January, July,
November (16 9 9, 7(?d, BPBM, CNC). Philippines:
Misamis Oriental [Mindanao Is.]. Dinawihan Gingoog, 26
km E Gingoog City, 100-300 m; Negros Oriental: 16 km
W Dumaguete; May, August (19, \6 , BPBM, ROM).
Singapore: Nee Soon Water Res.; Singapore Botanical
Garden; December (26 6, BPBM). Thailand: Ban Na;
Fang, April ( 1 9 , lc5,BPBM). Vietnam: 20 km S Da Lat,
1300 m, September {\6, BPBM).

[Papuan form] Indonesia: Seram [AmboinaJ: no date
and October (39 9, 1(5, MCZ); Irian Jaya: Biak Is.,
50-100 m, sweeping in mangrove swamp. May {\ 6 ,
BPBM). Papua New Guinea: Adelbert Mts: Wanuma,
800-1000 m; Banz, Wahgi Valley, 1500 m; Banz. 16 km

NW, 1700 m; Central Prov., 20 km SE Port Moresby;
Bisianumu, E of Port Moresby, 500 m; Guega, W of
Swart Valley, 1200 m; Kougel Riv., 1250 m; Kundiawa,
Chimbu Riv.: Morobe Dist.. Mindik, 1200-1600 m; Mt
Hagen, 25 km E; Mt Suckling, Mau 1 [?], 500-1000 m,
collected on Castanopsis; Nenguag, Asaro-Chimbu div.,
2500 m; W Highlands, Goiburung, E of Korn Farm,
1560-1650 m; W Highlands. Kamang nr Minj, 1840 m;
September to February, April to July (19 9 9, 6 c5 (5, AEI,
ANIC. BPBM); Bougainville: Kokure, nr Crown Prince
Range, 900 m; Kokure, 690 m; June (2 9 9 , 2c5 c? ,
BPBM); Goodenough Is.: 400 m, September (19, AEI).
Philippines: Davao Prov. [Mindanao Is.], Calian; Negros
Oriental [Negros Is.], L. Balinsasayao, bait traps with
chicken intestines; Camp Lookout, 16 km W Dumaguete,
300 m; May (29 9 , 2c5 c? , BPBM, MCZ, ROM).
Solomon Islands: Guadalcanal, Suta (Suta-Gold Ridge),
Jonapau Mt, 1000 m; Suta, 500-1200 m; June (29 9,
BPBM).

[Carolinean form] Caroline Islands: Palau Islands:
Arakabesan Is.; Auluptagel Is.; Aurapushekaru Is.;
Babelthuap Is.: Airai, Ngarsung, along streams; Airai,
Ngerimal Riv., treefem ravine; Gakip; Imeluk, Netkeng;
Melekeiok; Ngaremeskang, 25 m; E Ngatpang. 65 m,
light trap; Ngerchelong; Ngiwal, jungle; SW of Ulimang,
wooded peak; Ulimang; Koror Is., limestone ridge; Peleiu
Is., Amiangal Mt; Uruklethapel Is., (NE), Palau Ngerem,
180 m; January, April to July, September, December
(319 9, 17(5(5, BPBM, USNM). Yap Islands: Yap Is.:
Kolonia, along streams; Dugai, with ants; Mt Gillifits.
150 m; Mt Matade, 95 m; Ruul Dist.; nr Yaptown; Yap
Hill, behind Yaptown, 50 m; Yap Is.; June to August,
November to December ( 1 3 9 9 , 8 (5 (5 , BPBM, USNM).

Neolosbanus gressitti-group

The 5 species in this group all have the following: clypeal
margin weakly rounded, face sculptured in most species,
ovipositor enlarged along the entire length, first valvula
smooth and without ridges or teeth, and second valvula
with lateral teeth. Species arc widespread in the Indo-
Pacific region with none known from Australia. A single
specimen is known from Uruguay and the record, if valid,
represents a significant range extension for this genus.

(JROUP DESCRIPTION

Head 0.3-0.4x as long as wide; ocelli arranged in
obtuse subtriangle (not line), median ocellus meeting or
only slightly exceeding line drawn across anterior margin
of lateral ocelli; face smooth to complelciy rugulosc or
punctate; scrobal depression smooth and shallowly
impressed; occiput aciculate or glabrous. Clypeal margin
broadly rounded and only slightly produced over mouth-

parts, epistomal sulcus distinct. Malar depression absent
or reduced. Antenna 1 1- or I2-segmented; scape cylindri-
cal or expanded apically; anellus present or absent; funi-
cle 8-segmented in both sexes, segments with MPS in
both sexes.

Mesosoma with dorsum weakly sculptured to
scabrous; frenal area glabrous or carinate. angled 80 to 90
degrees to dorsum of mesosoma (Fig. 254). Mescpimeron
glabratc; femoral groove continuing vcntrally. dividing at
ventrolateral margin, and .separate sulcus continuing pos-
teriorly to meet base of mid coxa (Figs. 232-233);
mesepisternum with wedge-shaped sternaular sculpture in
some species. Prepcctus foveate and triangular dorsally.
gradually narrowed veiUraily, polishetl below upper trian-
gular lobe (ridgelike in some N. violarens). Forewing
2.3-2.6X as long as broad; speculum and basal area of
forewing bare; stigmal vein of forewing broad and at

113

most 1.5x as long as broad; postmarginal vein shorter
than stigmal vein and abruptly narrowed.

Giisicr u ith basal terga glabrous. Ovipositor thickened
along entire length and straight (Fig. 270); first valvula
glabrous, without ridges or teeth; second valvula broad,
glabrous medially, and apically with 5 to 6 sharp teeth on
lateral margin. Ms^^ and genitalia of male diagnostic for
species.

Neolosbanus gressitti (Watanabe) comb. nov.

Loshanus gressitti Watanabe, 1958:28, fig. 5. Caroline
Islands [USNM, examined].

TYPE MATERIAL

Holotype, 9 , "PONAPE: Mt Tamatamansakir/ 1 80 m. I-
19-'53." "Caroline Is./ Pac. Sci. Bd./ J. L. Gressitt."
"Type No./ 64015/ USNM." Caster and petiole missing.
Deposited in USNM.

Paratypes [examined]: Ponape: same data as holotype
except collected on 18. i. 1953, (BPBM); N.2, SE.
Nanponmal, 7. i. 1953, J. L. Gressitt, light trap (1 9.
EIHU).

DIAGNOSIS

Recognized by the following: lateral lobe almost smooth,
anellus absent, and femora completely yellow. This
species bears a strong resemblance in general form to
species of GoUumiella, especially in head shape, antenna,
femora, and wing venation.

FEMALE

Length, 1.7-1.8 mm. Dark brown to black; antenna,
coxae, and gaster dark brown; scape and legs yellow.
Wings hyaline, venation pale brown.

Head subcircular in frontal view; occiput transverse to
slightly emarginate; median ocellus separated from
occiput by more than its own diameter; LOL 1.2-1.5x
OOL. Face broadly rounded, polished with sparse setifer-
ous punctures; occiput glabrous, occipital carina weak,
not reaching dorsal eye margin. Eyes separated by
1.5-1.7X height of eye. Malar space 0.8-0.9x height of
eye, malar depression absent. Lateral margin of clypeus
sharply impressed. Antenna 1 1 -segmented; pedicel 1.5x
as long as broad, 1.5x wider than F2 at base; scape nar-
row, reaching ().6-0.8x distance to median ocellus; anel-
lus absent; flagellum 1.3-1.4x height of head; funicular
segments with numerous MPS; F2 3.7-3.9x as long as
broad, 1.3-1.6x F3, following segments subequal in
length; clava with 2 incompletely fused .segments.

Mesosoma dorsally with midlobe, axilla anteriorly,
and apical half of scutellum rugose-areolate; lateral lobe
smooth or very weakly carinate, axilla and anterior half of

scutellum widely spaced carinate. Scutellum slightly
longer than broad, rounded apically: frenal area complete-
ly polished; axillula glabrate, axillular sulcus indistinct.
Propodeum with irregular median sculpture and weak
median carina; callus with few sparse hairs. Coxae and
femora polished with few short setae; hind tibia with
sparse setae. Forewing 2.4-2. 5x as long as broad; stigmal
vein 1.5x as long as broad (speculum bare, not pilose as
figured in the original description).

Metasoma with petiole 1.8-l.9x as long as hind coxa,
I.5-1.7X as long as propodeum; petiole longitudinally cari-
nate.

MALE

Unknown.

DISTRIBUTION

Eastern Caroline Is. (G, Fig. 279).

Neolosbanus kokureanus sp. nov.

Figs. 176-177

TYPE MATERIAL

Holotype, 9 , "SOLOMON IS./ BOUGAINVILLE (S.)/
Kokure, 690 m./ June 12. 1956." "E. J. Ford. Jr./
Collector." "HOLOTYPE/ Neolosbanus/ kokureanus
Heraty." Deposited in BPBM.

Paratypes: Papua New Guinea: Bougainville: Kokure, nr
Crown Prince Range, 900 m., 9.vi.l956. J. L. Gressitt
(19, BPBM); New Guinea: Mt Kaindi, 1790 m,
20-23.vii.1977, J. L. Gressitt, malaise trap (Id, BPBM).

DIAGNOSIS

This species is closest to A', gressitti. Both species have
the femora yellow, face of female polished with sparse
pits (Fig. 176), frenal area polished medially (in female),
and face broadly rounded. N. kokureanus differs by: anel-
lus distinct (Fig. 177), lateral lobe of mesoscutum cari-
nate, and coloration strongly metallic blue-green.

FEMALE

Length, 2.1-2.2 mm. Body, including coxae, dark metal-
lic blue-green; legs and antenna yellow, antenna darker
apically. Wings hyaline, venation pale brown.

Head subtriangular. cheek broadly rounded; occiput
weakly emarginate; median ocellus separated from occip-
ital margin by more than its own diameter; LOL 1.5x
OOL. Face rounded, polished with few minute setiferous
punctures lateral to supraclypeal area; occiput glabrous,
occipital carina weak and extending just beyond lateral
ocellus. Eyes separated by 1 .4x their height. Malar space
0.7-().8x height of eye, malar depression weakly
impressed near base of mandible. Lateral margin of

114

clypeus deeply impressed; epistomal sulcus distinct.
Antenna 12-segmented; pedicel longer than broad, slight-
ly wider than F2; scape narrow, reaching median ocellus;
anellus present; flagellum 1.7x height of head; funicular
segments with sparse MPS; F2 5. Ox as long as broad.
1.2x F3, following segments subequal; clava appearing 1-
segmented.

Mesosoma with dorsum strong rugose-areolate; lateral
lobe transversely carinate; axilla longitudinally carinate.
Scutellum as long as broad, apically rounded; frenal area
glabrous medially; axillula glabrous, axillular sulcus
indistinct. Propodeum with prominent median carina
becoming areolate band dorsally; callus with sparse setae.
Coxae and femora smooth with sparse setae; hind tibia
moderately setose. Forewing 2.6x as long as broad; stig-
mal vein as long as wide.

Metasoma with petiole 2.1x as long as hind coxa. 2.2x
as long as propodeum; petiole longitudinally ribbed dorsally.
weakly sculptured sublaterally with weak ventral keel.

MALE

Length, 2.6 mm. Colour as in female except antenna com-
pletely yellowish brown. Median ocellus separated by
less than its diameter from occipital margin. Face with
cheek and frons next to eye shallowly punctate becoming
weakly rugose to punctate dorsally; occipital carina
prominent. Antenna 12-segmented; scape strongly
expanded apically but clublike and without flange; anel-
lus present; F2 l.lx F3, F3 1.2x F4, following segments
subequal in length. Scutellum with frenal area strongly
rugose. Petiole 1.9x as long as hind coxa, 2.7x as long as
propodeum. MSj^ broad and truncate. Genitalia broad,
paramere short, aedeagus broad.

DISTRIBUTION

Known only from Bougainville and New Guinea (K, Fig.
279).

ETYMOLOCY

From Kokure; referring to the type locality.

Neolosbanus anapetus sp. nov.

Figs. 186, 207-208, 233, 254, 270, 273-274

TYPE MATERIAL

Holotype. 9. "PHILIPPINES:/ Mt Province/ Mayoyao.
Ifugao/ 1200-1500 m/ 19.IX.1966." "H. M. Torrevillas/
Collector/ BISHOP." "HOLOTYPE/ Neolosbanus/
anapetus Heraty." Deposited in BPBM.
Paratypes: Mai,aysia: Sabah: W Coast Residency, Ranau.
13.2 km N. of Poring Hot Springs. .500 m. S-1 1.x. 1958.
T. C. Maa. sweeping (19. Id. BPBM); Ranau. .500 m.
28.ix-7.x.l958, T. C. Maa (19, BPBM); Tawan Dist..

Kalabakan. primary forest. 8-15. xi. 1958. T. C. Maa (19,
BPBM); Forest Camp, 9.8 km SW of Tenom. 19.xii.l962,
K. L. Kuncheria (19, BPBM); "Borneo West" (19,
BMNH). Philippines: Mt Province: Mayoyao, Ifugao,
1200-1500 m. 3,4,6-7, 10, 13. ix. 1966 (99 9 . 8(? c?,
BPBM); same locality, 27.viii.1966 {26 6. BPBM); La
Lune Mtns, Davao, Mindano, 5.vii C. S. Clagg (19,
MCZ); Davao. Mindano, Baker (19, MCZ); Los Banos,
Baker (Id. BPBM).

DIAGNOSIS

Recognized by the following: head and mesosoma dark
green, lower region of face strongly rounded, clypeus
flush with lower face (Fig. 207), scape of male with
strong inner flange apically (Fig. 186), and mesosoma
with dorsal sculpture shallow rugose-areolate.

