Bulletin of the
British Museum (Natural History)
Entomology series Vol48 1984
British Museum (Natural History)
London 1984
Dates of publication of the parts
No1 . . 26 January 1984
No2 23 February 1984
No3 28 June 1984
ISSN 0524-6431
Printed in Great Britain by Henry Ling Ltd, at the Dorset Press, Dorchester, Dorset
Contents
Entomology Volume 48
No 1 Gelechiid moths of the genus Mirificarma
By Linda M. Pitkin
No 2 Macronematine caddisflies of the genus Amphipsyche (Trichoptera:
Hydropsychidae)
By P. C. Barnard
No 3 A review of the genera of Indo-Pacific Encyrtidae (Hymenoptera:
Chalcidoidea)
ByJohnS. Noyes&M.Hayat
71
131
Bulletin of the
British Museum (Natural History)
Gelechiid moths of the genus Mirificarma
Linda M. Pitkin
Entomology series
Vol 48 No 1
26 January 1984
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World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.)
Trustees of the British Museum (Natural History), 1984
The Entomology series is produced under the general editorship of the
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ISBN 565 06000 7
ISSN 0524-6431 Entomology series
Vol 48 No 1 pp 1-70
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 26 January 1984
Gelechiid moths of the genus Mirificarma
'i
Linda M. Pitkin
I niviti >i . i iirvin
Department of Entomology, British Museum (Natural History), Cromwell Road, London
SW7 5BD
Contents
Synopsis 1
Introduction 1
Methods 2
Abbreviations of institutions 3
Acknowledgements 3
Host-plant relationships within Mirificarma 3
The systematic position of Mirificarma 5
Classification of Mirificarma species 6
The structure of the filament of the male genitalia 12
Mirificarma Gozmany 13
Key to the species of Mirificarma 16
Males 16
Females 17
The montivaga-group 18
The maculatella-group 19
The interuptella-group 29
References 45
Index 70
Synopsis
The Palaearctic genus Mirificarma is revised and three new species are described. Twenty-one species are
recognized and four synonyms are newly established; seven new generic combinations are included. Keys
to the species, and figures of the external features and genitalia of all the species are given. The systematic
position of Mirificarma is discussed and a provisional classification of the species based on a cladistic
analysis is provided, together with an account of the filament, a unique structure of the male genitalia of
Mirificarma. Biological data, as far as known, are given for all the species, and the host-plant relationships
within the genus are discussed.
Introduction
The genus Mirificarma consists of 21 species of Microlepidoptera from Europe and the
Mediterranean region, with one species introduced to the U.S.A. The moths are of average size
for Gelechiidae, with a wingspan usually between one and two centimetres. They vary in wing
pattern; some species have the fore wing with yellowish zig-zag markings, others have spots, a
median longitudinal stripe, or are almost uniform in colour. The larvae feed on Leguminosae.
Mirificarma cannot be defined readily on external characters and, since early microlepi-
dopterists relied on such characters, many species have been misplaced in other genera.
Hitherto, no comprehensive account of the genus has been given, although a partial revision by
Sattler (1960: 41-45) covered aspects of eight species.
The purpose of this study is to describe three new species, to include seven previously
misplaced species and to provide a key by which all the species can be identified. In addition, this
study examines the host-plant relationships within Mirificarma, describes the structure of the
filament, a character of the male genitalia unique to the genus, investigates the systematic
position of the genus within the Gelechiidae and proposes a classification of the species within
Mirificarma.
This study aims to resolve the confusion over the interpretation of the filament and its
Bull. Br. Mus. nat. Hist. (Ent.)48(l): 1-70 Issued 26 January 1984
2 L. M. PITKIN
associated structures. Moreover, examination of these structures has helped to indicate the
relationships oiMirificarma within the Gelechiidae.
Photographs of the wings of every species are reproduced but should only be used as a rough
guide for identification. The wing patterns of some species are very similar and, in a few cases,
considerable variation within species can make identification very difficult. However, there are
abundant distinct genitalic differences between Mirificarma species and, to enable accurate
determination, the key is based mainly on these.
The leguminous host-plants are of particular interest since they provide evidence to corrobo-
rate the division of the genus into species-groups. Two species, flavella and eburnella, feed on
plants grown as fodder-crops but neither is recorded as an agricultural pest in Europe. However,
eburnella has been introduced to the U.S.A. where it has caused severe damage to clover and
vetch crops (see p. 28).
Methods
The measurements given at the beginning of each description are the range in fore wing length of
the specimens examined, to the nearest half millimetre. Measurements which are far outside the
normal range for a species are given in parentheses, in addition to the range for the other
specimens examined of the species. The length of the apophysis anterior in the female genitalia
was measured from the apex of the apophysis to the level at which it joins the antrum. Since the
apophyses anteriores are sometimes of unequal length, the range in length given is derived from
the mean of the pair. The number of specimens examined is given in parentheses after this
measurement.
Standard methods of genitalia preparation were followed except that where sufficient
material was available, the vinculum was separated from the tegumen on one side of the male
genitalia and laid out flat beside the tegumen, in order that the complex structures should be
more clearly displayed in the photographs. This has also facilitated the examination of the
structures. For staining, mercurochrome was used. The terminology of the genitalia follows that
of Klots (1956) and Sattler (1979). The descriptions of the sacculus refer to the posterior portion
of this which is free from the vinculum.
The specimens examined are deposited in the British Museum (Natural History) unless
otherwise stated. Records from Mr E. Jackh, Bidingen are based on photographs of genitalia
preparations only. The data of type-material are given in full and sometimes include information
from the original description, additional to that from the specimen label; data from sources
other than these are given in square brackets. Data of specimens other than types are
abbreviated by omitting collectors' names and dates other than months. For widespread and
well-documented species localities are summarized under country, with province or region
where appropriate. The spelling of locality names follows The Times Atlas of the World
(Comprehensive Edn), 1968, as far as possible. The distribution records are based on material
examined unless otherwise stated.
Most of the specimens examined in this study bear my determination labels dated '1982' or
earlier. Any specimens with subsequent determination labels are not included.
Lectotype designations are made where necessary, in some cases for species which were based
on an unspecified number of specimens, even where only one original specimen is currently
known. Lectotypes are also designated for species described by Walsingham where he referred
to a 'type cf" and a 'type $' but did not specify a single 'type' or holotype. In these cases the
specimen selected is that listed first by Walsingham, usually the male. These lectotype
designations follow the practice discussed by Sattler (1976: 89).
Some of the photographs of the wings have been reversed.
The botanical nomenclature and classification follow Polhill & Raven (1981); European plant
names not included in that work follow Tutin et alii (1964-1980).
GELECHIID MOTHS 3
Abbreviations of institutions
BMNH British Museum (Natural History), London.
LN Landessamlungen fur Naturkunde, Karlsruhe.
MINGA Muzeul de Istorie Naturala 'Grigore Antipa', Bucharest.
MNHN Museum National d'Histoire Naturelle, Paris.
MNHU Museum fiir Naturkunde der Humboldt-Universitat, Berlin.
NM Naturhistorisches Museum, Vienna.
TM Termeszettudomanyi Muzeum, Budapest.
ZM Zoologisk Museum, Copenhagen.
Acknowledgements
I am most grateful to the following specialists for loan and donation of material, and for helpful
information: Mr L. Aarvik, As, Norway; Rev. D. Agassiz, Grays, Essex; Mr E. Arenberger,
Vienna; Mr K. Burmann, Innsbruck; Dr L. Gozmany, TM; Dr H. J. Hannemann, MNHU; Dr
R. W. Hodges, U.S.D. A. /National Museum of Natural History, Washington, D.C.; Mr E.
Jackh, Bidingen, West Germany; Mr O. Karsholt, ZM; Dr F. Kasy, NM; Dr J. Klimesch, Linz;
Dr J. Langmaid, Portsmouth, Hampshire; Dr E. S. Nielsen, ZM; DrT. Peet, Guernsey; Dr A.
Popescu-Gorj, MINGA; Dr J. Powell, University of California, Berkeley; Dr R. U. Roesler,
LN;DrP. Viette, MNHN.
The photographs were produced by the Photographic Unit, BMNH.
Host-plant relationships within Mirificarma
Gelechiid genera are often associated with particular host-plant groups, for example, Ornatival-
va Gozmany - Tamarix, Caryocolum Gregor & Povolny - Caryophyllaceae. The known
host-plants of Mirificarma larvae are all Leguminosae. It is sometimes difficult to verify
host-plant records since the plant or the moth may have been misidentified; for example, an
erroneous record oieburnella on Populus is based on a misidentification of the moth, which was
probably Schiffermuelleria formosella ([Denis & Schiffermiiller]). Additionally, changes in
botanical nomenclature can cause problems, such as the following examples. Cytisus scoparius
L. was known as Spartium scoparium by some authors and records of Spartium probably refer to
this plant and not to species currently in Spartium. Trifolium has been used broadly to cover
plants in other genera of the Trifolieae such as Medicago. Some 'host-plant' records are not
based on moths bred from larvae on the plant, but merely on the circumstantial evidence of the
presence of the moths around the plant, and may be erroneous. Such errors are often
compounded by one author copying another without providing confirmatory data or citing the
source of information.
The choice of host-plant does not seem to reflect the systematic relationships of Mirificarma
within the Gelechiidae. Leguminosae are utilized as larval host-plants by species in several
unrelated groups of the Gelechiidae, for example, Xystophora Heinemann (Aristoteliinae),
Syncopacma Meyrick (Anacampsinae), Dichomeris Hubner (Dichomerinae) and Anarsia
Zeller (Chelariinae). In the Gelechiinae, Leguminosae are known as host-plants of several
unrelated species in the tribe Gelechiini, including Lita solutella (Zeller), Chionodes lugubrella
(Fabricius) and Aroga aristotelis Milliere. Species of Gelechia Hubner, the apparent close
relative of Mirificarma, utilize a diverse range of host-plants, predominantly Salicaceae,
Juniperus and Rosaceae.
The host-plants of 11 species of Mirificarma are known and are all in the subfamily
Papilionoideae. The systematics of the Leguminosae are taken from Polhill & Raven (1981).
The Papilionoideae are divided into several groupings of tribes including the epulvinate series,
which comprises the temperate herbaceous tribes, which have lost the basal pulvinus of the leaf,
and the genistoid alliance, which includes the tribe Genisteae. Four species of Mirificarma have,
as yet, been recorded from only one host-plant, but seven species have been bred from two or
more plant species and eburnella and mulinella are each known from seven.
Three species-groups are here recognized in Mirificarma based on a number of morphological
L. M. PITKIN
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GELECHIID MOTHS 5
characters. The division of the two major species-groups is corroborated by host-plant differ-
ences. Host-plants of the montivaga-group are unknown. The host-plants of the maculatella-
group are all in the epulvinate series, in the tribes Loteae, Coronilleae, Trifolieae, Vicieae and
Galegeae, while the interuptella-group has Genisteae host-plants (see Table 1).
One species in the interuptella-group, cytisella, has Ononis as a host-plant, in addition to
several species of Genisteae. Although Ononis is a member of the Trifolieae, it has some
Genisteae-like morphological characters and it has been linked with the Genisteae by early
authors (see the discussion of the taxonomic history of Ononis by Kupicha, 1977: 134). Ononis
also has biochemical affinities with the Genisteae, to which it has similar isoflavenoids (see
Gomes etalii, 1981: 472, 473). It is possible that these characteristics make the plant acceptable
to the moth which otherwise selects Genisteae. In addition the host-plant must be acceptable to
the larva.
Although Mirificarma species within a species-group share host-plants of the same tribe or
group of tribes, there is little evidence that the close relationship of species is reflected in the
close relationship of their host-plants. In the maculatella-group, eburnella and flavella both
occur mainly on Trifolieae, including Trifolium. However, eburnella also occurs on Coronilleae,
the host-plant tribe of maculatella, a species to which eburnella is less closely related. M.
maculatella and its sister-species denotata use plants of different tribes, Coronilleae and
Galegeae respectively.
In the interuptella-group , the closely related mulinella and ulicinella both occur on Ulex,
although on different species, but mulinella shares Cytisus nigricans, Calicotome spinosa and
Genista with cytisella; Genista germanica and G. tinctoria with lentiginosella, and Cytisus
scoparius with interuptella. M. mulinella and ulicinella form a close group with cabezella which
differs considerably from these two in its host-plant Adenocarpus , an outlier in the Genisteae.
The distribution of a host-plant has not always restricted the distribution of the moth, which
may have adapted to different plants in different parts of its range. For example, cytisella was
described from specimens bred on Cytisus nigricans in Germany, and has subsequently been
bred on this, for example, by Klimesch (1961: 651) in Austria. C. nigricans does not occur in
France (Tutin etalii, 1968: 86) and cytisella has been recorded there on Calicotome and Ononis.
The ability of some Mirificarma species to utilize a range of host-plant genera suggests that they
have adapted to different hosts rather than that they have evolved in conjunction with their
hosts.
The systematic position of Mirificarma
Many species currently in Mirificarma were originally described in Gelechia, and subsequently
listed in this genus by Gaede (1937). However, Gelechia has been widely used to include many
unrelated species and has been referred to as the microlepidopterist's 'waste-paper box'
(Chambers, 1872: 147, and subsequent authors). Mirificarma is clearly separated from Gelechia
(sensu Saltier, 1960) by the presence of a filament on the male genitalia, as well as other
characters (see also remarks, p. 15).
Following the original placing of flavella in Acompsia Hiibner, the related species eburnella
(as formosella Hiibner) and pallidipulchra were also listed in this genus by Meyrick (1925: 142)
and Gaede (1937), possibly because Acompsia and Mirificarma have similar wing venation.
Acompsia and Mirificarma differ, however, in many characters including the male eighth tergite
and sternite which are laterally fused in the former but separated in the latter. The three species
have also been placed in Rhinosia Treitschke, either in the original descriptions or by
subsequent authors including Meess (in Spuler, 1910: 344), apparently because of a strong
superficial similarity to species then in Rhinosia but now transferred to Orophia Hiibner
(Oecophoridae). Orophia ferrugella ([Denis & Schiffermiiller]), formerly in Rhinosia, has
sometimes been misidentified as flavella (see p. 27).
In the original description, Mirificarma was placed between Neofaculta Gozmany and Lita
Treitschke, both currently in the subfamily Gelechiinae, without indication as to whether its
position was deliberate or arbitrary. Sattler (1960: 41) also placed it among genera currently in
the Gelechiinae although its placement between Lita and Aroga Busck was arbitrary.
O L. M. PITKIN
The subfamilies of Gelechiidae are mostly poorly defined and the relationships of the genera
still require much investigation. However, Mirificarma does appear to share some characters
with certain other genera within the Gelechiidae. The eighth abdominal tergite and sternite of
the male are separated into free flaps in Mirificarma. This is usual in the Gelechiinae (tribes
Gelechiini, Teleiodini and Gnorimoschemini) whereas the tergite and sternite are normally
fused laterally in the rest of the Gelechiidae. The long narrow scales of the coremata of the
eighth tergite of Mirificarma are similarly placed to those occurring in many Gelechiinae,
attached closely to the tergite. Coremata occur in a few genera outside the Gelechiinae, and are
probably homologous in some cases, such as in Deltophora Janse in the Aristoteliinae, although
in this genus the coremata are usually loosely attached to the tergite. The coremata of the male
eighth sternite of Mirificarma, on the dorsal membrane, consist of enlarged scales with a
rounded, often inflated structure. Some Gelechiinae have scales similarly situated, as seen in
Gelechia scotinella Herrich-Schaffer, G. hippophaella (Schrank), G. nigra (Haworth) and
Teleiodes myricariella (Frey), in which the scales are large and sometimes long although not
inflated. The scales are deciduous, consequently fresh preparations need to be made to assess
how widespread they might be in the Gelechiinae and whether they might occur outside the
Gelechiinae. The scales are most similar to those of Mirificarma in Psoricoptera gibbosella
(Zeller), currently in the Hypatiminae although with a number of similarities to Gelechia near
which it might be better placed.
In common with many Gelechiinae, the posterior apophyses of the female genitalia are often
long in Mirificarma, although this character is not restricted to this subfamily.
Within the Gelechiinae, Mirificarma does not appear to be closely related to the Gnorimos-
chemini, in which the signum of the female genitalia is a strong hook-like structure. The signum
of Mirificarma varies interspecifically but is never of this form. M. montivaga, which has the least
apomorphies of the genus, has a signum similar to that frequently seen in Gelechia and the
Teleiodini. This type of signum is also found in Neofriseria Sattler, currently placed in the
Gelechiini although not in close relationship with Gelechia, and Psoricoptera gibbosella,
currently in the Hypatiminae although, as mentioned above, probably more closely related to
Gelechia. Mirificarma does not show evidence of a close relationship with the Teleiodini, in
which the male genitalia have frequent modifications not shared by Mirificarma, such as a
narrow, elongate uncus. The male eighth segment varies in shape in the Teleiodini and usually
differs from that of Mirificarma. The gnathos is absent in some Teleiodini and if present, usually
also differs from that of Mirificarma.
The possession of a filament on the male genitalia distinguishes Mirificarma from all other
Gelechiidae. However, the pair of sclerites which support the filament in Mirificarma, extending
anteriorly from the base of the sacculus and valva and lying between the vinculum and tegumen,
appear to be homologous with similar structures in species of Gelechia, particularly G. sabinella
Zeller, G. nigra, G. rhombella ([Denis & Schiffermiiller]) and G. senticetella Staudinger. The
sclerites in these species are fused anteriorly, at the region from which the filament would arise in
Mirificarma. This region is usually membraneous in Gelechia, and never a well-sclerotized
swelling as in montivaga which has the least-developed filament of Mirificarma. A membraneous
sac is present around the filament and supporting sclerites of Mirificarma and this is sometimes
also present around the sclerites of Gelechia. Slight sclerotized extensions from the base of the
sacculus or valva may occur in other genera, such as Chionodes Hiibner, but it is the
development of these into the long sclerites that appears to be a synapomorphy of Mirificarma
and Gelechia.
Thus I consider Mirificarma to belong in the Gelechiinae, as delimited above, on account of
the structure of the male eighth abdominal segment. Within this subfamily I consider it to be
close to Gelechia, with which it shares the presence of the sclerites described above.
Classification of Mirificarma species
The cladogram (Fig. 2) is based on the list of characters (Table 2) of Mirificarma and its
presumed close relative Gelechia. Further material of Mirificarma and related genera would be
GELECHIID MOTHS
rhodoptera
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L. M. PITKIN
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17-19
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20-22
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montivaga
scissella
rhodoptera
denotata
maculatella
pall idipul chra
aflavella
flavella
eburnella
ocellinella
fasciata
lentiginosella
constricta
cytisella
monticolella
interuptella
flavonigrella
burdonella
cabezella
ulicinella
mulinella
Fig. 2 Cladogram derived from the character matrix for the species of Mirificarma. Numbers refer to
characters listed in Table 2. Black squares (with numbers below) denote presumed apomorphies, open
squares (with numbers above) denote presumed convergences, black circles (with numbers below)
denote presumed character-reversals.
required to resolve fully the relationships within Mirificarma. Autapomorphies of individual
species are not included except where they are presumed to involve character-reversal or
convergence. The females of four species are unknown. Intermediate states are scored as
apomorphies, with the exception of those discussed in the additional comments on characters 39
and 45. Apparent convergences occur in characters 9, 13, 14, 21, 22, 26, 34, 35, 40, 42-45.
GELECHIID MOTHS
Polarities of characters have been estimated by comparison with the out-group (Gelechia and
other Gelechiinae), but this has not always been possible and therefore the polarities of some
character states are dubious.
From the analysis of the characters examined (Figs 1 , 2) montivaga has the least apomorphies
of the genus, since it is plesiomorphic in characters 1-3, whereas these are apomorphic in all
other species of Mirificarma. These other species can be divided into two groups, the macu-
latella-group and the interuptella-group. The interuptella-group is defined by the apomorphic
states of characters 4-7 and 14-16, although the last three characters are reversed in some
species (see Figs 1,2). The maculatella-group is defined by the apomorphy of character 37 and
the reversal of 8. The division of the two species-groups is corroborated by their host-plant
relationships (see p. 5).
Within the maculatella-group, the relationships oirhodoptera are not fully resolved, hence the
trichotomy in Fig. 2. In order to resolve the relationships within the interuptella-group, reversal
of characters 14-16, 20-22, 24 and 25 has been presumed.
The 'Remarks' on each species that follow (p. 19 onwards) reflect the primary concern of this
paper, namely species diagnosis and therefore stress similarities and differences between species
but do not necessarily indicate their relationships.
Table 2 Characters used in the construction of a cladogram for Mirificarma.
Character
State
Plesiomorphic
Apomorphic
1 Male filament
2 Ductusbursae
3 Aedeagus
4 Vinculum
5 Uncus
6 Tegumen lateral margins
7 Membrane between papillae
anales and eighth
abdominal segment of
female
8 Hind edge of vinculum
9 Hind wing veins Rs and MI
10 Sacculus
very weakly developed
extremely long
bulbous in basal half
without sclerites extending
from saccus, although
sometimes with short faint
sclerites extending from
anterior margin of
vinculum (sclerites of
montivaga are more similar
to this than to the
apomorphic state)
large, slightly narrower
than tegumen
with pair of small rounded
processes (often weak in
montivaga)
with two invaginations
broadly projecting but
without distinctly
demarcated median
projection
separate
broad, at least basally
[this character has four
apomorphic states -
see also 20, 26, 33]
well developed
comparatively short
uniformly slender
with pair of sclerites or
single sclerite extending
from saccus towards hind
edge of vinculum
small, usually considerably
narrower than tegumen
(small but narrowing
evenly from tegumen
in monticolella)
comparatively even
with single invagination
with distinctly demarcated
median projection, usually
comparatively narrow, or
hind edge a narrow
projection
on common stalk
moderately slender, straight
or slightly curved, with
one or more small
irregular projections at apex
10
L. M. PITKIN
Table 2 - Continued
Character
State
Plesiomorphic
Apomorphic
11 Median projection of
hind edge of vinculum
12 Projection near apex
of aedeagus
13 Pair of membraneous
sacs between apophyses
anteriores and antrum
14 Antrum
15 Saccus
16 Aedeagus/tegumen
length ratio
17 Apophyses anteriores
18 Female eighth
abdominal segment
19 Filament extent
20 Sacculus
21 Uncus
22 Filament
23 Patch of sclerotization
at posterior margin of
female seventh
abdominal segment
24 Gnathos
25 Uncus
26 Sacculus
27 Aedeagus apex
28 Apophyses anteriores
29 Female eighth sternite
30 Filament
not sharply rectangular
relatively broad [this
character has three
apomorphic states -
see also 32, 37]
absent
comparatively short
short or moderately long
low (at most 1-3)
long (at least 0-6 mm)
slightly sclerotized
far beyond hind edge
of vinculum
broad, at least basally
[this character has four
apomorphic states -
see also 10, 26, 33]
not or scarcely
constricted at base
gently curved, slightly
helical or straight
[this character has
two apomorphic states -
see also 30]
faint, diffuse, merging
with median longitudinal
band or absent
a large hook, or complex
broad or not narrowing
from base towards apex
not stouter apically [this
character has four
apomorphic states -
see also 10, 20, 33]
not cylindrical
rod-like
without strongly contrasting
areas of sclerotization
narrowing evenly towards
apex [this character has
two apomorphic states -
see also 22]
sharply rectangular,
scarcely emarginate,
membraneous
slight, very narrow,
distinctly sclerotized
present
extremely long
extremely long
high (1-7-1 -8)
short (less than 0-6 mm)
well sclerotized (either
uniformly, or with patches
of very strong sclerotization)
at most a comparatively
short distance beyond hind
edge of vinculum
slender, moderately curved,
with even apex
constricted at base (more so
in constricta than in
lentiginosella)
with apical half kinked,
basal half straight
well defined
small simple hook-shape
small and narrowing from
base towards apex
stouter towards apex (club-
shaped or slender and
spatulate)
almost cylindrical
lobe-like
strongly sclerotized
laterally, contrasting with
weakly sclerotized median area
with basal half
dorsoventrally expanded
and laterally compressed
GELECHIID MOTHS
11
31 Posterior margin of
female seventh
abdominal segment
32 Projection near apex
of aedeagus
33 Sacculus
34 Signum
35 Signum
36 Filament-supporting
sclerites
37 Projection near apex
of aedeagus
38 Fore wing pattern
39 Median longitudinal
band of sclerotization
of female seventh
abdominal segment
40 Antrum
41 Fore wing pattern
42 Fore wing pattern
43 Gnathos
44 Saccus
45 Female frenulum
with, at most, diffuse
band of scales
small, sclerotized, evenly
tapering [this character
has three apomorphic
states - see also 12, 37]
broad, at least basally
[this character has four
apomorphic states -
see also 10, 20, 26]
present
with very many small
spines or without spines
not twisted
small, evenly tapering
[this character has
three apomorphic states
-see also 12, 32]
small spots or uniform
[this character has
three apomorphic states -
see also 41, 42]
if present, not or slightly
constricted
curved or constricted
towards apex
small spots or uniform
[this character has
three apomorphic states
see also 38, 42]
small spots or uniform
[this character has three
apomorphic states -
see also 38, 41]
curved
broad, at least basally
three setae
with distinct, dense band
of scales
weakly sclerotized, rounded
evenly slender, straight and
smooth (the particularly
slender sacculus of
mulinella and ulicinella
represents a further
development - 33x)
absent
with large spines (small
spines sometimes also
present)
twisted (weakly twisted in
flavella)
large, angular, hooked,
narrowing to a point
zig-zag yellowish markings
strongly constricted
straight and not constricted
towards apex
large spots
median longitudinal stripe
(broken or unbroken)
almost straight (short in
ulicinella, long in
fasciata)
relatively slender
usually two setae
Additional comments on the characters listed in Table 2
5 M. ocellinella and fasciata are intermediate, more similar to the plesiomorphic than to the
apomorphic state.
8 M. montivaga is classed as intermediate since the hind edge of the vinculum sometimes
approaches the apomorphic state.
9 This character is not very reliable since both states are found in some species, although one
state is always greatly predominant.
16 Aedeagus/tegumen length ratio is intermediate in lentiginosella (1-4-1-5).
18 This character is difficult to observe (since the sclerotization is more obvious in well-stained
specimens) but trends of difference between species can be observed. M. montivaga has the
12 L. M. PITKIN
least sclerotization. M. lentiginosella and cytisella have larger areas of sclerotization than in
species with the plesiomorphic state but smaller areas or weaker sclerotization than in the
other species with the apomorphic state.
19 The species with the longest filaments of the apomorphic state, in relation to tegumen length,
are marked as intermediate.
21 M. interuptella is intermediate between the two states, with the base of the uncus constricted,
usually slightly, in a minority of specimens.
22 M. interuptella is marked as intermediate since the filament is kinked throughout its length.
26 The saccus offlavella is occasionally stouter apically but is usually more uniformly stout than
in the apomorphic state.
35 It is impossible to assess this character in denotata and the other species in which the signum is
absent, or in eburnella which has a barely discernible signum.
37 The aedeagus projection of montivaga (plesiomorphic state) is always small in relation to
aedeagus width. M. rhodoptera and eburnella are intermediate with an aedeagus projection
of the derived shape but small. M. scissella is intermediate with a more rounded projection
than the derived state.
39 The derived state is not to be confused with the slight constriction of the band of
sclerotization in the several species marked as intermediate and not scored on the
cladogram.
40 M. denotata is classed as intermediate since its antrum narrows towards the apex more than
that oiflavella and eburnella. It is not appropriate to score this character in species with an
extremely short antrum.
42 In the species marked as intermediate both states are present in different specimens.
45 Too few specimens have been examined to be certain of the character state of every species,
since this character is sometimes very variable intraspecifically. As this character is
unstable, only the species which usually have two setae are scored as apomorphic. Species
with an equal occurrence of two and three setae or with a tendency towards three setae are
classed as intermediate and are not scored on the cladogram. This character seems to show a
trend towards a relationship between mulinella, ulicinella and cabezella, which have the
apomorphic state, although this is also shared by cytisella.
The structure of the filament of the male genitalia
In his original description of Mirificarma, Gozmany referred to the filament as 'a very long
filamental prong' accompanying the aedeagus. It is orientated longitudinally and is near to the
aedeagus in the undissected genitalia, but it has no physical connection with the aedeagus. The
aedeagus lies between the ventral surface and the dorsal membrane of the vinculum, whereas
the filament is more dorsal, lying between the dorsal membrane of the vinculum and the
tegumen. Saltier (1960: 41) compared the filament of Mirificarma with processes of the transtilla
in Bryotropha Heinemann and Filatima Busck, but these structures are not homologous with the
filament.
The filament is very weakly developed in montivaga, in which it is merely a slight sclerotized
swelling where the supporting sclerites meet. In all other Mirificarma species it is a long,
well-sclerotized tube, usually cylindrical, although narrowing gradually, posteriorly, and usually
very narrow at the posterior end; usually gently curved or, if very long, sometimes slightly
helical. Ocasionally the filament is straight, as in ocellinella, or modified in shape, for example
with a strong kink near the apex in lentiginosella and with a projecting lobe in constricta and
ulicinella. The filament varies in length between species, being shortest in relation to tegumen
length in flavonigrella and ulicinella. In the latter, the filament does not usually reach the
anterior of the tegumen and extends posteriorly only as far as the hind edge of the vinculum. The
filament is longest in relation to tegumen length in pallidipulchra, in which it extends very far
beyond the anterior of the tegumen and extends posteriorly beyond the uncus.
The filament is open anteriorly and has a small opening near the posterior end. This opening is
almost terminal in flavella and ocellinella whereas it is situated further from the posterior in
GELECHIID MOTHS 13
maculatella and cabezella. In scissella a second tiny posterior opening is present. In ocellinella
and denotata a tiny projection occurs from the posterior opening of the filament. In maculatella a
similar projection is situated more posteriorly. The filament surface of all species appears
smooth except for occasional minute spines.
The filament arises from a pair of lateral sclerotized strips, referred to here as filament-
supporting sclerites. These extend from the base of the sacculus and, to a less extent, the valva,
and meet anteriorly, at which point they fuse to form the filament. The filament is attached only
at its anterior, and the length of the sclerites thus varies with the anterior extent of the filament.
In many species the filament-supporting sclerites appear to be a smooth continuation of the
sacculus. The sclerites are usually parallel-sided over much of their length, at least in the species
in which they are long, but they are widened, often considerably and irregularly, towards the
sacculus and valva. This expanded area of the sclerite is often faintly rugose and may represent
an area of muscle attachment. This feature is not confined to Mirificarma since similarly placed,
although even fainter, rugose patches are present in Gelechia nigra. The filament-supporting
sclerites of Mirificarma are widened anteriorly, where they meet.
The filament and supporting sclerites are covered ventrally by a membraneous sac which is
attached to the sclerites and the anterior end of the filament. The sclerites are spiralled round
each other in some species and in these the membrane is similarly twisted. A membraneous sac is
usually also present dorsally, from the tegumen. These membranes are flimsy and the dorsal
membrane in particular tends to disintegrate during the preparation of genitalia slides. The
dorsal membrane is relatively well developed in cabezella and mulinella, for example, but it is
most obvious in ulicinella.
The function of the filament is unknown. There is some correlation between its length and that
of the female apophyses anteriores. There appears to be no correlation between the length of the
filament and that of the aedeagus such as the correlation discussed between the saccus and the
aedeagus (see p. 15).
MIRIFICARMA Gozmany
Helina Guenee, 1849: 411. Type-species: Carcina flammella Hiibner, [1825], by monotypy. [Junior
homonym of Helina Robineau-Desvoidy, 1830 (Diptera).]
Mirificarma Gozmany, 1955: 308, 309 [keys], 313. Type-species: Tinea maculatella Hiibner, 1796, by
original designation.
O", $ (4-5) 5-0-11-0 mm. Head without frontal modifications. Ocellus present. Proboscis well developed;
squamose, particularly at base. Maxillary palpus with four segments. Labial palpus recurved, first segment
much shorter than second; third segment approximately same length as second or, rarely, slightly shorter;
second segment without brush below or, rarely, with slight to moderate brush. Antenna without pecten on
scape. Metascutum with paired group of narrow hair-like scales. Fore wing sometimes with zig-zag pattern
of yellowish markings; sometimes with large or small dark spot across fold and another at end of cell;
sometimes with dark, median, longitudinal, broken or unbroken stripe; a few species without any of these
patterns. Fore wing with veins R 4 and R 5 on long common stalk (Fig. 3); distance R\-R2 slightly greater
than, to four times R^-R^', M^ and Cui separate. Hind wing with veins Rs and MI on common stalk or
separate (Figs 3,4). Frenulum of female with three or sometimes two long setae.
GENITALIA cf Eighth tergite and sternite separated into free flaps. Eightn tergite with pair of coremata
consisting of dense brushes of long, thin, hair-like scales, inserted laterally at anterior of tergite and
attached to ventral membrane of tergite. Dorsal membrane of sternite with coremata consisting of two
groups of large, rounded, often inflated, grape-like scales covering most of sternite. Genitalia withdrawn
inside eighth segment. Uncus rounded, often large. Gnathos well-sclerotized, usually simple hook.
Gnathos base with small membraneous area densely covered with microtrichia. Tegumen with or without
lateral pair of rounded processes. Anterior margin of tegumen with deep V- or occasionally U-shaped
emargination. Valva long, usually reaching uncus, slender, simple, slightly swollen at rounded apex.
Sacculus clearly separated from valva. Long pair of sclerotized strips present, extending from base of valva
and sacculus and meeting anteriorly. Long sclerotized tube (filament) present, arising from where pair of
sclerotized strips join; extending posteriorly. In montivaga, filament extremely small; merely slight
swelling of sclerotization. Hind edge of vinculum usually with medially emarginate median projection.
Vinculum extends far posteriorly, hind edge reaching anterior of gnathos in a few species. Vinculum with
14
L. M. PITKIN
Figs 3, 4 Wing venation of Mirificarma species, d* . 3, M. maculatella (Hiibner). 4, hind wing of M.
interuptella (Hubner).
or without pair of narrow sclerites or, exceptionally, single narrow sclerite, extending from saccus towards
hind edge of vinculum. Saccus ranging from very slight projection to extremely long. Aedeagus usually
cylindrical, slender; occasionally inflated basally; without cornuti. Projection near apex of aedeagus
ranging from scarcely discernible to large, hook-like. Anterior part of ductus ejaculatorius often with
sclerotized lamina.
GENITALIA. $. Dorsal and ventral abdominal surfaces sometimes form broad, median, longitudinal
sclerotized band. Posterior margin of seventh segment sometimes with more strongly sclerotized patches.
Anterior margin of seventh sternite usually unmodified but in one species (pallidipulchrd) with pair of
sclerites and membraneous sac. Papilla analis usually longer than broad. Invagination of membrane
between eighth tergite and papillae anales usually present, immediately opposite invagination, sometimes
present, of membrane between sternite and papillae anales. Eighth segment with predominantly lateral
areas of sclerotization; occasionally almost completely sclerotized. Apophysis anterior usually rod-like,
occasionally short , broad lobe ; length 0-2-1 -3mm; apophysis posterior two to six times length of apophysis
anterior in each species. Sclerotized antrum present, usually long, nearly as long as apophysis anterior to
much longer; occasionally extremely short. Ductus bursae often much shorter than corpus bursae,
sometimes longer, exceptionally six times length of corpus bursae. Ductus seminalis arises from posterior
part of ductus bursae, usually very near antrum. Corpus bursae with minute spines on inner surface
; GELECHIID MOTHS 15
sometimes sparse, sometimes extending into ductus bursae. Corpus bursae with or without signum.
Signum, if present, usually oval or round, usually spinose; spines, if present, directed inside corpus bursae;
signum sometimes curved inwards, never hook-like.
REMARKS. The fore wing venation of Mirificarma has been checked in nine specimens and is
similar to that of many Gelechiidae but appears to differ from that of many Gelechia species. In
Mirificarma the fore wing veins A/ 3 and Cu\ are separate whereas these are frequently on a
common stalk in Gelechia.
The coremata of the male eighth abdominal segment of Mirificarma are readily lost during the
preparation of genitalia slides although the scale bases can still be seen. The membraneous area
of the base of the gnathos is much smaller in Mirificarma than in Gelechia and the Gnorimosche-
mini in both of which it is an extensive sac. This structure is seen in some genera outside the
Gelechiinae, for example Dichomeris Hiibner and Acompsia. The strongly sclerotized hook-
shaped gnathos usual in Mirificarma occurs in various Gelechiidae but not in Gelechia.
Mirificarma is distinguished by the presence of the filament of the male genitalia (see also the
discussions on pp. 5, 6).
The narrow sclerites of the vinculum which extend from the saccus towards the hind edge of
the vinculum in the interuptella-group of Mirificarma are not usually present in the Gelechiidae,
although they occur in Psoricoptera gibbosella which may be related to Gelechia and Mirificarma
(see p. 6). In the maculate '//a-group the sclerotized anterior margin of the vinculum is
occasionally continued towards the median across the posterior of the saccus as a pair of very
faint short sclerites. Similar, though longer and very distinct, sclerites occur in the Gnorimosche-
mini. These sclerites, unlike those of the interuptella-group, do not appear to arise from the
saccus itself. The vinculum in Mirificarma is longest in interuptella, sometimes almost reaching
the gnathos base of this species.
The aedeagus is usually more slender than in Gelechia. There appears to be some correlation
between the length of the saccus and the aedeagus in Mirificarma. In relation to the tegumen
length, these structures are both exceptionally long in ocellinella and fas data; in lentiginosella
they are slightly shorter, although still longer than in the remaining Mirificarma species. The
saccus is shortest in rhodoptera and in this species the aedeagus is also the shortest. In the species
with the longest saccus the aedeagus exceeds 1-5 times the length of the tegumen and the
sclerotized saccus may serve to support the aedeagus where it extends beyond the tegumen
anteriorly. The aedeagus is attached to the anterior of the saccus by retractor muscles in the
representative of the Gelechiidae examined by Kuznetsov & Stekol'nikov (1978: 131) and there
is a similarly attached muscle in Mirificarma fasciata.
Sattler (1976: 93) mentions an apparent correlation between the length of the ductus
ejaculatorius of the male and the ductus bursae of the female in the species of Ornativalva
Gozmany, currently in the Gelechiini, in which these structures are exceptionally long.
Likewise, in Mirificarma, the ductus ejaculatorius and the ductus bursae are both considerably
longer in montivaga than in the rest of the genus. There is no obvious correlation in the other
Mirificarma species, in which both structures are shorter.
Mirificarma species with slight sclerotization of the female eighth sternite often have
membraneous areas between the apophyses anteriores and the antrum, although the apophyses
anteriores and the lateral margins of the antrum are usually continued into the sternite as narrow
sclerites, reaching the sclerotized areas of the sternite. In montivaga there is little sclerotization
of the sternite other than the continuation of the apophyses anteriores. The longitudinal band of
the female abdomen in some species of Mirificarma is faint and is best observed in well-stained,
freshly prepared specimens. I have included this feature in the species diagnoses only when it is
characteristic. The band is also present in the male but is less useful as a character for species
diagnosis.
The sclerotized antrum of Mirificarma is usually long, whereas it is usually very short if present
in Gelechia. Most Mirificarma species have a rounded, spinose signum. Only montivaga has a
signum of the type common in Gelechia, diamond-shaped with a pair of serrated-edged ridges.
16 L. M. PITKIN
BIOLOGY. The known host-plants of Mirificarma are all Leguminosae and are discussed under
each species and in the section on host-plant relationships. Although most of the species of
Mirificarma are not known to have any economic significance, eburnella has been reported as an
agricultural pest in the U.S.A., where it damages clover crops (see p. 28).
The egg stage and site of egg deposition on the host-plant are unknown. The larva makes a
spinning amongst the leaves of the host-plant or feeds in the shoots. In plants with small leaves,
such as Ulex, the larva usually feeds in the flowers. Where details are known, pupation usually
takes place in a slight cocoon amongst fallen leaves on the ground, although eburnella is
recorded as pupating between the leaves on the host-plant. There are few records of the number
of generations per year; two species are reputed to have only one generation but two are
recorded for eburnella. Species may breed continuously where climatic conditions permit as is
suggested by the wide range of dates on which ocellinella has been collected.
DISTRIBUTION. Mirificarma species are distributed in the Palaearctic region between 55 N and
30 S, extending east to c.45. One species, eburnella, also occurs in the U.S.A., where it has
been introduced.
Key to the species of Mirificarma
Males
1 Fore wing with large dark spots (Figs 9,10) 2
- Fore wing with small dark spots or different markings 3
2 Fore wing with large dark spot across fold narrowing gradually from middle towards costa;
spot at end of cell comparatively distinct from patch at costa (Fig. 10). Portion of aedeagus
apical to hook large (Fig. 65) maculatella (p. 23)
Fore wing with large dark spot across fold narrowing sharply from middle towards costa; spot
at end of cell merging with patch at costa (Fig. 9). Portion of aedeagus apical to hook small
(Fig. 64) denotata (p. 22)
3 Filament scarcely discernible (Fig. 60) montivaga (p. 18)
Filament well developed (Figs 61, 62, 64-82) 4
4 Uncus large, occasionally constricted at base but otherwise only slightly narrower than
tegumen (Figs 61, 62, 66-72). Tegumen with pair of rounded lateral processes as in Fig. 31, or
if absent, saccus extends far beyond tegumen anteriorly (Figs 70-72) 5
- Uncus small, and usually considerably narrower than tegumen; tegumen without rounded
lateral processes; saccus extends a comparatively short distance beyond tegumen anteriorly
(as in Figs 76, 79-82) 13
5 Tegumen without rounded lateral processes; saccus extends far beyond tegumen anteriorly
(Figs 70-72) 6
- Tegumen with pair of rounded lateral processes as in Fig. 31; saccus extends at most a short
distance beyond tegumen anteriorly (Figs 61 , 62, 66-69) 8
6 Filament strongly kinked near apex (Fig. 72) lentiginosella (p. 32)
Filament straight (Figs 70, 71) 7
7 Saccus of uniform width; basal half of gnathos increasingly very broad then narrowing sharply
to hook-shaped apical half (Figs 46, 71) ocellinella (p. 29)
Saccus spatulate, apex approximately twice width of base; gnathos slightly curved, basal half
unmodified (Figs 47, 70) fasciata (p. 31)
8 Filament not extending beyond apex of valva (Figs 61 , 68, 69) 9
- Filament extends beyond apex of valva (Figs 62, 66, 67) 11
9 Fore wing with zig-zag pattern of yellowish markings (Figs 11-14). Hind wing with veins Rs and
M j separate , as in Fig. 3 10
Fore wing without yellowish zig-zag markings. Hind wing with veins Rs and MI on common
stalk, as in Fig. 4 scissella (p. 20)
10 Sacculus with rounded, moderately broad apex (Fig. 68) fiavella (p. 26)
Sacculus with pointed, moderately slender apex (Fig. 69) eburnella (p. 27)
11 Fore wing without zig-zag pattern of yellowish markings. Saccus extremely short and very
broad (Figs 35, 36, 62) rhodoptera(p.20)
- Fore wing with zig-zag pattern of yellowish markings (Figs 11,12). Saccus moderately short and
broad (Figs 39, 40, 66, 67) 12
GELECHIID MOTHS 17
12 Filament extends far beyond apex of valva posteriorly, and very far beyond tegumen anteriorly
(Fig. 66) pallidipukhra (p. 24)
- Filament extends short distance beyond apex of valva posteriorly, and moderately beyond
tegumen anteriorly (Fig. 67) afiavella (p. 25)
13 Sacculus uniformly slender (Figs 73, 80-82) 14
- Sacculus broad, or broader apically than basally, club-shaped or spatulate; otherwise very
small, hook-shaped (Figs 57, 59, 74-79) 17
14 Uncus distinctly constricted at base; filament helical in apical half; sacculus gently S-curved
(Fig. 73) constricta (p. 33)
Uncus scarcely constricted at base; filament almost straight or curved dorsoventrally, not
helical ; sacculus almost straight (Figs 80-82) 15
15 Gnathos almost straight; sacculus does not reach gnathos arms (Fig. 81) ulicinella (p. 42)
- Gnathos distinctly curved ; sacculus reaches or extends beyond gnathos arms (Figs 80 , 82) 16
16 Filament stout, particularly basally, laterally compressed apically; median projection of hind
edge of vinculum high (Fig. 82) mulinella (p. 43)
- Filament very slender; median projection of hind edge of vinculum low (Fig. 80) cabezetta (p. 41)
17 Sacculus stoutly club-shaped (Figs 57, 76, 78, 79) 18
- Sacculus slender and spatulate (Fig. 77) or small and hook-shaped (Figs 74, 75) 20
18 Gnathos extremely short ; saccus very slender; filament extends to hind edge of vinculum or just
beyond (Figs 49, 76) monticolella (p. 37)
- Gnathos developed normally; saccus comparatively broad; filament does not reach hind edge
of vinculum (Figs 51, 52, 78, 79) 19
19 Sacculus scarcely extends beyond median projection of hind edge of vinculum (Fig. 79)
burdonella (p. 40)
- Sacculus extends far beyond median projection of hind edge of vinculum (Fig. 59)
fiavonigrella (p. 39)
20 Gnathos very large, without median spine; sacculus long, slender, spatulate (Fig. 77)
interuptella (p. 38)
- Gnathos small, with median, sometimes minute, spine; sacculus small, hook-shaped (Figs 74,
75) cytisella (p. 34)
Females
Note. The females ofscissella, monticolella and flavonigrella are unknown; constricta is also omitted since
the female genitalia are unknown.
1 Fore wing with large dark spots (Figs 9, 10) 2
Fore wing with small dark spots or different markings 3
2 Fore wing with large dark spot across fold narrowing gradually from middle towards costa; spot
at end of cell comparatively distinct from patch at costa (Fig. 10). Signum present; antrum
curved towards anterior, without indented anterior (Fig. 89) maculatella (p. 23)
- Fore wing with large dark spot across fold narrowing sharply from middle towards costa ; spot at
end of cell merging with patch at costa (Fig. 9). Signum absent or scarcely discernible;
antrum straight , with indented anterior (Fig. 88) denotata (p. 22)
3 Antrum same length as apophysis anterior or shorter (Figs 85-87 , 90-92 , 98) 4
Antrum extends beyond apophysis anterior (Figs 94, 96-103) 10
4 Antrum extremely short (Figs 85 , 98) 5
- Antrum comparatively long (Figs 86, 87, 90-92) 6
5 Signum a narrow ridge with faint surround (Fig. 98). Ductus bursae slightly longer than corpus
bursae or of similar length cytisella (p. 34)
- Signum elongate-oval to diamond-shaped with serrated edge (Fig. 85). Ductus bursae approx-
imately four to six times length of corpus bursae montivaga (p. 18)
6 Antrum almost straight, narrowing evenly towards anterior (Figs 92, 93). Fore wing with
zig-zag pattern of yellowish markings (Figs 13,14) 7
- Antrum curved or constricted towards anterior (Figs 86, 90, 91); if not curved and only very
slightly constricted (Fig . 87) , forewing without zig-zag pattern of yellowish markings 8
7 Antrum with rounded or scarcely indented anterior; abdomen with distinct, longitudinal,
median band (Figs 93, 109) eburnella (p. 27)
- Antrum with strongly indented anterior (Fig. 92); abdomen with, at most, very faint band
ttavella (p. 26)
8 Abdomen with pair of sclerites and membraneous sac at anterior margin of seventh segment
(Fig. 107) pallidipulchra(p. 24)
18 L. M. PITKIN
Abdomen with unmodified anterior margin of seventh segment 9
9 Signum with approximately three or four large spines projecting from one side, otherwise
smooth; seventh abdominal segment comparatively broad (Figs 91, 108) attavella(p. 25)
Signum covered with numerous spines of various sizes; seventh abdominal segment narrow
(Figs 86, 87, 105,106) rhodoptera (p. 20)
10 Antrum very long, coiled or strongly curved (Figs 94, 96, 97) 11
Antrum moderately long, not coiled or strongly curved (Figs 98-103) 13
11 Apophyses anteriores parallel, long; pair of membraneous sacs from sternite, between
apophyses anteriores and antrum (Figs 94, 96) 12
Apophyses anteriores diverging, short; without sacs between apophyses anteriores and antrum
(Fig. 97) lentiginosella(p.32)
12 Apophysis posterior three to four times length of apophysis anterior. Unsclerotized median
area of eighth sternite at least as wide as each lateral sclerotized area (Fig. 94) ocellinella (p. 29)
Apophysis posterior twice length of apophysis anterior. Unsclerotized median area of eighth
sternite narrower than each lateral sclerotized area (Fig . 96) fasciata (p . 3 1 )
13 Apophysis anterior lobe-shaped; eighth sternite strongly sclerotized laterally, contrasting with
weak sclerotization of median area (Figs 99, 100) 14
Apophysis anterior rod-like (Figs 93, 101-103); eighth sternite slightly more sclerotized
laterally than medially 15
14 Signum present; median area of eighth sternite without horizontal striations (Fig. 99)
interuptella (p. 38)
Signum not discernible; median area of sternite with horizontal striations (Fig. 100)
burdonella (p. 40)
15 Antrum straight , narrowing evenly towards anterior ; abdomen with median longitudinal band ,
very distinct in seventh segment (Figs 93, 109) eburnella (p. 27)
Antrum curved and constricted towards anterior (Figs 101-103); seventh abdominal segment
without median longitudinal band 16
16 Base of apophysis anterior with separate rounded lobe towards antrum (Fig. 101) cabezella (p. 41)
Base of apophysis anterior without separate lobe (Figs 102, 103) 17
17 Eighth tergite with pair of curved sclerotized lobes overlapping median longitudinal area
(Fig. 102) ulidnella (p. 42)
Eighth tergite without lobes (Fig. 103) nwlinclhi (p. 43)
The montivaga-group
Characters as described under montivaga.
Mirificarma montivaga (Walsingham) comb. n.
(Figs 1,2, 6, 32, 33, 60, 85, 104)
Gelechia montivaga Walsingham, 1904: 221. LECTOTYPE cf, ALGERIA (BMNH), here designated
[examined].
Gelechia pulverosella Zerny, 1936: 137, pi. 2, fig. 44. LECTOTYPE cf , MOROCCO (NM), here designated
[examined]. Syn. n.
O" , 7-5-8-5 mm. $ , 7-0-7-5 mm. Head cream. Labial palpus cream mottled with brown on outer surface.
Thorax, tegula and fore wing (Fig. 6) cream to light brown mottled with brown. Hind wing with veins Rs
and M\ separate as in Fig. 3.
GENITALIA cf (Figs 32, 33, 60). Uncus large, only slightly narrower than tegumen. Gnathos a moderately
large simple hook. Tegumen with pair of rounded lateral processes, sometimes very slight. Actual margin
of tegumen slightly less emarginate than sclerotized margin anteriorly; sclerotized margin distinctly
X-shaped medially. Sacculus moderately broad. Filament scarcely discernible; filament-supporting scler-
ites not twisted around each other. Hind edge of vinculum a broad projection, scarcely emarginate
medially. Vinculum short, with pair of very short faint sclerites converging from anterior edge. Saccus
narrowing slightly towards rounded apex, extending short to moderate distance beyond tegumen anterior-
ly. Aedeagus same length as tegumen plus uncus or slightly shorter. Aedeagus with bulbous basal half,
small to moderate projection near apex, and long ductus ejaculatorius.
GENITALIA $ (Figs 85, 104). Posterior margin of seventh abdominal segment with faint dorsal patch of
sclerotization and pair of narrow longitudinal stripes ventrally. Very flimsy, broad, rounded invagination
GELECHHD MOTHS 19
of membrane between eighth tergite and papillae anales, directly opposite less flimsy, long, narrow
invagination of membrane between sternite and papillae anales. Eighth segment with very weak sclejotiza-
tion, longitudinally wrinkled, minutely spined anteriorly. Apophysis anterior rod-like, length 0-8-0-9 mm
(2); apophysis posterior approximately three times length of apophysis anterior. Antrum slightly indented
at anterior and extremely short. Ductus bursae gently coiled, narrow; coiled extent approximately four to
six times length of oval or round corpus bursae. Signum very large, elongate oval to diamond shape, with
deep longitudinal indentation edged with pair of serrated-edged projecting ridges.
REMARKS. The frenulum was examined in the four females and consists of three setae in three of
these and two setae in one, on both wings. The X-shaped anterior margin of the tegumen is
particularly distinct in montivaga, although it occurs to a less degree in constricta and is very
weak in several other Mirificarma species.
M. montivaga is the only species of the genus in which the fore wing has no pattern other than
the mottling of the ground colour, although the markings are extremely small in monticolella,
lentiginosella and a few specimens of ocellinella. The ground colour is much darker in
lentiginosella. M. montivaga differs from the rest of the genus in genitalic characters including
the scarcely discernible male filament which is merely represented by a slight swelling between
the supporting sclerites, the shape of the signum and the extremely long ductus bursae of the
female. The long ductus bursae approaches that of rhodoptera, but this species has a much
longer antrum than montivaga.
M. montivaga was described from 3 cf , 3 9 from Algeria: El Kantara, all of which I have
examined. I designate the specimen bearing Walsingham's number 96484 as the lectotype. This
is referred to as the 'Type cf" on the specimen label and in the original description in which the
date of capture is given as 3. v. 1903; however, 1 cf , 1 9 > bearing Walsingham's paratype labels,
are labelled 9. v. 1903.
G. pulverosella was described from 8 cf , 5 9 from Morocco, of which I have examined 3 cf ,
1 9- Of these 1 cf , 1 $ bear the labels Type cf' and Type 9' respectively and I designate the
male as the lectotype. The 5 cf , 4 9 syntypes which I have not examined are from Tachdirt,
middle and end vii (Zerny) .
BIOLOGY. Host-plant unknown. Moths have been found from May to July.
DISTRIBUTION. Morocco and Algeria.
MATERIAL EXAMINED (including 6 cf , 3 9 genitalia preparations)
Lectotype cf (montivaga), Algeria: [Constantine province,] El Kantara, 3. v. 1903 (Walsingham) (genita-
lia slide no. 7124). Lectotype cf (pulverosella), Morocco: Haul ('Grosser') Atlas, c. 17 km SE. of Asm, in
highest Iminene Valley, Tachdirt, 2200-2900 m, ll-19.vii. 1933 (Zerny) (genitalia slide no. 11223; NM).
Morocco: 1 cf (pulverosella paralectotype), Haut Atlas, c. 70 km SW. of Marrakech, Goundafa area,
3 km above Kasbah Goundafa, at junction of Agoundi Valley in Fis Valley, 'n Zala, above Ijjoukak,
1500-1800 m, 21-29.vi.1933 (Zerny) (NM); 1 cf, Haul Atlas, Oukaimedene, 2500-2900 m, vii (coll.
Burmann, Innsbruck); 1 cf, 1 9 (pulverosella paralectotypes), Haut Atlas, Tachdirt, 2200-2900 m,
ll-19.vii.1933 (Zerny) (NM); 1 cf, Moyen Atlas, Val d'Ifrane, 1600-1700 m, vi (coll. Burmann,
Innsbruck). Algeria: 2 cf , 3 9 (montivaga paralectotypes), El Kantara, 3, 9. v. 1903 (Walsingham).
The maculatella-group
Cf, 9- Fre wing sometimes with zig-zag pattern of yellowish markings; without median longitudinal
stripe. Hind wing with veins Rs and M l separate or on common stalk.
GENITALIA cf . Uncus large, only slightly narrower than tegumen. Gnathos a large simple hook. Tegumen
with pair of rounded lateral processes. Filament-supporting sclerites usually crossed or spiralled round
each other. Hind edge of vinculum a low, broad, sclerotized projection, usually with broad emargination,
but without distinctly demarcated median projection. Vinculum without sclerites which extend from saccus
and approach hind edge of vinculum. Projection near aedeagus apex usually large, angular and narrowing
to point.
GENITALIA 9- Invagination of flimsy membrane between eighth tergite and papillae anales present,
immediately opposite invagination of slightly less flimsy membrane between sternite and papillae anales.
Eighth segment weakly sclerotized. Apophysis anterior rod-like, long. Signum, if well developed, with
large spines; small spines sometimes also present.
20 L. M. PITKIN
REMARKS. The filament-supporting sclerites of the male genitalia are more strongly spiralled
round each other in the species in which they are long, particularly maculatella and rhodoptera.
Although the vinculum does not have sclerites extending from the saccus and approaching the
hind edge of the vinculum, in a few species the anterior margin of the vinculum is continued
towards the median as faint, narrow, short sclerites on either side of the saccus. This is seen in
rhodoptera and pallidipulchra and, even more faintly, in one specimen of aflavella; in scissella
the anterior margin of the vinculum is sometimes continuous across the saccus posterior as a
narrow sclerite. The sclerotization of the eighth segment is uniformly weak or limited to smaller
areas than in most of the interuptella-group, but is always greater than in montivaga.
BIOLOGY. Host-plants: Trifolieae (other than Ononis); Loteae; Coronilleae; Galegeae; Vicieae.
Mirificarma scissella (Chretien) comb. n.
(Figs 1,2, 7, 34, 61)
Gelechia scissella Chretien, 1915: 319. LECTOTYPE cf , ALGERIA (MNHN), here designated [examined].
Cf , 7-0-7-5 mm. Head whitish cream. Labial palpus cream, mottled with brown at base. Thorax and tegula
cream mottled with brown. Fore wing (Fig. 7) cream to pale brown, mottled with brown; slightly darker at
extreme base and at costa in four-fifths. Dark brown spot across fold at one-third with trace at dorsal
margin. Small dark brown spot at end of cell. Hind wing with veins Rs and MI on common stalk as in Fig. 4.
GENITALIA cf (Figs 34, 61). Actual margin of tegumen less emarginate than sclerotized margin anteriorly.
Sacculus broad at base, slightly wrinkled at narrower apex. Filament very slightly curved, slender, not
reaching apex of valva posteriorly, extending short to moderate distance, beyond tegumen anteriorly.
Filament-supporting sclerites not or scarcely twisted. Hind edge of vinculum with shallow U- or V-shaped
emargination of sclerotization. Saccus usually moderately broad, scarcely narrower towards rounded
apex, extending short distance beyond tegumen anteriorly. Aedeagus approximately same length as
tegumen plus uncus, slender; apical projection moderately large, not distinctly pointed.
GENITALIA $. Unknown.
REMARKS. This species bears a superficial resemblance to rhodoptera, constricta and cabezella,
although the head is paler and the spot in the fold of the fore wing scarcely extends to the dorsal
margin; however, the only specimens known of scissella are in poor condition externally. It
differs from rhodoptera in the hind wing venation, and the shorter filament and longer saccus of
the male genitalia. It differs from constricta and cabezella in some genitalic characters, including
the presence of a pair of rounded processes on the male tegumen, since these last two species are
in the interuptella-group.
M. scissella was described from an unspecified number of specimens of which I designate as
lectotype the single type-specimen examined. The coating of debris on this specimen renders it
almost unrecognizable externally. One of the rounded processes of the tegumen is absent in the
lectotype.
BIOLOGY. Host-plant unknown. Moths have been found in April and May.
DISTRIBUTION. Algeria.
MATERIAL EXAMINED (including 2 cf genitalia preparations)
Lectotype cf , Algeria: [Constantine province,] Biskra, 7. v. 1907 (genitalia slide no. 43, LMP; MNHN).
Algeria: 1 cf , M'zab ('Uzab') country, Oued Nsa ('Nza'), iv; 1 cf , Hamman-es-Salahine, iv.
Mirificarma rhodoptera (Mann)
(Figs 1, 2, 8, 35, 36, 62, 63, 86, 87, 105, 106)
Gelechia rhodoptera Mann, 1866: 353, pi. 1 (B), fig. 10. Holotype cf, RUMANIA ('Turkey') (NM)
[examined].
Cf, 5-0-7-5 mm, $,5-5-7-0 mm. (Typical form, cf , 6-5-7-5. $, 6-0-7-0. Small form, cf , 5-0-6-5 mm. <j>,
5-5-6-0 mm.) Both forms. Head mid brown. Labial palpus mottled cream and brown; darker on outer
surface. Thorax and tegula mottled mid brown. Fore wing (Fig. 8) mottled light brown, very faintly
GELECHIID MOTHS 21
pink-tinged, and dark brown; darker at base and in apical third; ochreous tinge near base. Small cream
patch at costa at two-thirds extending diffusely to dorsal margin. Narrow, transverse dark brown spot
across fold at one-third extending to dorsal margin. Smaller dark brown spot at end of cell, constricted
medially, occasionally bisected. Both spots with ochreous cream surround. Very small diffuse dark brown
spot in fold at one-fifth. Hind wing with veins Rs and M\ separate as in Fig. 3.
GENITALIA cf (Figs 35, 36, 62, 63). Actual margin of tegumen less emarginate than sclerotized margin
anteriorly. Sacculus short, moderately broad, wrinkled. Filament slightly helical, slender, extending
posteriorly moderately beyond apex of valva, and very far beyond tegumen anteriorly. Hind edge of
vinculum with broad, shallow U- or V-shaped emargination of sclerotization. Saccus very broad, usually
with very shallow emargination at apex; extremely short, never reaching anterior margin of tegumen.
Aedeagus approximately three-quarters length of tegumen plus uncus, basal half very slightly swollen;
apical projection small.
GENITALIA $ (Figs 86, 87, 105, 106). Posterior margin of seventh abdominal segment with area of weak
sclerotization merging with progressively faint median longitudinal band. Seventh abdominal segment
comparatively narrow, anterior margin unmodified. Invagination of membrane between eighth sternite
and papillae anales indistinct, extremely long and narrow. Eighth sternite with lateral pair of weakly
sclerotized strips. Tergite and rest of sternite very weakly sclerotized, with minute spines mainly on sternite
extending into posterior of antrum. Apophysis anterior length in typical form 1-2-1-3 mm (3); small form
0-6-0-7 mm (3); apophysis posterior twice length of apophysis anterior in each form. Antrum scarcely
curved; anterior sometimes slightly indented, particularly in small form. Antrum of typical form very
broad posteriorly, constricted strongly to narrow anterior half; shorter than apophysis anterior. Antrum of
small form moderately broad posteriorly, constricted comparatively slightly for short distance at anterior;
almost as long as apophysis anterior. Ductus bursae approximately two to four times length of oval corpus
bursae. Signum covered with large and small spines; signum of typical form large, elongate kidney-shape,
strongly curved inwards; signum of small form comparatively small, oval to broad kidney-shape, scarcely
curved inwards.
REMARKS. One specimen has veins Rs and MI on a very short common stalk in one hind wing.
The frenulum, examined in seven females, consists of three setae in three of these, two setae in
one of these, and in three it consists of three setae on one wing and two on the other.
The specimens examined from Lakonia in Greece are smaller, on average, than the typical
form, particularly in the genitalia. The small form is typical externally except that the fore wing
markings tend to be relatively slightly smaller. Apart from the size, the female genitalia show
some consistent differences in structure from the typical form. The eighth sternite has slightly
longer, narrower sclerotized strips than in the typical form, the narrow anterior part of the
antrum is considerably shorter and the signum differs in shape. The saccus is variable in length
but tends to be more pronounced in the small form. It is possible that the Lakonia specimens
may prove to be a distinct species; however, the differences are comparatively slight for the
genus. There is insufficient material to determine whether the Lakonia population represents a
distinct subspecies oirhodoptera. The few specimens I have seen from other Greek localities are
of the typical form. A difference in the genitalia of one sex within a species is also discussed for
cytisella (see p. 34).
M. rhodoptera bears a superficial resemblance to scissella, constricta and cabezella. It differs
from all these in the hind wing venation, and the very long filament and very short saccus of the
male genitalia, and in addition differs from constricta and cabezella in many genitalic characters
since these two species are in the interuptella-group.
BIOLOGY. Host-plant unknown. Moths of the typical form have been found in June and July,
those of the small form in May and June.
DISTRIBUTION. Rumania, Greece, Turkey, Lebanon.
MATERIAL EXAMINED
Typical form (including 4 cf , 4 $ genitalia preparations)
Holotype c? , Rumania (Turkey'): Dobrogea ('Dobrudscha'), far outside Tulcea ('Tultscha'), middle
vi.1865 (Mann) (genitalia slide no. 11225; NM).
Greece: 1 9 , Parnassfosl Mts, vii (MNHU); 1 cf , 3 $, Pelop[6nnisos], near Kalavfrita], Zakhlorou, vi
22 L. M. PITKIN
(Coll. Klimesch, Linz). Turkey: 1 <j>, Mara ('Marasch'), vii (MNHU). Lebanon: 2 cf, N., Bcharre
('Becharre'), 1400 m, vi, vii (NM).
Small form (including 3 cf , 4 9 genitalia preparations)
Greece: 1 $ , Lakonia, 7 km SW. of Monemvasia, v; 4 cf , 5 $ , Lakonia, 5 km S. of Monemvasia, v, vi;
1 cf , Lakonia, Mt Taygetos, 1000 m, vi (all ZM).
Mirificarma denotata sp. n.
(Figs 1,2, 9, 37, 64, 88)
Cf, 7-5-8-5 mm. 9> 7-0-8-0 mm. Head light or occasionally mid brown. Labial palpus first and third
segments dark brown and cream; second segment predominantly cream. Thorax and tegula brown; tegula
base usually darker. Fore wing (Fig. 9) mottled brown and light pink-grey, ochreous tinge near base; apical
quarter to third predominantly dark brown, edged with diffuse, basal, transverse pink and cream streak
pronounced at costa; moderately dark brown patches slightly basal to streak at costa and dorsal margin,
and at base. Large transverse dark brown spot across fold at one-third extending to dorsal margin,
narrowing sharply from middle towards costa; dark brown spot at end of cell merging with patch at costa,
comparatively distinct from patch at dorsal margin. Both spots with narrow yellow-ochre surround. Hind
wing with veins Rs and MI on common stalk as in Fig. 4.
GENITALIA cf (Figs 37, 64). Actual margin of tegumen less emarginate than sclerotized margin anteriorly.
Sacculus short, moderately broad, flattened, slightly wrinkled at apex. Filament slightly helical, slender,
extending posteriorly short distance beyond valva, and well beyond tegumen anteriorly. Hind edge of
vinculum with moderately shallow U- or V-shaped emargination of sclerotization. Saccus broad basally,
narrow apically, extending slightly beyond tegumen anteriorly. Aedeagus approximately same length as
tegumen plus uncus. Aedeagus portion apical to hook small; apical hook moderately large.
GENITALIA 9 (Fig- 88). Posterior margin of seventh abdominal segment with area of weak sclerotization
mainly on tergite. Invagination of membrane between eighth sternite and papillae anales indistinct, long
narrow tube without sac. Flimsy membrane between eighth tergite and papillae anales invaginated to
broad rounded pouch. Eighth sternite with lateral pair of narrow, longitudinal sclerotized strips. Tergite
and rest of sternite very weakly sclerotized, with faint longitudinal wrinkles. Apophysis anterior length
0-9-1-1 mm (2); apophysis posterior approximately three times length of apophysis anterior. Antrum
almost straight, narrowing considerably towards indented anterior; approximately same length as apo-
physis anterior. Antrum with minute spines mainly in posterior half. Ductus bursae approximately twice
length of oval corpus bursae. Signum absent or scarcely discernible.
REMARKS. One specimen has the veins Rs and MI just separate in one hind wing. The frenulum
was examined in one female and consists of three setae. During the preparation of the male
genitalia, a tiny narrow projection was seen near the apex of the filament of one specimen,
similar to that visible in maculatella and ocellinella, although not obviously tubular as in the
latter.
M. denotata is very similar to maculatella in the fore wing pattern, but differs very slightly in
the form of the two large fore wing spots. The hind wing venation of denotata usually differs from
that of maculatella. The male genitalia are very similar to those of maculatella although the shape
of the extreme apex of the aedeagus differs; the female genitalia differ from those of maculatella
in characters including the shape of the antrum and the absence of a signum.
BIOLOGY. Host-plant: Galegeae: Astragalus lusitanicus Lamarck (type-specimens bred by
Walsingham). The moths emerged in late April and May.
DISTRIBUTION. Morocco.
MATERIAL EXAMINED (including 3 cf , 2 $ genitalia preparations)
Holotype cf, Morocco: [SE. of Tanger], El Fendek ('Fondak'), larva 28. iv. 1902, on Astragalus
lusitanicus, moth emerged 27. v. 1902 (Walsingham) (genitalia slide no. 22027).
Paratypes. 4 cf , 2 9 , same data as holotype, moths emerged 22-30. v. 1902.
GELECHIID MOTHS 23
Mirificarma maculatella (Hiibner)
(Figs 1-3, 10, 38, 65, 89)
Tinea maculatella Hiibner, 1796: 60, pi. 24, fig. 162 [legends to figs 161 and 162 are transposed].
LECTOTYPE cf , EUROPE (NM), here designated [examined].
Gelechia maculatella (Hiibner) Stainton, 1865: 228, pi. 7, fig. 3.
[Gelechia vicinella Douglas; Bruand d'Uzelle, 1858: 481; Disque, 1908: 65. Misidentifications.]
Cf , 7-0-9-5 mm. $, 7-5-9-0 mm. Head mottled mid brown. Labial palpus mottled cream or light brown,
and dark brown. Thorax and tegula mottled mid brown. Fore wing (Fig. 10) mottled brown and light
pink-grey; ochreous tinge near base; apical quarter to third predominantly dark brown, edged with diffuse,
basal, transverse pink and cream streak pronounced at costa; moderately dark brown patches slightly basal
to streak at costa and dorsal margin, and at base. Large transverse dark brown spot across fold at one-third
extending to dorsal margin, narrowing gradually from middle towards costa; dark brown spot at end of cell
comparatively distinct from patch at costa, usually merging with patch at dorsal margin. Both spots with
narrow ochreous surround. Hind wing with veins Rs and MI separate (Fig. 3).
GENITALIA d" (Figs 38, 65). Actual margin of tegumen less emarginate than sclerotized margin anteriorly.
Sacculus short, broad, slightly flattened, strongly wrinkled at apex and inner edge. Filament slightly
helical, slender, extending posteriorly well beyond apex of valva, and anteriorly, beyond tegumen. Hind
edge of vinculum with broad, moderately shallow U- or V-shaped emargination of sclerotization. Saccus
broad basally, narrow apically, extending a short distance beyond tegumen anteriorly. Aedeagus slightly
longer than tegumen plus uncus. Aedeagus portion apical to hook large; apical hook large.
GENITALIA $ (Fig. 89). Posterior margin of seventh abdominal segment with small area of weak
sclerotization mainly on tergite. Invagination of membrane between eighth sternite and papillae anales
very long narrow tube ending in pronounced, large oval sac. Eighth sternite with lateral pair of broad,
moderately short sclerotized strips. Tergite and rest of sternite very weakly sclerotized; tergite with faint
longitudinal wrinkles. Apophysis anterior length 0-9-1-0 mm (6); apophysis posterior approximately twice
length of apophysis anterior. Membraneous area between antrum and each apophysis anterior produced
into slight lobe, covered in minute spines. Antrum curved and narrower towards anterior; anterior not
indented; approximately 1-5 times to less than twice length of apophysis anterior. Antrum with minute
spines mainly in posterior half and extending into median area of sternite. Ductus bursae shorter than or of
similar length to round corpus bursae. Signum moderately large, round or oval; with spines, large around
edge.
REMARKS. In approximately 7 per cent of the specimens examined the veins Rs and M l are on a
common stalk in one or both hind wings. The frenulum of five out of six females examined
consists of three setae; in the remaining female it consists of three setae on one wing and two on
the other. During the preparation of the male genitalia, a tiny narrow projection was seen near
the apex of the filament, similar to those in denotata and ocellinella, although not obviously
tubular as in the latter.
The fore wing pattern of maculatella clearly distinguishes it from all other species except
denotata to which it is very similar. For differences from denotata see p. 22.
M. maculatella was described from an unspecified number of specimens. In the NM there is
one specimen from Mazzola's collection, here designated as lectotype, already labelled 'lecto-
type' by Sattler.
BIOLOGY. Host-plants: Coronilleae: Coronilla varia L. (moths bred by Miihlig, W. Germany;
Stainton, 1865: 228). C. emerus L. (Bruand d'Uzelle, 1858: 482, as Gelechia vicinella Douglas;
also subsequent authors).
The larva occurs in May and June feeding in two opposite leaflets spun together. It pupates in
a slight cocoon on the ground (Stainton, 1865: 228). Stainton records only one generation per
year. Prose (1957: 111) records the larva in July. Moths have been collected from June to
August.
DISTRIBUTION. France; Germany; Austria; Czechoslovakia; Yugoslavia; Turkey; Syria.
Additional records. Switzerland (Miiller-Rutz, 1914: 489); Italy (Hartig, 1964: 27); Poland
(Schille, 1931: 179); Hungary (Gozmany, 1958: 227); Albania (Rebel, 1931: 146); U.S.S.R.:
Ukraine (Piskunov, 1981: 674).
24 L. M. PITKIN
MATERIAL EXAMINED (including 8 cf , 6 $ genitalia preparations)
Lectotype cf , Europe (Mazzola) (genitalia slide no. 11221; NM).
France: 1 cf , 1 $, Paris; 1 cf , Basses-Alpes, viii. Germany: 1 cf , ; 1 cf , 1 $, 'N. Germany'. Germany
(West): 1 cf, 4 $, Hessen, vii; 1 cf, 2 $, Bayern. Germany (East): 8 cf, 11 $, Gera. Austria: 1 cf,
Nieder-Osterreich, vii. Czechoslovakia: 1 cf, Bohemia (NM). Yugoslavia: 1 cf, Croatia (NM); 1 $,
Dalmatia (NM); vi. Turkey: 1 cf , Konya, vii (NM); 1 cf , Amasya (MNHU). Syria: 1 $, NW., vi (NM). No
locality data: 15 ex.
Mirificarma pallidipulchra (Walsingham) comb. n.
(Figs 1,2, 11,39,66,90, 107)
Rhinosia pallidipulchra Walsingham, 1904: 269. LECTOTYPE cf, ALGERIA (BMNH), here designated
[examined].
Rhinosia striolella Turati, 1924: 166, pi. 6, fig. 11. LECTOTYPE cf , LIBYA (BMNH), here designated
[examined]. Syn. n.
Cf , 6-5-8-5 mm. $, 6-5-8-0 mm. Head cream to moderately light brown. Labial palpus cream frequently
tinged with brown ventrally. Thorax and tegula as head. Fore wing (Fig. 11) with alternating transverse
indistinct zig-zag patches of cream and ochre or occasionally deep yellowish brown; darker markings
usually narrow. Several very narrow yellowish or brown striations radiating from wing base towards apex.
Fore wing markings mostly weakly or moderately, occasionally strongly, contrasting. Hind wing with veins
Rs and Af t , separate as in Fig. 3.
GENITALIA cf (Figs 39, 66). Actual margin of tegumen usually coincides with sclerotized margin anteriorly.
Sacculus moderately slender in apical half; apex wrinkled. Filament slightly helical, slender, extending
posteriorly well beyond valva, and very far beyond tegumen anteriorly. Hind edge of vinculum with
V-shaped emargination of sclerotization. Saccus moderately broad, apex truncate or very slightly
emarginate; short, not quite reaching anterior of tegumen. Aedeagus approximately same length as
tegumen plus uncus; basal half very slightly swollen, with moderately large apical hook.