FEMALE

Length, 2.0-3.0 mm. Mesosoma black with strong green-
ish reflections, head dark green with reddish reflections,
metasoma dark brown to black; femora and apical seg-
ments of antenna dark brown; most of antenna, apex of
femora, tibiae, and tarsi yellowish brown. Wings hyaline,
venation pale brown.

Head subtriangular, cheek broadly rounded; occiput
broadly emarginate; median ocellus separated from
occiput by slightly less than its own diameter; LOL
0.9-1.3X OOL. Face strongly rounded in lower half,
punctation widely separated, interstices glabrate (Figs.
207-208); occiput weakly aciculate, occipital carina
extending laterally below dorsal margin of eye. Eyes sep-
arated by 1.4-1.7X their height. Malar space 0.7x height
of eye; malar depression shallow and poorly defined,
broader next to oral margin. Clypeal region polished and
covered by short setae with several elongate setae near
apex, lateral margin sharply impressed and clypeus flush
with lower face. Antenna 11- or 12-segmented; scape
slender, not reaching median ocellus; pedicel slightly
longer than broad, wider than F2 at base; anellus present,
but in various degrees of fusion with F2; flagellum
1.3-1.9X height of head; funicle with few MPS; F2
3.3^.3x as long as broad. 1.4x F3; clava 1 -segmented;
flagellomeres cylindrical and subequal in length.

Mesosoma with dorsum finely and shallowly rugose-
areolate; lateral lobe of mesoscutum transversely carinate;
axilla longitudinally carinate. Scutellum as long as broad,
base well separated from TSA; frenal area irregularly car-
inate; axillula glabrate. axillular sulcus obscured by sur-
face sculpture. Propodeum evenly rounded with shallow,
irregular rugose-areolate sculpture medially; callus with
several sparse, short setae. Femoral groove broad and
deeply foveatc to carinate. Coxae glabrous; femora weak-
ly coriaceous lo glabralc. Forewing 2.3-2.5x as long as
broad; sligmal \ciii Itroader than long.

115

Mciasoma with petiole 1. 7-2. Ox as long as hind coxa.
I.3-2.2X as long as propodcum; petiole longitudinally
ribbed, sublaterally weakly sculptured and with weak
ventral keel. Ovipositor as described for group (Fig. 270).

MALE

Length, 2.2-2.6 mm. Colour as in female. Antenna 1 1- or
12-segmented: scape expanded apically, inner margin
with protruding flange apically bordering broad excavat-
ed region below pedicel (Fig. 186); anellus absent or par-
tially fused to F2; flagellum 1.7x height of head; F2
2.2-2.9X as long as broad, 1.2x F3, distinctly broader
than F3, basal flagellar segments compressed and becom-
ing cylindrical distally, apex of each segment abruptly
enlarged. Petiole 2.1-3.1x as long as hind coxa, 2.2-2.6x
as long as propodeum. Ms, with weak constriction near
base (Fig. 273). Ms^^ rounded apically. Genitalia as in
Figure 274, paramere long.

VARIATION

Sculpture on dorsum of the mesosoma on the Sabah mate-
rial is finer and more closely spaced than any of the
Philippines specimens, but otherwise the material is iden-
tical.

DISTRIBUTION

Philippines and Malaysia (Sabah; A, Fig. 279).

ETYMOLOGY

From Latin anapetes, meaning expanded; referring to the
widened male scape.

Neolosbanus violaceus sp. nov.

Figs. 185,206,232

TYPE MATERIAL

Holotype, 9, "Baiyer R., N. Guinea/ Xn.24-26.1978/
1100 m. J. Sedlacek." "HOLOTYPE/ Neolosbanus/ vio-
laceus Heraty." Deposited in AEL A male was chosen as
holotype because neither female has a complete antenna.
Paratypes: Papua New Guinea: Guega, W of Swart
Valley, 1200 m, 15.xi.l958, J. L. Gressitt (19, BPBM);
Brown Riv., 22. v. 1958, E. J. Ford, Jr. (19, BPBM);
Baiyer Riv., 1 100 m, 24-26.xii.1978 (Id), 25.i-6.ii. 1979
(36 6), 25.ii-9.iii.1979 (36 6), J. Sedlacek (AEI);
Karimui, S of Goroka, 1000 m, 5.vi.l961, J. L. Gressitt
and M. Gressitt (1 (5, BPBM); Morobe Dist., Wau, Kunai
Ck, 1250 m, 26.viii.1963, J. Sedlacek, Malaise trap (lc5,
BPBM).

dia(;n()sis

Distinguished by the following: anellus large (Fig. 185),
scape of female slender and scape of male medially

expanded (Fig. 185), supraclypeal and clypeal areas later-
alls delimited by weak narrow groove (Fig. 206), cheek
not broadly rounded and not flush with clypeal region,
mesosoma with strong dorsal sculpture, and petiole of
male long and sinuate.

FEMALE

Length. 2.3-2.6 mm. Head, mesosoma. and petiole dark
blue with strong violaceous reflections; coxae, femora,
and gaster dark brown; antenna, tarsi, and apex of femora
yellowish brown; tibiae dark brown or yellowish brown
medially. Wings hyaline, venation brown.

Head subtriangular, cheek broadly rounded; occiput
weakly emarginate; median ocellus separated from poste-
rior margin by less than its own diameter; LOL l.Ox
OOL. Face weakly rounded, moderately setiferous-punc-
tate with interstices glabrate to weakly colliculate (Fig.
206); occiput weakly aciculate, occipital carina weak,
extending halfway between ocellus and eye margin. Eyes
separated by 1.5x their height. Malar space 0.7x height of
eye, malar depression narrow and shallowly impressed
next to oral margin. Clypeal region glabrous, lateral mar-
gin distinctly impressed. Antenna 12-segmented; pedicel
slightly longer than broad, wider than F2 at base; scape
narrow and cylindrical, almost reaching median ocellus;
anellus present or partially fused to Fl; F2 3.8-6.3x as
long as broad. 1.2x F3, F2 slender with dense, small setae
(rest of antenna missing).

Mesosoma with dorsum, including lateral lobe of
mesoscutum. scabrous to deeply rugose-areolate; posteri-
or half of axilla longitudinally carinate. Scutellum as long
as broad; frenal area irregularly carinate to areolate dor-
sally; axillula glabrous, axillular sulcus lacking.
Propodeum with irregular median carina in posterior half,
bordered by irregular medial band of areolate sculpture,
callus with sparse patch of setae. Femoral groove broad
and weakly foveate. Coxae and femora weakly sculptured
and with fine sparse setae. Forewing 2.4x as long as
broad; stigmal vein slightly longer than broad.

Metasoma with petiole 1.8-l.9x as long as hind coxa,
1.9-2.1X as long as propodeum; petiole longitudinally
ribbed, interstices of ribs weakly rugulose. (Ovipositor
hidden in sheath.)

MALE

Length. 2.1-3.1 mm. Colour as in female, hind tibia dark
brown to black medially, apices yellowish brown. LOL
0.9-1. 2x OOL. Antenna 1 1 -segmented: scape not reach-
ing median ocellus, swollen medially and flattened below
pedicel (Fig. 185); anellus minute; funicular segments
cylindrical to slightly compressed laterally, uniform along
entire length, subcqual in length and width; F2 l.l-1.3x
F3. slightly broader than F3: clava with 2 incompletely
fused segments. Propodeum with median carina bordered

116

by medial band of rugose or areolate sculpture. Petiole
2.3-2.8X as long as hind coxa, 3.0-3.3x as long as
propodeum. Ms^ rounded. Genitalia typical, paramere
narrow and elongate.

DISTRIBUTION

New Guinea (V, Fig. 279).

ETYMOLOGY

From Latin violaceus. meaning violet coloured; referring
to the colour of the head and mesosoma.

ADDITIONAL SPECIES

Two species, similar to N. violaceus, are known from
Malaya (X, Fig. 279), and apparently Uruguay. The sin-
gle specimen from Uruguay (Tacuarembo: 40 km NW
Tacuarembo, 10-16.ii.l963, J. K. Bouseman; AMNH)
represents an anomaly for the genus, which is otherwise
restricted to the Indo-Pacific. J. K. Bouseman (INHS) was

contacted regarding the validity of the label. He took part
in the 1963 expedition to Uruguay, and malaise trap
material was returned to AMNH for processing. This
specimen is covered with lepidopteran scales, suggesting
that it was collected in a malaise trap. I have seen no
other eucharitid material in the AMNH collection collect-
ed from the Indo-Pacific region, and chances for misla-
beling of the specimen would appear to be remote. I pre-
fer not to describe this species until the locality can be
verified. It is a distinct species and is closer to the male
from Thailand, described below, than to A', violaceus.

A single male specimen (not described because of its
poor condition) from Thailand (Trang Prov.,
Khaophoppha Khaochang, 200 m, 9-11.1.1964, G. A.
Samuelson, BPBM) differs in having a shorter petiole,
very slender scape and antennal segments, F2 as broad as
F3, hind coxa polished, femora yellow, mesoscutum
transversely carinate laterally and anteriorly, and Ms^ nar-
rowly rounded.

117

Summary

The Orascminao and Eucharitinac arc recognized as a
nionnphylclic group that fits the traditional concept of the
Eucharitidae (Burks, 1979: Graham, 1969; Heraty, 1985,
1989). Females of both subfamilies deposit their eggs in
or on plant tissue and are parasites of ant pupae through
the larval stage of the host. First-instar larvae possess sev-
eral character states that justify the monophyly of
Eucharitinac + Oraseminae. and provide reliable evidence
for determining phylogcnetic relationships with respect to
the Perilampidae. Other subfamily groupings proposed by
Boucek (1978, 1988) are not dealt with, and placement in
the Eucharitidae for these taxa is problematical. The
Echthrodapinae and Philomidinae are parasites of bee
pupae and Philomidinae have a planidiaform larva that is
distinct from those of Perilampidae and Eucharitidae and
should therefore not be included in Eucharitidae (Darling,
1992). The biology of species of Akapalinae is unknown.
Character states that are deemed ancestral for Oraseminae
+ Eucharitinac are either typical of most Chalcidoidea or
justify the monophyly of the 2 subfamilies. No character
states were found that would clarify relationships with the
other subfamilies proposed by Boucek.

Oraseminae were previously defined as a subfamily
based on the presence of an anelliform first flagellomere,
independent prepectus, constricted first gastral stemite,
and subapically expanded ovipositor. These character
states of adults that were used to support the earlier con-
cept of Oraseminae are either plesiomorphic (as recog-
nized in earlier studies) or homoplastic and uninformative
in defining relationships. In particular, I postulate that an
expanded ovipositor has arisen independently in at least 5
unrelated taxa (Oraseminae, Neoloshanus, Anorasema,
Schizaspidia, and Chalcura montana). The habit of
depositing eggs into plant tissue likely provides a func-
tional basis for convergent development of the ovipositor.
The constriction of the first gastral stemite (Ms.,) is ple-
siomorphic and shared with the Perilampidae. It is found
in all species of Oraseminae, and I propose that its loss is
an apomorphy of Eucharitinac.

The Oraseminae is recognized as a monophyletic
group that includes the genera Indosema, Orascma.
Orascmoipha, and Timiodenis. Certain species previous-
ly treated as Orasema were transferred to 2 new genera
that are included in the Eucharitinac. The morphological
limits of the Eucharitinac were expanded to include the
new genera Neoloshanus and Psilocharis. Both genera
are placed in a new tribe, the Psilocharitini. Anorasema +
Gollnniiclla + Eucharitinac .s.v. are interpreted as a mono-
phyletic group and placed together in a new tribe,
Eucharitini. The Eucharitinac .v. .v. (excluding Anorasema
and GoUumiella) is monophyletic as based on characters

treated in previous studies.

The biology of Oraseminae and Eucharitinac was
reviewed based on new larval collections and host records
for Orasema, Orasemorpha. and Neoloshanus. The con-
servative nature of the morphology of first-instar larvae
was supported in all of the taxa examined. A reexamina-
tion of larvae described by other authors removed evi-
dence of conflicting hypotheses of character states within
Oraseminae. Additional characters were used for defining
eucharitid relationships based on later immature stages
including the pupa, characters of eggs, characteristics of
oviposition, and ant host. The phylogeny of Eucharitidae
as based on an analysis of characters of adults was robust
with respect to larval characters. Inclusion of larval char-
acters along with characters of adults in the cladistic
analysis supported the realignment of Oraseminae and
Eucharitinac. Larval characters and within-nest behaviour
of Neoloshanus are the same as for other members of the
Eucharitinac. The previous classification of species of
Neoloshanus within Orasema would have provided a
false prediction of larval characters, behaviour within the
nest, and ant host.

There are strong correlations between ant hosts and
phylogcnetic relationships for Eucharitidae. Myrmicinae
are regarded as the ancestral host for Oraseminae.
Pheidole is the only known host in the Old World, but
New World species of Orasema are also found on other
Myrmicinae, Formicinae, and Ecitoninae. The known ant
host of Neoloshanus is Hypoponera, and Ponerinae are
postulated as the ancestral host for Eucharitinac. Within
Eucharitinac, several derived lineages have switched to
Formicinae. Either the Ponerinae and Myrmicinae are
treated as derived sister groups, or the Myrmicinae
(including Myrmeciinae -i- Pseudomyrmecinae) are treat-
ed as the sister group (excluding or including the fomii-
coid subfamilies) to the Ponerinae + army ants;
Formicinae are not closely related to Ponerinae (Baroni-
Urbani, 1989; Bolton, 1990; Holldobler and Wilson,
1990; Baroni-Urbani et al., 1992; Shattuck, 1992).
Coevolution or "tracking" of the host evolution cannot be
ruled out among basal groups of eucharitids. However, I
propose an invasive or colonizing form of host adaptation
with eucharitids overcoming host defences and radiating
within a particular ant lineage. The best evidence for this
strategy was observed in the Oheza-clddc, in which
females deposit their eggs in association with fruit or
offer egg secretions as an attractant for Camponotus
(Formicinae) (Heraty and Barber. 1990). Behaviour of the
planidial stage is recognized as a means of gaining access
to the host, and changes in behaviour may be responsible
for host shifts in Formicidae.