GENITALIA $ (Figs 90, 107). Abdomen with patchy, broad, median, longitudinal band constricted at
anterior margin of sixth sternite. Seventh abdominal sternite with pair of sclerites and membraneous sac at
anterior margin. Invagination of membrane between eighth sternite and papillae anales long, narrow,
funnel-shaped. Eighth sternite with lateral pair of weakly sclerotized, broad strips. Tergite and rest of
sternite very weakly sclerotized. Apophysis anterior length 0-9-1-0 mm (6); apophysis posterior approx-
imately three times length of apophysis anterior. Antrum curved or constricted towards anterior, slightly
shorter than or of similar length to apophysis anterior. Antrum with a few minute spines around posterior;
anterior sometimes very slightly indented. Ductus bursae shorter than oval or pear-shaped corpus bursae.
Signum large, spinose, particularly around edge, strongly curved inwards.
REMARKS. In approximately 7 per cent of the specimens examined the veins Rs and MI are on a
short common stalk in one hind wing. The frenulum was examined in five females and consists of
three setae. In the male genitalia the sclerotized anterior margin of the vinculum does not
coincide with the actual margin of the vinculum, although both are often indistinct. This occurs
in a few other Mirificarma species but in these one of the margins is much more distinct than the
other, whereas in pallidipulchra both margins appear equally faint. The constriction of the
female abdominal band of this species and aflavella is seen to a much less degree in some other
species (montivaga, rhodoptera, flavella, ocellinella and fasciatd) .
M. pallidipulchra closely resembles eburnella, aflavella and flavella in the fore wing pattern,
although its darker markings tend to be narrower, but it is distinguished from these species by
the very long filament of the male genitalia and the pair of sclerites with the membraneous sac of
the female abdomen. This last character is unique within the genus.
M. pallidipulchra was described from 23 specimens from Algeria: Hamman-es-Salahine,
18. iv, and El Kantara. I have examined 9 cf , 9 $ from El Kantara and designate as lectotype the
specimen bearing Walsingham's number 96479. This is referred to as the 'Type cf' on the
specimen label and in the original description. 1 cf , 1 $ of the remaining syntypes which I have
not examined are in the NM.
R. striolella was described from 4 cf from Libya, all of which I have examined. The specimen I
GELECHIID MOTHS 25
designate as lectotype, which bears Turati's 'Typus' label, was already labelled 'lectotype' by
Sattler.
BIOLOGY. Host-plant unknown. According to Walsingham (1904: 270) the moths are always
found among stems and root-crowns of Teucrium (polium L. ?) (Labiatae), but it seems very
unlikely that this is the host-plant. Moths have been found in March to May and July.
DISTRIBUTION. Algeria, Tunisia, Libya.
A record of this species from Egypt (Rebel, 1914: 268) is a misidentification of eburnella.
MATERIAL EXAMINED (including 9 d", 6 $ genitalia preparations)
Lectotype cf (pallidipulchra), Algeria: [Constantine province,] El Kantara, 5. v. 1903 (Walsingham)
(genitalia slide no. 22491). Lectotype cf (striolella) , Libya: Cyrenaica, Benghazi, 20. Hi. 1922 (Kriiger)
(genitalia slide no. 22490).
Algeria: 1 $, Oran, Aflou, vii; 1 cf, 1 $, Hassi Bahbah, v (NM); 8 cf, 9 $ (pallidipulchra
paralectotypes), El Kantara, 24.iv, 4, 5, 11. v. 1903 (Walsingham); 1 $, Constantine, Khenchela, v ?
Tunisia: 1 $, Tozeur, iv (MNHN); 1 $, Hammamet, v (NM). Libya: 1 cf, 4 $, Tripolitania, Gharyan
('Garian'),700m,iii,iv;5 $, Tripolitania, Tarhunah, 200 m,iv; 1 cf , 1 $?, Tripolitania, Bam WalId('Beni
Ulid'), 300 m, iv; 5 cf , 1 $, Tripolitania, Al Khums ('Horns'), 10 m, iv; 2 cf (striolella paralectotypes),
Cyrenaica, Benghazi, 18, 20.iii.1922 (Kriiger); 1 cf (striolella paralectotype), Cyrenaica, Benghazi,
'Merg', 9.iv (Kriiger).
Mirificarma afiavella (Amsel) stat. n.
(Figs 1,2, 12,40,67,91,108)
Rhinosiaflavella aflavella Amsel, 1935: 275. LECTOTYPE cf , ISRAEL (LN), here designated [examined].
Cf, 7-5-9-5 mm. $, 7-5-9-5 mm. Head moderately light golden-brown. Labial palpus mostly cream
dorsally, golden-brown ventrally. Thorax and tegula as head. Fore wing (Fig. 12) with alternating
transverse zig-zag patches of light golden-brown and mid brown, weakly contrasting. Hind wing with veins
Rs and MI usually on common stalk as in Fig. 4.
GENITALIA cf (Figs 40, 67). Actual margin of tegumen slightly less emarginate than sclerotized margin
anteriorly. Sacculus moderately broad, apex wrinkled. Filament gently curved; moderately stout in basal
half, tapering apically; extending posteriorly short distance beyond valva, and moderately far beyond
tegumen anteriorly. Hind edge of vinculum with V-shaped emargination of sclerotization. Saccus
moderately broad, narrowing slightly towards rounded apex; short, approximately reaching anterior of
tegumen. Aedeagus almost as long as tegumen plus uncus; with moderately large apical hook.
GENITALIA $ (Figs 91, 108). Abdomen with diffuse, broad, median, longitudinal band, slightly more
distinct at posterior margin of seventh segment, strongly constricted at anterior margin of sixth sternite.
Seventh abdominal segment comparatively broad, anterior margin unmodified. Invagination of membrane
between eighth sternite and papillae anales long, narrow, funnel-shaped. Eighth sternite with lateral pair
of broad sclerotized strips. Tergite and rest of sternite very weakly sclerotized. Apophysis anterior length
0-7-0-9 mm (5); apophysis posterior three times length of apophysis anterior. Antrum curved or
constricted anteriorly, of similar length to apophysis anterior; with minute spines posteriorly, extending
into median area of sternite. Antrum anterior sometimes very slightly indented. Ductus bursae one-
quarter to slightly more than one-third length of elongate pear-shaped corpus bursae. Signum with
approximately three or four large spines arising from sclerotized ridge on one side, otherwise smooth.
REMARKS. In approximately 25 per cent of the specimens examined the veins Rs and MI are
separate in one or both hind wings. The frenulum consists of three setae (five females
examined). In the female genitalia the ductus bursae merges gradually with the corpus bursae
and is distinguished only by its paler appearance in stained preparations.
This species resembles pallidipulchra, flavella and eburnella in fore wing pattern. It is
particularly close to flavella in this respect although the fore wing pattern is slightly less
contrasted. M. aflavella differs from these three species in the male genitalia, in which the
filament extends a short distance beyond the apex of the valva, and in the form of the signum of
the female genitalia.
M. aflavella was originally described as a subspecies of flavella, with which it is allopatric.
However, I consider that the genitalic differences between them are sufficient to separate them
26 L. M. PITKIN
as distinct species; moreover, aflavella appears to have even more features in common with
pallidipulchra than with flavella.
M. aflavella was described from a series collected by Amsel in Palestine, 1930: Tabgha, 12.iii;
Jordan, Salt, 7.iv, and Waldheim (c. 15 km E. of Haifa), 9.v (Amsel, 1935: 265). I have
examined 8 cf , 4 $ , from Tabgha, all of which probably belong to the type-series although some
lack Amsel's type-labels and are dated IS.iii instead of 12.iii. The specimen I designate as
lectotype, which bears Amsel's label 'Typus', was already labelled 'lectotype' by Sattler.
BIOLOGY. Host-plant unknown. Moths have been found from March to May.
DISTRIBUTION. Greece (Rodhos), Turkey, Israel (including Jordan west bank territories).
MATERIAL EXAMINED (including 4 cf , 5 $ genitalia preparations)
Lectotype cf , Israel: ('Palastina'), 10 km N. of Tiberias, En Sheva ('Tabgha'), 12.iii.1930 (Amsel)
(genitalia slide no. 17, LMP.; LN).
Greece: 1 $, Rodhos ('Rhodes'), Miramare [Hotel], iv. Turkey: 1 $, Gavur Daglari ('Amanus'),
'Schechle', v (NM); 1 cf , Gavur Daglari, Tarib', v (NM); 3 cf , 1 $ , Haleb, Shar Deresy ('Shar Devesy').
Israel: 1 cf, 3 $, ('Palestine'), Haifa, iii-v; 3 cf, 3 $ (paralectotypes) , ('Palastina'), Tiberias, En Sheva
('Tabgha'), iii.1930 (Amsel) (BMNH and LN); 4 cf , 1 $ (paralectotypes ?), Tiberias (including 3 cf , 1 $ ,
'See Genezareth'), En Sheva ('Tabgha'), iii, 15.iii.1930 (Amsel) (NM); 2 $, Palestine, plains of Jordan.
Mirificarma ffavella (Duponchel)
(Figs 1,2, 13,41,42,68,92)
Acompsia flavella Duponchel, [1844]: 512, pi. 89, fig. 7. Type(s), FRANCE: Bondy Forest, end of vi
(Begrand) [not traced].
Gelechia segetella Zeller, 1847: 847. LECTOTYPE cf, SICILY (BMNH), here designated [examined].
[Synonymized by Wocke, 1861: 115.]
Cf , 7-0-9-0 mm. $ , 7-5-9-0 mm. Head yellowish. Labial palpus cream to yellowish, sometimes tinged with
brown ventrally. Thorax yellowish, occasionally with faint brown median longitudinal stripe; tegula
yellow-ochre to golden-brown. Fore wing (Fig. 13) with alternating transverse zig-zag patches of yellowish
colour and deep yellow-tinged brown; moderately strongly contrasting. Hind wing with veins Rs and MI
separate as in Fig. 3.
GENITALIA cf (Figs 41, 42, 68). Actual margin of tegumen variable but usually slightly less emarginate than
sclerotized margin anteriorly. Sacculus with rounded, moderately broad apex. Filament moderately stout
at base, tapering and slightly curved towards apex; almost reaching apex of valva posteriorly, extending
slightly or scarcely beyond tegumen anteriorly. Hind edge of vinculum with median U- or V-shaped
emargination of sclerotization. Saccus broad to very broad, sometimes narrower at rounded apex, reaching
or extending slightly beyond tegumen anteriorly. Aedeagus same length as tegumen plus uncus or slightly
longer; with moderately large apical hook.
GENITALIA $ (Fig. 92). Abdomen with at most faint median, longitudinal band. Eighth sternite and tergite
very weakly sclerotized; sternite with faint lateral areas of sclerotization. Apophysis anterior length
0-8-1-0 mm (8); apophysis posterior approximately three times length of apophysis anterior. Antrum
almost straight, tapering, usually slightly, towards strongly indented anterior; slightly shorter than
apophysis anterior; without minute spines. Ductus bursae slightly shorter to longer than oval or round
corpus bursae. Signum small to medium-sized; spinose, particularly around edge.
REMARKS. The fore wing pattern varies from predominantly yellowish to predominantly brown.
The frenulum consists of three setae (five females examined). The sacculus of the male genitalia
tends to be longer than that in eburnella or aflavella. The saccus is usually relatively broader than
in eburnella although it is variable in both species.
M. flavella resembles eburnella, aflavella and pallidipulchra in the fore wing pattern,
particularly aflavella, although the fore wing pattern of flavella is usually more contrasted than in
aflavella. The male genitalia of flavella differ from those of eburnella by the rounded apex of the
sacculus, and from the other two species by the relatively short filament. The female genitalia
differ in having a straight antrum together with an indented apex.
M. flavella was described from an unspecified number of specimens from France: Bondy
GELECHIID MOTHS 27
Forest (near Paris). The type-specimens have not been traced and are not in the MNHN (Viette,
pers. comm.).
G. segetella was described from an unspecified number of specimens from Sicily: Siracusa. I
have examined 1 Cf, 1 9 syntypes and designate as lectotype the male already labelled
'lectotype' by Sattler.
BIOLOGY. Host-plants: Trifolieae: Trifolium pratense L. (moth bred by Frey, France). T. minus
Rehl. = procumbens G.G.' (the identity of this Trifolium species is not clear since minus is
currently a synonym of T. dubium Sibthorp and procumbens is a synonym of T. campestre
Schreber). Loteae: Lotus corniculatus L.
These host-plants are referred to by Lhomme ([1948-1949] : 653) , who states that the larva has
been found in May and June. Moths have been collected from April to July.
G. segetella was described from moths collected on the edges of wheatfields and on
chrysanthemum-like flowers (Compositae); however, there is no reason to suppose that the
latter could be its host-plant.
DISTRIBUTION. France, Corsica, Italy, Sardinia, Sicily, Greece, Crete, Cyprus, Algeria, Tunisia.
Additional records. Spain (Agenjo, 1968: [4]); Belgium (Lhomme, [1948-1949]: 652);
Greece: Attiki (Staudinger, 1871: 255).
The following records should probably be referred to aflavella. Israel: Jerusalem (Caradja,
1920: 112); 'Palestine'; Turkey ('Kleinasien') (Wocke, 1871: 300); Turkey ('Asia Minor'); Syria
(Rebel, 1901: 158; Meess (in Spuler, 1910: 344); Meyrick, 1925: 142); Turkey ('Asia Minor')
(Rebel, 1903:332).
A record of the species from Bulgaria: Rilo (Rebel, 1903: 332) is based on a misidentification
of Orophia ferrugella ([Denis & Schiffermiiller]) (Oecophoridae). A record oiflavella from
Yugoslavia: Dalmatia (Rebel, 1903: 332) may also be a misidentification of O. ferrugella.
MATERIAL EXAMINED (including 5 cf , 8 9 genitalia preparations)
Lectotype cf (segetella), Sicily: Siracusa, v. (Zeller) (genitalia slide no. 7135).
France: 1 $, ; 1 cf , Hie et Vilaine; 1 cf , Mayenne (MNHN); 1 cf, Calvados (MNHN); 1 $, Sarthe
(MNHN); 1 cf , Essonne (MNHN); 2 cf , 5 $, Paris; 4 cf , 1 $, Alpes-Maritimes; vi, vii. Corsica: 7 cf , vi.
Italy: 3 cf , 3 9 Toscana, vi, vii. Sardinia: 1 $, vi ? (NM). Sicily: 1 $ (segetella paralectotype), Siracusa, v.
Crete: 2 $ , v, vi (NM). Cyprus: 7 cf , 8 $ , iv, v. Algeria: 2 cf , Algiers province (MNHN); 2 $ , Constantine
province, vi. Tunisia: 1 $ . No locality data: 23 ex.
Mirificarma eburnella ([Denis & Schiffermiiller]) comb. n.
(Figs 1,2, 14,43,69,93, 109)
Tinea eburnella [Denis & Schiffermiiller], 1775: 140. Syntypes, AUSTRIA: Wien area (lost). NEOTYPE cf ,
AUSTRIA (NM), here designated [examined].
Tinea formosella Hiibner, 1796: 62, pi. 23, fig. 160. Syntypes, SWITZERLAND: Genf ('Geneve') [not traced].
[Junior primary homonym of Tinea formosella [Denis & Schiffermiiller], 1775: 140 (Oecophoridae).]
Carcina flammella Hiibner, [1825]: 410. [Objective replacement name for Tinea formosella Hiibner.]
[Synonymized by Zeller, 1847: 848.]
Gelechia rufeoformosella Bruand d'Uzelle, 1859: 652. [Objective replacement name for Tinea formosella
Hiibner.]
[Rhinosia palidipulchra Walsingham; Rebel, 1914: 268. Incorrect subsequent spelling of pallidipulchra
Walsingham. Misidentification.]
Mirificarma formosella (Hiibner) Capue, 1964: 17, figs 7, 8.
Cf, 5-0-7-5 mm. 9> 5-5-7-5 mm. Head cream. Labial palpus cream, frequently tinged with brown
ventrally. Thorax cream with faint, narrow, median, longitudinal, yellow-ochre stripe; tegula yellow-
ochre. Fore wing (Fig. 14) with alternating transverse zig-zag patches of cream and yellowish to ochre,
strongly contrasting. A few very narrow ochre striations radiating from wing base towards apex. Hind wing
with veins Rs and MI separate as in Fig. 3.
GENITALIA cf (Figs 43, 69). Actual margin of tegumen considerably less emarginate than sclerotized
margin anteriorly. Sacculus with pointed, moderately slender apex. Filament moderately slender, almost
straight, almost reaching apex of valva posteriorly, extending short to moderate distance beyond tegumen
28 L. M. PITKIN
anteriorly. Hind edge of vinculum with U- or V-shaped emargination of sclerotization. Saccus broad,
slightly narrower at rounded apex, extending short distance beyond tegumen anteriorly. Aedeagus same
length as tegumen plus uncus or slightly longer; apical hook usually moderately large.
GENITALIA $ (Figs 93, 109). Abdomen with distinct broad, median, longitudinal band, most prominent in
seventh segment. Eighth sternite and tergite very weakly sclerotized; sternite with faint lateral areas of
sclerotization. Apophysis anterior length 0-6-0-9 mm (10); apophysis posterior three to four times length
of apophysis anterior. Antrum almost straight, tapering, usually slightly, towards rounded or scarcely
indented anterior; of similar length to apophysis anterior or slightly longer; without minute spines. Ductus
bursae usually merging indistinguishably with elongate corpus bursae. Signum extremely small, sometimes
apparently absent.
REMARKS. In less than 2 per cent of the specimens examined the veins Rs and M l are on a short
common stalk in one or both hind wings. The frenulum consists of three setae (five females
examined). In the male genitalia, the uncus and tegumen posterior tend to be narrower than in
flavella. The corpus bursae of the female genitalia is extremely flimsy and sometimes very small.
M. eburnella closely resembles flavella, aflavella and pallidipulchra in the wing pattern,
although it is usually slightly more contrasted in eburnella. It differs from these in the pointed
apex of the sacculus together with the relatively short filament of the male genitalia, and the
straight antrum without an indented apex in the female genitalia.
M. eburnella was described from an unspecified number of specimens. Zeller (1847: 848)
refers to two specimens of this species in the Schiffermuller collection, which is now lost. I
designate a male neotype from the type-locality, Austria: Wien area.
BIOLOGY. Host-plants: Trifolieae: Medicago saliva L., M. lupulina L. (Lhomme, [1948-1949]:
652); M. polymorpha L. (Bur clover), Trifolium repens L. (variety - Ladino clover), T. hirtum
Hall (Rose clover); Vicieae: Vicia americana Muhlenberg (Purple vetch) (California Depart-
ment of Agriculture unpublished report); Coronilleae: Hippocrepis comosa L. (Lhomme,
[1948-1949]: 652).
The records of the California Department of Agriculture refer to the species in U.S.A.; the
other records are European. I have seen a Walsingham specimen bred from 'Trifolium'';
however, this name has been used broadly and may refer to Medicago. A record of the species on
Populus (Salicaceae) (Hofmann, 1875: 118) is erroneous since it is based on a misidentification
of the moth which was probably Schiffermuelleriaformosella [Denis & Schiffermuller] (Chretien
in Lhomme, [1948-1949]: 652).
The larva has been found in April and May, both in Europe (Lhomme, [1948-1949]: 652) and
in U.S.A. (California Department of Agriculture unpublished report). This species has caused
severe damage to clover in California, U.S.A., where the larva 'semiskeletonizes' leaves and
lightly spins two leaves together, pupating inside the folded leaves (Anonymous, 1969: 69).
Moths have been collected from March to July (also in August according to Piskunov, 1981:
674); in May and June in U.S.A. Hartig (1964: 51) states that there are two generations in Italy,
with the moths in June and August.
DISTRIBUTION. M. eburnella occurs in western, central and southern Europe south of c. 50 N.
and extends to North Africa, the Middle East and U.S.S.R.: Armeniya. It is also found in
U.S.A.: California where it is presumed to have been introduced.
According to the literature it has also been found in the following countries. Portugal
(Zerkowitz, 1946: 132); Belgium (Lhomme, [1948-1949]: 652); Netherlands (Lempke, 1976:
27); Sardinia (Hartig & Amsel, 1951: 86); Poland (Schille, 1931: 199); Czechoslovakia (Hruby,
1964: 293); Bulgaria (Rebel, 1903: 332); Crete (specimen in NM, Sattler, pers. comm.);
U.S.S.R.: European part (Piskunov, 1981: 674).
MATERIAL EXAMINED (including 7 cf , 10 $ genitalia preparations)
Neotype cf (eburnella), Austria: Nieder-Osterreich ('Austr. inf.'), [near Wien,] Klosterneuburg,
2.vi.l918 ('Freiberg') genitalia slide no. 11230; NM).
Spain: 6 cf , Granada; 1 cf , Mallorca; iv-vi. France: 1 cf , 1 $ , Sarthe (MNHN); 1 cf , SW. ; vii. Germany:
3 cf . Switzerland: 2 cf , 3 $ , Valais. Corsica: 19 cf , 7 $, v-vii. Italy: 2 cf , 1 9, Toscana; 1 cf , Lazio; 1 cf ,
Campania; iv-vi. Sicily: 5 cf , 4 $, 5 ex., iii-v. Austria: 1 cf , 2 $?, ; 1 Cf , Wien area (NM); vi. Yugoslavia:
GELECHIID MOTHS 29
1 Cf, 1 $, Slovenija;3 cf, 1 $, Croatia; 1 cf, 1 $, Dalmatia; 2cf, Bosna i Hercegovina; vi, vii. Hungary:
4 cf , Rumania: 6 cf , Timi, vi (BMNH; NM); 1 cf , Caras-Severin, vi? (NM). Albania: 2 cf , vii. Greece:
1 $, Levkas, vi (NM); 1 cf, Lakonia, iv (ZM); 1 cf, 1 $, Rodhos I., iv. Cyprus: 4 cT, 3 $, 1 ex., iv, v.
Turkey: 1 $ , Bursa (NM); 1 cf , Toros Daglari. Malta: 1 cf , 1 $ , iv. Morocco: 2 cf , 1 $ , iv, v. Algeria: 7 cf ,
3 $, Oran province; 2 $, Algiers province (BMNH; MNHN); 13 cf, 6 $, Constantine province; v, vi.
Tunisia: 3 cf, 3 $, iv, v. Egypt: 2 cf , ; 1 $, Cairo area, iv (NM) (Rebel, 1914: 268, as Rhinosia
palidipulchra Walsingham). Israel [including Jordan west bank territories]: 4 cf , 5 9, 'Palestine'; iv, v.
Lebanon: 4 cf , v. Syria: 2 $, N.; 2 cf , 2 $, NW. U.S.S.R.: 1 $, Armeniya, vii (NM). U.S.A.: 6 cf , 4 $,
California, v, vi. No locality data: 37 ex.
The iJiterupte//a-group
Cf, $ Fore wing without zig-zag pattern of yellowish markings; sometimes with median longitudinal
stripe. Hind wing with veins Rs and MI on common stalk as in Fig. 4.
GENITALIA cf . Uncus usually small or constricted at base; occasionally large. Gnathos a large or small
simple hook or modified in shape. Tegumen without pair of lateral processes. Filament-supporting sclerites
not crossed or spiralled round each other. Hind edge of vinculum a narrow projection or with distinctly
demarcated, usually narrow, median projection. Vinculum with pair of sclerites or single sclerite extending
from saccus and approaching hind edge of vinculum.
GENITALIA $. One invagination of membrane between eighth segment and papillae anales present,
comparatively sturdy, usually between tergite and papillae anales (between sternite and papillae anales in
interuptelld) . Eighth segment usually moderately to strongly sclerotized. Apophysis anterior rod-like or
very short, broad lobe; usually short. Signum, if present, with many small spines or without spines.
REMARKS. In some species occasional specimens have the veins Rs and M\ separate in one or
both hind wings. In mulinella the lateral edge of the male tegumen is sometimes wavy, producing
slight projections but not the distinct rounded processes of the maculatella-group. The scler-
otization of the female eighth segment is limited to small areas in fasciata and, particularly,
ocellinella; the degree of sclerotization is greater in lentiginosella and cytisella but still slightly
less than in the rest of the interuptella-group . The species in this group are more diverse in
genitalia structure than those of the maculatella-group.
BIOLOGY. Host-plants: Genisteae (cytisella also on Trifoliae: Ononis).
Mirificarma ocellinella (Chretien) comb. n.
(Figs 1, 2, 15, 16, 46, 71, 83, 94, 95)
Gelechia ocellinella Chretien, 1915: 317. LECTOTYPE $, TUNISIA (MNHN), here designated [ex-
amined].
Gelechia aurantiella Chretien, 1915: 317. LECTOTYPE , TUNISIA (MNHN), here designated [ex-
amined]. Syn. n.
Gelechia retamaeofoliella Dumont, 1931: 148. LECTOTYPE $, TUNISIA (MNHN), here designated
[examined]. Syn. n.
Cf , 9-0-11-0 mm. 9, 8-5-10-5 mm. Head cream to light brown. Labial palpus cream mottled with brown,
sometimes dark brown, particularly on outer surface; second segment with slight to moderate brush below.
Thorax and tegula cream to mid brown; tegula sometimes darker at base. Fore wing (Figs 15, 16) cream to
light brown, sometimes mottled with darker brown; usually with narrow longitudinal dark brown stripes
radiating to apical wing margin. Median stripe sometimes overlaid with broader median longitudinal band,
occasionally very dark. Tiny spot sometimes present at one-third to half, on costal edge of cell, and
occasionally at or near end of cell.
GENITALIA cf (Figs 46, 71). Uncus large, two-thirds to nearly three-quarters width of tegumen. Gnathos
basal half increasingly very broad then narrowing sharply to hook-shaped apical half. Actual margin of
tegumen slightly less emarginate than sclerotized margin anteriorly. Sacculus very slightly S-curved,
usually moderately slender, apex slightly wrinkled and with small irregular projections. Filament very
moderately stout, almost straight; extending beyond apex of valva posteriorly, and very far beyond
tegumen anteriorly. Tiny tube projecting from opening near filament apex. Hind edge of vinculum with
sharply rectangular weakly sclerotized, scarcely emarginate, median projection. Vinculum short, with pair
30 L. M. PITKIN
of almost parallel sclerites. Saccus parallel-sided, moderately slender, apex not or scarcely wider than base;
extremely long but not reaching anterior of filament. Aedeagus slightly less than twice length of tegumen
plus uncus; apex with very narrow, distinctly sclerotized, slightly projecting structure.
GENITALIA 9 (Figs 94, 95). Abdomen with at most faint median longitudinal band, scarcely more distinct at
posterior margin of seventh segment. Invagination of membrane between eighth tergite and papillae
anales funnel-shaped. Eighth sternite with lateral pair of relatively narrow longitudinal sclerotized strips;
tergite and rest of sternite weakly sclerotized and faintly wrinkled longitudinally. Unsclerotized median
area of sternite at least as wide as each sclerotized area. Apophyses anteriores parallel, rod-like, length
1-0-1-3 mm (7); apophysis posterior three to four times length of apophysis anterior. Pair of membraneous
sacs from sternite between apophyses anteriores and antrum, densely covered in minute spines. Antrum
gently coiled, anterior not indented; extremely long, coiled length approximately twice length of apophysis
anterior. Ductus bursae less than half length of oval or round corpus bursae. Signum strongly curved
inwards, elongate-oval, distinctly covered with tiny spinules; with spiny-edged plate, usually medially
indented, projecting obliquely inside bursa.
REMARKS. The fore wing pattern of ocellinella varies by degrees from strongly contrasted
markings to the comparatively uniform fore wing of the type-specimens of retamaeofoliella,
which lacks stripes. The frenulum of six out of seven females examined consists of three setae; in
the remaining female it consists of two setae. In the male genitalia the tiny tube near the apex of
the filament was not visible in two specimens; however, it is easily lost during the preparation of
the genitalia. The specimens examined show a wide but gradual range both in wing length and in
the female sternite sclerotization. The lectotype of G. retamaeofoliella is the smallest specimen
of ocellinella and has the shortest strips of sternite sclerotization. The signum of this specimen is
more elongated than others examined and the projection of the signum has no median
indentation.
M. ocellinella is the largest species of the genus. It closely resembles fasciata in the male and
female genitalia, but differs in the shape of the male gnathos and saccus, and to a less extent in
the female sternite sclerotization and signum.
M. ocellinella was described from an unspecified number of specimens from Tunisia. I
designate the single type-specimen I have examined as the lectotype. The collector is likely to
have been Chretien but may have been Oberthiir or Lucas (Chretien, 1915: 289).
G. aurantiella was also described from an unspecified number of specimens from Tunisia. I
have examined a single specimen labelled TYPE' [by Viette], 'Gelechia aurantiella'. It bears the
locality label 'Mansour', not 'Gafsa' as cited in the original description; however, there is a
locality of this name near Gafsa. Despite this discrepancy, I consider it to be a type-specimen
and designate it as the lectotype.
G. retamaeofoliella was described from a pair of specimens from Tunisia, both of which I have
examined. They are labelled 'Coll. D. Lucas, 1952' and in Lucas's handwriting: 'Gelechia
retamaeofoliella Dumont' on the female and 'Gelechia acupediella Frey' on the male. The
specimens are conspecific and the label on the male is incorrect. According to Dr P. Viette (pers.
comm.) the type-specimens, which are in the MNHN, belong to the Dumont Collection, not the
Lucas Collection, and Lucas may have replaced their original labels. I designate the female as
the lectotype.
The type-specimens of retamaeofoliella represent the small, poorly-marked form of ocel-
linella.
BIOLOGY. Host-plant: Genisteae: Retama raetam (Forskal) Webb & Berthelot (R. 'retem Webb')
(type-series of G. retamaeofoliella). The larva remains in a long silk tube at the base of the
food-plant during the day and comes out to feed on the leafy branches at night (Dumont, 1931:
149). It is fully grown at the beginning of March and the adults Dumont reared emerged in
September and October. Moths have been found in January, March to May and September to
November. A few females have been collected at light.
DISTRIBUTION. Morocco, Algeria, Tunisia, Libya, Jordan.
MATERIAL EXAMINED (including 7 d", 7 $ genitalia preparations)
Lectotype $ (ocellinella), Tunisia: Tozeur, 31.x. 1904 (genitalia slide no. 28, LMP. ; MNHN). Lectotype
GELECHIID MOTHS 31
$ (aurantiella), Tunisia: [Gafsa], Mansour, 11. xi. 1908 (genitalia slide no. 29, LMP; MNHN). Lectotype $
(retamaeofoliella), Tunisia: Metlaoui, larva on Retama 'retem Webb', moth emerged 9.x 1921 (Dumonf)
(genitalia slide no. 39, LMP; MNHN).
Morocco: 1 cf , Agadir, 'Rocksin', xi (NM). Algeria: 1 cf , S. Oran, Aflou region, x; 3 cf , Lambese, x.
Tunisia: 1 cf (retamaeofoliella paralectotype), Metlaoui, on Retama 'retem Webb' (MNHN); 1 $,
Kasserine to Thelepte road, iv (MNHN); 1 cT, 1 $, 1 ex., Bu Hadmah ('Bou Hedma'), iii, x, (MNHN);
6 Cf, 4 $, Maknassy, x, xi (MNHN). Libya: 1 $, W. ('occ.') Sirtica, Sawfajjin ('Sofeggin'), v; 1 cf , W.
('occ.') Sirtica, Al Qaddahiyah ('Gheddahia'), ix. Jordan: 1 $, Jordan Valley, Zarqa ('Zerqa') R. Colony,
c. 100 m below sea level, i.
Mirificarma fasdata sp. n.
(Figs 1,2, 17,47,70,96)
Cf, $,8-5 mm. Head white mottled with grey. Labial palpus white mottled with grey; mainly grey on outer
surface. Thorax mottled grey and white; tegula grey. Fore wing (Fig. 17) white and ochreous cream,
mottled with dark brown scales mainly near wing margins; very faintly and diffusely forming longitudinal
narrow stripes radiating from base towards apex. Brown median longitudinal band, very dark brown on
costal side. Very narrow, broken, dark brown stripe along fold to dorsal margin.
GENITALIA cf (Figs 47, 70). Uncus large, approximately two-thirds width of tegumen. Gnathos long and
only slightly curved; basal half unmodified, extreme apex a very slight hook-shape. Actual margin of
tegumen slightly less emarginate than sclerotized margin anteriorly. Sacculus straight, slender; extreme
apex with small irregular projection. Filament straight, very moderately stout; reaching apex of valva
posteriorly, extending far beyond tegumen anteriorly. Hind edge of vinculum with sharply rectangular,
weakly sclerotized, scarcely emarginate median projection. Vinculum short, with pair of parallel sclerites.
Saccus slender, parallel-sided except apex which is twice width of base; extremely long, extending beyond
filament anteriorly. Aedeagus slightly less than twice length of tegumen plus uncus; apex with very narrow,
distinctly sclerotized, slightly projecting structure.
GENITALIA $ (Fig. 96). Posterior margin of seventh abdominal segment with area of weak dorsal
sclerotization and pair of faint ventral patches merging with extremely faint median longitudinal band.
Invagination of membrane between eighth tergite and papillae anales conical. Eighth sternite with lateral
pair of broad longitudinal sclerotized strips; tergite and rest of sternite weakly sclerotized and very slightly
wrinkled longitudinally. Less sclerotized median area of sternite narrower than each sclerotized area.
Apophyses anteriores parallel, rod-like, length 0-7 mm (1); apophysis posterior twice length of apophysis
anterior. Small pair of membraneous sacs from sternite between apophyses anteriores and antrum, densely
covered in minute spines. Antrum coiled; anterior not indented; extremely long, actual length approx-
imately three to four times length of apophysis anterior. Ductus bursae less than half length of oval corpus
bursae. Signum moderately elongate-oval, with irregular surround and small inwards projection which is
medially indented; covered with tiny spinules.
REMARKS. The frenulum of the single female examined consists of two setae on one wing and
three on the other. The longitudinal fore wing stripe of this species gives it a superficial
resemblance to interuptella although the stripe is more diffuse than in the latter. However, it
differs considerably in the genitalia, particularly in the shape of the male filament, saccus and
sacculus, and in the female apophysis anterior and antrum. M. fasdata is also similar externally
to specimens of ocellinella with a fore wing stripe and the two species appear to be closely
related. It differs from ocellinella in the shape of the male gnathos and saccus and in the female,
to a lesser degree, in the sternite sclerotization and the signum shape.
BIOLOGY. Host-plant unknown. Moths have been found in December.
DISTRIBUTION. Spain.
MATERIAL EXAMINED (including 1 cf , 1 $ genitalia preparations)
Holotype cf , Spain: [Malaga,] San Pedro de Alcantara, xii.1972 (Ffennelt) (genitalia slide no. 22083).
Paratype. 1 $, same data as holotype.
32 L. M. PITKIN
Mirificarma lentiginosella (Zeller)
(Figs 1,2, 18, 44, 72, 97)
[Haemylis obscurella Hiibner; Treitschke, 1832: 240-241 (larva only). Misidentification.]
Gelechia lentiginosella Zeller, 1839: 198; Stainton, 1865: 64, pi. 2, fig. 3. LECTOTYPE $, GERMANY
(EAST) (BMNH), here designated [examined].
Cf , 6-5-8-5 mm. $ , 6-0-8-0 mm. Head mid to dark brown. Labial palpus dark brown with scattered cream
scales; paler on dorsal or inner surface and at segment apices. Thorax and tegula mid to dark brown. Fore
wing (Fig. 18) dark brown with scattered pinkish buff scales; small pinkish spot at costa, at two-thirds to
three-quarters, sometimes extended to basal margin. Very small darker brown spots with ochreous
surround; one in fold, one in cell, both approximately at one-third; one at end of cell. All spots indistinct.
GENITALIA cf (Figs 44, 72). Uncus large, slightly constricted at base, at this point half to two-thirds width of
tegumen, otherwise slightly narrower than tegumen. Gnathos a large simple hook. Actual margin of
tegumen slightly less emarginate than sclerotized margin anteriorly. Sacculus moderately curved inwards,
with evenly rounded apex; slender. Filament almost straight basally, without median lobe, with strong kink
at apex, very moderately stout; extending posteriorly to hind edge of vinculum or slightly beyond;
extending well beyond tegumen anteriorly. Hind edge of vinculum with low, weakly sclerotized median
projection not emarginate ventrally. Vinculum with pair of sclerites, parallel or converging and undulating
from saccus, almost reaching hind edge of vinculum. Saccus parallel-sided or broader at rounded apex,
very long, extending far beyond tegumen anteriorly. Aedeagus approximately 1-5 times length of tegumen
plus uncus, with moderately small apical projection.
GENITALIA 9 (Fig- 97). Abdomen with at most faint, patchy, median longitudinal band scarcely more
distinct at posterior margin of seventh segment. Invagination of membrane between eighth tergite and
papillae anales long and narrow. Eighth segment not usually strongly sclerotized. Sternite with lateral pair
of very broad, longitudinal sclerotized areas; median area of sternite less sclerotized, usually slightly, and
narrower than each lateral area of sternite. Tergite with medially expanded sclerotized area at anterior and
pair of tiny lateral patches at posterior. Apophyses anteriores diverging, rod-like, length 0-4-0-5 mm (8);
apophysis posterior four to five times length of apophysis anterior; without sacs between apophyses
anteriores and antrum. Antrum gently coiled or strongly curved, anterior not indented; extremely long,
coiled extent two to four times length of apophysis anterior. Ductus bursae usually not more than half
length of round corpus bursae. Signum small, oval, weakly sclerotized, slightly curved inwards, covered
with tiny spinules.
REMARKS. In six females examined, the frenulum consists of three setae, except for one female in
which it consists of four on one wing.