118

The number of species found in each geographic
region is summarized and presented along with a clado-
gram derived from the analysis of relationships at the
species-group level (Fig. 281). Only Orasema and
Psilocharis have representative species in all of the
regions of the Old World tropics. Orasemorpha,
Timioderus, and Indosema have restricted continental dis-
tributions, and almost all collections of Neolosharms were
confined to the Indo-Pacific region. These differences
occur because different geographical and evolutionary
histories affect lineages of supposedly different ages. Of
the 6 genera treated in this analysis, the distribution of
each species was limited, and only a few species were
found in more than one geographical subregion. The sin-
gle records of Psilocharis from Argentina and
Neoloshanus from Uruguay are difficult to dismiss, and
may be the result of a relict distribution, dispersal, human
transport (e.g., ships' ballast), or possibly mislabelled
specimens; however, in both cases the label data appear
to be accurate.

Phylogenetic hypotheses and present disjunctions
among Oraseminae and Psilocharitini support the broad
patterns proposed in Fig. 282. The Australasian fauna is
distinct from the Indo-Pacific fauna, and species of
Orasema, Psilocharis, and Neoloshanus in the Papuan
region appear to be derived from ancestral species in the
Indo-Chinese + Malayan subregions, as postulated also
by Gressitt (1982), Noonan (1985), and Schuh and
Stonedahl (1986). Species of Orasema and Psilocharis
found in the Australian region are phylogenetically isolat-
ed from the most closely related Indo-Chinese species
and have closer relationships to Ethiopian species.
Relationships between Orasemorpha, Timioderus, and
Indosema also suggest an early Ethiopian/Australian dis-
junction, supporting a Gondwanan pattern of distribution,
although only Orasema is found in South America (with
questionable records of Psilocharis and Neoloshanus).
Thus, Oraseminae and Psilocharis may have originated
during the late Cretaceous or early Eocene. Neoloshanus
is largely restricted to the Indo-Pacific region and may
have radiated eastward through to the Papuan subregion
some time after the Miocene uplift of islands in the Indo-
Pacific region (Noonan, 1985). Species in the Orasema

uichancoi-group exhibit a similar pattern of radiation to
Neoloshanus but are most closely related to species found
in the Malagasy region. The proposed faunal shifts
between the Ethiopian and Indo-Chinese regions, as
based on phylogeny, agree with patterns found in
Carabidae and Miridae (Noonan, 1985; Schuh and
Stonedahl, 1986).

The distribution of Neoloshanus suggests that the
northern limits of the Indo-Chinese subregion should be
extended north to include the islands Kyushu, southern
Honshu, and Tsushima (Figs. 279-280, 284). Orasema
koghisiana is the only orasemine known from New
Caledonia and the New Hebrides and it shows a closer
phylogenetic relationship to species from Australia rather
than from the Indo-Pacific region as proposed by Gressitt
(1956). Otherwise, the boundaries of subregions outlined
by Gressitt (1956) and Schuh and Stonedahl (1986) are
supported.

Revisionary studies were carried out on 6 genera of
Eucharitidae belonging to Oraseminae and Eucharitinae
in the Old World; 56 species are treated, of which 31
belong to the Oraseminae and 25 to the Psilocharitini
(Eucharitinae). Phylogenetic hypotheses are proposed for
species within each of the genera. This is the first com-
prehensive treatment of the Eucharitidae of the Old
World tropics. I do not expect it to encompass all of the
species in this area, and I expect that more species remain
to be discovered.

Prediction is the ultimate goal of any scientific
endeavour. Phylogenetic studies were used to develop a
new classification for Eucharitidae. The present classifi-
cation is correlated with information on morphology of
adults and immatures, behaviour, and host relationships,
and also with faunistic histories of the Old World tropics.
There are many gaps in our knowledge of geographical
distributions, host records, and immature stages, and rela-
tionships of Eucharitinae need to be more completely
resolved. New information on these groups can be used to
modify and refine the hypotheses as I have presented
them, and I would like to think that this work provides the
challenge to others to continue studies on this fascinating
group of insects.

119

Acknowledgements

This study was initialed at Texas A&M University and
completed at Carleton University and the Biosystematics
Research Centre (BRC). Ottawa. I thank my Ph.D. exam-
ining committee who reviewed a version of this manu-
script and offered helpful suggestions: J. B. Woolley,
R. A. Wharton. M. V. Sweet, and S. B. Vinson (all Texas
A&M University). I also thank G. Gibson and J. Huber
(BRC). C. Darling (ROM), and J. LaSalle and J. Noyes
(BMNH) for thoroughly reviewing the final version of
this manuscript and providing additional suggestions for
improvement. Numerous people offered assistance during
my stay at Texas A&M and I would like to thank
G. Zolnerowich, D. Judd, J. C. Schaffner, H. R. Burke,
M. Rose, and many others. My studies in Ottawa were
greatly enhanced by the support from J. Huber and
G. Gibson (BRC). R. S. Anderson (Canadian Museum of
Nature), and H. Howden and S. Peck (Carleton
University).

I am grateful to the individuals and institutions listed
in the materials sections for the loans of specimens. I
would also like to acknowledge the assistance on my col-
lecting and museum expeditions from the following: E. C.
Dahms (QMB); R. Storey (Department of Plant
Industries, Queensland); I. D. Naumann (ANIC); Liang-
Y. Chou, K.-C. Chou, and Soo-Min (TARI); R. Loo

(Taipei. Taiwan); H. S. Yong (University of Malaya.
Malaysia); Chaweewan Hutacharern and Surochai
Choldrumkul (Royal Forestry Department. Thailand);
S. I. Farooqi. Manickivasagam (lARI); J. S. Noyes and
Z. Boucek (BMNH). The Socio-Economic Unit of Mal-
aysia and the National Research Council ol Thailand gave
their permission to conduct research in their countries.

I also thank David Swofford and Dave Maddison for
making available test versions of PAUP and MACLADE
for use in my analyses.

This work was supported in part by a Snodgrass
Memorial Research Grant, a Natural Sciences and
Engineering Research Council of Canada (NSERC)
Postdoctoral Fellowship and a Smithsonian Postdoctoral
Fellowship to the author, and a United States National
Science Foundation Dissertation Improvement Grant
(BSR 8914680) to the author and J. B. Woolley.

I would like to give special thanks to Jim Woolley for
providing me with the environment that was needed to
complete this work. Over the years Zdenek Boucek has
provided assistance, encouragement, and some tremen-
dous material for which I am truly grateful. Finally, 1
would like to dedicate this work to my wife, Laura, and
my daughter, Joanne, for all that they have had to endure
over the course of this study.

120

Literature Cited

ALEXANDER. B.

1990 A preliminary phylogenetic analysis of sphecid
wasps and bees. Sphecos 20:7-16.
ANANTHAKRISHNAN, T. N.

1984 Bioecoiogy of thrips. Oak Park, Michigan, Indira
Publishing. 233 pp.
ARNETT, R. J., Jr., and G. A. SAMUELSON

1986 The insect and spider collections of the world.
Gainesville, Florida, E. J. Brill. 228 pp.
ASHMEAD, W. H.

1888 Descriptions of three new eucharids from Florida
with a generic table of the Eucharinae. Entomolo-
gica Americana 3: 1 86-1 88.
1892 Notes on the eucharids found in the United States.
Proceedings of the Entomological Society of
Washington 2:354-358.
1895 Some parasitic Hymenoptera from Baja California
and Tepic Mexico. Proceedings of the California
Academy of Sciences 5:539-555.
AXELROD, D. 1. and P. H. RAVEN

1982 Paleobiogeography and origin of the New Guinea
flora. In Gressitt, J. L., ed., Biogeography and ecol-
ogy of New Guinea, Vol. 2, pp. 919-945. The
Hague, Junk.
AYRE, G. L.

1962 Pseudometagea schwarzii (Ashm.) (Eucharitidae:
Hymenoptera), a parasite of Lasius neoniger Emery
(Formicidae: Hymenoptera). Canadian Journal of
Zoology 40:157-164.

BARONI-URBANI, C.

1989 Phylogeny and behavioral evolution in ants, with a
discussion of the role of behavior in evolutionary
processes. Ethology, Ecology and Evolution
1:137-168.

BARONI-URBANI. C. B. BOLTON, and P. S. WARD

1992 The internal phylogeny of ants (Hymenoptera:
Formicidae). Systematic Entomology 17:301-329.
BESMEAR. R.J.

1974 A chalcidoid planidium and thrips larvae in
Georgia. Journal of the Georgia Entomological
Society 9:265-266.
BLANCHARD. E.

1840 Histoire naturelle des inscctes 111. Paris, Dumeril.
672 pp.
BOLTON. B.

1990 Army ants reasses.sed: the phylogeny and classifi-
cation of the dory line section (Hymenoptera, For-
micidae). Journal of Natural History 24:1339-1364.

BOUCEK. Z.

1956 A contribution to the biology of Eticlutris aclscen-

clcns (F.) Hymenoptera. Acta Societatis Zoologicae

Bohemoslovenicae 20:97-99.
1978 A generic key to Pcrilampinac (Hymenoptera.

Chalcidoidea). with a revision of Kromhcinii.s n.

gen. and Euperilampiis Walker. Entomologica

Scandinavica 9:299-307.
1988 Australasian Chalcidoidea (Hymenoptera).

Wallingford: C. A. B. International. 832 pp.
BOUCEK, Z. and J. S. NOYES

1987 Rotoitidae, a curious new family of Chalcidoidea

(Hymenoptera) from New Zealand. Systematic

Entomology 12:407^12.
BRETHES, J.

1927 Hymenopteres Sud-Americains du Deutches

Entomologisches Institut: Terebrantia. Entomolo-

gische Mitteilungen (Berlin) 16:319-335.
BROWN, W. L.. Jr.

1954 Remarks on the internal phylogeny and subfamily

classification of the family Formicidae. Insectes

Sociaux 1:21-31.
1973 A comparison of the Hylean and Congo-West

African rain forest ant faunas. //; Meggers, B. J., A.

S. Ayensu, and W. D. Duckworth, eds.. Tropical

forest ecosystems in Africa and South America: a

comparative review, pp. 161-185. Washington,

D.C., Smithsonian Institution Press.
BRUES. C. T.

1919 A new chalcid fly parasitic in the Australian bull-
dog ant. American Entomological Society of

America 12:13-23.
1934 Some new eucharidid parasites of Australian ants.

Bulletin of the Brooklyn Entomological Society

29:201-207.
BUCHER, G. E.

1948 The anatomy of Monodontomerus dentipes Boh., an

entomophagous chalcid. Canadian Journal of

Research 26:230-281.
BURKS, B. D.

1979 Eucharitidae. //; Krombein, K. V., P. D. Hurd. Jr.,

D. R. Smith, and B. D. Burks, eds.. Catalog of

Hymenoptera in America north of Mexico. Vol. 1,

Symphyta and Apocrita (Parasitica), pp. 875-878.

Washington, D.C., Smithsonian Institution Press.

CAMERON, P.

1884 Hymenoptera. In Godman. F. D., and O. Sal i via,
eds., Biologia Centrali-Americana. Insecta
Hymenoptera (Familias Tenthredinidae — Chrysi-
didae), pp. 81-144. London. Taylor and Francis.
487 pp.

1909 A contribution to the knowledge of the parasitic
Hymenoptera of Argentina. Transactions of the
American Entomological Society 35:419^50.
CLANCY. D. W.

1946 The insect parasites of the Chrysopidao (Neu-
roptera). University of California Publications in
Entomology 7:403^96.

121

CLAUSEN. C. P

1923 The biology of Schizaspidia tenuicoinis Ashiiiead,
a eucharid parasite of Camponoius. Annals of the
EntoMiolojzical Society of America 16:193-217.

1928 The manner of oviposition and the pianidium of
Schizaspidia manipurensis n. sp. (Hymenoptera,
Eucharidae). Proceedings of the Entomological
Society of Washington 30:81-85.

1940a Entomophagous insects. New York, McGraw-Hill.
688 pp.

1940b The oviposition habits of the Eucharidae (Hymen-
optera). Journal of the Washington Academy of
Sciences 30:504-516.

1940c The immature stages of the Eucharidae. Proceed-
ings of the Entomological Society of Washington
42:161-170.

1 94 1 The habits of Eucharidae. Psyche 48:57-69.

DAHMS, E. C.

1983 A checklist of the types of Australian Hymenoptera
described by Alexandre Arsene Girault: II.
Preamble and Chalcidoidea species A-E with advi-
sory notes. Memoirs of the Queensland Museum
21:1-255.

1984 A checklist of the types of Australian Hymenoptera
described by Alexandre Arsene Girault: III.
Chalcidoidea species F-M with advisory notes.
Memoirs of the Queensland Museum 21:579-842.

1986 A checklist of the types of Australian Hymenoptera
described by Alexandre Arsene Girault: IV.
Chalcidoidea species N-Z and genera with advisory
notes and corrigenda. Memoirs of the Queensland
Museum 22:319-739.
DALLA TORRE, K. W. VON

1898 Catalogus Hymenopterorum hucusque descripto-
rum systematicus et synonymicus. Vol. V,
Chalcididae et Proctotrupidae. Leipzig. 598 pp.
DARLING, D. C.

1983a Systematic studies of the Perilampidae (Hymen-
optera: Chalcidoidea). Ph.D. diss., Cornell Univer-
sity. 278 pp.

1983b A review of the New World species of
Euperilampus (Hymenoptera: Chalcidoidea), with
notes about host associations and phylogenetic rela-
tionships. Questiones Entomologicae 19:1-40.