M. lentiginosella differs in the fore wing pattern of very small indistinct spots on a dark
background, although this is very occasionally approached by extremely dark specimens of the
externally variable mulinella. M. lentiginosella can be distinguished by the shape of the male
filament, which is straight basally, kinked apically and without a lobe, and by the very long
antrum together with the diverging apophyses anteriores in the female. This species appears to
be close to constricta with which it has some common features of the uncus, filament and
sacculus. However, the uncus and filament shape are similar in interuptella, although to a lesser
degree.
M. lentiginosella was described from a series of 25 specimens of which I designate as lectotype
the single specimen I have examined, which was already labelled 'lectotype' by Sattler. The
type-locality is neither stated in the original description nor on the lectotype label. The original
description attributes the species name to Tischer, and I have seen a record, in Tischer's
handwriting, of the species collected in Dresden (East Germany). It seems most likely that the
specimens from Dresden referred to in Tischer's record were sent to Zeller and that Zeller based
his description on these.
BIOLOGY. Host-plants: Genisteae: Genista tinctoria L. (larvae and moths bred by Bankes and
Ford, Great Britain; Stainton, 1865: 64); G. anglica L. (moths bred by Nielsen, Denmark;
Sorhagen, 1886: 186); G. germanica L., G. sagittalis L. (Sorhagen, 1886: 186); Laburnum
anagyroides Medicus (= Cytisus laburnum L.) (Miiller-Rutz, 1913-1914: 485).
Records of Centaurium erythraea erythraea Rafn (= Erythraea centaurium Persoon) (Gen-
GELECHIID MOTHS 33
tianaceae) (Lhomme, [1946-1948]: 607, 'according to certain authors') and Salix repens L. ?
(Salicaceae) (Sorhagen, 1886: 187) are extremely dubious as host-plants of this species.
Sorhagen attributes his record to Hartm[ann] but I have not been able to trace the reference.
The larva occurs in May and June, spinning together the young terminal shoots of the plant; it
pupates in a cocoon amongst leaves on the ground (Stainton, 1865: 64), from June to July
(Bradford, [1979]: 123). Moths have been collected from June to August (also in May according
to Mariani, 1943:167).
DISTRIBUTION. Great Britain, France, Denmark, Central Europe, Italy, Rumania.
Additional records. Spain (Agenjo, 1968: [4]); Netherlands (Lempke, 1976: 26); Sweden
(Krogerus et alii, 1971: 21); Poland (Schille, 1931: 176); Czechoslovakia (Nicked, 1908: 20);
Yugoslavia (Mariani, 1943: 167); Hungary (Gozmany, 1958: 227); Turkey (Klimesch, 1961:
648); U.S.S.R.: European part (Piskunov, 1981: 672); U.S.S.R.: Armeniya (Rebel, 1901: 14).
MATERIAL EXAMINED (including 8 cf , 8 $ genitalia preparations)
Lectotype $, [Germany (East): Dresden], larva on Genista tinctoria, ex viii (genitalia slide no. 22492).
Great Britain (England): 67 ex., ; 2 ex., Worcester: 15 cf, 13 $, 3 ex., Dorset; 1 ex., New Forest;
12 ex., Isle of Wight; 52 ex., Sussex; 8 ex., Kent; 7 ex., Essex; vii-viii. France: 1 cf, Basses Alpes;
1 <j>, Alpes-Maritimes, viii. Denmark: 3 cf, 1 <j>, Jylland, vi (BMNH; ZM). Germany (West): 4 cf,
1 $, Niedersachsen; 1 cf , Bayern, viii. Germany (East): 6 cf , 3 $, 1 ex., Gera. Switzerland: 1 cf , Zurich.
Italy: 1 $ , Toscana, viii. Austria: 2 cf , 1 9 , Wien, viii (NM). Rumania: 1 cf , 1 $ , Cluj, viii (MINGA). No
locality data: 34 ex.
Mirificarma constricts sp. n.
(Figs 1,2, 19,45,73)
Cf , 5-5-6-0 mm. Head mid brown. Labial palpus mottled cream and brown. Thorax and tegula mid brown.
Fore wing (Fig. 19) mottled mid brown, slightly darker at base. Dark brown spot across fold at one-third
extending to dorsal margin. Small dark brown spot at end of cell.
GENITALIA cf (Figs 45, 73). Uncus small, slightly more than half width of tegumen, strongly constricted at
base. Gnathos large, only moderately curved; extreme apex a very slight hook-shape. Actual margin of
tegumen slightly less emarginate than sclerotized margin anteriorly; sclerotized margin X-shaped medial-
ly. Sacculus moderately S-curved; with evenly pointed apex, moderately slender. Filament basal half
straight, very broad dorsoventrally and compressed laterally; median lobe projecting ventrally; apical half
more slender, kinked; extending beyond hind edge of vinculum posteriorly, and moderately beyond
tegumen anteriorly. Median projection of hind edge of vinculum with ventral V-shaped emargination;
dorsally deeply U-emarginate. Vinculum with pair of sclerites converging from saccus, almost reaching
hind edge of vinculum. Saccus moderately broad, parallel-sided or slightly bow-sided, with rounded apex;
extending slightly beyond tegumen anteriorly. Aedeagus same length as tegumen plus uncus or slightly
shorter; projection near apex slight.
GENITALIA $. Unknown.
REMARKS. M. constricta is very similar in fore wing pattern to cabezella and, to a less extent,
scissella and rhodoptera but differs in the constricted uncus and the shape of the filament. M.
constricta resembles lentiginosella in these male genitalic characters; however, the single known
female of constricta lacks the abdomen and it is impossible to be certain of its relationships.
BIOLOGY. Host-plant unknown. Moths have been found in August and October.
DISTRIBUTION. Northern Morocco.
MATERIAL EXAMINED (including 2 cf genitalia preparations)
Holotype cf , Morocco: Tanger, 45 m, 30.x. 1934 (Querci) (genitalia slide no. 7477).
Paratypes. Morocco: 1 cf , 1 $, Tanger, 45 m, 30. viii, 24.x. 1934 (Querci).
34 L. M. PITKIN
Mirificarma cytisella (Treitschke)
(Figs 1, 2, 5, 20, 21, 48, 74, 75, 98, 110)
Lita cytisella Treitschke, 1833: 99.
Cf , 6-0-8-0 mm. $ , 6-0-7-5 mm. Head white to cream, eye socket edged with dark brown anteriorly. Labial
palpus white to cream, with dark brown predominantly on outer surface of first segment and outer, basal
two-thirds of second segment. Thorax and tegula white to cream with scattered brown scales. Fore wing
(Figs 20, 21) white to cream, mottled, sometimes sparsely, with brown scales, usually slightly darker in
apical fifth. Costa darker at base. Indistinct, narrow, transverse cream streak basal to apical fifth. Brown
wedge-shaped spot, usually dark, across fold at one-third extending to dorsal margin. Smaller dark brown
spot, sometimes bisected, at end of cell. Dark spots often with narrow ochreous surround.
GENITALIA O" (Figs 48, 74, 75). Uncus small, narrowing progressively from tegumen, with tiny, pointed
median projection at apex. Gnathos small, hook-shaped, with median spine on anterior surface. Actual
margin of tegumen coincides with or slightly less emarginate than sclerotized margin anteriorly. Sacculus
very small, hook-shaped. Filament slightly, sometimes irregularly, curved; moderately slender, gradually
broader at base; almost reaching hind edge of vinculum or extending moderately short distance beyond,
posteriorly; extending short distance beyond tegumen anteriorly. Hind edge of vinculum projecting
slightly, medially; projection with truncate or irregular apex ventrally, slightly emarginate dorsally.
Vinculum with pair of sclerites parallel or slightly diverging from saccus. Saccus almost parallel-sided,
usually with truncate apex; almost reaching anterior of tegumen. Aedeagus of similar length to tegumen
plus uncus; projection near apex small.
GENITALIA $ (Figs 98, 110). Posterior margin of seventh abdominal tergite with well-defined crescent of
sclerotization. Invagination of membrane between eighth tergite and papillae anales funnel-shaped.
Eighth sternite sclerotized laterally; median area with sclerotization often weaker and patchy. Tergite very
weakly sclerotized. Apophysis anterior rod-like, length 0-4 mm (7); apophysis posterior approximately
three times length of apophysis anterior. Antrum extremely short. Ductus bursae usually of similar length
to, or slightly longer than, pear-shaped or oval corpus bursae. Signum a narrow, inwards-projecting ridge,
with faint surrounding area of sclerotization.
REMARKS. The frenulum of five out of seven females examined consists of two setae; in the
remaining two it consists of three setae on one wing and two on the other.
Specimens from Portugal, the extreme west of the range, differ in fore wing pattern from
specimens from France eastwards and were described as subspecies leonella. Examination of
material from intermediate localities, particularly in Spain, is necessary to decide whether
subspecific status is justified.
M. cytisella has two forms of male genitalia. In the typical form of c. cytisella, the filament
extends distinctly beyond the hind edge of the vinculum and is usually gently and evenly curved,
or almost straight excluding the base. In other specimens of cytisella, including c. leonella, the
filament scarcely extends beyond the hind edge of the vinculum and is usually more irregularly
curved. The spine on the gnathos is usually shorter in the typical form than in the other
specimens of c. cytisella, which I am referring to as the small form, or in leonella. The small form
of c. cytisella, and leonella, have smaller male genitalia than the typical form of c. cytisella. In the
female genitalia, the signum varies from small to large but without apparent correlation with the
two forms of male genitalia. I have assigned the females to each form of c. cytisella on the basis of
geographic association with the males.
It is possible that the small form of c. cytisella could be a separate subspecies from the typical
form, and might be more closely allied to leonella since the small form and leonella are similar in
genitalia structure. However, the small form has a widely disjunct distribution, populations
occurring both west and east of the typical form, thus it seems unsatisfactory to consider it a
distinct subspecies.
The fore wing spots of cytisella, other than in leonella, are usually more distinct than in the
other species of Mirificarma, except maculatella and denotata which have larger spots. M.
cytisella can be distinguished from the other species of the genus by the small hook-shaped
sacculus, the spine of the gnathos, and the form of the signum.
GELECHIID MOTHS
35
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T c
E -E* J>
J2
~8J
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u
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O
D
D
36 L. M. PITKIN
BIOLOGY. Host-plants: Genisteae: Cytisus nigricans L. (Treitschke, 1833: 100); Genista
(Schiitze, 1931: 120 attributes this to an untraced record by Disque); Laburnum anagyroides
Medicus (= Cytisus laburnum L.), Calicotome spinosa (L.) Link and Trifolieae: Ononis spinosa
spinosa L. (= campestris Koch & Ziz) (Lhomme, [1946-1948]: 592; the record of L. anagyroides
is based on the proximity of the moths to the plant and requires the confirmation of bred
specimens).
A record of Daphne (Thymelaeaceae) (Mariani, 1943: 168) as a host-plant is probably
erroneous.
The larva occurs in June, September and October within two or three spun leaves and the
pupal stage is from October to April (Treitschke, 1833: 100; Lhomme, [1946-1948]: 592;
Eckstein, 1933: 134). Moths have been collected from April to September.
DISTRIBUTION (Fig. 5). France, Germany, Corsica, Italy, Austria, Yugoslavia, Czechoslovakia,
Hungary, Albania, Greece, U.S.S.R.
Additional records from the literature: Spain (Agenjo, 1968: [4]); Switzerland (Miiller-Rutz,
1914: 489); Poland (Schille, 1931: 179).
Key to the subspecies
1 Spot across fold of fore wing dark, strongly contrasting with background c. cytisella (p. 36)
- Spot across fold of fore wing not strongly contrasting with background c. leonella (p. 37)
Mirificarma cytisella cytisella (Treitschke)
(Figs 5, 20, 48, 75, 98, 110)
Lita cytisella Treitschke, 1833: 99. LECTOTYPE $ , GERMANY (EAST) (TM), here designated [examined].
Gelechia cytisella Treitschke ab. roseella Hauder, 1918: 102. [Unavailable, infrasubspecific name.]
Cf, 6-0-8-0 mm. $, 6-0-7-5 mm. Fore wing (Fig. 20) with mottled brown darker at costa in four-fifths.
Wedge-shaped spot across fold dark brown, strongly contrasting with background, usually unbroken.
GENITALIA cf (Figs 48, 75). Filament almost reaches or extends beyond hind edge of vinculum.
Typical form Small form
Tegumen plus uncus length 1-2-1-3 mm (14) 1-0-1-1 mm (10)
Filament length 1-2-1-3 mm (14) 0-8-0-9 mm (10)
Filament length/tegumen plus uncus length 1-0 mm (14) 0-7-0-9 mm (12)
GENITALIA $ (Figs 98, 110). As described on p. 34.
REMARKS. The nominate subspecies differs from c. leonella in the more strongly contrasting fore
wing pattern of the former.
M. c. cytisella was described from an unspecified number of specimens. I designate as
lectotype the single type-specimen I have examined, which was already labelled 'lectotype' by
Karsholt.
BIOLOGY. Host-plants as described on p. 36. Moths of the typical form have been found from
April to August; those of the small form have been found from May to September.
DISTRIBUTION (see Fig. 5). Typical form: Germany, Austria, Yugoslavia, Hungary, Albania,
Greece. A pair of specimens from the Meyrick Collection, bearing the data 'France: Ardeche',
may have been mislabelled.
Small form: France, Corsica, Italy, U.S.S.R.
Form unidentified: Czechoslovakia.
MATERIAL EXAMINED
Typical form (including 14 cf , 5 $ genitalia preparations)
Lectotype $ (cytisella), Germany (East): Meissen, highlands, larva on Cytisus nigricans, ix, ex iv (von
Tischer) (TM).
France: 1 cf , 1 $, Ardeche. Germany: 3 cf , 1 $?, S. Germany (West): 1 $, Regensburg ('Ratisbon');
2 cf , Bavaria. Austria: 1 cf , [Karnten,] Ofen; 2 cf , 1 $, Ober-Osterreich, Linz area, iii ?, vii (LN; MM);
2 $, Nieder-Osterreich, Wiener Wald, Leopoldsberg, vi (NM); 1 cf , Modling; 1 $, Nieder-Osterreich,
GELECHIID MOTHS 37
Klosterneuburg, iv (NM); 1 cf, Nieder-Osterreich, Falkenstein, vi. Yugoslavia: 1 cf , Rijeka ('Fiume')
(NM); 1 $, Istra ('Istria'), 'Sin', iv (NM); 2 $, Zadar ('Zara'), viii (NM). Hungary: 1 cf, near Debrecen,
Ermihalyfalva, v (NM) (Rothschild, 1913: 80). Albania: 1 cf , [NE], Kula Ljums, vi (NM) (Rebel, 1931:
146). Greece: 1 cf, Lakonia, 5 km S. of Monemvasia, iv (ZM).
Small form (including 13 cf , 4 $ genitalia preparations)
France: 1 cf, Lot, Douelle, vii (MNHN); 1 cf, Ste Croix-Vallee-Francaise, vii (MNHN); 1 cf,
Hautes-Alpes, St Julien-en-B[eauchene], vii; 1 cf, Hyeres; 1 $, Hyeres, v (MNHN); 1 cf, 1 $,
Alpes-Maritimes, St Martin, 1500 m, vi. Corsica: 3 cf, 1 $, Evisa, 850 m, viii, ix (NM). Italy: 1 cf,
Piemonte, 'Valle di Poggio di' Casasco, v; 1 $, Piemonte, Monferato, Cardona, v; 1 cf, Trentino-Alto
Adige, Pietramurata, 250 m, viii; 1 $, Trentino-Alto Adige, Val Sarca, Pietramurata, 250 m, vii (all coll.
Jackh, Bidingen); 1 cf, Toscana, Fiesole, vii; 1 cf, Rome (MNHU). U.S.S.R.: 4 cf, [S. of Volgograd,]
Krasnoarmeysk ('Sarepta'), viii; 1 cf , Bol'shoy Kavkaz ('Caucasus'), Tbilisi ('Tiflis'), v.
Form unidentified (including 1 $ genitalia preparation)
France: 1 ?, Le Rozier, vii (MNHN); 1 $, Herault, 'St Guilhem-le-Dt' (MNHN). Italy: 2 , [Friuli
Venezia], Raibl (NM). Czechoslovakia: 2 $, Bohemia; 1 <j>, Praha ('Prag') (MNHU). No locality data: 4
Cf,6$.
Mirificarma cytisella leonella Amsel
(Figs 5, 21, 74)
Mirificarma [Gelechia] cytisella leonella Amsel, 1959: 156, 164, pi. 1, fig. 3. Holotype cf , PORTUGAL (LN)
[examined] .
Cf , 6-5-7-5 mm. Fore wing (Fig. 21) with mottled brown slightly darker in four-fifths. Wedge-shaped spot
across fold not strongly contrasting with background, usually broken.
GENITALIA cf (Fig. 74). Filament does not extend beyond hind edge of vinculum.
Tegumen plus uncus length 0-9-1-0 mm (3)
Filament length 0-7-0-8 mm (3)
Filament length/tegumen plus uncus length 0-8-0-9 mm (3)
GENITALIA $. Not examined.
REMARKS. M. c. leonella differs from the nominate subspecies in the less strongly contrasting
fore wing pattern of the former.
BIOLOGY. Host-plant unknown. Moths have been found in June (also in August according to
Amsel, 1959: 157).
DISTRIBUTION. Portugal.
MATERIAL EXAMINED (including 3 cf genitalia preparations)
Holotype cf , Portugal: [40 km N. of Porto,] Singeverga, vi.1953 (Monteiro) (only genitalia slide, no.
323c, Sattler, examined; LN) (labelled 'spp. lusitaniella Ams.').
Portugal: 3 cf (paratypes), Singeverga, vi.1953 (Monteiro) (LN); 1 cf (paratype), Montalegre (Mon-
teiro} (LN) (only genitalia slide examined).
Mirificarma monticolella (Rebel) comb, n., stat. n.
(Figs 1,2, 22, 49, 57, 58, 76)
Lita acuminatella f. monticolella Rebel, 1931: 147, 160. LECTOTYPE cf, ALBANIA (NM), here designated
[examined].
Cf , 6-5-7-0 mm. Head cream. Labial palpus third segment and apex of second segment cream, otherwise
mid brown mottled with cream on inner surface. Thorax and tegula cream mottled with brown. Fore wing
(Fig. 22) cream mottled with brown, predominantly cream in central area of wing; apex edged with small
dark brown spots. Small dark brown spots, sometimes indistinct: spot or streak in fold near base, spot in
fold at one-third, two on costal edge of cell and one at end of cell.
GENITALIA cf (Figs 49, 57, 58, 76). Uncus small, narrowing progressively from tegumen, without apical
projection. Gnathos extremely short, without median spine. Actual margin of tegumen coincides with
sclerotized margin anteriorly. Sacculus broad, particularly towards apex, club-shaped. Filament gently
38 L. M. PITKIN
curved, very moderately stout; posteriorly extending to hind edge of vinculum or just beyond, scarcely
extending beyond tegumen anteriorly. Hind edge of vinculum projecting medially with broad U-shaped
emargination of sclerotization. Vinculum with pair of sclerites. Saccus very slender, wider at base than at
pointed apex; not quite reaching anterior of tegumen. Aedeagus almost as long as tegumen plus uncus;
projection near apex minute.
GENITALIA $. Unknown.
REMARKS. The small dark spots at the apex of the fore wing are also present in many other
species but they are particularly distinct in monticolella and interuptella in which they contrast
with the light ground colour. In the male genitalia, the sclerites of the vinculum appear faint,
perhaps because the preparations are not stained.
This species bears a superficial resemblance to montivaga and some specimens of ocellinella
which also have a pale fore wing with sparse dark markings. However, monticolella is smaller
than ocellinella and montivaga has no fore wing markings other than the mottled ground colour.
M. monticolella has many genitalic differences from both species, including the size of the male
filament. M. monticolella can be distinguished from all other Mirificarma species by the
extremely short gnathos and the very slender, short saccus.
M. monticolella was described from 2 cf , both of which I have examined.
BIOLOGY. Host-plant unknown. Moths have been found in May or June.
DISTRIBUTION. Northern Albania.
MATERIAL EXAMINED (including 2 cf genitalia preparations)
Lectotype cf , Albania: 15 km NNE. of Kula e Lumes, Beshtriq ('Pashtriq'), 1896 m, 29.v.^.vi.l918
(Penther, Predota & Zerny) (genitalia slide no. 1491; NM).
Albania: 1 cf (paralectotype), same data as lectotype (NM).
Mirificarma interuptella (Hiibner)
(Figs 1, 2, 4, 23, 50, 77, 84, 99, 111)
Phfalaena] Tin[ea] interuptella Hiibner, 1793: 14, pi. 88. Type(s) [not traced].
Tinea interruptella Hiibner; Hiibner, 1822: 72. [Incorrect subsequent spelling of interuptella Hiibner.]
Anacampsis interrupta Curtis, 1827: no. 189, folio [2]. [Unjustified emendation of interuptella Hiibner.]
Cf, $,7-0-8-5 mm. Head white to cream. Labial palpus with dark apex; second segment with brown areas,
particularly externolaterally, sometimes dark, apex cream. Thorax white to cream, occasionally with thin,
median, longitudinal dark brown stripe. Tegula dark brown, sometimes with scattered cream scales. Fore
wing (Fig. 23) cream tinged with pale yellow; with scattered brown scales. Dark brown, median,
longitudinal band present, continuous but usually darker in fold and on costal edge of band from one-third
to apex. Fore wing apex distinctly edged with small dark brown spots.
GENITALIA cf (Figs 50, 77). Uncus small, half to two-thirds width of tegumen; sometimes slightly
constricted at base. Gnathos a large strongly-angled hook, without median spine. Actual margin of
tegumen slightly less emarginate than sclerotized margin anteriorly. Sacculus long, slender with spatulate
apex. Filament helical, very moderately stout; not extending posteriorly far beyond hind edge of vinculum,
scarcely extending beyond tegumen anteriorly. Hind edge of vinculum far-projecting, narrowing to
median V-shaped emargination. Vinculum with single sclerite. Saccus very slender, slightly deflected to
left in ventral view, extending slightly beyond tegumen anteriorly. Aedeagus slightly longer than tegumen
plus uncus; apex cylindrical with tiny projection.
GENITALIA $ (Figs 99, 111). Posterior margin of seventh abdominal segment with area of well-defined
sclerotization on tergite and pair of adjoining patches on sternite. Very slight, flimsy invagination of
membrane between eighth sternite and papillae anales. Eighth sternite strongly sclerotized laterally;
asymmetrical, with lobe projecting over median area from left side in ventral view; lateral areas distinctly
contrasting with membraneous median area. Median area of sternite without striations. Tergite consisting
of lateral longitudinal strips connected by narrow posterior horizontal strip, all strongly sclerotized;
otherwise absent. Apophyses anteriores diverging lobes, length approximately 0-3-0-4 mm (8), left shorter
than right in ventral view; apophysis posterior approximately three to four times length of apophysis
anterior. Antrum almost straight, sometimes deflected to one side, narrowing towards anterior; anterior
not indented; moderately long, extending beyond apophyses anteriores. Ductus bursae of similar length to
oval or round corpus bursae. Signum oval, curved inwards, covered with tiny spinules.
GELECHIID MOTHS 39
REMARKS. The small dark spots at the apex of the fore wing are also present in many other
species; as the spots contrast with the light ground colour of the wing in interuptella, they are
particularly pronounced. The frenulum in five females examined consists of three setae. The
genitalia are more asymmetrical than those of other Mirificarma species, particularly the male
saccus and the eighth sternite and apophyses anteriores of the female.
This species bears a superficial resemblance tofasciata and to specimens oimulinella in which
the fore wing ground colour is pale and the stripe is unbroken, although the dark fore wing stripe
of interuptella contrasts more strongly with the cream ground colour. M. interuptella is clearly
distinguished from both species by the form of the male filament, the slender sacculus with a
spatulate apex which is unique within the genus, and the apophysis anterior of the female.
M. interuptella appears to have some affinities to burdonella and flavonigrella, with which it
shares the aedeagus apex shape; in the female, the lobe-shaped apophysis anterior and the
degree of sclerotization of the eighth sternite resemble those of burdonella. M. interuptella is
distinguished from both species by characters including the sacculus shape and the large
gnathos, and from burdonella by the presence of the signum. The female of flavonigrella is
unknown.
Subsequent to Hiibner's original description, many authors have misspelt interuptella as
' inter ruptella' including Koc.ak (1982: 106) who incorrectly cites 'interruptella Hiibner, 1793' as a
junior primary homonym of interruptella de Villers (1789: 520) and proposes albicosta Haworth
as the replacement name. Coleophora albicosta Haworth is currently a valid species of
Coleophoridae (Bradley, 1966: 132). The identity ofPhalaena (Tinea) interruptella de Villers is
unknown.
BIOLOGY. Host-plants: Genisteae: Cytisus scoparius (L.) Link (as Spartium scopariuni) (Disque,
1908: 129); C. purgans (L.) Boissier, formerly in Genista (Lhomme, [1946-1948]: 594); Genista
(tinctoria L. , pilosa L. , germanica L. , or sagittalis L.) (Disque, 1908: 79).
The larva probably lives in the flowers (Lhomme, [1946-1948]: 594, attributes this informa-
tion to Peyerimhoff) although Schiitze (1931: 121) records it in August under leaves spun to the
twig (on C. scoparius). Sorhagen (1886: 187) records the larva in May. Moths have been
collected from March to July (also in August according to Sorhagen, 1886: 187).
DISTRIBUTION. Spain, France, central Europe, Czechoslovakia, Poland, North Africa.
Additional records. Portugal (Zerkowitz, 1946: 133); Belgium (Lhomme, [1946-1948]: 594);
Netherlands (Lempke, 1976: 26); Italy (Mariani, 1943: 167). Records of interuptella from Great
Britain are misidentifications oimulinella (see p. 43).
MATERIAL EXAMINED (including 11 cf , 10 $ genitalia preparations)
Spain: 1 cf , Avila (NM); 1 cf , 1 $, Teruel (NM); v-vii; 1 cf , 1 $, Granada (NM). France: 2 cf , 1 $,
Essonne; 1 cf , Lot; 2 <J>, Pyrenees-Orientales; 2 cf , 1 $, Basses Alpes; 6 cf , 11 $, Var; iv-vi. Germany:
1 Cf , 2 $, ; 1 cf , N. Germany (West): 1 ex., Baden-Wiirttemberg; 1 $, Niedersachsen, v (coll. Jackh,
Bidingen); 1 cf, Hamburg, vi (coll. Jackh, Bidingen). Germany (East): 1 $, Potsdam, vii (NM).
Switzerland: 1 cf (NM). Austria: 1 cf, 1 ?, Wien (NM). Czechoslovakia: 1 $, Plzen, v ? (NM). Poland:
5 Cf , 5 $ , Szczecin; 1 cf , Zielona Gora; v, vi ? Morocco: 2 cf , iii. Algeria: 1 Cf , 1 $ , Constantine province, v
? Tunisia: 1 cf , iv. No locality data: 33 ex.
Mirificarma fiavonigrella (Chretien) comb. n.
(Figs 1,2, 24, 51, 59, 78)
Gelechia flavonigrella Chretien, 1915: 318. LECTOTYPE cf, ALGERIA (MNHN), here designated
[examined].
Cf , 6-0 mm. Head cream. Labial palpus cream, mottled with dark brown mostly confined to outer surface
of first and second segments. Thorax cream; tegula dark brown. Fore wing (Fig. 24) mottled light brown
and cream, darker in apical fifth; costal margin brown, dark at base; with dark brown stripe along fold
separated by pale area from longitudinal, median, more diffuse brown stripe which extends from half to
near apex. Faint yellowish tinge scattered on fore wing.
40 L. M. PITKIN
GENITALIA cf (Figs 51, 59, 78). Uncus small, narrowing towards apex, approximately half width of
tegumen. Gnathos a small simple hook. Actual margin of tegumen coincides with sclerotized margin
anteriorly. Sacculus long, extending far beyond median projection of hind edge of vinculum; broad
apically, club-shaped, with inner edge slightly rugose towards apex. Filament curved although apical half
almost straight; basal half expanded dorsoventrally and compressed laterally, smooth; apical half
moderately, uniformly, slender. Filament not reaching hind edge of vinculum posteriorly, extending
beyond tegumen anteriorly. Median projection of hind edge of vinculum with shallow V-shaped emargina-
tion. Vinculum with pair of sclerites converging near hind edge of vinculum. Saccus moderately broad,
parallel-sided with rounded apex, barely extending beyond tegumen. Aedeagus slightly longer than
tegumen plus uncus; apex almost cylindrical with minute projection.
GENITALIA $. Unknown.
REMARKS. This species bears a very close superficial resemblance to some mulinella specimens in
which the fore wing has a light background and a broken stripe, but it can be distinguished from
mulinella by the club-shaped sacculus of the male genitalia. M. flavonigrella is very similar to
burdonella in external appearance and genitalia, and might prove to be no more than a
subspecies if further material becomes available. They are allopatric but this may not be
significant as both species are known from so little material. M. flavonigrella differs from
burdonella in the fore wing pattern since the two stripes are slightly less widely separated by a
pale area. The male genitalia differ from those of burdonella, mainly in the sacculus extending
far beyond the median projection of the hind edge of the vinculum, and in the shallower
emargination of this projection.
M. flavonigrella was described from an unspecified number of specimens of which I designate
as lectotype the single type-specimen examined, which was already labelled 'lectotype' by
Saltier.
BIOLOGY. Host-plant unknown. The moth has been found in May.
DISTRIBUTION. Algeria (Oran area).
MATERIAL EXAMINED (including 1 cf genitalia preparation)
Lectotype cf , Algeria: near Oran, Frenda, v.1911 (genitalia slide no. 471b, Sattler; MNHN).
Mirificarma burdonella (Rebel)
(Figs 1,2, 25, 52, 79, 100, 112)
Gelechia burdonella Rebel, 1930: 25. LECTOTYPE cf , CORSICA (NM), here designated [examined].
Gelechia bardonella Rebel; Gaede, 1937: 148. [Incorrect subsequent spelling.]
Cf , 5-5-6-5 mm. $ , 5-5-6-0 mm. Head cream to light brown. Labial palpus cream, mottled with dark brown
mostly confined to outer surface of first segment and basal part of second segment; apex dark brown.
Thorax cream to light brown, sometimes with scattered brown scales. Tegula usually brown in basal half,
pale apically. Fore wing (Fig. 25) mottled light brown and cream, usually darker in apical fifth and at costal
margin, with irregular dark brown stripe along fold. This stripe usually well separated by pale area from
longitudinal, irregular, dark brown stripe which is median or slightly costad, extending from almost half to
three-quarters or near apex, continuous, or, infrequently, broken. Faint longitudinal yellowish streak
sometimes present in costal half and fold line.
GENITALIA cf (Figs 52, 79). Uncus small, narrowing towards apex, slightly more than half to two-thirds
tegumen width. Gnathos a small simple hook. Actual margin of tegumen almost coincides with sclerotized
margin anteriorly. Sacculus scarcely extending beyond median projection of hind edge of vinculum; broad,
club-shaped, with inner edge slightly rugose. Filament slightly curved although apical half almost straight;
basal half expanded dorsoventrally, laterally compressed and wrinkled; apical half moderately, uniformly,
slender. Filament not reaching hind edge of vinculum posteriorly, extending a short distance beyond
tegumen anteriorly. Median projection of hind edge of vinculum with deep, narrow U- or V-shaped
emargination. Vinculum with pair of sclerites converging near hind edge of vinculum. Saccus moderately
broad, parallel-sided or narrowing slightly towards rounded apex, extending slightly beyond tegumen
anteriorly. Aedeagus slightly longer than tegumen plus uncus, slightly S-shaped; apex almost cylindrical
with tiny projection.
GELECHIID MOTHS 41
GENITALIA $ (Figs 100, 112). Posterior margin of seventh abdominal segment with area of well-defined
sclerotization on tergite, pair of patches sometimes present on sternite. Invagination of membrane
between eighth tergite and papillae anales long and narrow. Eighth sternite strongly sclerotized laterally,
wrinkled, distinctly contrasting with weakly sclerotized, horizontally striated median area bordered by pair
of narrow longitudinal sclerites. Tergite with pair of very small, lateral, sclerotized areas extending from
sternite, otherwise weakly sclerotized. Apophyses anteriores diverging lobes, length 0-2 mm (3); apo-
physis posterior approximately six times length of apophysis anterior. Antrum slightly curved and
constricted towards anterior; anterior not indented; moderately long, approximately two to three times
length of apophysis anterior. Ductus bursae at most one-third length of oval corpus bursae. Signum not
discernible.
REMARKS. The frenulum of two out of the three females examined consists of three setae; in the
remaining female it consists of three setae on one wing and two on the other.
Externally this species strongly resembles flavonigrella and some specimens of mulinella in
which the fore wing pattern has a pale background and a broken stripe, although in burdonella
the two stripes are slightly more widely separated. M. burdonella is clearly distinguished from
mulinella by the club-shaped sacculus, the lobe-shaped apophysis anterior and the degree of
sclerotization of the female eighth sternite. M. burdonella appears to be most closely related to
flavonigrella from which it can be distinguished by the extent of the sacculus.
M. burdonella was described from an unspecified number of syntypes from Corsica: Col de
Vergio and Evisa. I have examined 2 cf syntypes from Col de Vergio and 1 9 syntype from Evisa
and I designate a male as lectotype. All three specimens bear the label Type'.
BIOLOGY. Host-plant unknown. Moths have been found in August and September.
DISTRIBUTION. Corsica and Sardinia.
MATERIAL EXAMINED (including 6 cf , 3 $ genitalia preparations)
Lectotype cf , Corsica: Col de Vergio, 1460 m, at light ('Lichtfang'), 31.viii.1929 (Reisser) (genitalia slide
no. 11227; NM).
Corsica: 1 cf (paralectotype), Col de Vergio, 1460 m, 3.ix.l929 (Reisser) (NM); 3 Cf , 3 $ , Col de Vergio,
1450, 1460m, viii,ix(BMNH;NM); 1 $ (paralectotype), Evisa, 850 m,4.ix. 1929 (Reisser) (NM) ; 1 $, Col
de Sevi, 1000 m, ix. Sardinia: 1 cf , centr[al], valley ? ('vl.') Bruncu Spina, 1750 m, viii; 1 cf , S. ('merid.'),
'Musei', 120m, ix.
Mirificarma cabezella (Chretien)
(Figs 1,2, 26, 53, 54, 80, 101)
Gelechia maculatella f. cabezella Chretien, 1925: 245. Lectotype cf , SPAIN (MNHN), designated by Sattler
(1961: 86) [examined].
Mirificarma cabezella (Chretien) Sattler, 1960: 42; Agenjo, 1962: 159, pi. 2, fig. 7, pi. 3, fig. 8 [legends to
figs 7 and 8 are transposed on pi. 3].
Cf , 6-5-7-5 mm. $, 6-5-7-0 mm. Head mid brown, occasionally ochreous brown. Labial palpus mottled
dark brown and cream, usually predominantly cream on inner surface. Thorax and tegula as head. Fore
wing (Fig. 26) mottled brown, darker in apical half; small area of dark brown at base; ochreous tinge near
base. Faint cream spot at costa at three-quarters extending diffusely to dorsal margin, usually crossed
medially by faint longitudinal ochreous dash. Dark brown spot across fold at one-third, extending diffusely
to dorsal margin; another at end of cell, usually constricted or bisected, frequently merging with dark
background. Both spots sometimes diffuse; with narrow ochreous surround. Very small, indistinct dark
brown spot in fold near base.
GENITALIA cf (Figs 53, 54, 80). Uncus small, nearly half width of tegumen; scarcely constricted at base;
apex without projection. Gnathos a moderately large, distinctly curved, simple hook. Actual margin of
tegumen slightly less emarginate than sclerotized margin anteriorly. Sacculus very long, extending beyond
gnathos arms, almost straight, uniformly moderately slender, smooth. Filament almost straight, very
slender; extending posteriorly approximately to hind edge of vinculum; extending a short distance beyond
tegumen anteriorly. Median projection of hind edge of vinculum low with indistinct dorsal V-shaped
emargination ; very slight median ventral emargination. Vinculum with slightly bowed pair of sclerites from
saccus. Saccus usually slightly narrower towards truncate or irregular apex, extending slightly beyond
42 L. M. PITKIN
tegumen anteriorly. Aedeagus same length as tegumen plus uncus; projection near apex moderately large,
weakly sclerotized, rounded.
GENITALIA $ (Fig. 101). Posterior margin of seventh abdominal segment without sclerotized area but with
distinct dense band of scales. Invagination of membrane between eighth tergite and papillae anales
funnel-shaped. Most of eighth segment sclerotized. Median longitudinal areas of sternite and tergite
slightly less sclerotized than surrounding areas, slightly sunken. Tergite median longitudinal area bordered
by pair of faint, narrow sclerites. Apophysis anterior rod-like, with separate, large, rounded lobe at base
towards antrum; length 0-3-0-4 mm (4); apophysis posterior three to four times length of apophysis
anterior. Antrum slightly curved and constricted towards anterior; anterior not indented; moderately long,
longer than but less than 1-5 times length of apophysis anterior. Posterior half of antrum less sclerotized
than anterior half. Ductus bursae not more than half length of oval corpus bursae. Signum not discernible.
REMARKS. The female frenulum consists of three setae. The band of scales at the posterior
margin of the female abdomen, distinct in this species and in ulicinella and mulinella, is also
present, although more diffuse, in some other species. The weaker sclerotization of the
posterior half of the antrum is distinct in cabezella, although it also occurs, less noticeably, in
other species including mulinella, ulicinella and burdonella.
The Moroccan male differs slightly in genitalia from Spanish males. The filament is slightly
shorter, particularly the part apical to the tiny opening near the apex, and the saccus is almost
parallel-sided, whereas it narrows slightly, towards the apex, in the Spanish specimens.
This species bears a superficial resemblance to rhodoptera, scissella and, particularly, to
constricta. It differs from rhodoptera and scissella in many genitalic characters since these two
species are in the maculatella-group, and from constricta in the shape of the uncus, sacculus and
filament. M. cabezella appears to be closely related to mulinella and ulicinella. They share the
uniformly slender, straight sacculus of the male, and the distinct, dense band of scales of the
posterior margin of the female abdomen. M. cabezella differs in wing pattern and by the very
slender filament of the male genitalia and the presence of the lobe at the base of the apophysis
anterior of the female.
BIOLOGY. Host-plant: Genisteae: Adenocarpus hispanicus (Lamarck) (type-series bred by
Chretien).
Chretien (1925: 245) found the larvae in the shoots in June; the moths emerged in September.
Moths have been collected in August and October.
DISTRIBUTION. Spain and Morocco.
MATERIAL EXAMINED (including 4 d", 4 $ genitalia preparations)
Lectotype cf , Spain: [Segovia,] La Granja ('San Ildefonso'), larva on Adenocarpus hispanicus, vi.1902,
ex. ix. 1902 (Chretien & Dumont) (genitalia slide no. 3562, Viette; MNHN).
Spain: 3 cf , 2 $ (paralectotypes), data as lectotype, ex viii, ix.1902 (Chretien & Dumont) (MNHN; LN);
1 $, , viii (MNHN). Morocco: 1 cf 1 $, Moyen Atlas, Val d'Ifrane, 1500-1600 m, x (coll. Burmann,
Innsbruck).
Mirificarma ulicinella (Staudinger)
(Figs 1,2, 27, 55, 81, 102)
Gelechia ulicinella Staudinger, 1859: 240; Milliere, 1863: 325, pi. 38, figs 8-10. LECTOTYPE cf , SPAIN
(MNHU), here designated [examined].
Cf , 5-5-6-5 mm. $ , (4-5) 5-0-6-0 mm. Head ochreous cream. Labial palpus cream mottled with brown on
outer surface, usually predominantly brown towards base; predominantly cream on inner surface; apex
brown. Thorax and tegula ochreous cream slightly mottled with brown; tegula usually very dark brown at
base. Fore wing (Fig. 27) mottled dark and light brown; with paler yellow-ochre areas situated as follows:
dorsal margin at base; along fold; median longitudinal stripe from one-third to three-quarters or near apex,
adjoining fold, irregular, constricted or broken, crossed by indistinct transverse stripe or blotch at
two-thirds or nearer apex; longitudinal stripe near costa from base to one-third. Two small dark brown
spots on median stripe at one-third and one-half, sometimes indistinct.
GELECHIID MOTHS 43
GENITALIA cf (Figs 55, 81). Uncus small, less than one-third to less than half-width of tegumen; scarcely
constricted at base; apex with tiny, pointed, median projection. Gnathos short, only slightly curved,
extreme apex a small hook-shape. Actual margin of tegumen coincides with sclerotized margin anteriorly.
Sacculus not reaching gnathos arms, straight, uniformly very slender, smooth. Filament curved, not
spiralled; extremely stout basally, laterally compressed and dorsoventrally expanded in apical half,
narrowing towards apex; extending posteriorly almost to hind edge of vinculum or slightly beyond; not
usually reaching anterior of tegumen. Median projection of hind edge of vinculum with heart-shaped
dorsal emargination, not emarginate ventrally. Vinculum with strongly bowed pair of sclerites from saccus.
Saccus broad at base, narrowing towards truncate apex; reaching anterior of tegumen. Aedeagus same
length as tegumen plus uncus or slightly shorter; projection near apex moderately large, weakly
sclerotized, rounded.
GENITALIA $ (Fig. 102). Posterior margin of seventh abdominal segment without sclerotized area but with
distinct dense band of scales. Invagination of membrane between eighth tergite and papillae anales broad,
almost truncate. Most of eighth segment sclerotized; median longitudinal areas of sternite and tergite
slightly less sclerotized than surrounding areas. Tergite strongly sclerotized, with pair of gently curved
lobes overlapping narrow median longitudinal area. Apophysis anterior rod-like, enlarged at base towards
antrum, but without separate lobe; length 0-3-0-4 mm (5); apophysis posterior three times length of
apophysis anterior. Antrum slightly curved and constricted towards anterior; anterior not indented;
moderately long, approximately 1-5 times length of apophysis anterior. Ductus bursae considerably
shorter than oval corpus bursae. Signum not discernible.
REMARKS. The frenulum was examined in five females and consists of two setae. This species
appears to be closely related to mulinella and cabezella with which it shares genitalic characters
including the sacculus shape and the distinct band of scales at the posterior margin of the female
abdomen. It differs in its slightly curved gnathos and shorter sacculus of the male and in the form
of the eighth tergite of the female. The fore wing of ulicinella, with yellow-ochre areas not in a
transverse zig-zag pattern, differs from all other Mirificarma species.
M. ulicinella was described from four specimens from Spain: Granada. I have examined 1 cf ,
1 9 and designate as lectotype the male which was already labelled 'lectotype' by Saltier.
BIOLOGY. Host-plant: Genisteae: Ulex parviflorus Pourret (= australis Clemente; = provincialis
Loisel) (type-series bred by Staudinger).
The larva has been recorded from October to April in the flowers; it pupates at the base of the
plant in a shell of dried leaves, the adult emerging in August; there is only one generation per
year (Milliere, 1863: 326; Lhomme, [1946-1948]: 566, Staudinger, 1859: 241). Moths have been
collected at light and bred in September (also in May and June according to Lhomme,
[1946-1948]: 566).
DISTRIBUTION. Spain and France.
Additional record. Italy: Liguria (Mariani, 1943: 166).
MATERIAL EXAMINED (including 7 cf , 5 $ genitalia preparations)
Lectotype cf , Spain: Granada, iv, larva on Ulex australis, moth emerged ix (Staudinger) (genitalia slide
no. 476d, Sattler; MNHU).
Spain: 1 $ (paralectotype), same data as lectotype (MNHU); 23 cf , 7 $, Granada province, Sierra de
Alfacar, 1500 m, ix. France: 1 cf , Provence. No locality data: 1 cf .
Mirificarma mulinella (Zeller)
(Figs 1,2, 28-30, 56, 82, 103)
Gelechia mulinella Zeller, 1839: 199; Stainton, 1865: 96, pi. 3, fig. 3. LECTOTYPE $, GERMANY (EAST)
(BMNH), here designated [examined].
[Anacampsis interrupta Curtis, 1827: no. 189, folio [2]. Unjustified emendation of interuptella Hiibner.
Misidentification.]
[Anacampsis interrupted (Hiibner) ?; Stephens, 1829: 197; Curtis, 1829 [-1831]: [91]. Incorrect subse-
quent spelling of interuptella Hiibner. Misidentifications.]
[Anacampsis interrupted (Hubner); Stephens, 1834: 215; Westwood, 1845: 191, pi. 107, fig. 3. Incorrect
subsequent spelling of interuptella Hubner. Misidentifications.]
Gelechia caminariella Fuchs, 1902: 323. Holotype $. GERMANY (WEST): [Rheinland-Pfalz,] Rheingau,
44 L. M. PITKIN
near Bornich, Rheinberge, Rieslingberg, around Sarothamnus, viii. 1878 (Fuchs) [not traced]. [Synony-
mized by Rebel, 1930:25.]
Gelechia carminariella Fuchs; Eckstein, 1933: 134. [Incorrect subsequent spelling of caminariella Fuchs.]
Gelechia nigraesilvae Amsel, 1950: 27, figs 1, 2. Holotype cf, GERMANY (WEST) (LN) [examined].
[Synonymized by Karsholt & Nielsen, 1976: 33.].
Cf , 6-0-7-5 mm. $ (4-5) 5-5-7-5 mm. Head cream to light brown. Labial palpus mottled brown and cream;
apex dark brown; apex second segment usually mostly cream. Thorax and tegula light to mid brown often
mottled with dark brown. Tegula slightly darker brown towards base. Fore wing (Figs 28-30) mottled
brown, mixed with cream or light brown in apical third and occasionally also in posterior third. Dark brown
spot usually present, at end of cell. Small dark brown spots sometimes present, at one-third, one in fold,
one in cell. Dark brown, median, longitudinal stripe sometimes present, from one-third to apex, another in
fold line. These stripes merge to form single diffuse band, or slightly separated by pale area.
GENITALIA cf (Figs 56 , 82) . Uncus small , half to almost two-thirds width of tegumen ; scarcely constricted at
base; apex without projection. Gnathos a moderately large, distinctly curved, simple hook; extreme apex
usually a very slight hook-shape. Actual margin of tegumen coincides with sclerotized margin anteriorly.
Sacculus long, reaching or extending beyond gnathos arms, straight, uniformly very slender, smooth.
Filament straight or gently curved, not spiralled; stout, particularly basally, laterally compressed in apical
half; not reaching hind edge of vinculum posteriorly, extending a short distance beyond tegumen
anteriorly. Median projection of hind edge of vinculum high with shallow, ventral U- or V-shaped
emargination. Vinculum with pair of sclerites slightly bowed or diverging from saccus. Saccus slender,
almost parallel-sided with rounded or irregularly truncate apex; extending moderately well beyond
tegumen anteriorly. Aedeagus slightly longer than tegumen plus uncus; projection near apex moderately
large, weakly sclerotized, rounded.
GENITALIA 9 (Fig. 103). Posterior margin of seventh abdominal segment without sclerotized area but with
distinct dense band of scales. Invagination of membrane between eighth tergite and papillae anales small,
broadly conical or funnel-shaped . Most of eighth segment sclerotized ; median longitudinal areas of sternite
and tergite slightly less sclerotized than surrounding areas; median longitudinal area of sternite slightly
sunken. Tergite without pair of curved lobes. Apophysis anterior rod-like, without separate lobe at base
towards antrum, slightly constricted near apex; length 0-3-0-5 mm (9); apophysis posterior three to four
times length of apophysis anterior. Antrum slightly curved and constricted towards anterior; anterior not
indented; moderately long, usually 1-5 times to twice length of apophysis anterior. Ductus bursae usually
one-quarter to less than half length of oval or round corpus bursae. Signum not discernible.
REMARKS. The frenulum of mulinella consists of two setae, or two setae on one wing and three on
the other. In the male genitalia, the filament basal half varies in stoutness. The fore wing pattern
of this species varies by degrees from almost uniform to a pale background contrasting with a
dark broken or unbroken stripe. The contrasted form tends to occur particularly in southern
France and southern England. The specimens with an unbroken stripe bear a strong superficial
resemblance to interuptella although the contrast of pale and dark areas is never as great in
mulinella as it is in interuptella. Specimens with a broken stripe on a pale background sometimes
closely resemble flavonigrella and burdonella, although the more apical stripe is slightly longer
than in burdonella.
M. mulinella can be distinguished from these by its uniformly slender sacculus and its rod-like
apophysis anterior. It appears to be closely related to ulidnella and cabezella, since the genitalia
are similar, although the fore wing pattern differs. The male genitalia differ from those of
ulidnella in the shape of the gnathos, and from cabezella in the shape of the filament and the
median projection of the hind edge of the vinculum. In the female genitalia, mulinella can be
distinguished from ulidnella by the form of the eighth tergite and from cabezella by the absence
of lobes at the base of the apophysis anterior.
M. mulinella was described from 2 d", 4 9 from East Germany: Dresden and Poland: Glogow
('Glogau'). I designate as lectotype the single type-specimen examined, from Dresden, which
was already labelled 'lectotype' by Saltier.
BIOLOGY. Host-plants: Genisteae: Ulex europaeus L. (Stainton, 1865: 228); Cytisus scoparius
(L.) Link (formerly in Sarothamnus) (specimens bred by Pitkin and Saltier, England; Stainlon,
1865: 228); C. nigricans L. (Harlig, 1964: 27); Genista germanica L. (Sorhagen, 1886: 187);
Calicotome spinosa (L.) Link (Lhomme, [1946-1948]: 593).
GELECHIID MOTHS 45
In Great Britain, the only published host-plant records for mulinella are Ulex and C.
scoparius; however, I have seen specimens bred by Agassiz from a cultivar of Genista tinctoria L.
This plant is the host of the only other species of Mirificarma in Great Britain, lentiginosella.
Specimens of mulinella have also been bred by Langmaid (pers. comm.) from Lupinus arbor eus
Sims in Great Britain (Hampshire).
Outside Great Britain, Spartium has been recorded as a host-plant by Mariani (1943: 167);
however, this should probably be referred to C. scoparius, known as Spartium scoparium by
some authors including Stainton (1867: 26). A record of Bartsia aspera (Brotero) Lange
(Scrophulariaceae) (Zerkowitz, 1946: 133) as a host-plant of mulinella is very dubious.
The larva occurs from April to early May; on Ulex and C. scoparius it makes a small hole in a
bud that is not fully open and feeds on the interior of the flower, before repeating the process in
another flower. It pupates on the ground in a slight cocoon amongst leaves (Stainton, 1865: 96).
The larva is found on the leaves instead of the flowers on L. arboreus, which, unlike Ulex and C.
scoparius, does not have reduced leaves (Langmaid, pers. comm.).
The larva is also found in June (Sorhagen, 1886: 187). The pupal stage is in May and June
(Bradford, [1979]: 123). Moths have been collected from July to November, also in February in
North Africa.
DISTRIBUTION. Europe west of 20E, north to Denmark, extending south to North Africa
(Algeria, Tunisia). The published distribution extends further north, to Orkney Is. (Wolff,
1971: 161) and Norway (Opheim, 1978: 27). This widespread species is found further north than
any other Mirificarma species.
Additional records. Portugal (Zerkowitz, 1946: 133); E. Ireland (Meyrick, 1895: 603);
Channel Is.: Jersey (Saltier, pers. comm.); Belgium (Lhomme, [1946-1948]: 593); Netherlands
(Lempke, 1976: 26); Sweden (Krogerus etalii, 1971: 21); Switzerland (Muller-Rutz, 1914: 486);
U.S.S.R.: European part (Piskunov, 1981: 674).
MATERIAL EXAMINED (including 13 cf , 11 $ genitalia preparations)
Lectotype $ (mulinella), Germany (East): Dresden ('mis. Tischer') (genitalia slide no. 22493). Holotype
Cf (nigraesilvae), Germany (West): Baden-Wiirttemberg, Schwarzwald, Villengen district, Buchenberg,
22.vii.1947 (Amsel) (genitalia slide no. 827; LN).
Great Britain (England): 77 ex., ; 3 ex., Cheshire; 1 $, Salop; 2 cf, 1 $, Wiltshire; 7 cf, 3 $,
Hampshire; 1 $, 3 ex., Sussex; 6 cf , 5 Q, London; 1 $, Lincolnshire; 1 cf , Norfolk; 33 ex., Kent; vii-ix.
Channel Is.: 1 ex., Guernsey (coll. Peet, Guernsey). Spain: 1 cf, 1 $, Gerona, ix. France: 1 $,
Basses-Pyrenees; 1 $, Pyrenees-Orientales; 1 $, Alpes-Maritimes; viii, ix ? Denmark: 1 cf , NE. Jylland
(ZM); 1 9,Sjaelland(ZM);viii. Germany: 1 cf, 1 $, ; 3 cf, Berlin, vi, vii. Germany (West): 1 cf, 1 <j>,
Hamburg (coll. Jackh, Bidingen); 1 C?, Nordrhein-Westfalen (coll. Jackh, Bidingen); 1 cf , Hessen ?; viii.
Italy: 2 cf, 1 $, Liguria, ix (coll. Jackh, Bidingen). Sardinia: 2 $, ix. Sicily: 1 cf. Austria: 1 cf (NM).
Poland: 1 <j>, Szczecin (NM); 1 cf, 2 9, Wroclaw (BMNH; NM). Yugoslavia: 2 cf, 3 $, Dalmatia; ix-xi
(NM). Algeria: 1 $, Constantine province, x. Tunisia: 1 cf , 1 $, ii, x (NM). No locality data: 70 ex.
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GELECHIID MOTHS 47
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GELECHIID MOTHS
49
8
11
Figs 6-11 Wings of Mirificarma species. 6, M. montivaga (Walsingham). 7, M. scissella (Chretien). 8, M.
rhodoptera (Mann). 9, M. denotata sp. n. 10, M. maculatella (Hiibner). 11, M. pallidipulchra (Walsing-
ham).
50
L. M. PITKIN
13
14
15
16
17
Figs 12-17 Wings of Mirificarma species. 12, M. aflavdla (Amsel). 13, M. flavella (Duponchel). 14, M.
eburnella ([Denis & Schiffermiiller]). 15, 16, variation in M. ocellinella (Chretien). 17, M. fasciata sp. n.
GELECHIID MOTHS
19
20
21
22
23
Figs. 18-23 Wings of Mirificarma species. 18, M. lentiginosella (Zeller). 19, M. constricta sp. n. 20, M.
cytisella cytisella (Treitschke). 21, M. cytisella leonella Amsel. 22, M. monticolella (Rebel). 23, M.
interuptella (Hiibner).
52
L. M. PITKIN
25
28
Figs 24-29 Wings of Mirificarma species. 24, M. flavonigrella (Chretien). 25, M. burdonella (Rebel). 26,
M. cabezella (Chretien). 27, M. ulicinella (Staudinger). 28, 29, variation in M. mulinella (Zeller).
GELECHIID MOTHS
53
Fig. 30 Wings of Mirificarma mulinella (Zeller) .
valva
hind edge of
vinculum
sacculus
narrow
scjerite of
vinculum
filament
v
filament- supporting
sclerite
uncus
gnathos
process
of tegumen
tegumen
Fig. 31 Schematic diagram of the cf genitalia of Mirificarma combining some features of the different
species-groups.
54
L. M. PITKIN
36
38
39
40
44
Figs 32-45 Saccus of Mirificarma species. 32, 33, variation in M. montivaga (Walsingham). 34, M.
scissella (Chretien). 35, M. rhodoptera (Mann), typical form. 36, M. rhodoptera (Mann), small form. 37,
M. denotata sp. n. 38, M. maculatella (Hiibner). 39, M. pallidlpulchra (Walsingham). 40. M. aflavella
(Amsel). 41, 42, variation in M. flavella (Duponchel). 43, M. eburnella ([Denis & Schiffermiiller]). 44,
M. lentiginosella (Zeller). 45, M. constricta sp. n. Scale = 0-25 mm.
GELECHIID MOTHS
55
46
47
50
51
57
58
L
59
Figs 46-59 Mirificarma species. 46-56, saccus, scale = 0-25 mm. (46) M. ocellinella (Chretien); the
specimen figured is shorter than average. (47) M. fasciata sp. n. (48) M. cytisella cytisella (Treitschke).
(49) M. monticolella (Rebel). (50) M. interuptella (Hubner). (51) M. flavonigrella (Chretien). (52) M.
burdonella (Rebel). (53) M. cabezella (Chretien), Spain. (54) M. cabezella (Chretien), Morocco. (55)
M. ulicinella (Staudinger). (56) M. mulinella (Zeller). 57-59, scale = 0-25 mm. (57) left sacculus of M.
monticolella (Rebel). (58) aedeagus of M. monticolella (Rebel). (59) sacculi and hind edge of vinculum
of M. flavonigrella (Chretien).
56
L. M. PITKIN
/
/* I
\
I
',a
Figs 60-63 Genitalia of Mirificarma cf. 60, M. montivaga (Walsingham). 61, M. scissella (Chretien). 62,
M. rhodoptera (Mann), typical form. 63, aedeagus of M. rhodoptera (Mann), small form.
GELECHIID MOTHS
57
66
Figs 64-66 Genitalia of Mirificarma d". 64, M. denotata sp. n. 65, M. maculatella (Hiibner). 66, M.
pallidipulchra (Walsingham).
58
L. M. PITKIN
n
\
69
Figs 67-69 Genitalia of Mirlficarma cf. 67, M. aflavella (Amsel). 68, M. flavella (Duponchel). 69, M.
eburnella ([Denis & Schiffermuller]).
GELECHIID MOTHS
59
Figs 70-72 Genitalia of Mirificarma cT. 70, M. fasciata sp. n. 71, M. ocellinella (Chretien). 72. M.
lentiginosella (Zeller); the uncus of the figured specimen is broader than average.
60
L. M. PITKIN
Figs 73-76 Genitalia of Mirificarma cf . 73, M. constricta sp. n. 74, M. cytisella leonella Amsel. 75, M.
cytisella cytisella (Treitschke), typical form. The major difference between Figs 74 and 75 is indicated by
arrows. 76, M. monticolella (Rebel); some features which were indistinct in this photograph have been
outlined.
GELECHIID MOTHS
61
Figs 77-79 Genitalia ofMirificarma tf. 77, M. interuptella (Hiibner). 78, M.flavonigrella (Chretien). 79,
M. burdonella (Rebel).
62
L. M. PITKIN
80
81
Figs 80, 81 Genitalia of Mirificarma cf. 80, M. cabezella (Chretien). 81 , M. ulicinella (Staudinger).
GELECHIID MOTHS
63
83
84
- .
Hi
Figs 82-84 Mirificarma C?. 82, genitalia of M. mulinella (Zeller). 83, 84, eighth abdominal segment with
coremata. (83) M. ocellinella (Chretien). (84) M. intemptella (Hiibner).
64
L. M. PITKIN
88
Figs 85-88 Genitalia of Mirificarma $. 85, M. montivaga (Walsingham). 86, M. rhodoptera (Mann),
typical form. 87, M. rhodoptera (Mann), small form. 88, M. denotata sp. n. Scale = 0-1 mm and applies
to signa only.
GELECHIID MOTHS
65
91
Figs 89-91 Genitalia of Mirificarma $. 89, M. maculatella (Hiibner). 90, M. pallidipulchra (Walsing-
ham). 91, M. aflavella (Amsel). Scale = 0-1 mm and applies to signa only.
66
L. M. PITKIN
92
93
95
Figs 92-95 Genitalia of Mirificarma $. 92, M. flavella (Duponchel). 93, M. eburnella ([Denis &
Schiffermuller]). 94, 95, M. ocellinella (Chretien) and variation in the signum. Scale = 0-1 mm and
applies to signa only.
GELECHIID MOTHS
67
*V-JNJ
Figs 96-98 Genitalia of Mirificarma $. 96, M. fas data sp. n.91,M. lentiginosella (Zeller) . 98, M. cytisella
cytisella (Treitschke). Scale = 0-1 mm and applies to signa only.
L. M. PITKIN
100
101
Figs 99-103 Genitalia of Mirificarma $ . 99, M. intemptella (Hiibner). 100, M. burdonella (Rebel). 101,
M. cabezella (Chretien). 102, M. ulicindla (Staudinger). 103, M. mulinella (Zeller). Scale = 0-1 mm and
applies to signa only.
GELECHIID MOTHS
69
104
105
106
/
108 *** J
109
111
Figs 104-112 Posterior abdominal segments of Mirificarma $ . 104, M. montivaga (Walsingham). 105, M.
rhodoptera (Mann), typical form. 106, M. rhodoptera (Mann), small form. 107, M. pallidipulchra
(Walsingham). 108, M. aflavella (Amsel). 109, M. eburnella ([Denis & Schiffermiiller]). 110, M. cytisella
cytisella (Treitschke). Ill, M. intemptella (Hiibner). 112, M. burdonella (Rebel).
70
L. M. PITKIN
Index
Principal references are in bold; invalid names are in italics.
AcompsiaS, 15
aflavella 17, 18, 20, 24, 25,
26-28
albicosta 39
Anacampsinae 3
Anarsia 3
Aristoteliinae 3, 6
aristotelis 3
Aroga 3, 5
aurantiella 29, 30
Bryotropha 12
burdonella 17, 18,39,40,41,
42,44
cabezella 5, 12, 13, 17, 18, 20,
21,33,41,42-44
caminariella 43, 44
Caryocolum 3
Chelariinae 3
Chionodes 3, 6
Coleophoridae 39
constricta 12, 17, 19, 20, 21, 32,
33,42
cytisella 5, 12, 17, 21, 29, 34, 36
Deltophora 6
denotata 5, 12, 13, 16, 17,22,
23,34
Dichomeriinae 3
Dichomeris3, 15
eburnella3,5,12, 16-18,
24-26, 27, 28
fasciata 11, 15, 16, 18,24,29,
30,31,39
ferrugellaS, 27
Filatima 12
flammella 13, 27
flavella 5, 12, 16, 17, 24, 25, 26,
27,28
flavonigrella 12, 17, 39, 40, 41,
44
formosella [Denis &
Schiffermuller] 3, 27, 28
formosella Hubner 5, 27
Gelechia3,5,6,9, 13, 15
Gelechiidae3,6, 15
Gelechiinae3,5,6, 9, 15
Gelechiini 3, 6, 15
gibbosella 6,15
Gnorimoschemini 6, 15
Helina Guenee 13
Helina Robineau-Desvoidy 13
hippophaella 6
Hypatiminae 6
interrupta 38, 43
interuptella5,9, 12, 15,17, 18,
20, 21, 29, 31, 32, 38, 39, 43,44
lentiginosella 5, 11, 12, 15, 16,
18,19,29,32,33,45
leonella 34, 36, 37
Lita 3, 5
lugubrella 3
maculatella 5, 9, 13, 15-17, 19,
20, 22, 23, 29, 34
Mirificarma 3, 5, 6, 8, 9, 12, 13,
15,16
monticolella 17, 19, 37, 38
montivaga 5, 6, 9, 11-13,
15-17, 18, 19, 20, 24, 38
mulinella3,5, 12, 13, 17, 18,29,
32, 39-42, 43, 44, 45
myricariella 6
Neofaculta 5
Neofriseria 6
nigra 6, 13
nigraesilvae 44
obscurella 32
ocellinella 11-13, 15, 16, 18, 19,
22-24,29,30,31,38
Oecophoridae 5, 27
Ornativalva3, 15
Orophia5,27
pallidipulchra 5, 12, 14, 17, 20,
24, 25-28
Psoricoptera 6,15
pulverosella 18, 19
retamaeofoliella 29, 30
Rhinosia 5
rhodoptera 9, 12, 15, 16, 18, 19,
20,21,24,33,42
rhombella 6
roseella 36
rufeoformosella 27
sabinella 6
Schiffermuelleria 3, 28
scissella!2, 13, 16, 17,20,21,
33,42
scotinella 6
segetella 26, 27
senticetella 6
solutella 3
striolella 24
Syncopacma 3
Teleiodes 6
Teleiodini 6
ulicinella5,12, 13,17, 18,42,
43,44
vicinella 23
Xystophora 3
British Museum (Natural History)
Milkweed butterflies: their cladistics and biology
P. R. Ackery & R. I. Vane- Wright
The Danainae, a subfamily of the Nymphalidae, contains only some 150 species, yet aspects of
their biology have stimulated far more attention than can be justified by species numbers
alone. In recent years, an expansive literature has grown, considering aspects of their
courtship and pre-courtship behaviour, migration, larval hostplant associations, mimicry and
genetics. The popularity of danaines among biologists can certainly be attributed to this
combination, within one small group, of so many of the factors that make butterflies such an
interesting group to study. The obvious need to place this wealth of biological data within an
acceptable systematic framework provided the impetus for this volume.
Started eight years ago within the conventions of evolution by natural selection, and
Hennig's phylogenetic systematics, the book is now largely about natural history (what the
insects have and do, where they live and how they develop) and natural groups - as revealed
by a form of analysis approaching that practised by the new school of 'transformed cladistics'.
The authors have prepared a handbook that will appeal to a wide range of biologists, from
museum taxonomists to field ecologists.
1984, 448 pp (approx. ), 12 pp colour, 73 b/w plates, line and graphic illustrations, maps, extensive
bibliography. ISBN 565 00893 5
Titles to be published in Volume 48
Gelechiid moths of the genus Mirificarma.
By Linda M.Pitkin.
Macronematine caddisflies of the genus Amphipsyche (Trichoptera: Hydropsychidae)
By P. C. Barnard.
A review of the genera of In do- Pacific Encyrtidae (Hymenoptera: Chalcidoidea).
By John S. Noyes & M. Hayat
Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk
Printed in Great Britain by Henry Ling Ltd, Dorchester
6Nauj,
Bulletin of the
British Museum (Natural History)
Macronematine caddisflies of the genus
Amphipsyche (Trichoptera:
Hydropsychidae)
P. C. Barnard
Entomology series
Vol 48 No 2 23 February 1984
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology,
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^v
. /rt X*
v
I f
World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.)
1 J- 1LJ I__M
Trustees of the British Museum (Natural History), 1984
V
^. .
The Entomology series is produced under the general editorship of the
Keeper of Entomology: Laurence A. Mound
Assistant Editor: W. Gerald Tremewan
ISBN 565 06001 5
ISSN 0524-6431
British Museum (Natural History)
Cromwell Road
London SW7 5BD
Entomology series
Vol48No2pp71-130
Issued 23 February 1984
Macronematine caddisflies of the genus
Amphipsyche (Trichoptera: Hydropsychidae)
N^P|, U\<
P. C. Barnard
Department of Entomology, British Museum (Natural History), Cromwell Road, London
SW7 5BD
Contents
Synopsis 71
Introduction 71
Abbreviations of depositories 73
Acknowledgements 73
Taxonomic method 73
Classification of the Macronematini 74
Key to Old World genera of Macronematini 74
Amphipsyche McLachlan 76
Geographical distribution 78
Biology 78
Cladistic analysis 80
Check-list of Amphipsyche species 85
Key to species of Amphipsyche 85
Thepro/wta-group 86
The apicalis-group 94
The meridiana-group 101
References 127
Index 130
Synopsis
In this revision of the genus Amphipsyche McLachlan 22 species are recognized, of which two are described
as new. One new generic and ten new specific synonyms are established, and one species is transferred to
Amphipsyche from Protomacronema Ulmer. Eight lectotypes are designated. Keys are given to the Old
World genera of the tribe Macronematini and to the species of Amphipsyche. The classification of the
species is based on a cladistic analysis, and some of the evolutionary and zoogeographical implications of
the analysis are discussed.
Introduction
Amphipsyche McLachlan is an Old World genus of caddisflies having netspinning larvae that are
frequently found in fast freshwater streams throughout the Afrotropical and Oriental regions.
Some species have figured prominently in recent freshwater pollution and impoundment
studies, and at least one species is a predator on larvae of Simulium Latreille (Diptera).
Many of the recent ecological studies on tropical freshwater habitats are the result of the
pressing need for knowledge of the effects of man's activities, the most obvious of which is direct
pollution of the water by chemical or other agents. Although most Trichoptera are very sensitive
to such pollutants and tend to disappear even at low levels of contamination, there is a selective
response shown by different species of caddis. Resh & Unzicker (1975) have stressed that it is
important to be able to identify the organisms at the specific level for this kind of study. Another
important influence of man is the damming of rivers for hydroelectric or irrigation schemes.
Although the ecology of such impounded water is usually studied, the effects on the regulated
river itself are less well known and the few existing reports suggest that the natural watercourse
may be altered for a considerable distance downstream of the impoundment. The release of
Bull. Br. Mus. not. Hist. (Ent.) 48 (2): 71-130 Issued 23 February 1984
72 P. C. BARNARD
water rich in zooplankton from these dams leads to large populations of filter-feeding organisms
such as Hydropsychidae, and among these Amphipsyche has often been reported as reaching
pest proportions. However, it should be noted that the discharge of cold hypolimnial water from
dams can suppress the populations of such organisms immediately below the impoundment
(Stanford & Ward, 1981). Simulium, another filter-feeder, can also occur in large numbers in
such habitats, and various insecticides such as DDT have been used to control populations of the
S. damnosum complex, the vector of onchocerciasis. The effects of these control agents on
non-target organisms is always monitored, and because Amphipsyche occurs at similar sites it
has often figured prominently in such studies (Corbet, 1958; Statzner, 1981). Amphipsyche
scottae is also known to be a predator of Simulium (Chutter, 1968).
The identification of the organisms collected in all such freshwater studies, especially in the
tropics, is always a major problem. Scott (1975) stated that the larval stages of less than 15 per
cent of the African Trichoptera were known, and the corresponding figure for Asia must be
considerably lower. Such identification relies on the correct association of larvae and adults,
which often depends on long-term collecting programmes and rearing in the field; equally
important is the provision of reliable keys for the identification of adults. The netspinning larvae
of the Hydropsychidae are often one of the most abundant groups of macro-invertebrates in
running water, and as part of a continuing study of the subfamily Macronematinae this paper
deals with the adults of the genus Amphipsyche, in the tribe Macronematini. The species in this
genus are superficially very similar to each other, and they also resemble species oiAethaloptera
Brauer, in the Polymorphanisini (Barnard, 1980); I have frequently found these two genera
confused in collections. Ulmer's (1907) monograph of the subfamily is still useful for some
genera such as Macronema Pictet, but not for Amphipsyche; of the 22 species currently
recognized only two were known to Ulmer. Kimmins (1962; 1963) described several African
species, but new characters have been discovered in some of these.
The keys here provided to the Old World genera of Macronematini, and to the species of
Amphipsyche, are based on external characters as far as possible, but several species are known
only from males and critical examination of the genitalia is often necessary. Using a cladistic
analysis of the species of Amphipsyche the genus is divided into three main species-groups.
Some of the evolutionary and zoogeographical implications of this classification are discussed,
and it is intended to apply this approach to other genera of the Macronematinae and ultimately
to test the current generic and tribal groupings within the whole subfamily.
The methods of preparation and drawing of specimens are virtually the same as in the revision
of the Polymorphanisini (Barnard, 1980). Temporary glycerine preparations of male and female
genitalia were used for examination, and denuded wings were drawn from dry-mounted slide
preparations wherever possible.
The scale lines on the figures represent the following lengths: wings 1-0 mm; maxillary palps
0-25 mm; legs 0-5 mm; genitalia 0-25 mm. All other features illustrated have their scale indicated
on the figure. The arrows on some figures indicate features referred to in the keys or in the
species descriptions.
The nomenclature of wing veins and genitalia components follows Schmid's (1980) broadly
based study. This means that some of the names previously used in the Polymorphanisini
revision are now changed. Thus the aedeagus is here termed the phallotheca, and the gonopods
are now called the inferior appendages. The wing venation terminology is unchanged, except
that the apical forks are labelled I to V. Thus fork R 2 is now fork I, fork R 4 is fork II, fork M l is
fork III, fork M 3 is fork IV, and fork C la is fork V. These forks are the same in both the fore and
hind wing (except that fork IV never occurs in the hind wing of Trichoptera).
No attempt has been made to homologize the endothecal spines of Amphipsyche males with
those seen in some other genera. They are thus given the arbitrary names of dorsal, mid and
ventral spines, according to their level of insertion on the apex of the phallotheca. The
phallocrypt pocket may be homologous with the similar structure seen in some other families of
Trichoptera (Nielsen, 1957), but its ontogeny is unknown.
Under the heading 'Material examined' for each species are listed only the total numbers of
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 73
each sex, the countries of collection and institutions holding the material. Full collection data are
given only for type-specimens. Where there is further information on the distribution of a
species which is not apparent from the list of material examined, this is noted in the correspond-
ing 'Remarks' section.
Abbreviations of depositories
BMNH British Museum (Natural History), London, U.K.
IP Institut fur Pflanzenschutzforschung, Eberswalde, D.D.R.
IRSNB Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium
MCZ Museum of Comparative Zoology, Harvard University, U.S.A.
MNHN Museum National d'Histoire Naturelle, Paris, France
MNHU Museum fur Naturkunde der Humboldt-Universitat, Berlin, D.D.R.
MRAC Musee Royal de 1'Afrique Centrale, Tervuren, Belgium
NAC Nanjing Agricultural College, Nanjing, China
NM Naturhistorisches Museum, Vienna, Austria
RNH Rijksmuseum van Natuurlijke Historic, Leiden, The Netherlands
RSM Royal Scottish Museum, Edinburgh, U.K.
USNM National Museum of Natural History, Smithsonian Institution,
Washington D.C., U.S.A.
ZI Zoological Institute, Lund, Sweden
ZM Zoologisches Museum, Hamburg, B.R.D.
ZSI Zoological Survey of India, Calcutta, India
Acknowledgements
I am very grateful to the following for the loan of specimens: Mr R. Danielsson, ZI; Dr J.
Decelle, MRAC; Dr O. S. Flint, Jr, USNM; Dr D. C. Geijskes and Dr P. H. van Doesburg,
RNH; Dr A. Kaltenbach, NM; Dr J. Legrand, MNHN; Dr G. Marlier, IRSNB; Mr A. F.
Newton, MCZ; Prof. Dr H. Striimpel, ZM; Prof. Tian Li-xin, NAC; Frau H. Wendt, MNHU. I
also thank the following for information: Dr S. K. Ghosh, ZSI; Ake Lilliestam, Kungliga
Biblioteket, Stockholm; Dr Bo Tjeder, ZI; the Director, IP. Dr K. M. F. Scott kindly sent me
the manuscript of the Amphipsyche section of her paper on the Hydropsychidae of southern
Africa.
Taxonomic method
Although the cladistic method of classification is often taken to be equivalent to Hennig's (1966)
phylogenetic systematics, Platnick (1979) has pointed out that there is no necessary connection
between cladistics and the process of evolution: a cladogram can be constructed simply by
studying the pattern of the distribution of characters in a group of organisms. Although this
'transformed' cladistic approach has been criticized by several authors (e.g. Beatty, 1982) on the
grounds that the claimed evolutionary neutrality is actually counter-productive, Platnick argued
that cladistic methods are simply attempts to discover natural groups by analysing their
characters, which is surely the aim of taxonomy in general.
One of the difficulties with Hennig's phylogenetic method is that the taxonomist has to make a
priori decisions about the polarity of character states, and to sort them into apomorphies and
plesiomorphies on the basis of outgroup comparisons. This is a crucial step in the construction of
a phylogeny, because groups can be recognized only on the basis of synapomorphies. Inevitably,
some of these decisions on the polarity of character states are very hard to make, because the
taxonomist has to assume at least some of the evolutionary history of the group before he starts.