1986 Revision of the New World Chrysolampinae
(Hymenoptera: Chalcidoidea). Canadian Entomo-
logist 18:913-940.

1988a Comparative morphology of the labrum in
Hymenoptera: the digitate labrum of the
Perilampidae and Eucharitidae (Chalcidoidea).
Canadian Journal of Zoology 66:281 1-2835.

1988b A review of the genus Kromheinus (Hymenoptera:
Perilampidae) with a reexamination of generic lim-
its and phylogenetic relationships and the descrip-
tions of generic limits and the descriptions of two
new species. Journal of the New York Entomolo-
gical Society 96:63-8 1.

1992 The life history and larval morphology of

Apcrilcimpus (Hymenoptera: Chalcidoidea:
Philomidinae), with a discussion of the phylogenet-
ic affinities of the Philomidinae. Systematic
Entomology 17:331-339.

DARLING. D. C. and T. MILLER

1991 Life history and larval morphology of
Chrysolampus sisymhrii (Hymenoptera: Chalci-
doidea: Chrysolampinae) in western North
America. Canadian Journal of Zoology 69:
2168-2177.

DAS, G. M.

1963 Preliminary studies on the biology of Orasema
assc'ciator Kerrich (Hym., Eucharidae) parasitic on
Pheidole and causing damage to leaves of tea in
Assam. Bulletin of Entomological Research
54:373-378.

DAVIS, L. R., Jr., and D. P. JOUVENAZ

1990 Obeza floridana, a parasitoid of Camponotus
ahdominalis floridanus from Florida (Hymen-
optera: Eucharitidae, Formicidae). Florida Ento-
mologist 73:335-337.

DE SANTIS, L.

1968 Una nueva especie de ''Orasema" del Uruguay
(Hymenoptera: Eucharitidae). Revista Sociedad
Entomologica Argentina 31:1-3.

EADY, R. D.

1968 Some illustrations of microsculpture in the
Hymenoptera. Proceedings of the Royal Entomo-
logical Society 43:66-72.

FABRICIUS.J. C.

1 804 Systema Piezatorum. Brunsvigae, Reichard. 439 pp.
FAHRINGER, J. and F. TOLG.

1912 Beitrage zur Kenntnis der Lebensweise und
Entwicklungsgeschichte einiger Hautfliigler.
Verhandlungen des naturforschenden Vereins in
Brunn 50:242-269.

GAHAN. A. B.

1940 A contribution to the knowledge of the Eucharidae
(Hymenoptera: Chalcidoidea). Proceedings of the
United States National Museum 88:424-458.
GAHAN. A. B. and M. M. FAGAN

1923 The type species of the genera of Chalcidoidea or
chalcid flies. Bulletin of the United States National
Museum 124:1-173.
GEMIGNANI, E. V.

1933 La familia "Eucharidae" (Hymenoptera: Chal-
cidoidea) en la republica Argentina. Anales del
Museo Nacional de Historia natural de Buenos
Aires 37:477^93.

1937 Nueva nota sobre la familia Eucharidae (Hy-
menoptera: Chalcidoidea). Anales del Museo Ar-
gentino de Ciencias Naturales "Bernardino Riva-
davia" 39:157-166.

1947 Nuevas especies de la familia Eucharidae (Insecta,

122

Hymenoptera: Chalcidoidea). Comunicaciones del
Miiseo Argentine de Ciencias Naturales "Bernard-
ino Rivadavia" 1:1-15.

GHESQUIERE, J.

1946 Contributions a I'etude des Microhymenopteres du
Congo beige X-XI. Revue de Zoologie et de
Botanique Africa 39:367-373.

GIBSON. G. A. P.

1985 Some pro- and mesothoracic structures important
for phylogenetic analysis of Hymenoptera. with a
review of terms used for the structures. Canadian
Entomologist 117:1395-1443.

1986 Evidence for monophyly and relationships of
Chalcidoidea. Mymaridae, and Mymarommatidae
(Hymenoptera: Terebrantes). Canadian Entomo-
logist 118:205-240.

1989 Phylogeny and classification of Eupelmidae, with
revision of the world genera of Calosotinae and
Metapelmatinae (Hymenoptera: Chalcidoidea).
Memoirs of the Entomological Society of Canada
149:1-121.

1990 Revision of the genus Macioneura in America
north of Mexico (Hymenoptera: Eupelmidae).
Canadian Entomologist 122:837-873.

GIRAULT, A. A.

1913a More new genera and species of chalcidoid
Hymenoptera from Paraguay. Archiv fiir Natur-
geschicte 79, Abt. A, Hft. 6:51-69.

1913b New genera and species of chalcidoid Hymenoptera
from the South Australia Museum: Adelaide.
Transactions of the Royal Society of South Aus-
tralia 37:67-1 15.

1915 Australian Hymenopteran Chalcidoidea Eucharidae.
Memoirs of the Queensland Museum 4:225-237.

1922 New chalcid flies from eastern Australia (Hymen-
optera). Insector Inscitae Menstruus 10:100-108.

1929 Notes on and descriptions of chalcid wasps in the
South Australian Museum. Transactions of the
Royal Society of South Australia 53:309-346.

1932 New pests from Australia, X. Reprinted in Gordh.
G., A. Menke. E. C. Dahms, and J. Hall (1979), The
privately printed papers of A. A. Girault. Memoirs
of the American Entomological Institute
28:286-291.

1934 Eucharidae, Cynipidae, Proctotrupidae et Thy-
sanoptera nova australiensis. Reprinted in Gordh,
G., A. Menke, E. C. Dahms, and J. Hall (1979), The
privately printed papers of A. A. Girault. Memoirs
of the American Entomological Institute
28:306-307.

1936 Terror-errors, and novitates of pterygota (or earth
realities not state-bound). Reprinted in Gordh, G.,
A. Menke, E. C. Dahms. and J. Hall (1979), The
privately printed papers of A. A. Girault. Memoirs
of the American Entomological Institute
28:322-325.

1940 New genera and species of Australian Elasmidae
and Eucharitidac. Revisla de la Socicdad
Entomologica Argentina 10:321-326.

GOODPASTURE. C.

1975 Comparative courtship behavior and karyology in
Monodontomerus (Hymenoptera: Torymidae).
Annals of the Entomological Society of America
68:391-397.
GORDH, G.

1979 Superfamily Chalcidoidea. In Krombein. K. V.. P.
D. Hurd, Jr., D. R. Smith, and B. D. Burks, eds..
Catalog of Hymenoptera in America north of
Mexico, Vol. 1, Symphyta and Apocrita (Parasi-
tica), pp. 743-748. Washington, D.C., Smithsonian
Institution Press.
GRAHAM, M. W. R. de V.

1969 The Pteromalidae of north-western Europe
(Hymenoptera: Chalcidoidea). Bulletin of the
British Museum (Natural History), Entomology
Supplement 14:1-908.
GRESSITT, J. L.

1956 Some distribution patterns of Pacific island faunae.

Systematic Zoology 6:12-32.
1982 Zoogeographic summary. In Gressitt, J. L., ed.,
Biogeography and ecology of New Guinea, Vol. 2,
pp. 897-919. The Hague, Junk.

HABU, A.
1960

HARRIS, R.
1979

HEDQVIST,
1978

HEPPNER, J
1982

HERATY.J.
1985

1986

1989

1990

1992

A revision of the Chalcididae (Hymenoptera) of
Japan with descriptions of sixteen new species.
Bulletin of the National Institute of Agricultural
Sciences, Tokyo, Series C, 11:13 1-358.

A.
A glossary of surface sculpturing. Occasional
Papers in Entomology, State of California
Department of Food and Agriculture 28: 1-31.

K.J.

Some Chalcidoidea collected in the Philippines,
Bismark, and Solomon Islands. 2. Eucharitidae,
with keys and check-lists to Indo-Australian genera
(Insecta, Hymenoptera). Steenstrupia 4:227-248.

. B. and G. LAMAS
Acronyms for world museum collections of insects,
with an emphasis on Neotropical Lepidoptera.
Bulletin of the Entomological Society of America
28:305-315.

M.
A revision of the Nearctic Eucharitinae (Hy-
menoptera: Chalcidoidea: Eucharitidac). Proceed-
ings of the Entomological Society of Ontario
85:61-103.

Pseiidochakura (Hymenoptera: Eucharitidae): a
New World genus of chalcidoids parasitic on ants.
Systematic Entomology 1 1:183-212.
Morphology of the mesosoma of Kapala
(Hymenoptera: Eucharitidae) with emphasis on its
phylogenetic implications. Canadian Journal of
Zoology 67:1 15-125.

Classification and evolution of the Oraseminae
(Hymenoptera: Eucharitidac I. Ph.D. diss.. Texas A
& M llnivcrsity. 336 pp.
Revision of the genera GoUumiclla Hcdqvisi and

123

Anoicisiniii Boufck (Hymenoptera: Eucharitidae).
Invertebrate Taxonomy 6:.SS3-6()4.

HERATY. J. M. and K. N. BARBl-R

1990 Biology of Oheza floridana and Pscudochakura
gihhosci (Hymenoptera: Eucharitidae). Proceedings
of the Entomological Society of Washington
92:248-258.

HERATY. J. M. and D. C. DARLING

1984 Comparative morphology of the planidial larvae of
Eucharitidae and Periiampidae (Hymenoptera:
Chalcidoidea). Systematic Entomology 9:309-328.

HERATY. J. M.. D. P. WOJCIK. and D. P. JOUVENAZ

1993 Species of Oiascniu parasitic on the Solcnopsis
Saevissima-comp\ex in South America (Hymen-
optera: Eucharitidae, Formicidae). Journal of
Hymenoptera Research 2:169-182.

HEYDON. S. L.

1989 Review of Nearctic Rhinocoelia and CalUmerismus
with a discussion of their phylogenetic relationships
(Hymenoptera: Pteromalidae). Journal of the New
York Entomological Society 97:347-357.

HOLLDOBLER. B. and E. O. WILSON

1990 The ants. Cambridge, Belknap Press of Harvard
University Press. 732 pp.

HOWARD, L. O.

1896 On the Chalcididae of the island of Grenada, B. W.
I. Journal of the Linnean Society 86:129-178.

1897 Report on the Chalcididae of the subfamilies
Chalcidinae, Eucharinae, Perilampinae, Encyrtinae,
Aphelininae, Pireninae, Elasminae, and Elachis-
tinae. Journal of the Linnean Society 6, Zoology
26:79-108.

HUBER, J. T.

1988 The species groups of Gonatoceriis Nees in North
America with a revision of the sulphwipes and ater
groups (Hymenoptera: Mymaridae). Memoirs of the
Entomological Society of Canada 141:1-109.

HUSAIN, T. and M. AGARWAL

1983 Indosema indica gen. et sp. nov. (Hymenoptera:
Eucharitidae) described from India. Bolletino del
Laboratorio di entomologia agraria "Filippo Sil-
vestri" 40: 103-107.

JOHNSON, J. B., T. D MILLER, J. M. HERATY, and
F. W. MHRICKEL
1986 Observations on the biology of two species of
Orasenia (Hymenoptera: Eucharitidae). Proceed-
ings of the Entomological Society of Washington
88:542-549.

KEAST. A.

1981 Distributional patterns, regional biotas, and adapta-
tions in the Australian biota: a synthesis. /;; Keast,
A., ed.. Ecological biogeography of Australia, Vol.
3, pp. 1891-1997. The Hague. Junk.
KERRICH, G. J.

1963 Descriptions of two species of Eucharitidae damag-
ing tea, with comparative notes on other species
(Hymenoptera: Chalcidoidea). Bulletin of Ento-
mological Research 54:365-371.
KIRBY, W. P.

1886 A synopsis of the genera of the Chalcidoidea, sub-
family Eucharinae, with descriptions of several new
genera and species of Chalcidoidea and Tenthre-
dinidae. Journal of the Linnean Society, Zoology
20:28-37.
KISTNER, D. H.

1979 Social and evolutionary significance of social insect
symbionts. In Hermann, H. R.. ed.. Social insects,
Vol. 1, pp. 340-414. New York, Academic Press.
KONIGSMANN, E.

1978 Das phylogenetische system der Hymenoptera. Teil
3: Terebrantes (Unterordnung Apocrita). Deutsche
Entomologische Zeitschift 23:253-279.

LAING, J. E. and J. M. HERATY

1981 The parasite complex of the overwintering popula-
tion of Epihlcma scuddcriana (Lepidoptera:
Olethreutidae) in Southern Ontario. Proceedings of
the Entomological Society of Ontario 1 12:59-67.
LASALLE, J.

1987 New World Tanaostigmatidae (Hymenoptera:
Chalcidoidea) Contributions of the American
Entomological Institute 23:1-181.

ISHIL T.

1932 Some Philippine Eucharids with notes on their
oviposition habits. Bulletin of the Imperial
Agricultural Experimental Station, Nishigahara
3:203-212.
ISHIi, T. and S. NAGASAWA

1941 Description of a new eucharid from Palau Islands.
Tenthredo 3:291-294.

JOHNSON. D. W.

1988 Eucharitidae (Hymenoptera: Chalcidoidea): biology
and potential for biological control. Florida
Entomologist 71:528-5.37.

MADDISON. W. P.. M. J. DONOGHUE. and D. R. MADDISON

1984 Oulgroup analysis and parsimony. Systematic
Zoology 33:83-193.
MANN, W. M.

1914 Some myrmecophilous insects from Mexico.
Psyche 21:171-184.
MATSUDA, R.

1980 The insect head. Memoirs of the American
Entomological Institute 4: 1-334.
MICHENER. C. D.

1969 Immature stages of a chalcidoid parasite tended by
allodapine bees (Hymenoptera: Periiampidae and
Anthophoridac). Journal of the Kansas Entomolog-
ical Society 42:247-250.