There is thus an element of circularity in the process, because one cannot make such
assumptions about characters used to produce a phylogeny, and then use that phylogeny to draw
independent conclusions about the evolution of the group. Platnick (1979) argued that the
'plesiomorphic' state of a character is really the more general one, in that it is found in more
74 P. C. BARNARD
groups than the 'apomorphic', or less general, state. The group possessing the more specialized
character state is therefore contained within the group showing the more general state, and this
gives rise to the nested sets and subsets which form the hierarchical classification. This is an
important concept, in that it avoids the idea that plesiomorphic and apomorphic states are
alternatives: it also clearly shows why a group based on plesiomorphies alone cannot be a natural
one because it would be recognized only by the absence of characters. Thus the production of a
cladogram does not depend on the reconstruction of the evolutionary history of the group, but
on the differentiation of more general characters from less general ones. The hierarchical
structure of the cladogram is therefore a result of the inter-nested sets of unique characters, each
delimiting a natural group.
The test of whether the taxonomist has correctly identified the level of generality of a
character is whether or not it is congruent with other characters at higher and lower levels.
Instead of making decisions about the polarity of character states, one has only to distinguish the
presence of a character from its absence, the latter being hypothesized as the more general
condition. This highlights the problem of using the loss of a character to delimit a group.
Phylogeneticists would decide that a loss character may be apomorphic by a priori outgroup
reasoning, whereas transformed cladists would discover the level of generality of the 'loss' by its
congruence with all the other characters examined. In practice, however, it is preferable to use
presence characters to recognize groups, because without ontogenetic data it is hard to
distinguish the secondary loss of a character from its absence at a more general level, unless
there is a high degree of congruence.
Having produced a cladistic classification without any assumptions of evolutionary history or
speciation mechanisms, the taxonomist is then free to use the cladogram to infer something
about the evolution of the group being studied, by hypothesizing a phylogenetic tree. Following
the cladistic analysis of Amphipsyche I therefore discuss some of the phylogenetic and
zoogeographic implications of the cladogram. The use of the transformed cladistic method and
its application in biogeography are discussed in detail by Nelson & Platnick (1981).
Classification of the Macronematini
The current classification of the subfamily Macronematinae was discussed in a previous paper
(Barnard, 1980). Of the two constituent tribes, the Polymorphanisini is almost certainly
monophyletic, despite being delimited by loss characters. The adults are recognized by the loss
of the mouthparts, and the larvae by the loss of the stridulatory organs on the head and fore legs
(Scott, 1975). However, the tribe Macronematini lacks any diagnostic characters and is probably
not monophyletic, although certain generic groups can be distinguished within it. For example,
Macrostemum Kolenati, Amphipsyche and Protomacronema Ulmer can be grouped on both
adult and larval characters (Scott, 1975), the most noticeable larval character being the raised
carina on the head. The Neotropical genus Blepharopus Kolenati probably belongs here too
(Flint & Wallace, 1980) although the carina is only poorly developed. On the other hand, the
larvae of Leptonema Guerin-Meneville and Macronema s.str. (Flint & Bueno Soria, 1982) have
no carina, but Leptonema and Macrostemum adults are often very similar superficially. More
study is needed to clarify the validity of this tribe , but the group is retained here for convenience .
Key to Old World genera of Macronematini
1 Discoidal cell present in fore wing, but sometimes very small (Fig. 1) 2
Discoidal cell absent in fore wing (Fig. 3) 5
2(1) RI in hind wing ends on R 2 +i, joined to Sc by short cross-vein (Fig. 9) 3
R\ in hind wing fuses with Sc (Fig. 8) 4
3 (2) In fore wing, base of Rs entire (Fig. 1) PSEUDOLEPTONEMA Mosely
In fore wing, base of Rs obsolete, joined to RI by cross-vein (Fig. 2)
TRICHOMACRONEMA Schmid
4 (2) Maxillary palp with second segment longer than third (Fig. 4) LEPTONEMA Guerin-Meneville
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
75
III
Figs 1-3 1, Pseudoleptonema sp. cf , fore wing; 2, Trichomacronema sp. cf , fore wing; 3, Leptopsyche
gracills McLachlan cf , fore and hind wings.
Maxillary palp with third segment longer than second (Fig. 5) MACROSTEMUMKolenati
5 (1) Fork III in both wings with stalk (Fig. 3) LEPTOPSYCHE McLachlan
Fork III in both wings sessile (Figs 6,7) 6
6 (5) cf: anal area of fore wing strongly dilated (Fig. 12); $: Sc in hind wing ends on costal margin
(Fig. 18) AMPHIPSYCHE McLachlan
Cf : anal area of fore wing not dilated (Fig. 6); $ : Sc in hind wing fuses with R } to end on /? 7 + 3
(Fig. 7) PROTOMACRONEMAUlmer
76
P. C. BARNARD
AMPHIPSYCHE McLachlan
Amphipsyche McLachlan, 1872: 68. Type-species: Amphipsyche proluta McLachlan, by monotypy.
Phanostoma Brauer, 1875: 69. Type-species: Phanostoma senegalense Brauer, by monotypy. [Synony-
mized by Martynov, 1935: 201.]
Amphipsychella Martynov, 1935: 201. Type-species: Amphipsychella extrema Martynov, by original
designation and monotypy. Syn. n.
Figs 4-9 4, Leptonema sp. , maxillary palp; 5, Macrostemum sp. , maxillary palp; 6, Protomacronema sp.
Cf, fore wing; 7, Protomacronema sp. $, hind wing; 8, Macrostemum sp. cf, hind wing; 9,
Pseudoleptonema sp. cf , hind wing.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
77
Small to medium sized species, wing length cf 8-20 mm, $ 6-15 mm, yellowish or brownish in colour,
rarely with markings on head or thorax. Antenna up to three and a half times wing length in cf , up to twice
wing length in $ ; flagellar segments numerous (75-100 in cf , 45-70 in $), always elongate. Head with two
pairs of setigerous warts in cf , hind pair indistinct, only one pair in $; genae in apicalis-group flat, with
silverish pubescence. Maxillary palp with fifth segment usually very long and secondarily articulated, but
sometimes reduced or even entirely fused with fourth segment. Spur formula basically 1.4.4, but often
reduced to 0.4.4, 0.4.3, 0.4.2, 0.3.2 or 0.2.2. Tibia and tarsus of mid leg broad and flat in $ . Wing-coupling
mechanism consists of single row of curved macrotrichia on costal margin of hind wing, enaging on anal
fold of fore wing (Fig. 10). Discoidal cell absent in fore and hind wings ('false' discoidal cell formed by
secondary fusion of R 4 and R 5 in fore wing of apicalis); median cell present in fore wing, usually absent in
hind wing (present in magnd). In fore wing R\ and Rs often sinuous near anastomosis; fork I always stalked,
fork II usually sessile, but stalked in apicalis-group. Sc in hind wing ends on costal margin, joined to RI by
cross-vein. cf fore wing with strong dilated anal area.
Cf genitalia with elongate two-segmented inferior appendages; phallocrypt pocket, associated with base
of inferior appendages, and pre-anal appendages present in proluta-group only. Phallotheca usually with
broad base, narrow stem and bulbous apex, with up to three pairs of endothecal spines. $ eighth sternite
partially divided into two sclerites.
REMARKS. Within the Macronematini, Amphipsyche seems most closely related to the African
genus Protomacronema. Both genera have a very similar wing venation, although Protomac-
ronema males do not have the dilated anal margin of the fore wing seen in Amphipsyche, and in
the female hind wing Sc fuses with R^ to end on R 2 +i, instead of ending on the costal margin. The
male genitalia are also superficially similar, Protomacronema having a pair of endothecal spines
similar to those in the African species of Amphipsyche, but a detailed study of Protomacronema
is needed in order to clarify the relationships of these two genera.
Amphipsyche and Phanostoma Brauer have always been considered as being closely related,
and have usually been separated on the spur formula. Martynov (1935: 201) synonymized them
on the grounds that the species within Amphipsyche showed such variation in the number of
spurs that the two genera were essentially the same. This was not accepted by all later authors
(e.g. Ulmer, 1951) but eventually Kimmins (1962) showed that the spur formula of A.
senegalensis (the type-species of Phanostoma) had been wrongly described, and that the
distinction between the genera could no longer be maintained. Phanostoma is available as a
subgeneric name for the meridiana-group recognized in the current study, but such a formal
subdivision of the genus does not seem necessary. Kimmins also suspected that Amphipsychella
Martynov was a synonym of Amphipsyche, and although I have seen no specimens of A.
extrema, I am confident that this synonymy is correct.
10
0.5mm .
Fig. 10 Amphipsyche berneri cf , wing coupling mechanism on costa of hind wing.
78 P. C. BARNARD
Geographical distribution
Most species of Amphipsyche are restricted to the Old World tropics. The meridiana-group has
representatives throughout the Afrotropical region, Madagascar, India and Sri Lanka, and
through mainland South East Asia to Java, Borneo and the Philippines. The apicalis-group is
restricted to S. India, Burma, Thailand, Vietnam, West Malaysia, Sumatra and Borneo, and the
proluta-group occurs only in India, China and the Amur region of the U.S.S.R. Some
zoogeographical implications of these distributions are discussed below (p. 84).
A. senegalensis is the most widespread African species, being found throughout almost the
whole of the Afrotropical region, whereas the other African species have very restricted
distributions (Fig. 119). Similarly, meridiana is a very widespread species throughout India, Sri
Lanka and South East Asia as far east as Java, although this distribution is apparently disjunct
(Fig. 104). Most of the other species in the meridiana-group, and in the other groups, have more
restricted distributions. The unique occurrence of proluta in central and northern China
northwards to the Amur region of the U.S.S.R. shows an interesting parallel with Aethaloptera
evanescens (McLachlan) in the Polymorphanisini (Barnard, 1980). There is a third species in the
Macronematinae, Macrostemum radiatum (McLachlan), with a similar distribution, although
this species extends into Siberia (like A. evanescens) and also occurs in Japan.
Biology
The first account of the immature stages of a species of Amphipsyche was by Hafiz (1937), who
described the larva and pupa of meridiana (as indicd) from material collected near Calcutta.
Ulmer (1957) gave detailed descriptions of Javan and Sumatran larvae and pupae of meridiana,
but these vary in some features from Hafiz' account. Hafiz described the larval head as being
uniformly dark brown, whereas Ulmer described (and figured) a pair of yellow flecks extending
from the eyes onto the frontoclypeus. I have examined larvae recently collected from Java and
they match Ulmer's figures of the head markings, so this feature may represent a genuine
difference between the populations in India and Indonesia. The two descriptions also vary in the
gill formula (allowing for the fact that the two authors used slightly different terminology for
some gills) but here the recently collected Javan material matched exactly Hafiz' description of
Indian specimens. Further information is needed to determine whether this species is polymor-
phic or whether the two populations are perhaps subspecifically distinct.
The larva of A. proluta was described by Lepneva (1947: redescribed, 1970). Despite the two
species being in different species-groups it is apparent that the larvae of meridiana and proluta
resemble each other very closely, the main difference being that in proluta the yellow head
markings fuse to form a continuous transverse band. The gill formula of proluta matches that of
the recently collected specimens of meridiana from Java.
The larva of the African species senegalensis was first described from Ugandan material by
Hickin (1955). Jacquemart (1957) gave a further detailed account of this species from Lake
Edward (Zaire), but it should be noted that in his description the legends (and numbers) of the
figures of the prothorax and mesothorax have been transposed, and the metathorax is figured
upside-down. Ulmer (1963) described his Egyptian larval material as curvinerve, here con-
sidered a synonym of senegalensis. Ulmer's description seems to differ slightly from those of
Hickin and Jacquemart, but he made no direct comparisons with these earlier accounts, and
without seeing material from these different areas one cannot draw any conclusions. Ulmer gave
the gill formula for his 'curvinerve'' specimens, which is quite different from those of meridiana
and proluta, but as neither Hickin nor Jacquemart described the gills of senegalensis, further
comparison is impossible. Moreover, Ulmer's specimens may have represented ulmeri Kim-
mins, and not 'curvinerve'. The pupa of senegalensis was first figured and briefly described by
Gibbs (1973), with a detailed description by Marlier (1978). Several aspects of the biology of A.
scottae have been described in papers by Chutter and Scott (see below) and a full description of
the larva appears in Scott (in press).
Although the larvae of only these few species have been described in any detail, there is
sufficient in common between them to recognize some generic characters. This has been done by
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 79
Lepneva (1970), Gibbs (1973) and Scott (1975; in press), all of whom give characters sufficient to
distinguish Amphipsyche larvae from those of other macronematine genera, especially Macros-
temum (as Macronema), Leptonema and Protomacronema . Scott (1975) has demonstrated that
the larvae of Protomacronema and Amphipsyche seem to show a close relationship between
these two genera, thus confirming the evidence suggested by the adult characters (see 'Remarks'
p. 77).
Ulmer (1957) separated the larvae of Amphipsyche and Phanostoma on the form of the hind
tarsal claw. This was described as half the length of the tarsus and pointed in Amphipsyche
meridiana, and only one-third the tarsal length and blunt in Phanostoma. This character was
later figured in 'Phanostoma curvinerve' (Ulmer, 1963). However, if Hickin's (1955) and
Jacquemart's (1957) figures of senegalensis are accurate, the claw is also half the tarsal length and
pointed in this species. It is possible that the short, blunt claw in Ulmer 's specimens is due to
excessive abrasion on a rocky substrate (which is known to affect both the anal and tarsal claws in
other species of Trichoptera).
Habitats
Larvae of Amphipsyche are generally found in fast-flowing rivers on a stony substrate. Hickin
(1955) also recorded A. senegalensis in Lake Victoria, but the larvae were near the outfall of the
Nile and were therefore still in fast water. Scott (1970) found the same species in Lake Kariba, in
a deep bay at the mouth of a stream, and Seshadri (1955) described the mass occurrence of A.
meridiana in the very rapid water near the sluice gates of a reservoir.
Chutter (1963) described the ecological requirements of A. scottae in some detail. The species
was found on the Vaal River in South Africa, immediately below the man-made Vaal Barrage.
Here the larval population dropped in winter and built up again in September-November,
presumably in direct response to the increase in zooplankton populations, although some larvae
were present all the year round. Further down the same river Chutter (1968) found that most
adults of this species were caught in January, when the larval populations were again low. The
gut contents of some larvae showed that they are apparently omnivorous, feeding on algae as
well as on insects such as Simulium larvae.
Boon (1979) discovered populations of A. meridiana below the artificial Lake Rawapening on
the River Tuntang in central Java. Many organisms have difficulty in living in such a regulated
river which is subject to sudden large changes in both water level and current speed. Parts of the
substrate of this river are formed from vesicular volcanic lava, and large numbers of meridiana
larvae live in the vesicles in the rock. Boon has suggested four advantages of this habitat: (1) the
spacing of the vesicles enforces the spacing of the larvae, both within the same species and
between the other two hydropsychid species in the same river, thus preventing overcrowding;
(2) the fairly deep vesicles give protection against predation; (3) the larvae are protected from
being dislodged during high water levels; (4) they are protected from desiccation during
low water levels. Moreover, meridiana larvae also construct very tough feeding nets, which are
more resistant to damage than those of most Hydropsychidae and also do not collapse in low
current speeds or even when exposed at low water. Boon also showed that larvae apparently
co-operate in building large communal nets, which is unusual in this family. It there-
fore seems that meridiana is a particularly adaptable species, and this may be linked
to its widespread distribution through India, South East Asia and Indonesia. The fact that
senegalensis has been found in both rivers and lakes (albeit always in fast water) suggests
that it too may be an adaptable species, possibly accounting for its widespread Afrotropical
distribution.
Corbet (1958) studied the fauna of the Victoria Nile below the Owen Falls Dam, subsequent
to the use of DDT to eliminate populations of the Simulium damnosum complex in an effort to
control onchocerciasis. Trichoptera in general are very sensitive to DDT, and the previously
large populations of A. senegalensis disappeared entirely from the treated stretch of river
immediately after the addition of the insecticide. Over a year later the populations of
senegalensis were still very small, despite the chance of recolonization from unaffected popula-
80 P. C. BARNARD
tions immediately upstream. Corbet showed that this kind of insecticidal treatment can have
long-term effects on many such macroinvertebrates as well as on the fish which rely on them for
food.
'Pest' species
Where Amphipsyche larvae have colonized the fast, zooplankton-rich water immediately below
man-made reservoirs and impoundments, either the larvae or the adults have sometimes
reached 'pest' proportions. Seshadri (1955) gave an account ofmeridiana larvae occurring below
the sluice gates of a reservoir in India. Here the adults were the problem, flying in enormous
numbers every night between September and November, swarming around the street-lights and
causing a great nuisance to people living nearby. Seshadri vividly describes how 'By about 8 P.M.
it was a remarkable sight to see these insects in their millions dashing against lamps, and
dropping to the ground so as to cause considerable annoyance to passers-by and vehicles. This
went on throughout the night and every morning, to the Town Sanitary Staff fell the task of
cleaning up the streets and removing basket loads of dead insects, especially from under the
fluorescent lamps, where they formed shallow heaps several inches thick and many square feet in
extent.' The larval nets were found encrusting the rocks for a few hundred yards downstream of
the sluice gates and at times of low water the decaying stranded larvae were 'emanating a foul
stench all over the entire locality'. Hickin (1955) described the 'peculiar sickly odour' of the dead
bodies of vast numbers of adults of senegalensis which, together with a species of Cheumatop-
syche Wallengren, had been swarming around a light at the Ripon Falls, Lake Victoria. Adults
of senegalensis, together with a mayfly, were the main insect nuisance at the lights of the Owen
Falls Dam (Uganda) according to Corbet (19580), and he also (19586) reported that larvae of
senegalensis and two Cheumatopsyche species occasionally occurred in such numbers as to
obstruct filters in the same dam.
Flight activity
The flight period of Amphipsyche species, like that of most Trichoptera, is virtually continuous
in the tropics, but more noticeably seasonal in the more temperate regions. For example, A.
meridiana adults are found in virtually every month of the year, whereas scottae adults, in South
Africa, have been captured mainly from December to March.
Corbet & Tj0nneland (1955) studied the flight activity of different species of Trichoptera
throughout the night. The general pattern was of two peaks, one at dusk and one at dawn,
although not every species exhibited both peaks of activity. A. senegalensis was exceptional in
flying throughout the night, with no recognizable peaks; this species also flies in daylight. Only
females of senegalensis were caught, which led Corbet (1966) to postulate that this species may
be parthenogenetic. However, light-trap catches often show abnormal sex ratios because of
differential attraction to light, and the material examined in the present study contains
appreciable numbers of males of this species, many caught at light. Corbet's claim therefore
seems unjustified, although facultative parthenogenesis cannot be ruled out.
Cladistic analysis
The list of characters used in the analysis is given below, grouped under broad morphological
divisions. They are all 'presence' characters (see p. 74) and the state of the absence of the
character is given in parentheses after each one. In the data matrix (Table 1) their presence is
indicated by a plus sign and their absence by a dash, and the order of characters is re-arranged to
show how the groupings were constructed to produce the cladogram (Fig. 11). Two apomorphic
loss characters could have been used in this analysis, but are unnecessary. The meridiana-group
can be recognized by the loss of the fore tibial spurs and by the hind spurs being reduced to two or
three. The spur formula of 1.4.4 in theproluta- and apicalis-groups is demonstrably plesiomor-
phic for the genus (by outgroup comparison with the other genera in the tribe) but the presence
of character 21 is sufficient to distinguish the meridiana-group, making such phylogenetic
reasoning unnecessary. The list of characters used is as follows.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 81
Head
1 Vertex with dark brown markings (no markings)
2 Genae flat with silverish pubescence (rounded with no pubescence)
3 Fifth segment of maxillary palp simple (annulated)
4 Fourth and fifth segments of maxillary palp fused (separate)
Thorax
5 Mesoscutellum with pair of dark markings (no markings)
Fore wing
6 Anal margin dilated in male (margin straight)
7 Fork II stalked (sessile)
8 'False' discoidal cell present (absent)
9 Fork I with dark marking (no marking)
10 Series of dark spots at wing apex (no spots at apex)
11 Sc-Ri cross-vein with dark marking (no marking)
12 Diagonal marking proximal to anastomosis (no marking)
Hind wing
13 Af 3+ 4-Cw la cross-vein present (absent)
Male genitalia
14 Ninth segment with lateral row of setae (only dorsal row present)
15 Phallocrypt pocket present (absent)
16 Basal segment of inferior appendage broad distally (narrow distally)
17 Basal segment of inferior appendage entirely broad (entirely narrow)
18 Inferior appendage with median setigerous projection on inner side (no setigerous projection)
19 Ventral apex of phallotheca produced (apex rounded)
20 Ventral apex of phallotheca pointed (apex rounded)
21 Phallotheca with ventral median groove meeting gonopore (no groove)
22 Eversible endotheca present (no endotheca)
23 Base of phallotheca flattened dorso-ventrally (base rounded)
24 Base of phallotheca extended into two pointed lobes (base rounded)
25 Base of phallotheca broadly triangular (base rounded)
26 Stem of phallotheca thickened in lateral view (stem narrow)
27 Ventral endothecal spines present (absent)
28 Mid endothecal spines present (absent)
29 Dorsal endothecal spines present (absent)
30 Dorsal leaf-like lobes on phallotheca (lobes absent)
31 Mid endothecal spines blunt, rod-like (spines pointed)
32 Mid endothecal spines very long and thickened (spines short and narrow)
33 Mid endothecal spines fused (spines paired)
34 Mid endothecal spines sharply up-turned (spines straight or only slightly curved)
Three species were not included in the cladistic analysis. A. bengalensis and extrema were
omitted because I was unable to examine material, and delicata because males of this species are
unknown (male genitalic characters constitute over half the characters used in the analysis). A.
bengalensis and extrema belong to the meridiana-gioup on the basis of their spur formulae (that
of bengalensis probably being wrongly quoted - see p. 112). The male genitalia of bengalensis,
which seem to have only the mid endothecal spines present, modified into blunt rods, also place
the species in this group, presumably near to sinhala. Little can be surmised about extrema as it is
known only from Martynov's figures of the female, but the highly reduced spur formula and
shortened maxillary palps would suggest that it may also be closely related to sinhala. Although
specimens of delicata have been examined, the affinities of the species are doubtful as it
possesses none of the non-genitalic characters used in the analysis; it also has the intermediate
spur formula of 0.4.4. I suspect that it belongs in the proluta-group; it cannot belong in the
apicalis-group because it lacks characters 2, 7 and 9, and its Chinese distribution makes its
inclusion in the meridiana-group unlikely, though not impossible.
82
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P. C. BARNARD
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MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
83
Incongruencies in the cladogram
Although the cladogram shown is the most parsimonious one that can be constructed from the
available data, there are a number of apparent incongruencies in the data matrix (Table 1) to
which attention is drawn. Character 11 delimits a distinct group of African species (Fig. 11) but
this wing-marking also occurs in senegalensis and exsiliens. However, its appearance in
senegalensis is not consistent, and in exsiliens it is part of the broader stripe across the
anastomosis. Character 19, the produced ventral apex of the phallotheca, delimits a large
section of the meridiana-group, and also occurs in apicalis, but the phallotheca of this species is
different in all other respects. Character 30, the presence of leaf-like lobes on the phallotheca,
occurs in both meridiana and gratiosa, but each lobe in gratiosa is distinctive in bearing a spine at
its tip. Thus each of these apparent incongruencies probably arises from the non-homology of
the character, and no real doubt is cast on the validity of the groups suggested by the cladogram.
Character 13, the M 3+4 -CMi a cross-vein in the hind wing, occurs independently in instabilis
and proluta, and can thus be considered convergent for these two species.
The remaining three incongruencies are of more interest in that they may highlight some real
difficulties. Character 14, the presence of a row of lateral setae on the ninth abdominal segment
of the male, is used to combine the proluta- and apicalis-groups. However, the character is
absent in gratiosa and distincta, yet it occurs independently in pellucida of the meridiana-group.
Only the discovery of further characters at the same level of generality can test the validity of this
9
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Fig. 11 Cladogram of Amphipsyche species.
+ African spp.
84 P. C. BARNARD
group. Character 4, the fusion of the 4th and 5th segments of the maxillary palp, delimits the
berneri-corbeti-fuscata subgroup, yet this character is also seen in instabilis. Similarly, character
3, the lack of secondary articulation on the 5th segment of the maxillary palp, appears to delimit
the 'African' subgroup of the meridiana-group (excluding senegalensis)\ however, this character
does not occur in ulmeri but is seen in magna. Again, these incongruencies may indicate
incorrect groupings in the cladogram, or else possible homoplasy ; it seems reasonable to suggest
that character 4 in instabilis and character 3 in magna have each arisen independently, as this
tendency for simplification and reduction of the maxillary palps is well known in other genera of
the Macronematinae.
Thus there is still scope for further testing of the groups hypothesized in the cladogram, and
the discovery of more characters, perhaps in the larvae, is needed to confirm or modify these
groupings. Such extra data will show whether the incongruencies arise from non-homology of a
character, from the usage of the character at the wrong level of generality, or whether
homoplasy can be demonstrated in this genus.
It will be noted that several of the individual species do not have autapomorphies indicated on
the cladogram. All of these species are easily recognized, as is demonstrated in the key to
species, but they do not have easily described unique features which will differentiate them from
all other species in the genus rather than merely from their nearest neighbour. Thus berneri and
corbeti can be distinguished from each other by the lobes of the male tenth segment, which are
broad and divergent in berneri but narrow and sub-parallel in corbeti. Neither of these character
states would distinguish one or other species from all others in the genus, and their use would
generate confusion and repetition in the cladogram.
Evolutionary and zoogeographical considerations
The geographical distributions of the main species-groups and some of their components have
been outlined earlier (p. 78). In brief, the three species-groups have noticeably different
distributions, although all three overlap to some degree. It is interesting to try and deduce
something of the evolutionary history of the genus from these data, coupled with the informa-
tion from the cladogram (Fig. 11). For these purposes the cladogram can be considered as a
phylogram, showing degrees of common ancestry (Nelson & Platnick, 1981: 171), although
branching points do not denote actual speciation events or real ancestors.
The current distribution of the genus in Africa, Madagascar, India and South East Asia
suggests that it arose when the African, Malagasy and Indian land-masses were still closely
associated, namely before the end of the Cretaceous (Smith, Hurley & Briden, 1981). Each of
these three components carried its own fragmented group of species, and the further dispersal
and evolution of the genus would have occurred when India became linked with South East Asia
(Late Eocene/Oligocene).
This kind of model seems more useful than postulating the origin of the genus in one of the
three main areas of its distribution (Africa, India, South East Asia) with subsequent long-range
dispersal to the other two. Several other macronematine genera have similar widespread
distributions, such as Polymorphanisus Walker, Aethaloptera and Macrostemum (the latter also
occurring in the New World), and their long-range dispersal seems similarly unlikely in view of
their relatively limited powers of flight and their restriction to certain freshwater habitats. Some
other genera of the subfamily are restricted to one continent; Protomacronema, which may be
the sister-group of Amphipsyche is found only in Africa, and future studies on these genera
should indicate whether or not this model is satisfactory.
The meridiana-group of Amphipsyche has representatives in Africa, Madagascar, India and
South East Asia, but it is important to note that not all the African species form a monophyletic
group. A. senegalensis was apparently an early coloniser of Africa, where it is now the most
widespread species (Fig. 119), but all the other African species (including pellucida from
Madagascar) form a monophyletic group with allopatric distributions. Each member of this
sub-group is individually sympatric with senegalensis, which tends to confirm their more distant
relationship with that species (Nelson & Platnick, 1981: 384). The apicalis- and proluta-groups
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 85
have no African representatives but both occur in India and South East Asia. It thus seems likely
that both arose from a common Indian ancestral species, and that each group later dispersed and
speciated throughout South East Asia, along with some members of the meridiana-group.
Check-list of Amphipsyche species
Although most of the species in this list are arranged according to the relationships suggested by
the cladistic analysis (Fig. 11), they are not phyletically sequenced (Wiley, 1981: 211). This is
partly because three species, bengalensis, delicata and extrema, were omitted from the clado-
gram owing to lack of suitable material; these species would be sedis mutabilis sensu Wiley.
Moreover, the cladogram is not of the simple asymmetrical 'pectinate' type which lends itself to
this sequencing convention without the proliferation of formal subgroup names.
AMPHIPSYCHE McLachlan parva Banks
Phanostoma Brauer meridiana Ulmer
Amphipsychella Martynov syn. n. nirvana Banks syn. n.
pro/ute-group vedana Banks syn. n.
pro/ufa McLachlan propinqua Ulmer syn. n.
paraproluta Hwang syn. n. indica Martynov syn. n.
bifasciata Navas tricalcarata Martynov
distincta Martynov sigmosa Navas syn. n.
delicata Banks sinhala sp. n.
ap/ca/is-group bengalensis Martynov
apicalis Banks extrema (Martynov) comb. n.
exsiliens sp. n. pellucida (Navas) comb. n.
gratiosa Navas instabilis Kimmins
petiolata Ulmer plicata (Jacquemart) syn. n.
minima Banks syn. n. ulmeri Kimmins
pubescens Kimmins syn. n. scottae Kimmins
meridiana-group fuscata Kimmins
senegalensis (Brauer) corbeti Kimmins
curvinerve (Navas) syn. n. berneri Kimmins
magna Banks
Key to species of Amphipsyche
Of necessity this key is based largely on features of the male genitalia, which are often the only reliable way
of distinguishing species in this genus; moreover, the females of nine of the species are unknown. However,
geographical distribution and external characters such as wing venation and spur formulae are used where
feasible, so that isolated female specimens can be identified as far as possible.
1 Spur on fore tibia absent
Spur on fore tibia present 16
2 (1) Four spurs on hind tibia; $ ; RI in fore wing ends on Sc (Fig. 39) (O" unknown) delicata (p. 93)
Two or three spurs on hind tibia; R^ in fore wing ends on wingmargin 3
3 (2) c?:phallotheca with three pairs of endothecal spines (Fig. 82) 4
Cf : phallotheca with only two pairs of endothecal spines or less 5
4 (3) Very large species, fore wing 15-20 mm; pair of round markings on mesoscutellum(Fig.
87) (Philippines) magna (p. 102)
Small species, fore wing 8 mm; no markings on mesoscutellum ($ unknown)
(Borneo) parva (p. 105)
5 (3) Indian or Sri Lankan species 6
African or Malagasy species
6 (5) 9 : maxillary palps very short (Fig. 116); only one or two spurs on mid tibia (cf
unknown) extrema (p. 112)
Maxillary palps unmodified; four spurs on mid tibia
7 (6) SpursO.4.2 8
Spurs 0.4. 3 or 0.4. 4 meridiana (p. 106)
86 P. C. BARNARD
8 (7) cf : endothecal spines rod-like, longer than breadth of phallotheca stem (Fig. 117) (9
unknown) (India) bengalensis (p. Ill)
Cf : endothecal spines much shorter than breadth of phallotheca stem (Fig. 98) (Sri
Lanka) sinhala (p. 105)
9 (5) Hind tibia with three spurs; cf : base of phallotheca extended into two pointed lobes
(Fig. 134) (Malagasy species) pellucida (p. 1 15)
Hind tibia with two spurs ; cf : phallotheca of other shape ( Af rotropical species) 10
10 (9) cf: endothecal spines absent (Fig. 124) senegalensis (p. 112)
Cf : at least one endothecal spine present 11
11 (10) Cross-vein present between M 3+4 and Cw la in hind wing (Figs 140, 147); cf: mid
endothecal spines fused into single structure (Fig. 145) instabilis (p. 1 17)
No such cross-vein in hind wing; cf : mid endothecal spines paired 12
12 (11) cf : fifth segment of maxillary palp fused with fourth (line of fusion visible infuscata, Fig.
167); apex of phallotheca rounded (Fig. 175) 13
Cf : fifth segment of maxillary palp distinct (sometimes reduced in length); apex of
phallotheca pointed (Fig. 159) 15
13 (12) cf : diagonal fuscous marking on fore wing (Fig. 165); base of phallotheca narrow (Fig.
169) (9 unknown) fuscata (p. 123)
Cf : no diagonal wing marking; base of phallotheca broadly triangular (Fig. 175) 14
14 (13) cf : lobes of tenth segment broad and divergent in dorso-ventral view (Fig. 180) (9
unknown) (Ghana) berneri (p. 127)
Cf: lobes of tenth segment narrow, subparallel in dorso-ventral view (Fig. 176) ($
unknown) (Uganda) corbeti (p. 125)
15 (12) Cf : mid endothecal spines gently curved dorsally (Fig. 159) (South Africa) . . scottae (p. 121)
Cf: mid endothecal spines turned abruptly dorsally (Fig. 154) (9 unknown)
(Sudan) ulmeri (p. 120)
16 (1) Genae flat with silverish pubescence; fork II stalked in fore wing (Fig. 54) 17
Genae rounded with no pubescence ; fork II sessile in fore wing (Fig . 1 2) 20
17 (16) 'False' discoidal cell in fore wing (enclosing corneous spot) (Fig. 43) (India) . . . apicalis (p. 94)
No 'false' discoidal cell 18
18 (17) cf : phallotheca with eversible endotheca (Figs 59, 60) (9 unknown) (Burma) exsiliens (p. 96)
Cf : phallotheca without eversible endotheca 19
19 (18) Fore wing with striking pattern of five dark brown spots with other paler brown
markings (Fig. 61); cf: phallotheca with large dorsal leaf-like lobes (Fig. 65)
( 9 unknown) gratiosa (p. 98)
Fore wing with only one dark brown spot in fork I (Fig. 68); cf : phallotheca with no
dorsal lobes (Fig. 72) petiolata (p. 99)
20 (16) Cross-vein present between M 3+4 and Cw la in hind wing (Fig. 12) proluta (p. 86)
No such cross-vein in hind wing 21
21 (20) Vertex of head, antennal scape and pedicel with dark brown markings (Fig. 30)
distincta (p. 89)
No markings on head ($ unknown) bifasciata (p. 89)
The pro/ufa-group
Genae rounded, with no pubescence. Fork II sessile in fore wing. Spurs usually 1.4.4 but subject to
reduction. Fifth segment of maxillary palp long and secondarily annulated. cf interior appendages with
basal segment broad, at least distally; phallocrypt pocket present; pre-anal appendages present though
small (absent in distincta). Phallotheca lacking endothecal spines. Ninth segment with two rows of setae
(dorsal and lateral) in lateral view.
India, China and U.S.S.R. (Amur region).
Amphipsyche proluta McLachlan
(Figs 12-21)
Amphipsyche proluta McLachlan, 1872: 70. Lectotype cf, U.S.S.R. (BMNH), designated by Kimmins,
1957ft: 105 [examined].
Amphipsyche paraproluta Hwang, 1957: 387. Holotype cf, CHINA: Jiangsu Prov., Nanjing, 15.viii.1956
(Hwang) (NAC) [not examined]. Syn. n.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
115
129
Figs 126-129 Amphipsyche senegalensis $. 126, wing venation; 127, mid and hind legs; 128, eighth
sternites; 129, maxillary palp.
and Ulmer (1963) (as curvinerve); the pupa was described by Gibbs (1973) and Marlier (1978).
In addition to the distribution records below, Navas (1923) also recorded this species from
Madagascar, but this is unconfirmed.
MATERIAL EXAMINED
Lectotype d" of senegalensis , Senegal: 1869 (Steindachner) (NM). Lectotype $ oi curvinerve, Egypt:
Cairo, 20.vii.1916 (Alfieri) (USNM).
79 cf, 194 $, 3 larvae, 3 pupae, Chad, Sudan, Ethiopia, Ghana, Nigeria, Cameroun, Zaire, Uganda,
Tanzania, Zambia, Malawi, Zimbabwe, South Africa (Transvaal) (BMNH, IRSNB, MRAC, RSM,
USNM, ZI).
Amphipsyche pellucida (Navas) comb. n.
(Figs 130-139; distribution, Fig. 119)
Protomacronema pelluddum Navas, 1923: 26. Holotype $, MADAGASCAR (MNHN) [examined].
Cf . Antenna 45 mm, with c. 80 segments. Fore wing 15-16 mm. Body yellowish brown; basal antennal
segments annulated with dark brown, apical segments fuscous. Fore wing very pale yellow, with faint
darker stripe from R { to M in line with M-Cu\ cross-vein. Venation as in Fig. 130. Spurs 0.4.3 (Fig. 131).
Maxillary palp 5-segmented, 5th segment not secondarily annulated, shorter than segments 1-3 combined
(Fig. 132).
$. Antenna 15 mm, with c. 60 segments. Fore wing 11-13 mm. Coloration as in cf . Fore wing with no
markings, venation as in Fig. 136. Spurs 0.4.3 (Fig. 137). Maxillary palp similar to that of cf (Fig. 139).
GENITALIA cf (Figs 133-135). Ninth segment broadly rounded laterally. Base of phallotheca narrow, with
116
P. C. BARNARD
Figs 130-135 Amphipsyche pellucida cf . 130, wing venation; 131, mid and hind legs; 132, maxillary palp;
133, genitalia, lateral view; 134, phallotheca, lateral view; 135, genitalia, ventral view.
basal corners produced into pointed lobes. Stem of phallotheca narrow, apex produced into an elongate
lobe. Only mid endothecal spines present, very short and blunt. Inferior appendage slender and sinuous,
terminal segment clearly differentiated.
GENITALIA $ (Fig. 138). Eighth sternites oval, each sclerite almost symmetrical, with all corners rounded.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
87
. \
16
Figs 12-17 Amphipsyche proluta cf . 12, wing venation; 13, legs; 14, maxillary palp; 15, genitalia, lateral
view; 16, phallotheca, lateral view; 17, genitalia, ventral view.
O". Antenna c. 30 mm, with c. 80 segments. Fore wing 11-14 mm. Body pale yellowish brown, antenna pale
yellow, narrowly annulated with brown, becoming more fuscous towards antennal apex. Fore wing very
pale yellow, with slightly darker marking around Sc-R\ cross- vein, sometimes extending further onto
anastomosis. Venation as in Fig. 12; cross-vein present between M 3+4 and Cwi a in hind wing. Spurs 1.4.4;
P. C. BARNARD
pre-apical spurs on hind tibia very short (Fig. 13). Maxillary palp 5-segmented, 5th segment secondarily
annulated, over 1-5 times length of segments 1-4 combined (Fig. 14).
$. Antenna c. 12 mm, with c. 50 segments. Fore wing 8-10 mm. General coloration as in cf , fore wing
with no dark markings. Venation as in Fig. 18; M 3+4 _CM la cross-vein in hind wing, as in cf. Spurs
1.4.4 (Fig. 19). Maxillary palp similar to that of cf, but 5th segment approximately same length as
segments 1-4 combined (Fig. 21).
GENITALIA cf (Figs 15-17). Ninth segment narrow laterally, pre-anal appendages present as small
setigerous projections on tenth segment (Fig. 15). Phallocrypt pocket relatively broad and rounded(Fig.
15). Basal segment of inferior appendage broad apically only, terminal segment partly differentiated.
Phallotheca slender with narrow base; apex truncate, with slightly pointed lobe dorsally.
GENITALIA $ (Fig. 20). Eighth sternites broad, squarish; thickened inner margins wide, extending far down
inner edges.
REMARKS. Although easily identified by the male genitalia, both sexes of this species can be
distinguished from the rest of the proluta-group by the M 3+4 _Cw la cross-vein in the hind wing.
The only other species to show this character is instabilis, in the meridiana-group.
Hwang (1957) described paraproluta as differing from proluta in the male genitalia. However,
it seems that he had only McLachlan's original description on which to base his comparison, and
both McLachlan's description and figure of proluta are very poor. McLachlan saw only dried
material, and the pointed valves, which he thought were the intermediate appendages, are
apparently the tenth segment. McLachlan's (1878: 352) redescription of the species was rather
better, with a clearer figure; here he noted the setigerous pre-anal appendage as 'a distinct
tooth'. McLachlan's type-series of three males and two females is extant in the BMNH, although
Kimmins (19576) did not mention this in his lectotype designation.
21
Figs 18-21 Amphipsyche proluta $ . 18, wing venation; 19, legs; 20, eighth sternites; 21, maxillary palp.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 89
Although Hwang's (1957) description was apparently based on one male holotype, I was lent a
male specimen from NAC labelled as paratype, which had identical data to the type. Whether or
not this specimen has any type-status, it is an important 'topotypic' specimen, and examination
of it confirmed the synonymy of paraproluta with proluta.
Three specimens examined (MNHN, MCZ) are labelled 'Hanlseon', apparently from China,
according to Navas (1914). However, this is an impossible combination of letters for a Chinese
place-name, and the label data must have been mis-copied from another, presumably hand-
written, source. I tentatively suggest that the 'Is' is a misreading of the single (handwritten) letter
'k', and the final 'n' a misreading of 'u'. This would give the more plausible transliteration of
'Hankeou', the French spelling of Han-kow, for material collected by the Frenchman, de
Guerne.
The larva of proluta was described by Lepneva (1947; 1970).
MATERIAL EXAMINED
Lectotype cf of proluta, U.S.S.R.: 'Amur Land' (Maack) (BMNH).
19 Cf, 8 <j>, U.S.S.R.: Amurskaya (2 cf, 2 $ paralectotypes of proluta), China (1 cf 'paratype' of
paraproluta) (BMNH, MCZ, MNHN, NAC).
Amphipsyche bifasciata Navas
(Figs 22-27)
Amphipsyche bifasciata Navas, 1931a: 7. Holotype cf , CHINA: 'meridionale' (Bris) (lost).
[Amphipsyche proluta McLachlan; Banks, 1940: 207; Mosely, 1942: 361. Misidentifications.]
Cf . Antennal length unknown (both specimens damaged). Fore wing 10-15 mm. Antennal segments pale
yellow, with golden brown annulations. Head, thorax and abdomen yellowish brown. Fore wing pale
yellow, shaded pale brown at apex and with darker brown stripe across anastomosis. Venation as in Fig. 22;
fork I in fore wing approximately equal in length to its stalk. Spurs 1.4.4 (Fig. 24). Maxillary palp
5-segmented, 5th segment secondarily annulated, longer than segments 1-4 combined (Fig. 23).
$. Unknown.
GENITALIA cf (Figs 25-27). Ninth segment relatively narrow laterally; pre-anal appendages present as
small setigerous projections on tenth segment. Phallocrypt pocket elongate and sac-like in lateral view,
shield-shaped in ventral view (Fig. 27). Basal segment of inferior appendage very broad, viewed laterally
and ventrally; terminal segment clearly differentiated. Phallotheca slender, apex truncate, with pair of
pointed unsclerotized lobes (superficially resembling endothecal spines); dorsally a single pointed lobe.
REMARKS. Well-marked examples of this species are easily recognized by the wing markings, but
the single male examined from Szechwan has lost virtually all these markings, and is easily
confused with proluta, hence Banks's (1940) misidentification.
According to the original description the holotype of bifasciata should be in Navas's
collection, now in Barcelona, but when this was examined in 1979 the type was apparently
missing (T. R. New, pers. comm.). However, Navas's illustration of the wing is sufficient to
identify the species; it is obvious from his figure that the type is a male, although Navas does not
mention this.
MATERIAL EXAMINED
2 cf , China (BMNH, USNM).
Amphipsyche distincta Martynov
(Figs 28-38)
Amphipsyche distincta Martynov, 1935: 196. 6 cf syntypes, INDIA: Madhya Pradesh, river at Mandla,
Nerbudda Survey (Pruthi) (lost from ZSI).
Cf. Antenna up to 25mm, with c. 80 segments. Fore wing 8-10 mm. Body pale yellowish brown;
setigerous warts on vertex of head dark greyish brown; antennal scape and pedicel with dark brown
longitudinal stripe on dorsal surface (Fig. 30); front femur dark brown. Fore wing pale yellow, no dark
markings. Venation as in Fig. 28; in fore wing fork IV with short stalk; in hind wing Sc not reaching wing
90
P. C. BARNARD
27
Figs 22-27 Amphipsyche bifasciata of. 22, wing venation; 23, maxillary palp; 24, legs; 25, genitalia,
lateral view; 26, phallotheca, lateral view; 27, genitalia, ventral view.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
91
30
31
32
Figs 28-34 Amphipsyche distincta d". 28, wing venation; 29, mid and hind tibiae; 30, head, dorsal view;
31, maxillary palp; 32, genitalia, lateral view; 33, phallotheca, lateral view; 34, genitalia, ventral view.
92
P. C. BARNARD
margin, fork V very narrow. Spurs 0.3.2 (Fig. 29). Maxillary palp 5-segmented, 5th segment secondarily
annulated, longer than segments 1-4 combined (Fig. 31).
$. Antenna up to 10 mm, with c. 50 segments. Fore wing 6-8 mm. Coloration as in cf . Venation as in
Fig. 35; in fore wing R\ ends on Sc, fork IV sessile. In hind wing Cu\ not forked, 2A absent. Spurs (Fig. 36)
and maxillary palp (Fig. 37) as in cf .
GENITALIA cf (Figs 32-34). Ninth segment broad laterally, pre-anal appendages absent. Phallocrypt pocket
rounded in lateral view (Fig. 32), broadly triangular in ventral view. Inferior appendage broad and sinuous
in ventral view, terminal segment not differentiated. Phallotheca broad basally, with narrow stem; apex
triangular with no obvious appendages or lobes.
GENITALIA 9 (Fig. 38). Eighth sternites broad, with squarish corners; thickened inner margins extending
barely half length of sclerites.
REMARKS. This aptly named species is so distinctive that it is easily recognized; no other species
has markings on the head and antennae (the only other species with any body markings is
magna, with a pair of spots on the mesoscutellum).
Banks (1939) redescribed distincta, drawing attention to the dark front femur in both sexes.
Martynov had described all the legs as being pale, but since all six syntypes are apparently lost
(Ghosh, in lift. ) this character cannot be checked on Martynov's material. Banks also stated that
fresh specimens were 'plainly greenish'.
Martynov listed as the syntypes '4 cf ' followed by '2 cf ' with identical data; one of these may
y
Figs 35-38 Amphipsyche distincta $ . 35, wing venation; 36, mid and hind tibiae; 37, maxillary palp; 38,
eighth sternites.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
93
be an error for '9'- No descriptions of the female are given in his text, but this is also true of his
description ofindica (= meridiana), where both males and females make up the type-series.
A neotype designation does not seem necessary for such an easily recognizable species.
MATERIAL EXAMINED
19 cf , 28 9, India (MCZ, USNM).
Amphipsyche delicata Banks
(Figs 39-42)
Amphipsyche delicata Banks, 1939: 58. LECTOTYPE $ , CHINA (MCZ), here designated [examined].
Cf . Unknown.
$. Antenna over 12 mm, with more than 50 segments (all specimens damaged). Fore wing 6-8 mm.
Antennal segments pale yellow, slightly annulated with brown. Head, thorax and abdomen yellowish
brown. Fore wing very pale yellow with no markings, venation as in Fig. 39. RI in fore wing ends on Sc.
Spurs 0.4.4 (Fig. 40). Maxillary palp 5-segmented, 5th segment secondarily annulated, longer than
segments 1-4 combined (Fig. 41).
Figs 39-42 Amphipsyche delicata <j>. 39, wing venation; 40, mid and hind tibiae; 41, maxillary palp; 42,
eighth sternites.
94 p. c. BARNARD
GENITALIA $ (Fig. 42). Eighth sternites rounded, bluntly pointed posteriorly; thickened inner edge
relatively narrow.
REMARKS. The relationships of this species are in some doubt, mainly because no males have yet
been discovered. Banks (1939) compared it with minima (= petiolata), from which it differs in
venation, and with distincta, which he distinguished by the dark fore femora. It seems to be most
closely related to distincta, with which it shares the venational feature of /?! ending on Sc in the
fore wing; this may be a synapomorphy for this pair of species. However, the placing of this
species in the proluta-group is complicated by its unique spur formula of 0.4.4. It cannot belong
in the apicalis-group because of the sessile fork II in the fore wing, and its Chinese distribution
and unmodified maxillary palps make its inclusion in the meridiana-group unlikely.
The specimen here designated as lectotype was labelled 'type' by Banks, but not so published.
MATERIAL EXAMINED
Lectotype $, China: Hainan Tao I., Chung Kon, 18.vii.1935 (Gressitt) (type no. 23470, MCZ).
6 $ (paralectotypes), China (MCZ).
The apicalis-group
Genae flat, with silverish pubescence. Fore wing with fork II stalked, dark marking in fork I. Spurs always
1.4.4. Fifth segment of maxillary palp long and secondarily annulated (except $ apicalis). cf inferior
appendages slender; phallocrypt pocket and pre-anal appendages absent. Phallotheca lacking endothecal
spines. Ninth segment with two rows of setae as mproluta-group.
India, Burma, Thailand, Vietnam, West Malaysia, Sumatra and Borneo (Sarawak).
Amphipsyche apicalis Banks
(Figs 43-53)
Amphipsyche apicalis Banks, 1939: 56. LECTOTYPE cf , INDIA (MCZ), here designated [examined].
Cf . Antenna over 20 mm, with more than 50 segments (broken in both specimens examined). Fore wing
12-13 mm. Antennal segments pale golden brown, becoming more fuscous towards apex, slightly
annulated with brown. Head, thorax and abdomen yellowish brown. Fore wing golden yellow, with dark
brown spot in fork I, glabrous brown streaks at wing apex proximal to dark spot and across anastomosis
(Fig. 43). In fore wing 'false' discoidal cell enclosing corneous spot at base of R 4+5 , fork I with short stalk.
R 2 +3 fused in hind wing. Spurs 1.4.4, pre-apical spurs on hind tibia short (Fig. 44). Maxillary palp
5-segmented, 5th segment secondarily annulated, longer than segments 1-4 combined (Fig. 45).
$ . Antenna over 12 mm, with more than 50 segments (broken in both specimens examined). Fore wing
9-10 mm. Body coloration as in cf . Fore wing yellowish brown with pale brown spot on wing margin in fork
I and another at anterior end of anastomosis (Fig. 50). Fore wing with 'false' discoidal cell as in cf , but not
so clearly defined. Spurs 1.4.4, one pre-apical spur on hind tibia very small (Fig. 51). Maxillary palp
5-segmented, 5th segment not annulated, approximately equal in length to segments 1 and 2 combined
(Fig. 52).
GENITALIA cf (Figs. 46-49). Ninth segment with enlarged rounded side-pieces. Phallotheca with slender
stem, enlarged apically to form two ventrally directed subtriangular processes; apical process with fine
teeth at ventral apex, and hingeing dorsally to form a simple eversible endotheca (Figs 48, 49). Inferior
appendage with pronounced setigerous projection mid-dorsally; terminal segment not clearly differenti-
ated.
GENITALIA $ (Fig. 53). Eighth sternites with sharp indentation in posterior edge.
REMARKS. Superficially this species is very similar to exsiliens and petiolata, but is easily
distinguished by the characteristic shape of the male phallotheca (Fig. 48).
Banks (1939) was unsure whether the two females from Coimbatore belonged to this species,
but after examination of his material there seems little doubt that they are correctly placed here.
Banks apparently overlooked the 'false' discoidal cell in the females (which is admittedly less
obvious than in the males), and he also did not notice the very small pre-apical spur on the hind
tibia. The faint wing markings in the female are a reduced form of the pronounced male pattern;
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
95
Figs 43-49 Amphipsyche apicalis cf . 43, wing venation; 44, mid and hind tibiae; 45, maxillary palp; 46,
genitalia, lateral view; 47, genitalia, ventral view; 48, phallotheca, lateral view; 49, phallotheca with
endotheca everted.
96
P. C. BARNARD
Figs 50-53 Amphipsyche apicalis 9 . 50, wing venation; 51, mid and hind tibiae; 52, maxillary palp; 53,
eighth sternites.
this is also seen \npetiolata, and probably also in the closely related gratiosa and exsiliens, but the
females of the last two species have yet to be discovered.
The specimen designated as lectotype was labelled 'type' by Banks, but not so published.
MATERIAL EXAMINED
Lectotype Cf , India: Mysore, Shimoga, R. Tunga, 1865' [560 m], at light, lO.vi. [not iv as stated by
Banks] [year unknown] (Nathan) (type no. 22677, MCZ).
1 Cf (paralectotype), 2 <j>, India (MCZ).
Amphipsyche exsiliens sp. n.
(Figs 54-60)
Cf. Antennal length unknown (broken in all specimens). Fore wing 12-14 mm. Body yellowish brown,
back of head and dorsal surface of thorax brown; antennal segments pale yellow, narrowly annulated with
brown. Fore wing pale yellow with brown spot in fork I and another centred on Sc-Ri cross-vein; pale
brown stripe across anastomosis and very pale shading at wing apex. Venation as in Fig. 54. Spurs 1.4.4
(Fig. 55). Maxillary palp 5-segmented, 5th segment secondarily annulated, longer than segments 1-4
combined (Fig. 56).
$. Unknown.
GENITALIA cf (Figs 57-60). Ninth segment broad laterally. Base of phallotheca broadly triangular, stem
slender with triangular apex. Eversible sac-like endotheca present, connective membrane bearing many
small spines (Fig. 60). Inferior appendage slender, terminal segment scarcely differentiated.
REMARKS. This species is easily distinguished from the other members of the apicalis-group by
the rounded sac-like endotheca, which is more mobile than that of apicalis. In the latter species it
hinges through only a few degrees, but in exsiliens it can be invaginated almost entirely inside the
phallotheca apex, or hinged through almost 180 to lie dorsal to the apex (Figs 59, 60).
MATERIAL EXAMINED
Holotype cf , Burma: Tenasserim Valley (Doherty) (BMNH).
Paratypes. 2 cf , data as holotype (BMNH).
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
97
Figs 54-60 Amphipsyche exsiliens cf . 54, wing venation; 55, legs; 56, maxillary palp; 57, genitalia, lateral
view; 58, genitalia, ventral view; 59, phallotheca, lateral view; 60, phallotheca with endotheca everted.
98
P. C. BARNARD
Amphipsyche gratiosa Navas
(Figs 61-67)
Amphipsyche gratiosa Navas, 1922: 62. Holotype cf , VIETNAM: Tonkin, Hag Song, vii.1918 (lost).
C?. Antenna 25 mm, with c. 85 segments. Fore wing 10-12 mm. Body yellowish brown, antennal segments
narrowly annulated with brown. Fore wing pale yellow with striking pattern; pale brown streak across
whole width of wing proximal to anastomosis; five dark brown spots in apical area, partially linked to pale
brown areas in apical forks; venation as in Fig. 61. Spurs 1.4.4 (Fig. 62). Maxillary palp 5-segmented, 5th
segment secondarily annulated, much longer than segments 1-4 combined (Fig. 63).
. Unknown.
63
66
67
Figs 61-67 Amphipsyche gratiosa cf. 61, wing venation; 62, legs; 63, maxillary palp; 64, genitalia, lateral
view; 65, phallotheca, lateral view; 66, phallotheca, dorsal view; 67, genitalia, ventral view.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 99
GENITALIA cf (Figs 64-67). Ninth segment relatively narrow laterally. Base of phallotheca elongate,
flattened; stem narrow; apex with pair of leaf-like lobes, each bearing single spine on inner surface; pointed
dorsal lobe proximal to these apical lobes. Inferior appendage slender; setigerous projection midway on
inner surface in ventral view; terminal segment not differentiated.
REMARKS. This species is easily recognized by its prominent wing pattern. The unusual genitalia
are also diagnostic, and suggest no close affinities with the other species in the apicalis-group. I
have assumed that the dorsal lobes are not modified endothecal spines, despite their superficial
similarity to those structures, because no other species in the genus has spines with lobate bases.
There is certainly a close superficial similarity between the phallotheca of this species and that of
meridiana for example, although all other critical characters of this species definitely place it in
the apicalis-group. One male examined has a 'false' discoidal cell in the left fore wing only, a
character otherwise seen only in apicalis.
The type of this species should be in the Navas collection (now in Barcelona) but is apparently
missing (T. R. New, pers. comm.); however the species is easily recognizable from Navas's
figure. In addition to the distribution records below, the type was collected in Vietnam.
MATERIAL EXAMINED
4 cf , Burma, Thailand (BMNH).
Amphipsyche petiolata Ulmer
(Figs 68-77)
[Amphipsyche proluta McLachlan; Ulmer, 1910: 55; 1913: 79. Misidentifications.]
Amphipsyche petiolata Ulmer, 1930: 434. Lectotype $ [listed as 'holotype' by Weidner, 1964: 67], JAVA
(ZM), designated by Ulmer, 1951: 197 [examined].
Amphipsyche minima Banks, 1931: 395. LECTOTYPE $, WEST MALAYSIA (BMNH), here designated
[examined]. Syn. n.
Amphipsyche pubescens Kimmins, 1955: 387. Holotype cf, BORNEO (BMNH) [examined]. Syn. n.
Cf . Antenna c. 35 mm, with c. 80 segments. Fore wing 9-11 mm. Body yellowish brown; posterior part of
vertex and dorsal surface of mesothorax brown. Antenna pale yellow, annulated with brown, segments
becoming more fuscous towards apex. Fore wing pale golden yellow, with pale brown apex, brownish
stripe across anastomosis and dark brown spot in fork I; venation as in Fig. 68. Spurs 1.4.4 (Fig. 69).
Maxillary palp 5-segmented, 5th segment secondarily annulated, about three times length of segments 1-4
combined (Fig. 70).
$ . Antenna 10 mm, with c. 45 segments. Fore wing 7-8 mm. General coloration as in cf . Fore wing very
pale yellow, pale brown stripe across anastomosis, sometimes slight brown marking in fork I; venation as in
Fig. 74. Spurs 1.4.4 (Fig. 75). Maxillary palp 5-segmented, 5th segment secondarily annulated, about twice
length of segments 1-4 combined (Fig. 77).
GENITALIA cf (Figs 71-73). Ninth segment broadly rounded laterally. Base of phallotheca narrow,
rectangular, narrow stem abruptly right-angled. Tip of phallotheca globose with bifurcate membranous
process arising from apical depression, abruptly up-turned at apex (Fig. 72). Inferior appendage narrow
with setigerous median projection on inner surface, terminal segment not differentiated.
GENITALIA 9 (Fig. 76). Eighth sternites narrow, inner thickened edges slightly incurved; posterior margin
produced as rounded point.
REMARKS. The male of this species can be distinguished from others in the apicalis-group by the
form of the phallotheca. The apical rod-like process superficially resembles an endothecal spine
but it is a membranous median structure. Within this species-group the only other known female
is that of apicalis, which has differently shaped eighth sternites and a short apical segment of the
maxillary palp.
I have taken Ulmer's subsequent (1951) listing of the Javan syntype as 'type' as a lectotype
designation. Weidner (1964) listed this specimen as the holotype, but this is incorrect as Ulmer's
original description was based on three syntypes.
It is not clear from Banks's (1931) description of minima how many specimens constituted the
type-series. Of the two extant syntypes (with identical data, and both labelled 'type' by Banks)
100
P. C. BARNARD
70
72
Figs 68-73 Amphipsyche petiolata cf . 68, wing venation; 69, legs; 70, maxillary palp; 71 , genitalia, lateral
view; 72, phallotheca, lateral view; 73, genitalia, ventral view.
the MCZ specimen was labelled 'paratype' by H. H. Ross in 1965. 1 have therefore designated
the BMNH syntype as lectotype. Ulmer (1951) remarked on the similarity of minima to
petiolata.
Mosely identified the type-material of pubescens as petiolata, but Kimmins decided that it
represented a distinct species on the grounds that Ulmer had not mentioned an apical wing spot
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
-I
101
Figs 74-77 Amphipsyche petiolata $ . 74. wing venation; 75, legs; 76, eighth sternites; 77, maxillary palp.
in the description of petiolata. However, Ulmer's species was described from females only, and
the wing markings are very faint in this sex.
In addition to the distribution records below, Ulmer (1930) also recorded this species from
Sumatra.
MATERIAL EXAMINED
Lectotype $ of petiolata, Java: Wonosobo, iv.1909 (Jacobson) (ZM). Lectotype $ of minima, West
Malaysia: Kedah, nr Jitra, catchment area, 9.iv.l928 (Pendlebury) (BMNH). Holotype d" oipubescens,
Borneo: Sarawak, foot of Mt Dulit, junction of Rivers Tinjar and Lejok, 20.viii.1932 (Hobby & Moore)
(BMNH).
3 cf, 5 $, West Malaysia (1 $ paralectotype of minima); Borneo: Sarawak (1 cf, 2 $ paratypes of
pubescens) (BMNH, MCZ).
The meridiana-group
Genae rounded, with no pubescence. Fork II sessile in fore wing. spurs on fore leg; mid spurs sometimes
reduced to 3 or 2; hind spurs always reduced to 3 or 2. Fifth segment of maxillary palp often reduced, or
fused with 4th segment, c? inferior appendages slender; phallocrypt pocket and pre-anal appendages
absent. Phallotheca with up to three pairs of endothecal spines. Ninth segment with single (dorsal) row of
setae in lateral view.
Africa, Madagascar, India, Sri Lanka, Nepal, Cambodia, West Malaysia, Sumatra, Java,
Borneo, Philippines.
102
P. C. BARNARD
Amphipsyche magna Banks
(Figs 78-88)
Amphipsyche magna Banks, 1939: 58. Holotype cf , PHILIPPINES (MCZ) [examined].
Cf (holotype only). Antennae missing, described by Banks (1939) as 'pale, tips of joints dark'. Fore wing
20mm. Head, thorax and abdomen pale yellowish brown, mesoscutellum with two round dark brown
Figs 78-83 Amphipsyche magna cf . 78, wing venation; 79, mid and hind tibiae; 80, maxillary palp; 81.
genitalia, lateral view; 82, phallotheca, lateral view; 83, genitalia, ventral view.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
103
87
Figs 84-88 Amphlpsyche magna. 84. 9 wing venation; 85, $ mid tibia, 86, $ maxillary palp; 87,
thorax, dorsal view; 88, $ eighth sternites.
markings (Fig. 87). Fore wing elongate, yellowish brown with no markings. Venation as in Fig. 78; closed
median cell in hind wing formed by M 2 -M 3+4 cross-vein. Spurs 0.4.2 (Fig. 79), not 1 .4.2 as stated by Banks.
Maxillary palp 5-segmented, 5th segment short, not secondarily annulated (Fig. 80).
9 (single example). Antennal length unknown (specimen damaged). Fore wing 15 mm. Coloration as in
Cf, with similar round markings on mesoscutellum. Basal antennal segments pale yellow, narrowly
annulated with brown. Venation as in Fig. 84; closed median cell in hind wing as in cf Spurs 0.4. [? 2] (hind
legs missing) (Fig. 85). Maxillary palp 5-segmented, 5th segment shorter than in cf (Fig. 86).
GENITALIA cf (Figs 81-83). Ninth segment broadly rounded laterally. Base of phallotheca strongly
flattened dorso-ventrally, apex rounded. Three pairs of endothecal spines present; dorsal pair directed
ventrally, mid and ventral pairs curved dorsally. Inferior appendage thin and strongly sinuous; terminal
segment moderately clearly differentiated.
GENITALIA $ (Fig. 88). Eighth sternites subrectangular, much longer than broad; inner thickened margins
broad.
104
P. C. BARNARD
REMARKS. Despite the unlikely sounding combination of names, it seems that magna, the largest
species in the genus, andparva, one of the smallest, are sister species. They are, of course, easily
separable by the great disparity in size, and magna is particularly easy to recognize by the
mesoscutellar markings and hind wing median cell in both sexes. Both magna andparva have the
full complement of three pairs of endothecal spines and both have the unusually shaped
phallothecal base, which is elongate and strongly flattened in lateral view.
This is the first description of the female of magna. The two striking external characters of the
species, the mesoscutellar markings and the hind wing median cell, are the same in each sex, as is
the reduced condition of the maxillary palps.
MATERIAL EXAMINED
Holotype cf , Philippines: Luzon, Del Carmen, 15. xi. 1927 (Uichanco) (type no. 23471, MCZ).
1 $, Philippines: Luzon (USNM).
91
92
Figs 89-93 Amphipsyche parva c?. 89, wing venation; 90, mid and hind tibiae; 91, genitalia, lateral view;
92, phallotheca, lateral view; 93, genitalia, ventral view.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 105
Amphipsyche parva Banks
(Figs 89-93)
Amphipsyche parva Banks, 1920: 354. Holotype d", BORNEO (MCZ) [examined].
Cf (holotype only). Antenna c. 25 mm with c. 80 segments. Fore wing 8 mm. Antennal segments pale
yellow, slightly annulated with brown. Head, thorax and abdomen yellowish brown. Fore wing very pale
yellow, almost colourless, with no markings. Venation as in Fig. 89: RI in fore wing strongly sinuous, fork I
in hind wing clearly stalked. Spurs 0.4.2 (Fig. 90). Maxillary palps missing.
$. Unknown.
GENITALIA cf (Figs 91-93). Ninth segment only slightly broadened laterally. Base of phallotheca strongly
flattened dorso-ventrally, apex rounded. Three pairs of endothecal spines present; dorsal pair short,
curved latero-ventrally; mid pair strongly curved dorsally; ventral pair long and almost straight, slightly
directed dorsally. Inferior appendage thin and sinuous, terminal segment scarcely differentiated.
REMARKS. A. parva and magna are the only two species in the genus to possess three pairs of
endothecal spines. A. parva is easily distinguished from magna (its sister species) by its small size
and lack of thoracic markings. Little can be surmised about the maxillary palps of this species
(which are missing in the holotype): although these are often reduced in the meridiana-group,
this is not invariably the case, and the shortened apical segment in the sister species magna may
be a unique character within this species-pair.
MATERIAL EXAMINED
Holotype cf , Borneo: Mindai, vi.1882 (Grabowsky) (type no. 10886, MCZ).
Amphipsyche sinhala sp. n.
(Figs 94-103)
Cf . Antenna c. 35 mm, with c. 85 segments. Fore wing 10-12 mm. Body pale yellowish brown; basal
antennal segments narrowly annulated with pale brown, apical segments fuscous. Fore wing very pale
yellow with no markings; venation as in Fig. 94. Spurs 0.4.2 (Fig. 95). Maxillary palp 5-segmented; 5th
segment secondarily annulated, approximately equal in length to segments 1-4 combined (Fig. 96).
$. Antenna c. 14 mm, with c. 65 segments. Fore wing 7-8 mm. Coloration as in cf . Venation as in Fig.
100. Spurs 0.4.2 (Fig. 101). Maxillary palp as in cf , but 5th segment slightly shorter than segments 1-4
combined (Fig. 103).
GENITALIA cf (Figs 97-99). Ninth segment broad laterally. Phallotheca with moderately narrow stem,
broadly truncate at apex. Mid endothecal spines very short, rod-like (Fig. 98); dorsal and ventral
endothecal spines absent. Inferior appendage moderately narrow, slightly sinuous; terminal segment
clearly differentiated.
GENITALIA $ (Fig. 102). Eighth sternites narrow, outer borders strongly sloping, posterior border broadly
pointed.
REMARKS. A. sinhala is apparently restricted to Sri Lanka, and the only other species reported
from that country is meridiana. They can be separated easily on genitalic differences in both
sexes, as well as on the spur formula; meridiana always has at least three spurs on the hind tibia in
both sexes. Moreover, there is little overlap in size, sinhala being a noticeably small species. A.
sinhala also resembles bengalensis, but is distinguished by the much shorter endothecal spines.
MATERIAL EXAMINED
Holotype cf , Sri Lanka: Panamure, 15-21. x. 1970 (Flint) (USNM).
Paratypes. Sri Lanka: 16 cf , 36 $ , data as holotype (all in USNM except 2 cf , 2 $ in BMNH) ; 2 cf , 18 $ ,
Sella Kataragama, Menik Ganga, 24.x. 1970 (Flint) (USNM).
106
P. C. BARNARD
Figs 94-99 Amphipsyche sinhala cf . 94, wing venation; 95, mid and hind tibiae; 96, maxillary palp; 97,
genitalia, lateral view; 98, phallotheca, lateral view; 99, genitalia, ventral view.
Amphipsyche meridiana Ulmer
(Figs 104-1 14)
Amphipsyche meridiana Ulmer, 1909: 134. LECTOTYPE $ , JAVA (RNH), here designated [examined].
[Phanostoma sp. Betten, 1909: 234.]
Amphipsyche nirvana Banks, 1913: 236. Holotype cf , INDIA (MCZ) [examined]. Syn. n.
Amphipsyche vedana Banks, 1913: 235. Holotype 9, INDIA (MCZ) [examined]. Syn. n.
Ampsipsyche [sic] propinqua Ulmer, 1927: 177. LECTOTYPE cf, CAMBODIA (MNHU), here designated
[examined]. Syn. n.
[Amphipsyche proluta McLachlan; Navas, 1931ft: 91; 1934: 227. Misidentifications.]
Amphipsyche indica Martynov, 1935: 199. 8 syntypes, INDIA: 1 d", Bihar, Mokameh, at light; 1 cf , Bihar,
Dinapore, at light (Annandale); 2 CT, 2 $>, Bihar, Pusa, 5-10.xi.1915 (Gravely); 2 cf, E. Bengal,
Damukdia Ghat, at light on board steamer, 30. vi. 1908 (2 syntypes in ZSI, the other 6 lost) [not
examined]. Syn. n.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
107
Figs 100-103 Amphlpsyche sinhala $ . 100, wing venation; 101, mid and hind legs; 102, eighth sternites;
103, maxillary palp.
Amphipsyche tricalcarata Martynov, 1935: 197. Holotype $, INDIA: Orissa, Puri district, Bhubaneswar,
4-6.xi.1912 (Gravely) (lost from ZSI). [Synonymized with indica by Schmid, 1958: 107.]
Amphipsyche sigmosa Navas, 1935: 105. LECTOTYPE d", INDIA (MNHN), here designated [examined].
Syn. n.
C?. Antenna c. 40 mm with up to 100 segments. Fore wing 13-15 mm. Body pale yellowish brown, antennal
segments pale golden brown. Fore wing pale golden yellow, sometimes with pale brown marking behind
R\-Rs cross-vein. Venation as in Fig. 105; RI in fore wing strongly sinuous both proximal and distal to
anastomosis. Spurs 0.4.3 or 0.4.4 (pre-apical spurs on hind tibia always very small) (Fig. 106). Maxillary
palp 5-segmented, 5th segment secondarily annulated, approximately equal in length to segments 1-4
combined (Fig. 107).
$. Antenna c. 18 mm, with c. 70 segments. Fore wing 8-12 mm. Coloration as in cf ; dark marking on
fore wing always absent. Venation as in Fig. Ill; fork IV in fore wing occasionally stalked. Spurs (Fig. 112)
and maxillary palp (Fig. 114) as in cf .
GENITALIA cf (Figs 104, 108-109). Ninth segment moderately broad laterally. Phallotheca elongate, with
narrow stem. Dorsal endothecal spines absent, in their place a pair of semi-membranous leaf-like lobes,
variable in shape (Fig. 109). Mid and ventral endothecal spines short, varying in relative length; ventral
pair occasionally lost. Inferior appendage narrow and sinuous, terminal segment moderately well
differentiated.
GENITALIA 9 (Fig. 113). Eighth sternites broad and squarish with broadly rounded corners.
108
P. C. BARNARD
-0
70
Fig. 104 Variation in cf genitalia of Amphipsyche meridiana throughout its range.
REMARKS. A. meridiana is the most common and widespread of the Asian species in the genus.
The male is easily recognized by the dorsal leaf-like lobes on the phallotheca, but the female may
be confused with sinhala unless it is examined closely; the hind pre-apical spurs are always
extremely small, thus the spur formula may be taken erroneously as 0.4.2.
The large number of synonyms of this species is partly a result of its morphological variability
over a wide geographical range. Banks (1913) said that Betten's (1909) "Phanostoma sp.' was the
same as, or very similar to, nirvana, and Martynov (1935) said that it was identical with his new
species indica. Banks (1939) apparently regarded indica as a synonym of nirvana, as he placed
the name indica in parentheses after nirvana. Meanwhile, Ulmer (1927) had compared
propinqua with nirvana (but not with his own species meridiana) but later (1951) noted that
nirvana, propinqua and meridiana were all very similar. Thus the Indian species were long
considered as being synonymous, but the synonymy with meridiana was not suspected, partly
because of the geographical separation (meridiana being described from Java) and partly
because the male of meridiana was not described until 1951.
The extent of variation in the male phallotheca is shown semi-diagrammatically in Fig. 104.
The mid endothecal spines are moderately consistent in size and form throughout the whole
range, although in the single known male from Bombay they are very short and broad, and bent
abruptly upwards. However, the ventral endothecal spines vary greatly, and there seems to be a
correlation with distribution, such that in the most eastern specimens they are much longer than
the mid spines, whereas in the western (Indian) populations they are usually much shorter, and
even lost in specimens from west India (and also occasionally Sri Lanka). There is in fact a
discontinuity in the distribution of this species, with no specimens known from countries
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
109
Figs 105-110 Amphipsyche meridiana d" 105, wing venation; 106, mid and hind tibiae; 107, maxillary
palp; 108, genitalia, lateral view; 109, phallotheca, lateral view; 110, genitalia, ventral view.
between India and Cambodia. There may be justification for considering the two populations as
subspecifically distinct, in which case the Indian subspecies would have to be named meridiana
nirvana, with the nominate subspecies in South East Asia, but I do not propose such a formal
division at present.
110
P. C. BARNARD
113
Figs 111-114 Amphipsyche meridiana
sternites; 114, maxillary palp.
Ill, wing venation; 112, mid and hind tibiae; 113, eighth
I do not believe that the 'paratype' of meridiana mentioned by Weidner (1964) has any
type-status. Although from the type-locality, it bears a printed label with the date 'Dec. 1908'. A
'Paratype' label has also been attached, bearing the hand-written date '8.1907', not in Ulmer's
hand, to conform to the published type-data. However, the other labels do not match those on
the two remaining syntypes in Leiden; of the original three syntypes mentioned by Ulmer in the
RNH, one has apparently been lost (Geijskes, in lift.).
Of the three syntypes oipropinqua described by Ulmer (1927) I have examined the two males
in MNHU and designate as lectotype the one labelled 'type' by Ulmer. The third male syntype,
now a paralectotype (IP, not examined), lacks its abdomen (Director, IP, in lift.).
I was informed by Ghosh (in litt.) of the apparent loss of six syntypes of indica, and of the
holotype of tricalcarata from the ZSI. The female syntypes oisigmosa (from Khandala) are also
apparently lost, so the sole remaining male syntype in the MNHN is here designated as
lectotype.
The larva of this species was described by Hafiz (1937, as indica), and by Ulmer (1957). There
are some differences between these two descriptions, both in the gill formulae and in the head
markings. Specimens from Java that I have examined differ slightly in gill counts from Ulmer's
description, even though his material was also from Java (and Sumatra). Some aspects of the life
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
111
history are described by Seshadri (1955) and Boon (1979) - see p. 79. Some specimens in the
BMNH, received via the Commonwealth Institute of Entomology, were captured on paddy-
fields in India, but any economic significance of this is unknown.