124

MITTER, C. and D. R. BROOKS

1983 Phylogenetic aspects of coevolution. //; Futuyma,
D. J., and M. Slatkin, eds., Coevolution, pp. 65-98.
Sunderland, Mass., Sinauer Associates.

NOONAN. G. R.

1985 The influences of dispersal, vicariance, and refugia
on patterns of biogeographical distributions of the
beetle family Carabidae. /// Ball, G. E., ed.,
Taxonomy, phylogeny and zoogeography of beetles
and ants, pp. 322-350. Series Entomologica 33.
Dordrecht, Junk. 514 pp.

PARKER, H. L.

1937 The oviposition habits of Stilbula cynipiformis

Rossi (Hymenoptera: Eucharidae). Proceedings of

the Entomological Society of Washington 39:1-3.
1942 Oviposition habits and early stages of Orasema sp.

Proceedings of the Entomological Society of

Washington 44:142-145.

REICHENSPERGER, A.

1913 Zur Kenntnis von Myrmecophilen aus Abessinien,
I. Zoologische Jahrbiicher, Abteilung fiir
Systematik. Geographie und Biologie der Tiere
35:185-218.
REIMER, N. J.

1988 Predation on Liothrips ulrichi Karny (Thysan-
optera: Phlaeothripidae): a case of biotic interfer-
ence. Environmental Entomology 17: 132-134.
RICHARDS, O. W.

1977 Hymenoptera. Introduction and key to families.
Handbooks for the Identification of British Insects
6:1-103. London, Royal Entomological Society.
RIEK, E. F.

1970 Hymenoptera (Wasps, bees, ants). /// The insects of
Australia, pp. 867-959. Melbourne, Melbourne
University Press.
RISBECJ.

1952 Contribution a Tetude des Chalcidoides de
Madagascar. Memoires de ITnstitut scientifique de
Madagascar, Ser. E, 2:1-^49.
1958 Chalcidoides nouveaux d'Afrique du Sud.
Occasional Papers, National Museum of Southern
Rhodesia, Bulawayo 3. no. 22B: 147-162.

SCHAUFF. M. E.

1984 The Holarclic genera of Mymaridae (Hymenoptera:
Chalcidoidea). Memoirs of the Entomological
Society of Washington 12:1-67.

1991 The Holarclic genera of Enlcdoninae (Hymen-
optera: F^ulophidae. Contributions of the American
Entomological Institute 26: 1 109.
SrnUII, R. T and G M. .STONF.DAHI,

1986 Historical biogeography in the hulo-Pacific: a
cladistic approach. Cladistics 2:337-355.

SHATTUCK, S. L.

1992 Higher classification of the ant subfamilies
Aneuritinae, Dolichoderinae and Formicinae (Hy-
menoptera: Formicidae). Systematic Entomology
17:199-206.
SMITH, H. S.

1912 Technical results from the gipsy moth parasite labo-
ratory. IV. The chalcidoid genus Perilampus and its
relations to the problem of parasite introduction.
United States Department of Agriculture Technical
Series 19:33-69.

1917 The habit of leaf-oviposition among the parasitic
Hymenoptera. Psyche 24:63-68.
SNEHALATHA, S. and T. C. NARENDRAN

1992 On a new species of Orasema Cameron
(Hymenoptera: Eucharitidae), with a key to Indian
species. Journal of the Bombay Natural History
Society 89:355-357.
SNODGRASS, R. E.

1935 Principles of insect morphology. New York,
McGraw-Hill. 667 pp.

1956 Anatomy of the honey bee. Ithaca, Comstock Publ.
Associates, Cornell University Press. 334 pp.
STEYSKAL, G.

1980 Errors in nomenclatural grammar. //; Krombein et
al.. Catalog of Hymenoptera in America north of
Mexico (1979). Sphecos 3:43-46.
STRAND. R.

1942 Miscellanea nomenclatoria zoologica et palaeonto-
logica X-XII. Folia Zoologica et Hydrobiologica
ll:386--*02.
SWOFFORD. D. L.

1990 PAUP: Phylogenetic Analysis Using Parsimony,
Version 3.0n. Computer program distributed by the
Illinois Natural History Survey, Champaign,
Illinois.

TAYLOR. R. W.

1978 Nolhontyrmecia macrops: a living-fossil ant redis-
covered. Science 201:979-985.

VAN PELT, A. F.

1950 Orasema in nests of Pheidolc dentata Mayr
(Hymenoptera: Formicidae). Entomological News
71:161-163.
VANDER MEER, R. K.. D. P. JOUVENAZ, and D. P. WOJCIK

1989 Chemical mimicry in a parasitoid (Hymenoptera:
Eucharitidae) of fire ants (Hymenoptera: Formici-
dae). Journal of Chemical Ecology 15:2247-2261.

WALKER. F.

1839 Monographia Chalciditum, Vol. 2. London. 100 pp.

1846 Part I — Chalcidiles. List of the specimens of
hymenopterous insects in the collection of the
British Museum. Pt. I chalcidiles. Pt. 2 additional
species. London. 237 pp.

1862 Mr. F. Walkers notes on chalcidiles and characters

12.5

(if uiulcscribod species. Transactions of ihe Ento-
mological Society of London 1:380-385.

WARD. P. S.
1990

The ant subfamily F'seudomrymecinae (Hymen-
optera: Foniiicidae): generic revision and relation-
ship to other formicids. Systematic Entomology
15:449^89.
WATANABE. C.

1958 Hymenoptera: Eucharitidae. Insects of Micronesia
19:19-34. Honolulu, Bishop Museum.
WATERSTON. J.

1916 Notes on African Chalcidoidea, IV. Bulletin of Ent-
mological Research 6:413^23.
WHEELER. G. C. and E. W, WHEELER

1937 New hymenopterous parasites of ants (Chalci-
doidea, Eucharidae). Annals of the Entomological
Society of America 30:171-172.
WHEELER. W. M.

1907 The polymorphism of ants with an account of some
singular abnormalities due to parasitism. Bulletin of
the American Museum of Natural History 23:1-93.
1910 Ants: their structure, development and behavior.
New York, Columbia University. 663 pp.
WILLIAMS. R. N. and W. H. WHITCOMB

1973 Parasites of fire ants in South America. Proceedings
of the Tall Timbers Conference on Ecology,
Animal Control and Habitat Management 5:49-59.

WILSON. E. O.

1971 The insect societies. Cambridge, Belknap Press of
Harvard University Press. 584 pp.

WILSON, E. O.. F. M. CARPENTER, and W. L BROWN. Jr.

1967 The first Me.sozoic ants. Science 157: 1038-1040.

WILSON. T. H. and T. A COOLLY

1972 A chalcidoid pianidium and an entomophilic nema-
tode associated with the western flower thrips.
Annals of the Entomological Society of America
65:414^18.

WOJCIK, D. P.

1988 Survey for biocontrol agents in Brazil — a final
report, with comments on preliminary research in
Argentina. Proceedings of the Imported Fire Ant
Conference (1988):50-62.
WOOLLEY.J. B.

1988 Phylogeny and classification of the Signiphoridae
(Hymenoptera: Chalcidoidea). Systematic Entomo-
logy 13:465-501.

YOSHIMOTO, C. M.

1977 Revision of the Diparinae (Pteromalidae: Chal-
cidoidea) from America north of Mexico. Canadian
Entomologist 109:1035-1056.

126

Figures

127

r-E

C

c

HZ

C

C
C

k:

CHRYSOLAMPINAE
PERILAMPINAE

TIMIODERUS
INDOSEMA
ORASEMORPHA
ORASEMA ST
ORASEMA AS
ORASEMA GL
ORASEMA KO
ORASEMA VA
ORASEMA Ul
ORASEMA CO

PSILOCHARIS AE
PSILOCHARIS AF
PSILOCHARIS TH
NEOLOSBANUS GE
NEOLOSBANUS PU
NEOLOSBANUS TO
NEOLOSBANUS PA
NEOLOSBANUS GR

GOLLUMIELLA
ANORASEMA

STILBULA

OBEZA

PSEUDOCHALCURA

STILBULOIDA

EUCHARIS

PSEUDOMETAGEA

PROPSILOGASTER

TRICORYNA

SCHIZASPIDIA

KAPALA

ISOMERALA

CH MONTANA

AUSTEUCHARIS

CHALCURA

O
U

>

m

>
m

■D
(/>

|—

o
o

>

m

c
o

>

DO

m

c
o

>

H
Z
>

m

Fig. I. Strict consensus cladogram for reduced data set using adult characters and egg shape (characters
1-48; Table 2). Group names based on consensus cladogram in ligure 2. Length = 156, 18 trees, consistency
index = 0.5 1 , retention index = 0.79.

128

CHRYSOLAMPINAE
PERILAMPINAE

13'
1421

15'

21

5021

3 32' 42 43^ 44^

38' 41

■ •■•■•

TIMIODERUS

82 35' 38°
1 32 1 7 223 23 24 36 48 !»•■> 9 •••• INDOSEMA

1 2 14''152 25 27'30 33 433
♦ ORASEMORPHA

• • ■

39' 49 592 60

• •

37 62

36 433

-137 44'

28

J 15020233

■O

48 58 443

Lk>.^>

15° 43'

6 10 202 28 36 37

3

26'
46
61

9' 29' 393

45'

1 1 292 46

92

• I • O

49 12 35' 36

■O

10 12 42 43' 44

{1

• I •

58 42 442

ORASEMA ST
ORASEMA AS
ORASEMA GL
ORASEMA KO
ORASEMA VA
ORASEMA Ul
ORASEMA CO

PSILOCHARIS AE
PSILOCHARIS AF
PSILOCHARIS TH
NEOLOSBANUS GE
NEOLOSBANUS PU
NEOLOSBANUS TO
NEOLOSBANUS PA
NEOLOSBANUS GR

O
>

m

z
>
m

w

r-
O
O

X

>

19' 45'

■O

• •

18 23 36
— O^

■ 262 •37 595 Q3^i
O 49 52 I 18 36

40 42

r<MH>-
4" 20^452

2 18° 32'

■O

5- 352 45°
I I I I I

■O

58 8^ 16 27' 34 56

• I I •

15-^ 34 36 59

oJi631°35250

47'l__#-#-_

50^53

72 8334 43'47'l

17 27'

■I 1 I 120" 30 32°

20° 62

5^ 17
-•— I-

7' 17

593

• • 0-0

312 432 420 440

20^ 222 30 3^3 3g

42 47'

GOLLUMIELLA

m

c

ANORASEMA

0

X

STILBULA

>

33

OBEZA

PSEUDOCHALCURA
STILBULOIDA

m

c
0

>

m

EUCHARIS

>

PSEUDOMETAGEA

5

— i

TRICORYNA

z

PROPSILOGASTER

AUSTEUCHARIS

SCHIZASPIDIA

KAPALA

ISOMERALA

CH MONTANA

CHALCURA -1

-

6 72 20" 37

Fici. 2. Strict consensus cladogram for complete data set (characters 1-62; Table 2). Length = 183. 20 trees,
consistency index = 0.53. retention index - 0.79. Character states listed and described in Table 2 and
Appendix 1 . Multiple states for tenninal OTUs and outgroup states are not shown. Superscripts refer to states
of niultistale characters, otherwise derived slates are assumed to be I . Characters marked by a bar had tiiffer-
ent distributions on different tree topologies. Squares indicate consistency index of 1.0, black circles are
derived states that are unique for the clade but that show homoplasy on the tree, gray circles show homo-
plasy within the clade, and open circles indicate reversals (shading also applies to bars).

129

SaoCDOCDO OO CD O CZ) a O C=3 CD
OCX

Figs. 3-6. Oiascma. 3. Oviposition scars on Acacia, O. sp. nr hakcri. Texas, U.S.A. 4. Egg in oviposition
puncture. 5. Planidia of O. viridis on immature thrips. 6. First instar of O. xanihopits. dorsal and lateral
aspect of unfed planidium. Scale in mm. Abbreviations: T = tergite.

130

Figs. 7-12. Oiasema. 7. O. xaiuhopus on Solcnopsis invicta: a, external planidium (arrow) on second-inslar
host; b. distended first instar internal in mature host. 8-9. O. sp. nr costaricensis on Pheidole dentata: 8.
Distended first insiar internal in mature host; 9. First instar external on host pupa. 10. O. xaiuhi^piis early
third instar on host pupa. 1 1 12. O. m costaricensis: 1 1. third-instar larva; 12. 9 pupa. Scale in mm.

\}\

132

Figs. 17-26. 17-25. Neoloshanu.s palf>rayci: 17. Mature egg; 18. Freshly deposited egg in oviposition cham-
ber in leaf tissue: 19. .Scarified plant tissue surroundinj: mature egg; 20. Partially distended first-instar larva
attached externally to lateral region ol head ot host ant: 21. Second instar: 22. Second instar on prepupal
stage of host within cocoon: 23. Second instar on partially developed pupa of host: 24. Third instar; 25. 9
pupa. 26. Neoloshanus f^cmma. 9 pupa. Scale in mm.

33

FiCiS. 27-38. Timioderus. 27. 7. cicuminatus. habitus. 9. 28. T. peridcntatus. head, 9. 29. 7". aciiminatus.
ovipositor. 30-31. T. ramosus: 30. Head, 6\ 31. Ovipositor. 32. T. rcfrin^cns. head. 9: frontal view (a);
mandible closeup (b). 33-34. T. ramosus. head, 9: 33. Frontal view; 34. Dorsal view. 35. T. coronula,
propodcuin and motasoma in lateral view.c?. 36-37. 7". peridcntatus: 36. Mesothorax in dorsal view. 9; 37.
Petiole in dorsal view, 9. 38. /'. coroinila. head in lateral view, 6 . Abbreviations; IT = lateral teeth; MS = malar
.space length; Ms, = second meta.somal stemite; SAR = subapical ridge; IV = first valvula, 2V = second valvula.

134

Figs. 39-49. Timioderus. 39. T. peridentatus, partial head in posterolateral view, 9 . 40. T. ramosus, left
mandible, 6. 41. T. refringens. oral fossa showing labrum (LB), 6. 42. T. acuminatus. labrum and
mandibles. 43. T. refringens, forewing. 9. 44. T. ramosus, forewing. 9. 45. T. refringens. antenna, 6 . 46-47.
T. ramosus, antenna: 46.9; 47. d. 48-49. T. acuminatus, antenna; 48. 9; 49. 6 . Abbreviations: CC = costal
cell; F = flagellomere 2; FLG = flagellum; FUN = funicle; HP = humeral plate; LB = labrum; MV = marginal
vein; PMV = postmarginal vein; SMV = submarginal vein; SV = stigma) vein.

135

Figs. 50-54. Indosema indica. 50. Mesothorax in dorsal view, 9. 51. Habitu.s. 9. 52. Ovipositor in lateral
view. 53. Petiole in dorsal view, 9. 54. Posterior region of mesosoma and metasoma in lateral view, 6.
Abbreviation: GS = gonostylus.

136

^"^"^c,.

Figs. 55-68. Oioscmoipha. 55. O. xeiuades. head in frontal view, 9. 56. O. triilcniaia. lower face and
mandibles, 9. 57. O. xcniades. head and mesothorax in dorsal view, 9. 58. O. thdcniata. petiole in dorsal
view. 59-61. O. xeniades: 59. Head in lateral view. 9; 60. Forewing, d; 61. Ovipositor (a) and habitus (b).
62. O. myrmicae, antenna, 9 (holotype). 6.1. O. varideiitala. nicsothorax in dorsal view, 9 (holotypc). 64. O.
pyttalus. antenna, 9. 65. O. sparsepilosits. antenna. S . 66-67. O. pyttalus: 66. Head in frontal view, 9 (holo-
type): 67. Petiole in dorsal view. 9. 68. O. spaiscpilosus. forewing, 9. Abbreviations: AX = axilla: FL = fre-
nal line: FR - freniini: l.L = lateral lobe: ML = niidlobc of mcsosculum: NOT = notaulus: SCT = .scutellum;
SSS = scutoscutellar sulcus: TSA = transscutal articulation.

37

Figs. 69-79. Orasemorpha. 69. O. sparsepilosus, head and mesosoma in dorsal view. 9. 70-71. O. chhoies:
70. Mesothorax in dorsal view, cJdiololype); 71. Antenna, 9(HT off. viridis). 72. O. ilicleniaia. antenna. 9
(iectotype). 7.\ O. ciihoies. head in frontal view. c5(hololype). 14-19. O. didentata: 74. Ventral region of
gaster, 9; 75. Petiole and first gastral slernite in ventral view; 76. Head in frontal \iew, 9; 77. Head and
mesosoma in dorsal view, 9; 78. Posterior region of mesosoma and metasoma. 6 \ 79. Forewing.9.
Abbreviations: AR = anterior region; C = constriction; Ms. = second metasomal sternite; PRP = propodeum;
PET = petiole.

138

Figs. 80-86. Orasema. 80. O. communis, head in frontal view. 9. 81. O. seyrini. head in frontal view. 9
(HT). 82. O. communis, ovipositor in lateral view. 83. O. seyrifyi, head and mesosoma in lateral view, 9
(HT). 84-85. O. communis, antenna: 84.9; 85. d. 86, O. seyrigi, forcwing.

139

cClODoaixii

87

Figs. 87-97. Orasema. 87. O. itichancoi. antenna (lectotype). 88. O. ishii. antenna. 9. 89-92. O. nijiulosa:
89. Antenna, 9; 90. Head in frontal view, d; 91. Meso.soma and metasoma in lateral view, 6\ 92. Head and
mesothorax in dorsal view, d(holotype). 93. O. riif>iil()sa, ovipositor in lateral view. 94. O. uichancoi. base of
petiole in dorsal view. 95. O. pronwcea, head in lateral view, 6 . 96-97. Forewing: 96. O. ruf^ulosa, 6\ 97. O.
uichamni, forewing, 9 (lectotype). Abbreviations: MD = malar depression; OCG = ocellar-ocular groove.

140

Figs. 98-105. Oiascma stiiatosoma. 98. Head and mcsosoma in lateral view, 9 (holotype). 99. Ovipositor in
lateral view. 100. Antenna. 6. 101. Head and mesothorax in dorsal view. 9 (holotype). 102. Head in frontal
view. 6. 103. Genitalia and Ms, in ventral view. 104. Forewing.9. 10.5. Petiole in dorsal view. 9.
Abbreviations: AED = aedeagus; DG = digitus: PM = paramere.

141

/Ill

Figs. 106-116. Oiascma. 106-107. Male genitalia in ventral view: 106. O. uicluimoi: 107. O. ko{>hisiana.
108-109. Antenna: 108. O. koghisiana, 9: 109. O. initiator. 9. 1 10. (9. koi>hisiana. forewing. 9. 111-112.
O. asscciator. 9: 111. Mesothorax in dorsal view; 112. Head in frontal view. 113-116. O. nii^ra. 9: 113.
Posterior region of mesosoma and metasoma; I 14. Head in frontal view: 1 1!^. Head in lateral view; 1 16.
Ovipositor in lateral view.

142

Figs. 1 17-125. Orasenui. 117-1 18. (9. initiator, forewing: 117. 6; 1 18.9. 1 19. O. i>lahia, base of forewing.
9. 120-121. Base of petiole in dorsal view: 120. O. kofihisiana. 9; 121. O. synempora. 9. 122. O. valiiius.
forewing. 9. 123-12.'>. Antenna. 9: 123. O. synempora: 124. O. \o/.i,'/h.v; 125. O. glabra. Abbreviation: SP =
speculum.

143

Figs. 126-132. Orasema. 126-128. O. i^lahra. 9 : 126. Habitus; 127. Head in frontal view: 128. Ovipositor in
lateral (a) and dorsal (b) views. 129. O. synempora, head and mesothorax in dorsal view. 9. 130. O. i>lahra.
head and mesothorax in dorsal view, 9. 131.0. vali;iits. genitalia. 6 . 132. O. i>l(ihra. base of petiole in dor-
sal view. Abbreviations: AX AS = axillular sulcus; 1 V-2V = first and second vaivulae.

144

Figs. 133-143. Psilocharis. 133. P. pacifica, head in frontal view, 9. 134. P. monilicera, head in lateral
view, S . 135-136. P. afro. 6: 135. Head in lateral view; 136. Head in dorsal view. 137. P.pacifica. head and
mesothorax in dorsal view. 138-143. Antenna: 138. P. joanncac. 9; 139. P. llwocles, 9; 140. P. afro. 9;
141. P. thcinles. 6: 142. P. hypena (Taiwan). 6: 143. P. hypcna (Borneo). 6 . Abbreviation: OC = occipital
carina.

143

Figs. 144-151. Psilocharis. 144-145. Base of petiole in dorsal view, 9: 144. P. acnii^ma; 145. P. theocles.
146-148. Forewing,?: 146. P. theocles: 147. P. Iiypcna: 148. P. afro. 149-150. P. afni: 149. Apical margin
of hypopygium in ventral view. 9; 150. Ovipositor in ventral view. 151. P. theocles. 6 genitalia.

146

Figs. 152-161. Neohshamis. 152. N. wushcanus. lateral habitus,?. 153. N. sicmma. head in trontal view, 9.
154-157. yv. wushcanus: 154. Head in frontal view, ?; 155. Head and mesothorax in dorsal view, 9; 156.
Metanotum and propodcum in rear view. 9; 157. Antenna, 6. 158. N. apoanus. antenna. 6 . 159. N. ficmnui.
base of petiole in dorsal view. 9 . 160. N. wushcanus. forewing, 9 . 161 . /V. apoanus. forewing margin, (5 .

147

Figs. 162-169. Neoloshanus. 162. N. laevicep.s. forewing,9. 163. A', laeviceps, 6 genitalia and Ms« in ventral
view. 164-165. Forewing: 164. A', violaccus. 6: 165. N. i}(ili>r(i\ri. 9. 166-169. N. purpureovcniris: 166.
Antenna, 9; 167. Basal articles of antenna. 6\ 168. Antenna, c?; 169. Forewinu. 9.

148

Figs. 170-175. Neoloshanus townesi: 170. Pedicel and basal flagellomere. 9 . 171. Antenna. 9. 172.
Mctasoma, 9. 173. d genitalia. 174. Ovipositor in lateral view. 175. Forewing, 9.

149

Figs. 176-186. Neoloshanus. 176-1.77. N. kokiiieamis. 9: 176. Head in tronlal view; 177. Head in lateral
view. 178-179. Head in dorsal view, 9: 178. N. pilosns; 179. N. sloreyi. 180-186. Antenna: 180. N. pilosus.
6; 181. A', palgravei. 6\ 182. N. palgravei. 9; 183. N. laeviceps, 9; 184. N. laeviceps, 6; 185. N. violaceus,
6; 186. N. amipetus. 6 . antenna (a) with enlargement of basal articles (b).

150

Figs. 187-192. Head in frontal view. 187. Orasemorpha erihotes, 9. 188. Orasema uichaiuoi. 9 . 189-190.
Orascma koi>hisiana. 9. 191. Orasema initiator, 9. 192. Orasema valijiiis. 9 . Abbreviations: A = anlc-
clypcus; CLY = clypcus; gl = glabrate; re = reticulate; ru = rugose.

LSI

Figs. 193-198. Psiloclmns. head in rroniai view. 193. P.joaiuwae. 9: 194. P. tlwocles. 9. 195. P. afra. 9.
196. P. hypena (Borneo), 9. 197-198. P. hypena (Taiwan), 9. Abbreviations: A = antecylpeu.s; AS = ante-
clypeal seta; CLY - clypeus; LS = labral seta; gs = glabrous.

152

Figs. 199-204. Neoloshanus. head in frontal view. 199. N. tnwnesi. 9. 200. N. laniccps. 9. 201. /V. /nil-
iiiavci (Malayan) 9. 202. N.pilosus. 9. 203-204. A', ncpalcnsis. 9.

153

FuiS. 203-210. Ncoloshaiuis. head in tronlal view. 205. N. pal^inivi'i (Australian). 9. 206. N. violacciis. 6.
207-208. N. aiuipcius. 6: 207. Head: 208. Closciip of face lateral to toruliis. 2(W-210. A'. piirpweoYcnins.
9: 209. Head: 210. Closeup of clypeal area. Abbreviations: C = clypeogenai sulcus. CLY = elypeus. E =
epislomal sulcus, \- = I'rontogenal sulcus: pp = pilose-punclate.

154

Figs. 21 1-216. Antennal flagcllomcrcs. 21 1-212. Orasema kof^hi.siana. (5.213. Psilocluiiis iheocles. 9 . 214.
Neolo.shaniis palf>ravei. 9. 213-216. Neoloshanus anapclus. Abbreviations: F!~6 ~ napcllomercs 1-6; MPS
= multiporous plate sensillum; FED = pedicel.

LS.S

FuiS. 217-222. Mcsosoma in lateral view. 217. Orascmorpha crihotes. 9. 218. Oiascnia uichancoi. 9. 219.
Orascma ko}>hisiaiui. 6 . 220. Oiascnia initiator, 9. 221. Oiascma valgius, 9. 222. Psilocharis afra. 9.
Abbreviations: ca = callus; fo = foveate; im = imbricate; pre = prepectus; PS = pronotal sulcus; pst = proepi-
stemum.

156

Figs. 223-228. Mesosoma in lateral view. 22.3. Psilocharis thcoclcs, 9 . 224. Psilocharis hypeiia (Taiwan).
9. 225. Psilocharis hypcihi (Borneo), 9. 226. Ncoloshanus towiicsi. 9. 227. Ncoloslianus Uicviccps. 9. 228.
Neoloshanus pal^ravei (Malayan) 9. Abbreviations: ac = aciculale; CN = callar nib; fg = femoral groove:
MD = malar depression; SA = stemauiar area.

157

Figs. 229-234. Neoloshanus. me.sosoma in lateral view. 229. N. pitosiis. V. 230. N. iicpalciisis, 9. 231. N.
paliiiavei (Australian). 9. 232. N. violaccus, 6. 233. N. anapctus. 6. 234. N. puipureovenlris, 9.
Abbreviation: fo = foveate.

158

Figs. 235-240. Mesosoma in dorsal view. 235. Orasema uichancoi. 9. 236. Orasema koi>hisiana. 9 . 237.
Orasema initiator. 9. 238. Orasema valgius, 9. 239. Psilocharis hypena (Taiwan), 9 . 240. Psilocharis
hypena (Borneo), 9 . Abbreviation: ar = areolate.

59

Figs. 241-246. Neoloshanus. mesosoma in dorsal view. 241. N. townesi, 9. 242. N. laeviceps. 9. 243. A'.
pilosus, 9 . 244. N. palgiavei (Au.stralian), 9 . 243. N. anapetus, 6 . 246. N. purpweoventris, 9 .

160

Fi(is. 247-255. Propodeum. 247. Oiascma koi^hi.siana. 9. 248. Oiascma valiiius. 9. 249. Psilocluirls hypena
(Taiwan), 9. 250. Neoloshanus lownesi, 9. 251. Neoloshanus laeviceps. 9. 252. Neoloshaiuis nepalensis. 9.
253. Neoloshanus palgiavei (Australian), 9. 254. Neoloshanus anapctus. 6. 255. Neoloshanus purpureoven-
tris, 9. Abbreviations: fr - frcnum; prp - pri)podeum.

161

Figs. 256-261. 256-237. Orusemorpha eiihotcs. 9: 256. Petiole and lirst gastial sternite in ventrolateral
view; 257. Apex of gaster in subventral view. 258-259. Orasema nichancoi, 9: 258. Gaster in ventrolateral
view; 259. Ovipositor. 260-261. Orasema initiator, 9: 260. Gaster in ventrolateral view; 261. Ovipositor.
Abbreviations: ar = anterior region; cr = crenulate; Ms, = second metasonial sternite; pet = petiole.

162

Figs. 262-267. 262-263. Orasema val^ius. 9: 262. Gaster in ventrolateral view: 263. Ovipositor. 264-263.
Psilocharis thcocles. 9: 264. Apex of gaster; 263. Tip of ovipositor. 266-267. Neoloshanus palf^ravci
(Australian), 9: 266. Gaster in lateral view; 267. Ovipositor.

163

Figs. 268-274. Ncoloshamis. 268-269. A', laeviccps. 9: 268. Metasoma in lateral view; 269. Ovipositor in
lateral view. 270. N. aimpetus. ovipositor. 271-272. A', purpureoveniris, 9: 271. Gaster; 272. Tip of oviposi-
tor. 273-274. N. anapetus. 6: 273. Base of gaster in lateral view; 274. Genitalia in subventral view.
Abbreviations: aed = aedeagus, dg = digitus; Ms, = second metasonial stemite; pni - paramerc.

164

TIMIODERUS

/ / /

ORASEMORPHA

Fig. 275. Geographic distribution and phylogeny of Orasemorpha, Timioderus. and Indosema. Letters refer
to species discussed in text. A = T. acuminatus; C = T. coromda\ D = O. dideiuata; E = O. erihotes: G = O.
goethei; I = /. indica: M = O. myrmicae; N = T. peridentatus; O = T. ramosus; ? = O. pyttalus; R = T. refrin-
gens\ S-0. sparsepilosus: T - O. tridenlala: V - O. varidentata; X = O. xeniades.

165

uichancoi

ORASEMA

valgius (y)

koghisiana

Fig. 276. Geographic distribution and phylogeny of Oiaseina. Lower-case names refer to species-groups.
Letters refer to species discussed in text. Question marks refer to questionable species assignments. Solid line
encloses the distribution of the O. uichancoi-group. Abbreviations: A = assectafor. B = bouceki; C = commu-
nis; E = seyrigi: F = fraudulenta: G - glabra; H = ishii., I = initiator; K = koghisiana; M = synempora; N =
nigra; P = promecea: R = rugulosa; S = striatosoma; U = uichancoi; V = valgius.

166

^T^

®

: -^x®^ V ^

ENMHADPJT

^4J

PSILOCHARIS

Fig. 277. Geographic distribution and phylogeny of Psilocharis. Lower-case names refer to species-groups.
Letters refer to species names discussed in text. Abbreviations: A = afra\ D = dahmsi; E = aenigma; H =
hypena; J = joanneae: M = monilicera: N = pentellci: P = pacifica: T = theocles.

167

Sct

U A G P S W •<*<!'

NEOLOSBANUS

V

Fig. 278. Geographic distribution and phylogeny of the Neoloshanus gemma- and purpureoveniris-
groups. Letters refer to species names discussed in text. Question marks refer to questionable species
assignments. Abbreviations: A = apoanus; G = gemma; P = piirpureovennis: S = sroreyi: U = wusheanus;
W = watanahei.

168

^vD

y ®

©

8r

i TLIPN<^GKAVX

NEOLOSBANUS

Fig. 279. Geographic distribution and phylogeny of the Neolosbanus gressilti- and /jw/.ij/y/i'c/ -groups
(except yv. pali>ravei). Letters refer to species names discussed in text. Question marks refer to question-
able species assignments. Abbreviations: A - anapetus; G = gressitti; I = taiwancnsis; K = kokurcaims\ L
= laeviceps; N - nepalensis; P = pilosus; T = townesi; V = violaceus; X? = sp. nr violaceus.

169

ALGERIA

I®

tt.-

V

Fig. 280. Geographic distribution of Neolosbanus palgravei. Letters refer to population assignments dis-
cussed in text. Abbreviations: A = Australian; C = Carolinean; M = Malayan: P = Papuan.

170

NUMBER OF SPECIES IN EACH SUBREGION

ETHIOPIAN

5

2

2

1

1

2

1

4

INDO-CHINESE

1

1

2

2

2

1

1

4

MALAYAN

1

2

3

2

PHILIPPINE

1

1

1

1

1

PAPUAN

(2)

3+

(1)

2

1

5

2

POLYNESIAN

1

1

1

2

AUSTRALIAN

9

1

1

PALAEARCTIC

2?

TOTAL

5

1

9

7

2

3

1

2

1

8

3

3

6

4

cc

LJJ
Q
O

LU

O

Q

LU

Z)

<

CC

«a'

CD

Q-

Q_

O

X

CO

CO

CO

CO

Q-

Z)

1—

<J)

_l

<

o

cn

Z)

D

Z)

Z)

UL

C/)

<

o

>

•^

DC

<

X

z:

z:

z

z

U
LU

<

nr

<

LU
CO

<

<

2

<

<

<

<

CD
CO

<

CD
CO

<

CD
CO

<

CD
(0

LU

LU

UJ

UJ

UJ

o

O

o

o

O

<

CO

cn

CO

CO

CO

o

_j

_l

_l

_l

<

<

<

<

<

_l

O

o

o

o

O

o

O

DC

O

CE

o

cc
O

DC

o

C/)

Q-

LU

LU

LU

z

LU

Z

T

Fig. 281. Number of species of Oraseminae and Psilocharitini represented in each of eiglit biogeographic
zones recognized by Gressitt (1956). and arranged according to the phylogeny presented in Figure 2. A
species may occur in more than one region and numbers do not necessarily add up to the total number of
species. Bracketed values for the Papuan subregion indicate species found only in northern Australia. Of the
two questionable Palaearctic records, one is from Algeria and the other a single record from Tokono Shima
Is., Japan.

71

^^^

Fig. 282. General distributions of Oraseminae and Psilocharitini in the Old World. Elements are divided into
Ethiopian, Malagasy, Indo-Pacific, and Australian regions, with each region represented by a different shad-
ing pattern. Derivation of species within each region is discussed in the text. Arrow indicates the proposed
direction of evolution of species in the Indo-Pacific region; the most highly derived species are found in the
southeast.

172

Appendix 1 : Character Matrix Used for Phylogenetic Analyses

Taxon names refer to genera, and the following abbreviations to species groups discussed in the text. Numbers correspond to character
states listed in Table 2. Character states are ordered from presumed plesiomorphic to apomorphic states. Question marks denote miss-
ing data. Stacked values for character state refer to multiple states within taxon.

CHARACTERS

TAXON

12 3 4 5 6 7 8

1 1 1
9 0 12

1111
3 4 5 6

1112
7 8 9 0

2 2 2 2

12 3 4

22222333

5 6 7 8 9 0 12

3 3 3 3

3 4 5 6

3 3 3 4

7 8 9 0

4444444445555555555666
1234567890123456789012

CHRYSOLAMPINAE 0007700000070000

PER I LAMP I NAE

TIMIOOERUS

INDOSEMA
ORASEMORPHA

ORASEHA ST

ORASENA UI

OftASEMA CO

ORASENA KO
ORASENA VA

ORASENA GL
ORASENA AS

PSILOCHARIS AE
PSILOCHARIS AF

0007700000070100
0 0 0?

0 0 17

1

1117

0 0 17

8

0 0 17

0 0 18

9

0 0 18

0 0 17
0 0 17

0 0 17
0 0 17

0 0 0 7
0 0 0 7

7 0 0 1

1 2

7 0 0 0

7 0 0 1
8

7 0 0 2

8 0 0 0

1

8 0 0 0

7 7 0 0

7 0 0 2

7 7 0 0

7 0 0 2

7 10 0

7 10 0

PSILOCHARIS TH 00077100

NEOLOSBANUS GE
NEOLOSBANUS PU
NEOLOSBANUS TO

NEOLOSBANUS PA
NEOLOSBANUS CR

GOLLUMIELLA

ANORASENA
STILBULA

OBEZA

PSEUDOCHALCURA

STILBULOIDA
EUCHARIS

AUSTEUCHARIS

TRICORYNA

PROPSILOGASTER

PSEUDOMETAGEA

CHALCURA

SCHIZASPIDIA

KAPALA

ISOMERALA
CH MONTANA

0 0 7 7
0 0 0 9
10 0 8

0 0 0 8

0 0 0 8

1 1

0 0 0 7

1
0 0 17
1119

1

1

1 9

1 8
9
7 9
7 8
9
10 19

7 10 0

7 10 0

7 10 0
8

7 10 0

8 10 0

1

7 10 0

7 0 0 0

9 0 0 0

3

8 0 0 3

7 0 2 0
3
7 0 2 3
7 0 0 0
8
7 0 0 0

0 0 0 0
0 0 0 0

0 0 0 0

0 0 0 0

1

0 0 0 0

1

0 0 0 0
0 0 0 0

10 0 0
0 0 0 0

110 0
110 0

0 10 0

1

2 110
0 111
0 0 10

1

0 0 11
0 0 10

1

0 0 0 0

0 0 0 0
0 0 0 0

0 2 0 0

2 3 1

2 4 2 0

12 0 0

1

12 10

12 10

12 0 0

12 0 0

12 10

12 10

12 10

12 10

12 10

0 0 0 0
1

0 0 3 0

1

10 0 1

10 0 1

0 0 0 1

1

0 0 0 1

0 0 0 0

1 1

0 0 0 1

1

0 0 0 2

0 0 0 1

0 0 0 1

0 0 0 1

0 10 2

0 10 2

000000100000

0 0 0 0

13 11

13 11

10 0 0

10 0 0

10 0 0

10 0 0

10 0 0

10 0 0

10 0 0

10 0 0

10 0 0

10 0 0

0 0 2 0

7 0 0 0

10 10

0 0 0 0

7 0 0 0

7 0 0 0

7 0 0 0

7 0 0 1

0 0 0 1

7 0 0 0

7 0 0 0

0 0 0 1

0 0 0 1

0 0 0 2

0 0 0 1

0 10 1

0 0 0 1

0 0 0 1

0 0 0 1

0 0 0 0

1

0 0 0 1

0 0 0 0

1

0 0 0 1

0 0 0 1

10 0 0

10 0 0

0 0 10

1

0 0 0 0

1

0 0 11

1

10 0 1

0 0 0 0

1 1

0 0 0 0

1

0 0 0 0

0 0 0 0

1 1

2

0 10 0

1 2

0 0 0 0

0 10 0

0 10 0

1

0 2 10

0000002170000000000070

000000201

2
1 12 2 0 0

0 2 10
0 0 0 0 0 2 10

10 0 0

0 0 0 0

0 0 0 1

0 0 0 1

0 0 0 1

0 0 0 1

12 10

12 10

2

12 10

12 10

12 3 1

0 2 3 1

1210010210000001 10000001 1231

12 10

12 10

12 10

12 10

12 10

0 10 2

0 10 2

0 10 2

0 10 2

0 10 2

10 0 0

10 0 0

10 0 0

10 0 0

10 0 0

7 0 0 1

0 0 0 1

0 0 0 1

0 0 0 1

7 0 0 1

2 0 0 0

2 0 0 0

2 0 0 0

2 0 0 0

2 0 0 0

0 0 0 1

0 0 10

0 0 0 1

0 0 0 1

0 0 0 1

121000111010770000000001

12 10
12 0 0

0 0 0 0 12 0 1

0 0 0 0

19 9 0 0 0
7 9

1 7
9
10 17

10 19

10 19

10 7 9
10 7 9

7 0 10

8

7 0 0 0

B 1

7 12 0

7 0 2 0
9

7 0 2 0
3

7 0 2 0

9 0 0 0

0 0 0 0

0 0 0 0

0 0 0 0

1

0 0 0 0

0 0 0 0

0 0 0 0

0 0 0 0

1

0 0 0 0

1

0 0 0 0

0 0 0 0

0 0 0 0

12 0 1

1

12 0 0

12 2 0
2

12 0 0

12 0 0

12 0 0

12 10

2 2

12 0 0

12 0 0

12 0 0

12 0 0

12 0 0

0 111

0 12 3
1

112 3

112 3

0 12 3

0 0 2 4

1 1

0 12 4

0 0 2 0

3

10 2 3

0 0 2 0

1

0 12 4

0 12 3

4

0 12 5

0 12 5

0 12 3

10 0 0

110 0

110 0

110 0

110 0

110 0

110 0

110 0

110 0

110 0

1

110 0

110 0

12 0 0

12 0 0

110 0

0 10 0
12 0 0

12 10

12 0 0

'210
110 0

7 10 0

7 10 0

7 10 0

110 0

110 0

1110

1110

1110
7 110

0 0 0 0

0 0 11

0 0 11

0 0 0 1

0 0 11

0 10 0

0 0 0 2

0 0 0 1

0 0 0 1

0 0 0 1

0 0 0 2

0 0 2 2

1

0 13 2

0 13 2

0 0 0 2

0 0 0 0

0 0 2 1

0 10 1
1

112 1

110 1

0 10 1

1

0 0 0 0

0 0 0 0

0 0 0 0

0 10 1

0 0 0 0

1 1

0 0 0 0

1

0 0 0 1

0 0 0 1

0 0 0 0

12 2 1

12 2 1

12 2 1

12 2 1

12 2 1

0 2 2 1

1

0 2 2 0

12 2 1

12 2 1

12 2 1

12 2 1

0 2 2 1

1

0 2 2 1

0 2 2 1

0 2 2 1

0 2 2 1

1

12 2 1

0 2 2 1

0 2 2 1

1

0 2 2 1

0 2 2 1

113 2 0 0

1 12 2 0 0

0 13 10 0

0 12 3 0 0

0 1110 0

3

0 12 2 0 0

0 12 2 0 0

0 13 2 0 0

0 13 2 0 0

0 0 0 0 0 0
0 0 0 0 10

1
0 0 0 0 10

0 0 0 0 0 1

0 0 0 0 0 1

0 1110 1

0 1110 1

0 10 2 0 1

1 0 ?
2

2 0 7
2 1 7

2 7?

2 1 0

2 7 7

2 7?

2 1 0

2 1 7

7 1 0

2 1 7

2 7 7

2 7 7

2 7 7

2 7 7

2 1 7

2 1 1

2 7 7

10 0 1

7 7 7 7

7 7 7 7

7 7 7 7

7 7 7 7

7 0 0 1

7 7 7 7

7 7 7 7

7 7 7 7

7 7 7 7

2 0 0 1

1 1 1

7 7 7

7 ? ?

7 7?

7 7 7

1 1 0

7 7 7

7 7 7

7 7 7

7 7 7

1 1 0

7 ? ? 7 7 7 7

1 0 1

7 7 7

7 7 7

7 7 7

7 7 2

7 0 2

7 7 7

7 7 7

7 7 7

7 7 7

1 0 2

1

7 7 7

0 0 0

1

7 7 7

? ? ?

7 7 7

1 7 7

1 1 ?

7 7 7

7 7 7

7 1 1

7 7 7

1 1 i

7 7 7

7 7 7 7

7 7 7 7

7 7 7 7

7 7 7 7

7 7 11

2 111

7 7 7 7

7 7 7

7 7 7

7 7 7

7 7 7

i 7 7

1 1 1

7 7 7

7 7 7

7 7 7

7 7 7

7 7 7

7 7 7

1 1 3

7 7 7

7 7 7

7 7 7

0 7?

? 7 7

7 1 0

0 1 0

7 7 7

0000112177777777777727

0 10 0 11

0 0 0 0 0 1

0 0 0 0 2 1

0 0 0 0 2 1

0 0 0 0 7 1

0 0 0 0 2 1

0 0 0 0 2 1

0 0 0 7 7 1

0 0 0 7 7 1

0 0 0 0 2 1

0 0 0 0 0 1

0 0 2 0 2 1

1 1

0 0 0 0 2 1

0 0 0 0 2 1

0 10 0 2 1

2 1 7

2 1 0

2 1 0

0 1 0

1 7 7

2 1 1

2 7 1

2 7?

2 7?

2 1 1

2 1 1

2 1 ?

2 1 1

2 1 ■>

1 1 7

7 7 7?
2 10 1

7 7 7
1 1 1

2 10 1110

3 10 0

7 7 7?
2 111

2 111

7 7 7 7

1 1 1
1

7 7?

1 1 1

1 1 1

? 7 7 7

2 111

2 17 1

7 111

2 111

7777

2 111

7 ? 7

1 1 1

1 1 1

1 1 1

1 1 1

7 7'

1 1 1

7 7 7

1 1 5

1 1 7

1 1 5

7 7 7

1 1 4

1 7 4

7 7 7

7 7 7

1 1 4

1 7 3

1 1 3

1 1 3

7 7 7

1 1 ?

777
0 2 0

7 2 7

0 2 0

777
0 2 0

0 7'

7 7 7

7 7?

0 2 1

0 2 0

0 1 0

2
0 2 0

777

' 2 ■>

\1}

Appendix 2: New Synonymies

Appendix 5: New Taxa Described

Cienera

Orasema Cumeron. 1884

Parasemora Gemignani. 1933

Species

Tiiuiodents refrini'ens Waterston, 1916
Orascimi viriJicyaneci Risbec. 1958

Oiascmorpha tridentaia (Girault), 1915
Eucharomorpha wheeleri Brues, 1934

Orasemorpha eribotes (Walker), 1839
Eucharomorpha diibia Girault, 1913a
Eucharomorpha fuscipes Girault, 1913a
Eucharomorpha partiglabra Girault, 1 940
Eucharomorpha viridis Girault. 1913a

Neolosbanus palgravei (Girault), 1922

Psilogasler nishidai Ishii and Nagasawa, 1941

Losbamis petersoni Hedqvist, 1978

Orasema indica Snehalatha and Narendran, 1992

Appendix 3: New Combinations

Orasemorpha myrmicae {Epimetagea)
Orasema fraudulenta (Psilogaster)
Psilocharis the odes (Orasema)
Neolosbanus gemma (Orasema)
Neolosbanus purpureoventris (Eucharis)
Neolosbanus laeviceps (Psilogaster)
Neolosbanus palgravei (Orasema)
Neolosbanus gressitti (Losbanus)
Stilbula ranomafanae (Orasema)

Appendix 4: Lectotypes Designated

Eucharomorpha didentata Girault (Orasemorpha)

Eucharomorpha fuscipes Girault (Orasemorpha)

Eucharomorpha partiglabra Girault (Orasemorpha)

Eucharomorpha wheeleri Brues (Orasemorpha)

Orasema communis Risbec

Orasema palgravei Girault (Neolosbanus)

Orasema pheidolophaga Girault

Orasema uichancoi (Losbanus)

Psilocharis fraudulenta Reichensperger (Orasema)

Genera

Neolosbanus
Psilocharis

Species

acuminatus, Timioderus
coronula, Timioderus
peridentatus , Timioderus
ramosus, Timioderus

sparsepilosa . Orasemorpha

bouceki, Orasema
glabra. Orasema
ishii. Orasema
koghisiana, Orasema
nigra, Orasema
promecea. Orasema
rugulosa. Orasema
striatosoma. Orasema
synempora. Orasema

aenigma, Psilocharis
afra, Psilocharis
dahmsi. Psilocharis
hypena, Psilocharis
joanneae. Psilocharis
monilicera, Psilocharis
pacifica. Psilocharis
pent el la. Psilocharis

anapetus. Neolosbanus
apoanus. Neolosbanus
kokureanus. Neolosbanus
nepalensis, Neolosbanus
pilosus . Neolosbanus
sloreyi. Neolosbanus
taiwanensis, Neolosbanus
townesi, Neolosbanus
violaceus, Neolosbanus
watanabei. Neolosbanus
wusheanus. Neolosbanus

174

ROYAL ONTARIO MUSEUM LIFE SCIENCES PUBLICATIONS

INSTRUCTIONS TO AUTHORS

Authors should prepare Iheir manuscripls carefully
according to the following instructions; failure to do so
will result in the manuscript's being returned to the author
for revision. All manuscripts are considered on the under-
standing that they arc not currently offered for publication
elsewhere.

1. General Papers for publication are accepted from
ROM staff members and research associates, and
from researchers reporting on work done with ROM
collections. Monographs on the flora and/or fauna of
Ontario by authors not affiliated with the ROM may
be considered for publication. Financial contributions
towards publication may be required. Authors are
expected to write clearly and concisely and to omit
any material not essential for an understanding of the
main theme of the paper.

2. Format Manuscripts (including captions, syn-
onymies, literature cited, and tables) should be typed
double-spaced on standard letter-size paper with a 4-
cm (1") margin on all sides. Three photocopy copies
should be submitted to the Head of Publications and
Print Services; the original should be retained by the
author. The submission should include a separate
sheet giving the author(s) names and affiliations, the
title of the publication, the series if applicable, the
number of typed pages, the number of tables, and the
number of plates and figures. Manuscripts should
normally be organized in the following order: con-
tents, abstract, introduction, materials and methods,
results, discussion, conclusions, summary (if manu-
script is long), acknowledgements, appendices, and
literature cited. Authors are encouraged to include
foreign-language translations of the summary, if
appropriate. Main headings should be centred; sub-
headings should be left-justified to the text margin.
The first line of the first paragraph in each new sec-
tion should not be indented. A single space only
should be left between sentences. Material intended
to be typeset as italic must be provided as italic, not
as underlined roman type. Literature citations in the
text should be in one of the following forms: "Jones
(1994)"' or "(Jones, 1994)" or "(Smith, 1990:71-79,
fig. 17)."

3. Standard Sources The primary authority on ques-
tions of format and style is the Guide to Authors and
Editors, available from ROM Publications and Print

Services. For matters not covered in the guide, con-
sult the CBE Style Manual. Other standard sources
are as follows: for English spelling. The Concise
Oxford Dictionary; for Canadian place names and
coordinates. Canada Gazetteer Atlas; for the spelling
of geographic names. The Times Atlas of the World.

4. Abstract All papers must be preceded by a short,
factual abstract, about one per cent of the text in
length. The abstract may be followed by four to six
key words in parentheses.

5. Taxonomy The name of a taxon should be given in
full in headings, at the beginnings of paragraphs, and
at its first occurrence in the text. Give the authority
and date, if appropriate, with the first mention of
each taxon, but not thereafter. Taxonomic papers,
particularly synonymies, should follow the layout in
the Guide to Authors and Editors. International
Codes of Biological Nomenclature must be followed.

6. Literature Cited A complete list of references, in
alphabetical order of authors, must be given at the
end of the paper. When two or more works of one
author are cited, they should be listed chronological-
ly. The names of journals should not be abbreviated.
For correct bibliographic form, see the Guide to
Authors and Editors.

7. Tables All tables should be typed on separate
sheets and numbered consecutively in arabic numer-
als in the order of their first mention in the text. Mark
the location of each table in the margin of the text.

8. Plates, Figures, and Text-figures Illustrations
may be designated according to the conventions of
the author's discipline; in some disciplines grouped
photographs of .scientific subject matter are common-
ly termed Plates, while line drawings and locality and
other illustrations that occupy a full page or less are
Text-figures. Usage must be consistent throughout
the paper. A full-page illustration with its caption
should be sized to fit an area of 17.3 X 22.75 cm
(6.8" X 9") for a Contribution; for Occasional
Papers, the area is 14.1 x 21.2 cm (5.5" X 8.3"). If
captions are lengthy, they may be placed on the fac-
ing page. A scale or magnification factor should be
included. Authors are reminded that when illustra-
tions are reduced, magnification factors will change,
and that they are responsible for the conversion. For
details, see the Guide to Authors and Editors.

176

ROYAL ONTARIO MUSEUM LIFE SCIENCES CONTRIBUTIONS

Lite Sciences Contributions are a numbered series

148 Shallow-Water Hydroids of Bermuda:

of scientific publications of varied subject matter.

The Athecatae

Most recent contributions include the following:

Dale R. Calder

1988, 11 2 pp., ill., $24.50

156 Conodonts of the Lower Border Group and

ISBN 0-88854-339-5

Equivalent Strata (Lower Carboniferous) in

Northern Cumbria and the Scottish Borders, U.K.

147 Biostratigraphy and Palaeontology of the

Murk A. Purnell

Scollard Formation, Late Cretaceous and

1992. 80 pp.. ill., $19.95

Paleocene of Alberta

ISBN 0-8854-405-7

Loris S. Russell

1987, 23 pp., ill., $7.00

155 Revision of the World Species of

ISBN 0-88854-338-7

Spalangiopeltu (Hymenoptera: Chalcidoidea:

Pteromalidae: Ceinae)

146 Stipatocrimts, a New and Unusual Camerate

D. Christopher Darling

Crinoid from the Lower Silurian of Western

1991.48 pp., ill., $11.00

New York

ISBN 0-88854-395-6

James D. Eckert and Carlton E. Brett

1987, 17 pp., ill.. $6.00

154 Shallow-Water Hydroids of Bermuda:

ISBN 0-88854-336-0

The Thecatae. Exclusive of Plumularioidea

Dale R. Calder

145 An Annotated Checklist of the Fishes of the

1991, 144 pp.. ill., $24.50

Chagos Archipelago, Central Indian Ocean

ISBN 0-88854-354-9

Richard Winterbottom, Alan R. Emery, and

Eriing Holm

153 Silurian Trilobites from the Northern Yukon

1987, 240 pp.. ill.. $50.00

Territory

ISBN 0-88854-329-8

Rolf Ludvigsen and Ronald P. Tripp

1990, 64 pp.. ill.. $12.95

144 Revision of the Caddisfly Genus Psilotreta

ISBN 0-88854-349-2

(Trichoptera: Odontoceridae)

Charles R. Parker and Glenn B. Wiggins

152 The Type Species of the Ordovician Trilobite

1987, 55 pp.. ill., $12.00

Genus Isotelus: I. gigas Dekay, 1824

ISBN 0-88854-332-8

David M. Rudkin and Ronald P. Tripp

1989, 24 pp., ill., $10.25

143 Phylogeny. Speciation, and Palaeoecology of

ISBN 0-88854-345-X

the Early Carboniferous (Mississippian)

Conodont Genus Mestognathus

151 The Structure of the Call Note System of

Peter H. von Bitter, Charles A. Sandherg. and

the WarblingVireo

Michael J. Orchard

Daryl Howes-Jones and Jon C. Barlow

1986. 115 pp., ill., $25.00

1988,40 pp.. ill.. $11.00

ISBN 0-88854-319-0

ISBN 0-88854-343-3

142 Review of the Gobioid Fishes of the Chagos

150 Late Cretaceous-Early Tertiary

Archipelago. Central Indian Ocean

Dinoflagellates and Acritarchs from the Kashi

Richard Winterbottom and Alan R. Emery

Area, Tarim Basin. Xinjiang Province, China

1986, 82 pp.. ill.. $15.00

Mao Shaozhi and Geoffrey Norris

ISBN 0-88854-320-4

1988. 100 pp., ill., $25.00

ISBN 0-88854-334-4

A catalogue of ROM publications in print is

available from

149 Occurrence of the Cladid Inadunate Crinoid
Thalamocrinus in the Silurian (Wenlockian) of
New York and Ontario
George C. Mclnlosh and Carlton E. Brett
1988, 20 pp.. ill.. $7.75

University of Toronto Press
10 St. Mary Street, Suite 700
Toronto, Ontario M4Y 2W8
Canada

ISBN 0-88854-342-5

Tel. (416) 978-2229 Fax (416) 978-4738

ISBN 0-88854-4 1 2-X ISSN 0384-8 1 59