MATERIAL EXAMINED
Lectotype $ of meridiana, Java: Batavia, viii.1907 (Jacobson) (RNH). Holotype cf of nirvana, India:
Bengal, Pusa, at light, 23.iii.1908 (type no. 11755, MCZ). Holotype $ of vedana, India: Bengal, Pusa,
15. ix. 1907 (type no. 11757, MCZ). Lectotype cf of propinqua, Cambodia: Mekong, Pnom-Pech, i.1914
(Friederichs) (MNHU). Lectotype cf of sigmosa, India: Bombay, Lonawla [= Lonavla, = Lonauli],
9.x. 1934 (Benavent) (MNHN).
229 Cf, 332 9, c. 75 larvae, 2 pupae, India, Sri Lanka, Nepal, Cambodia (1 cf paralectotype of
propinqua), West Malaysia, Sumatra, Java (1 9 paralectotype and 1 $ as 'paratype' of meridiana; see
'Remarks' above) (BMNH, MCZ, MNHU, RNH, USNM, ZM).
Amphipsyche bengalensis Martynov
(Figs 117, 118)
Amphipsyche bengalensis Martynov, 1935: 201. 2 cf syntypes, INDIA: Bengal, Calcutta, at light, 19. vi. 1907
(Hodgart) (ZSI) [not examined].
Cf (from Martynov, 1935). 'Body pale yellow. Antennae yellow, with narrow dark annulations. Anterior
wings pale. R[/?i] curved in its apical portion even more strongly than in A. indicum [= meridiana] ... In
posterior wings first (apparently false) fork sessile, not pedicellate; . . . formula of spurs 1.4.2 [probably
0.4.2]. Length of body 6mm.'
9- Unknown.
GENITALIA cf (Figs 117, 118) (from Martynov, 1935). '10th [9th] segment as in [meridiana], but its
side-lobes appear to be somewhat broader. Lower end-lobe of the penis [phallotheca] broader, distinctly
excised in the middle and curved upwards (if seen from side); upper leaf-like lobes lacking, in their place is
an oval elevation, behind which are situated two stick-shaped appendages [mid endothecal spines];
underside of the penis thickened before its lower end-lobe.'
REMARKS. This species seems to be most closely related to sinhala, but the male genitalia are
different, assuming Martynov's figures to be accurate. Although Martynov gave no wing-length
for bengalensis, this would also seem to be a larger species than sinhala, whose body length is
116
117
118
Figs 115-118 115, 116, Amphipsyche extrema ?, (115) wing venation; (116) palps. 117, 118, Amphip-
syche bengalensis cf , (117) phallotheca, lateral view; (118), phallotheca, dorsal view. (After Martynov.)
112 P. C. BARNARD
only 4-5 mm. The 6 mm body length of bengalensis suggests that it is of a similar size to
meridiana. The spur formula of this species should almost certainly be 0.4.2; Martynov probably
mistook an apical seta on the fore tibia for a spur.
The two syntypes in the ZSI are damaged (Ghosh, in litt.) and I was unable to examine them.
The species is known only from these two specimens from Bengal.
Amphipsyche extrema (Martynov) comb. n.
(Figs 115, 116)
Amphipsychella extrema Martynov, 1935: 202. 2 syntypes, INDIA: 1 $ , Bengal, Calcutta, Eden Garden, at
light, 26.V.1912 (Gravely); 1 $, Bengal, Calcutta, v.1915 (Gravely) (both lost from ZSI).
Cf . Unknown.
$ (from Martynov, 1935). Tale yellow. Antennae very slender, yellowish, with narrow darker
annulations. Maxillary palpi very short, not reaching eyes; 2-4 joints subequal; 5th joint shorter than 3rd
and 4th combined, its distal half slender and but very indistinctly annulated; labial palpi also very short
[Fig. 116] . . .In anterior wings [Fig. 115] three false veinlets are seen between C and Sc; 1st apical fork a
little longer than its pedicel and somewhat approximated to R [R\]', R long, slightly arcuate; . . . 4th apical
fork with a short pedicel. RS 1+2 in posterior wings simple, not united at its base with RS 3 . Abdomen pale.
Length of body 5-5 mm.' [From generic diagnosis of Amphipsychella] 'Spurs 0.2(1). 2, the outer spur on the
median legs reduced, indistinct.'
REMARKS. It is difficult to comment on the relationships of this species, as it is known only from
the female which I have not examined; apparently both syntypes are lost from the ZSI (Ghosh,
in litt.). However, the highly reduced spur formula, and the shortened maxillary palp place it in
the meridiana-group. It would seem most closely related to bengalensis, meridiana and sinhala,
but can be distinguished from these other Indian species by the spur formula and the maxillary
palp.
Amphipsyche senegalensis (Brauer)
(Figs 120-129; distribution, Fig. 119)
Phanostoma senegalense Brauer, 1875: 71. Lectotype cf , SENEGAL (NM), designated by Kimmins, 1962: 86
[examined].
Phanostoma curvinerve Navas, 1927: 214. LECTOTYPE $ , EGYPT (USNM), here designated [examined].
Syn. n.
Amphipsyche senegalensis (Brauer) Kimmins, 1962: 85.
Cf . Antenna c. 40 mm, with c. 75 segments. Fore wing 11-17 mm. Body pale yellowish brown, antenna
narrowly annulated with brown. Fore wing pale yellow, often with brownish wedge-shaped pterostigmal
marking; venation as in Fig. 120. Spurs 0.4.2 (Fig. 121). Maxillary palp 5-segmented; 5th segment-
secondarily annulated, slightly longer than segments 1 and 2 combined (Fig. 122).
$. Antenna c. 15 mm, with c. 65 segments. Fore wing 9-12 mm. General coloration as in cf Fore wing
usually unmarked, rarely with pale brown pterostigmal marking; venation as in Fig. 126; Rs in fore wing
strongly sinuous. Spurs 0.2.2, 0.3.2 (Fig. 127) or 0.4.2. Maxillary palp similar to that of cf ; 5th segment
approximately equal to 1 and 2 combined (Fig. 129).
GENITALIA cf (Figs 123-125). Ninth segment broadly rounded laterally. Base and stem of phallotheca
narrow, apex bluntly rounded; no endothecal spines present. Inferior appendage slender; terminal
segment moderately well differentiated.
GENITALIA $ (Fig. 128). Eighth sternites broadly rounded; slight indentation in middle of posterior edge,
and outer posterior corners produced.
REMARKS. This species is easily distinguished from the other African species by the complete
absence of endothecal spines in the male. The female can probably be distinguished by the very
sinuous Rs in the fore wing, but this cannot be confirmed until the females of all the African
species have been discovered. A. senegalensis has a very wide distribution, being found in
virtually every country in the Afrotropical region except in the south-west and along the east
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
113
berneri
..-...- v '"'.' 'Vv ulmeri 0\
i. ';..'; senegalensis '-..' '.;.': .'.'.' '.''.'.'
(.'': '''.'.'.'.':':'. ''.''' corbeti '.;
scottae
Fig. 119 Distribution of the African species oiAmphipsyche.
coast (Fig. 119); this distribution coincides closely with the distribution of permanent waters in
Africa (Gourou, 1970). The species is often caught in very large numbers, especially at light.
The synonymy of curvinerve with senegalensis was suspected by Kimmins (1962); the only
remaining syntype of curvinerve, from the Alfieri collection, now in the USNM, has genitalia
indistinguishable from those of typical senegalensis. Ulmer (1963) retained the name curvinerve
when describing the larva of this species, but he admitted that he could not separate the two
species. Ulmer had seen no females from West Africa to compare with his Egyptian examples,
and he rightly suspected that his Sudanese specimens represented a different species: this was
described as ulmeri by Kimmins (1962).
The female figured by Savigny (1813) from Egypt is certainly this species, though not named;
this was the first published figure of a species olAmphipsyche.
The larva of senegalensis was described by Hickin (1955), Jacquemart (1957), Marlier (1962)
114
P. C. BARNARD
Figs 120-125 Amphipsyche senegalensis cf . 120, wing venation; 121, mid and hind legs; 122, maxillary
palp; 123, genitalia, lateral view; 124, phallotheca, lateral view; 125, genitalia, dorsal view.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
111
Figs 136-139 Amphipsyche pellucida $ . 136, wing venation; 137, mid and hind legs; 138, eighth sternites;
139, maxillary palp.
REMARKS. Although this species clearly belongs in the 'African' section of the meridiana-group,
it has no close affinities with any other species. It is the only species found in Madagascar, and
morphologically the form of the phallotheca renders it easily identifiable. This is the first time
that the male has been described.
Navas (1923) mis-read the type-locality of pellucida as 'Maeratanana'; this also applies to
other species described in the same paper.
MATERIAL EXAMINED
Holotype 9> Madagascar: Maevatanana [no further data] (MNHN).
3 cf , 13 $. Madagascar (BMNH, USNM).
Amphipsyche instabilis Kimmins
(Figs 140-150; distribution, Fig. 119)
Amphipsyche instabilis Kimmins, 1963: 126. Holotype cf , ETHIOPIA (BMNH) [examined].
Phanostoma plicata Jacquemart, 1963: 363. LECTOTYPE cf , ZIMBABWE (ZI), here designated [ex-
amined]. Syn. n.
O" . Antenna up to 33 mm, with c. 90 segments. Fore wing 11-14 mm. Body pale yellowish brown, antennal
118
P. C. BARNARD
U2
U5
Figs 140-146 Amphipsyche instabilis C". 140, wing venation; 141, mid and hind tibiae; 142, maxillary
palp; 143, genitalia, lateral view; 144, genitalia, ventral view; 145, phallotheca, lateral view; 146,
phallotheca, dorsal view.
segments becoming gradually more fuscous towards apex, narrowly annulated with brown. Fore wing pale
yellow with no markings; venation as in Fig. 140; cross-vein present between M 3+4 and Cw la in hind wing.
Apical venation often irregular in both fore and hind wings. Spurs 0.4.2 (Fig. 141). Maxillary palp
4-segmented, apical segment short and not secondarily annulated (Fig. 142).
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
119
9 (allotype only). Antennal length unknown (specimen damaged). Fore wing 9 mm. Coloration as in cf ,
venation as in Fig. 147; cross-vein between A/ 3+4 and Cu la in hind wing as in cf . Spurs 0.2.2 (Fig. 148).
Maxillary palp 4-segmented (Fig. 149), similar to that of cf .
GENITALIA cf (Figs 143-146). Ninth segment only moderately broad laterally. Base of phallotheca very
large and rounded, stem short and greatly thickened; dorsal clavate process present, with curved stem,
apex covered with fine spines (Fig. 145); on either side of process a similarly spinose triangular lobe. Mid
endothecal spines fused to form single median spine, very thick and curved dorsally. Lower apex of
phallotheca elongate, curved dorsally, extreme apex globular. Inferior appendage narrow, slightly sinuate;
terminal segment moderately well differentiated.
GENITALIA 9 (Fig- 150). Eighth sternites broad and squarish, with rounded indentation in posterior edge;
inner thickened edges narrow.
REMARKS. Although this species is closely related to the other African species of the genus, it is
easily recognized. Externally, both sexes are easily identified by the extra cross-vein between
M 3+4 and Cw la in the hind wing which is unique amongst the African species, though paralleled
inproluta, the type-species of the genus. The male genitalia are highly distinctive; the elongate
dorsal process and single median endothecal spine are not found in any other species. The only
known female closely resembles the female of senegalensis , which is found in the same locality,
but apart from the venational character already mentioned, it can easily be distinguished by the
4-segmented maxillary palp and the squarish eighth sternites.
The specimen here designated as lectotype ofplicata was labelled 'type' by Jacquemart, and
the paralectotypes as 'paratypes', but these designations were not pubished.
Although the descriptions ofinstabilis andplicata were published in the same year, there is no
doubt that Kimmins's appeared first. His paper was officially published on 20th February 1963
and, according to data in the BMNH Entomology Department Library, it was definitely
available before the end of that month. The book in which Jacquemart's paper appeared has no
exact date of publication, but the copy in the BMNH Zoology Department Library was received
U8
Figs 147-150 Amphipsyche instabilis $. 147, wing venation; 148, mid and hind tibiae; 149, maxillary
palp; 150, eighth sternites.
120
P. C. BARNARD
on llth October 1963. Further enquiries to the ZI revealed that their copy was received on 25th
November 1963 (Tjeder, in litt.), and the copy in the Kungliga Biblioteket, Stockholm (the
Swedish copyright library) was not received until February 27th 1964 (Lilliestam, in litt). It
seems certain, therefore, that the description oiplicata was not published until at least October
1963, and that the name instabilis has priority.
MATERIAL EXAMINED
Holotype cf of instabilis, Ethiopia: Dawa River, 12 km N. of Hudat, 12. iv. 1961 (Tj0nneland) (slide
preparation, BMNH). Lectotype cf of plicata, Zimbabwe: Victoria Falls, 16. v. 1951 (slide preparation,
ZI).
38 cf , 1 $ , Ethiopia (18 cf paratypes of instabilis and 1 $ allotype inadvertently labelled as 'cf paratype'
by Kimmins), Zimbabwe (5 d" paralectotypes of plicata), Zambia (BMNH, IRSNB, USNM, ZI).
Amphipsyche ulmeri Kimmins
(Figs 151-154; distribution, Fig. 119)
[Phanostoma senegalense Brauer; Ulmer, 1923: 19 (partim - specimens from Sennar only); 1924: 2.
Misidentifications . ]
Amphipsyche ulmeri Kimmins, 1962: 89. Holotype cf , SUDAN (NM) [examined].
151
153
Figs 151-154 Amphipsyche ulmeri cf . 151, wing venation; 152, genitalia, ventral view; 153, genitalia,
lateral view; 154, phallotheca, lateral view.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 121
Cf (holotype only). Antenna 32 mm, with c. 70 segments. Fore wing 14 mm. Antennal segments pale
golden brown, annulated with dark brown. Head, thorax and abdomen yellowish brown. Fore wing pale
yellow with indistinct brownish shading across anastomosis; venation as in Fig. 151. Spurs 0.4.2. Maxillary
palp as in senegalensis (cf. Fig. 122).
9- No specimens seen, but see 'Remarks' below.
GENITALIA cf (Figs 152-154). Lateral part of ninth segment broad and squarish. Base of phallotheca broad,
extreme apex bluntly pointed; mid endothecal spines long and stout, curved abruptly dorsally (Fig. 154).
Inferior appendage strongly sinuous in ventral view, terminal segment moderately well differentiated.
REMARKS. A. ulmeri and scottae are closely related, despite their widely separate distributions
(Fig. 119) , both species having very similar male genitalia. However, ulmeri can be distinguished
by the sharply up-turned mid endothecal spines.
Kimmins (1962) mentioned the existence of females from the type-locality, Sennar, in
Ulmer's collection. Subsequently Ulmer (1963) described these females as being distinguishable
from Egyptian females of curvinerve (= senegalensis) by the less sinuous Rs in the fore wing and
the spur formula of 0.3.2. It is quite probable that these are females of ulmeri, but the spur
formula is not significant, as Kimmins (1963) showed that there is great variation in the spurs of
female senegalensis, 0.3.2 occurring in that species also.
MATERIAL EXAMINED
Holotype cf , Sudan: Sennar, 18-27. ii. 1914 (Ebner) (NM).
Amphipsyche scottae Kimmins
(Figs 155-164; distribution, Fig. 119)
Amphipsyche scottae Kimmins, 1962: 93. Holotype cf , SOUTH AFRICA (BMNH) [examined].
Cf . Antenna c. 45 mm, with c. 95 segments. Fore wing 16-19 mm. Body yellowish brown, antenna narrowly
annulated with brown, segments becoming more fuscous towards apex. Fore wing pale yellow, with slightly
darker area along costa and near Sc-Ri cross-vein, often indistinct. Venation as in Fig. 155. Spurs 0.4.2
(Fig. 156). Maxillary palp 5-segmented, 5th segment approximately equal in length to segments 1 and 2
combined, not secondarily annulated (Fig. 157).
$. Antenna c. 20 mm, with c. 70 segments. Fore wing 14-15 mm. General coloration as in cf ; fore wing
with no darker markings. Venation as in Fig. 161. Spurs 0.4.2 (Fig. 162). Maxillary palp similar to that of
Cf , but 5th segment shorter (Fig. 164).
GENITALIA cf (Figs 158-160). Ninth segment broadly rounded laterally. Base of phallotheca broadly
triangular, with pronounced corner on dorsal side. Ventral apex forming pair of rounded lobes; extreme
apex bluntly pointed (Fig. 159). Mid endothecal spines long, curved dorsally. Inferior appendage slender,
terminal segment moderately well differentiated.
GENITALIA $ (Fig. 163). Eighth sternites broad and squarish, with slight indentation in middle of posterior
edge. Inner thickened margin extending far towards anterior edge.
REMARKS. A. scottae most closely resembles ulmeri in that the males of both species have the tip
of the phallotheca bluntly pointed in lateral view, but scottae can be distinguished by the gently
curved mid endothecal spines, which are sharply up-turned in ulmeri.
The larva of this species was described by Scott (in press), and some aspects of its biology are
mentioned by Chutter (1963; 1968); see p. 79.
MATERIAL EXAMINED
Holotype cf, South Africa: Natal, Wilge R., 5 miles [8 km] below Harrismith, 10.ii.1959 (slide
preparation, BMNH).
12 Cf , 2 $ , South Africa (12 cf , 1 $ paratypes) (BMNH).
122
P. C. BARNARD
Figs 155-160 Amphipsyche scottae C? . 155, wing venation; 156, mid and hind tibiae; 157, maxillary palp;
158, genitalia, lateral view; 159, phallotheca, lateral view; 160, genitalia, ventral view.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
123
Figs 161-164 Amphipsyche scottae $. 161, wing venation; 162, mid and hind legs; 163, eighth sternites;
164, maxillary palp.
Amphipsyche fuscata Kimmins
(Figs 165-170; distribution, Fig. 119)
Amphipsyche fuscata Kimmins, 1963: 128. Holotype cf , ETHIOPIA (BMNH) [examined].
Cf . Antenna up to 35 mm, with c. 85 segments. Fore wing 12-17 mm. Body pale yellowish brown; basal
12-15 segments of antenna yellowish brown, remainder of flagellum fuscous. Fore wing pale yellow, with
fuscous streak running obliquely from costal margin proximal to anastomosis; hind margin fuscous (these
markings may be very faint). Venation as in Fig. 165. Spurs 0.4.2 (Fig. 166). Maxillary palp short, 4th and
5th segments imperfectly separated, 5th segment narrow apically, not secondarily annulated (Fig. 167).
$. Unknown.
GENITALIA cf (Figs 168-170). Lateral lobe of ninth segment somewhat squarish. Base of phallotheca
narrow and rounded, apex elongate, produced into a bifid lobe. Mid endothecal spines long, curved
upwards.
REMARKS. Well-marked specimens of this species can be recognized by the oblique wing-
marking, but identification of specimens with faint markings depends on the male genitalia, and
here the close similarity with several other African species is apparent. However , fuscata differs
124
P. C. BARNARD
Figs 165-170 Amphipsyche fuscata d". 165, wing venation; 166, mid and hind tibiae; 167, maxillary palp;
168, genitalia, lateral view; 169, phallotheca, lateral view; 170, genitalia, ventral view.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE
125
from scottae and ulmeri in having the apex of the phallotheca rounded, not pointed, and from
berneri and corbeti in having a narrow base to the phallotheca. A.fuscata is further distinguished
by the unique maxillary palps, with the 4th and 5th segments only partly fused. Kimmins (1963)
noted the colour of the palps, but did not notice their unusal form.
MATERIAL EXAMINED
Holotype d", Ethiopia: Koka Dam, 29.iii.1964 (Tj0nneland) (slide preparation, BMNH).
62 cf , Ethiopia (60 cf paratypes), Sudan (BMNH).
Amphipsyche corbeti Kimmins
(Figs 171-176; distribution, Fig. 119)
Amphipsyche corbeti Kimmins, 1962: 89. Holotype cf , UGANDA (BMNH) [examined].
Cf. Antenna up to 30 mm, with c. 85 segments. Fore wing 11-12 mm. Body pale yellowish brown, thorax
pale brown dorsally, antenna pale greyish brown. Fore wing pale yellowish brown, usually with pale brown
shading around Ri~Rs cross-vein. Venation as in Fig. 171. Spurs 0.4.2 (Fig. 172). Maxillary palp
4-segmented, terminal segment narrow and slightly elongate, but not secondarily annulated (Fig. 173).
$. Unknown.
Figs 171-176 Amphipsyche corbeti cf . 171, wing venation; 172, mid and hind tibiae; 173, maxillary palp;
174, genitalia, lateral view; 175, phallotheca, lateral view; 176, genitalia, ventral view.
126
P. C. BARNARD
GENITALIA cf (Figs 174-176). Base of phallotheca broadly triangular, stem thickened, apex forming pair of
lobes ventrally; mid endothecal spines long, curved dorsally. Inferior appendage slender, terminal
segment scarcely differentiated.
REMARKS. This species closely resembles berneri and, to a lesser extent, ulmeri. It differs from
ulmeri in the thicker stem of the phallotheca and the longer endothecal spines, and from berneri
in the lobes of the tenth segment. These lobes are narrower in dorsal view in corbeti, and do not
diverge. There are also slight differences in the apex of the phallotheca, best seen in ventral
view.
Figs 177-182 Amphipsyche berneri cf . 177, wing venation; 178, mid and hind legs; 179, maxillary palp;
180, genitalia, ventral view; 181, phallotheca, ventral view; 182, genitalia, lateral view.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 127
MATERIAL EXAMINED
Holotype cf, Uganda: Northern Province, Victoria Nile, Karuma Falls [no date] (Corbet) (slide
preparation, BMNH).
33 Cf paratypes, Uganda (BMNH).
Amphipsyche berneri Kimmins
(Figs 177-182; distribution, Fig. 119)
[Phanostoma senegalense Brauer; Kimmins, 1957a: 13 (partim - specimens from Gold Coast [Ghana]
only). Misidentification.]
Amphipsyche berneri Kimmins, 1962: 91. Holotype cf , GHANA (BMNH) [examined].
Cf . Antenna c. 25 mm, with c. 85 segments. Fore wing 11-12 mm. Body coloration uncertain (both
specimens originally preserved in alcohol). Fore wing yellowish brown, with darker marking centred on
R\-Rs cross-vein; venation as in Fig. 177. Spurs 0.4.2 (Fig. 178). Maxillary palp 4-segmented, terminal
segment scarcely longer than 3rd (Fig. 179), not secondarily annulated.
$. Unknown.
GENITALIA cf (Figs. 180-182). Base of phallotheca broadly triangular; ventral apex forming pair of lobes;
mid endothecal spines long and curved dorsally. Inferior appendage slender, terminal segment not clearly
differentiated.
REMARKS. This species is very similar to corbeti, both externally and in the form of the genitalia.
It can be distinguished by slight differences in the shape of the apex of the phallotheca and by the
lobes of the tenth segment. These are broader and more divergent in dorsal view in berneri,
although it should be noted that in the paratype (figured here) the lobes are less divergent than in
the holotype figured by Kimmins (1962: 91, fig. 26). The holotype is now mounted laterally as a
permanent slide preparation.
Kimmins did not comment on the 4-segmented maxillary palps, although these are clearly
visible in his slide preparation of the holotype.
MATERIAL EXAMINED
Holotype cf , Ghana: Volta R., Senchi, l.viii.1950 (Berner) (slide preparation, BMNH).
1 cf paratype, Ghana (BMNH).
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Index
Synonyms are in italics; principal references are in bold. References to Table 1 (p. 82), the cladogram
(p. 83) and the check-list (p. 85) are not included.
Aethaloptera72,78,84
Amphipsyche 71, 75, 76
Amphipsychella 76, 77, 112
apicalis77,83,94,96,99
apicalis-group 77, 78, 80, 81, 83,
84, 94, 96, 99
bengalensis 81, 105, 111,112
berneri 77, 84, 125,126, 127
bifasciata 89
Blepharopus 74
Cheumatopsyche 80
corbeti 84, 125, 127
curvinervelS, 79, 112, 113, 115,
121
delicata81,93
distincta83,86,89,94
exsiliens83,94,96
extrema76, 81, 112
fuscata 84, 123
gratiosa83,96, 98
indica 78, 93, 106, 108, 110, 111
instabilis83,84,88, 117
Leptonema 74, 79
Leptopsyche 75
Macronema 72, 74, 79
Macronematinae 72, 74
Macronematini 72, 74
Macrostemum 74, 75, 78, 79, 84
magna 77, 84, 92, 102
meridiana 78, 79, 80, 83, 93, 99,
105, 106, 111, 112
meridiana-group 77, 78, 80, 81,
83,84,85,88,94,101,105,
112,117
minima 94, 99
nirvana 106, 108, 109
paraproluta 86, 88
parva 104, 105
pellucida84, 115
petiolata 94, 96, 99
Phanostoma 76, 77, 79, 108
plicata 117, 119, 120
Polymorphanisini 72, 74
Polymorphanisus 84
proluta76,78,83,86,89,99,
106, 119
proluta-group 77, 78, 80, 81, 83,
84, 86, 88, 94
propinqualQ6,10S, 110
Protomacronema 74, 75, 77, 79,
84
Pseudoleptonema 74
pubescens 99, 100
scottae 72, 79, 80, 121,125
senegalensis 76, 78, 79, 80, 83,
84, 112, 119, 121
sigmosa 107, 110
Simulium71,72, 79
sinhala81,105, 108, 111,112
tricalcarata 107, 110
Trichomacronema 74
ulmeri78,84, 113,120, 121,
125, 126
vedana 106
British Museum (Natural History)
Blue Butterflies of the Lycaenopsis-group
J. N. Eliot and A. Kawazoe
The most wide-spread member of the Lycaenopsis-group is known and loved in Britain as the
Holly Blue, in North America as the Spring Azure and in Japan as Ruri Shijimi (the Small
Lapis Lazuli). In appearance and behaviour it is typical of the group, which attains its
maximum diversity and abundance in the mountains of South East Asia and New Guinea.
Hitherto the systematics of the group have been in a state of confusion. In this work 112
species are recognised divided among 21 genera. These are defined mainly on characters of the
genitalia, which are figured for the males of each, and the females of most, species. There are
keys to, and descriptions of, the genera, subgenera, species and subspecies, including 8 new
genera and 27 new species. The new species and those not previously figured are illustrated in
the plates at the rear of the book, and references are given to published figures of the
remaining species. Finally, there is a complete bibliography, enabling the original descriptions
of all the taxa to be traced.
John Eliot is the author of many papers of a taxonomic nature and was the reviser of the
third edition of Corbet & Pendlebury's classic work "The Butterflies of the Malay Peninsula"
first published half a century ago. His contribution to zoology has been recognised by the
presentation of the Stamford Raffles Award of the Zoological Society of London and the
J. H. Bloomer Award of the Linnean Society of London.
Akito Kawazoe has published many taxonomic papers, mainly in Japanese journals, and is
best known in Europe as the senior author of "Coloured Illustrations of the Butterflies of
Japan", a work notable not only for its superb coloured plates but also for numerous black and
white drawings illustrating structural characters. His skill as artist and microscopist is again
demonstrated in this book by more than two hundred figures of genitalia which will prove
indispensable to a proper understanding of the Lycaenopsis-group.
Blue butterflies of the Lycaenopsis-group. J. N. Eliot & A. Kawazoe
1983, 296pp, 18 plates include 6 in colour, 560 text figures. (0 565 00860 9)
28.00
Titles to be published in Volume 48
Gelechiid moths of the genus Mirificarma.
By Linda M.Pitkin.
Macronematine caddisflies of the genus Amphipsyche (Trichoptera: Hydropsychidae)
By P. C. Barnard.
A review of the genera oflndo- Pacific Encyrtidae (Hymenoptera: Chalcidoidea).
By John S. Noyes and M. Hayat
Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk
Printed in Great Britain by Henry Ling Ltd, Dorchester
Bulletin of the
British Museum (Natural History)
A review of the genera
of Indo-Pacific Encyrtidae
(Hymenoptera: Chalcidoidea)
John S. Noyes & M. Hay at
Entomology series
Vol 48 No 3 28 June 1984
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INDO-PACIFIC ENCYRTIDAE 133
work is required. Conversely, where we agree it should give workers a greater degree of
confidence in any proposed taxonomic changes. There will thus also be a measure of duplication
between our two works, but hopefully this has been kept to a minimum.
Contributions to knowledge of the fauna of other areas of the Indo-Pacific region have
included papers by several authors, e.g. Ashmead (19050, b), Girault (I919a,b; 1920c), Gahan
(1927&), Ferriere (1931), Eady (19600,6), Kerrich (1963; 1967; 1978) also Subba Rao (1970;
1973; 1978) and Trjapitzin (1965) on the fauna of South East Asia, whilst Perkins (1910), Swezey
(1946), Fullaway (1946), Timberlake (1920; 1924; 1941) and Beardsley (1969; 1976) contributed
papers on the fauna of the Hawaiian Islands and other Pacific islands.
The present study recognises 263 described genera and 977 described species of Encyrtidae as
occurring in the Indo-Pacific region. The types, or reliably determined specimens of virtually all
of the described species known from the area, have been examined by either one or both of us.
This includes examination (by JSN) of nearly all of the types of the species described by Girault
from South East Asia and Australia. We have also examined a great deal of unidentified
material collected from all over the region.
The relationship between faunas of the various component areas of the Indo-Pacific region
and other zoogeographical areas in terms of distribution of the genera is summarised in Tables 1
and 2. Here the relationship between these areas is indicated by the number of genera with
distributions which are restricted to a particular type. For example, five genera are distributed
from the Indian subcontinent to Australasia excluding Australia (Table 1; line 3 column 4, or,
line 4 column 3), but nine from the Indian subcontinent to Australia (Table 1; line 2, column 4,
or, line 4, column 2); similarly six genera are distributed from Australasia (excluding Australia)
to Africa and Europe (Table 2; line 3, column 3), but four are restricted to India and the
Palaearctic and Afrotropical regions (Table 2, line 7, column 4). As might be expected, the
Australian fauna has a strong relationship with that of the Oriental region, there being at least 11
genera known only from Australia, through India to the Afrotropical region and at least a
further 30 genera found in South East Asia and Australia only. Sixteen genera have been found
only in India and 60 only in Australia, but this probably reflects the activities of collectors rather
than actual distribution. It is apparent that the relationship between the Australian and
Neotropical faunas is not as strong as suggested previously (Noyes, 1980), although there are
some genera known only from the Australasian and Neotropical regions, e.g. Austroencyrtus,
and from the Oriental, Australasian and Neotropical regions, e.g. Meniscocephalus .
Table 1 The relationships between the encyrtid faunas of the component areas of the Indo-Pacific
region (given in numbers of genera).
Geographical region
Pacific
Australia
Australasia
(excluding
Australia)
Indian
subcontinent
Continuous distribution to
Pacific only
Continuous distribution to
Australia only
Continuous distribution to
Australasia only (excluding
Australia)
Continuous distribution to
the Indian subcontinent
5
1
1
4
1
60
10
9
1
10
16
5
4
9
5
16
Keys to genera found in other zoogeographical regions have been published by Trjapitzin
(19710) for the Palaearctic region, Trjapitzin & Gordh (19780,6) for the Nearctic region,
Prinsloo & Annecke (1979) for the Afrotropical region, and Noyes (1980) for the Neotropical
region.
134 J. S. NOYES & M. HAYAT
Table 2 The relationships between the encyrtid faunas of the component areas of the Indo-Pacific
region (given in numbers of genera) and other zoogeographical areas.
Geographical region Pacific Australia Australasia Oriental
(excluding
Australia)
Continuous distribution to Europe
and Africa 11
Continuous distribution to Europe
excluding Africa 113
Continuous distribution to Africa
excluding Europe 2 11 6 7
Stated region plus Palaearctic
only 3
Stated region plus Neotropics
only 1 5
Stated region plus New World
only 6
Stated region, Palaearctic,
and Africa only 4
Cosmopolitan genera 50, introduced or probably introduced genera 10, other distribution patterns 15
Notes on generic review
Classification
Currently there are two basic systems of classification of the Encyrtidae in use. Most previous
authors (Erdos & Novicky, 1955; Hoffer, 1955; Compere & Annecke, 1960; Tachikawa, 1963;
Kerrich, 1967) have divided the family into three subfamilies: Arrhenophaginae, Antheminae
and Encyrtinae, the last mentioned containing almost all known genera. In the present work we
follow Trjapitzin (I973a,b) who recognises only two subfamilies, the Tetracneminae and the
Encyrtinae, which can be separated as follows.
Tetracneminae. Paratergites present or at least represented by a membranous strip which connects the
outer plates of the ovipositor to the sides of the last gastral tergite, either along its length or at the base near
the cereal plates only. Linea calva of forewing with undifferentiated margins and filum spinosum almost
always absent. Hypopygium triangular and always reaching apex of gaster. Mandibles with all teeth
apically acute (except Doliphoceras siccus Prinsloo & Annecke from southern Africa).
Encyrtinae. Paratergites almost always absent (present in some Trechnites and Cercobelus}. Linea calva
of forewing generally with setae on proximal side longer and stronger than those on distal side. Filum
spinosum almost always present. Hypopygium often short and subrectangular (not reaching more than half
way along gaster) but often triangular and reaching apex of gaster. Mandibles sometimes with a broadly
truncate edge or tooth.
Trjapitzin divides the Tetracneminae into 12 tribes and the Encyrtinae into 36 tribes. We feel
that many of these tribes are unnecessary and occasionally they are even placed by Trjapitzin in
the wrong subfamily, e.g. Mirini, Neodiscodini, Rhinoencyrtini. Even so his study is the most
detailed to date (although it is based mainly on the Palaearctic Fauna whilst encyrtids are a
predominantly tropical group), therefore we have attempted to place, as far as possible, the
Indo-Pacific genera according to his proposed classification. At the same time we have
commented on several tribes and subtribes which require some modification. A new system of
tribal classification is not proposed here since this is beyond the scope of the present work; the
genera are arranged alphabetically, although a summary of their possible systematic positions in
relation to Trjapitzin's classification is given on p. 353.
Taxonomic changes
Unless otherwise stated, the new generic and specific synonymies and the new combinations
have resulted from the examination of relevant type-material. Generally, if genera are here
INDO-PACIFIC ENCYRTIDAE 135
synonymised without comment, the relevant type-species are so close morphologically as to be
difficult to separate even at specific level. This usually applies only to genera described by
Girault from Australia. For new combinations, comments are limited to those species where we
feel that this is necessary, since to discuss each proposed new combination would greatly and
unnecessarily increase the length of the text.
Notes on key
The encyrtid genera are not easily keyed into distinct groups such as subfamilies, tribes etc.,
therefore the key deals with all genera together and may thus prove very daunting to the user
because of its length. We have tried to overcome this by dividing the key into groups of not more
than 27 couplets. Each group is entered from one of the first 44 couplets and is delimited by
couplet numbers in bold type. The genera in each group are not necessarily related. Thus, to
arrive at a generic name, it should not be necessary to run any specimen through more than 29
couplets and generally fewer than that.
Some of the characters used to separate groups of genera, e.g. relative widths of scape,
position of apex of hypopygium, relative length of funicle segments, forewing hyaline or
infuscate etc. , can be rather weak or ambiguous. For instance, it is not always easy to be certain
whether a wing is truly hyaline or slightly infuscate; however, in several such instances, a species
has been keyed out to the relevant genus via both alternatives. Some of the couplets are
complex. It is therefore possible for the user to make a wrong decision and go hopelessly wrong
unless both alternatives of a couplet have been carefully read and understood before a decision is
made to go one way or the other. The key is almost entirely artificial, therefore it must be
stressed that all determinations should be confirmed by comparison of the relevant material with
a reliable generic description. It must also be remembered that a specimen does not necessarily
belong to an undescribed genus if it does not run easily through the key or if it runs directly to a
genus to which the user knows that it cannot belong. Even if the key is used correctly it is likely
that only a small majority of species will run to the correct genus. This is because it is doubtful
whether the present review covers more than a very small fraction of the species which actually
occur in the region and which can be placed in already recognised genera.
It is inevitable that there has to be some degree of simplification in a work with as large a
coverage as this, and which deals with many poorly worked genera; this is particularly so with
regard to the separation of some of the genera within taxonomically difficult groups, e.g.
Anagyrini, Cheiloneurini, Habrolepidini and Microteryini (subtribe Syrphophagina) (see com-
ments under relevant genera). Such simplification has been necessary in order to complete the
key and to avoid making it difficult to use.
Finally, the males are not keyed to genera because those of a very large number of
Indo-Pacific genera are not known; also our experience has shown that most entomologists do
not attempt to place unknown males to genus.
Notes on terms and measurements
Unless otherwise stated in the captions, the figures were drawn directly from slide-mounted
material using a drawing tube attachment on a compound microscope, therefore relative
measurements can be taken directly from these figures. However, such measurements must not
be made where the points of reference for these were not equidistant from the objective of the
microscope when these drawings were made, e.g. relative width of scape (since the scape is
rarely absolutely flat on a slide mounted specimen), relative distance of antennal toruli from
mouth margin, relative length of malar space to eye length, POL to OOL, etc. These
measurements are only reliable if taken from a dry, card-mounted specimen.
Head (Figs 1-4)
Antennal clava. Composed of one to three segments. If more than one segment then these are
separated by partial or complete sutures and are not as clearly separated as the funicle segments.
136 j. s. NOYES & M. HAYAT
The apex of the clava has a sensory part which is indicated by an area of micropilosity and/or
microtubules and/or a sieve-plate structure (these are individually only visible on a good slide
preparation examined at high magnification). This sensory part is easily seen on dry-mounted
material, is usually flattened and may either be transverse, oblique or a narrow horizontal strip.
If it is large it gives the clava a truncate appearance, thus the clava may appear transversely or
obliquely truncate as opposed to apically rounded. Often a slide-mounted antenna which is
apically obliquely truncate will appear to be apically rounded; this may either result from the
clava not being correctly orientated or the sensory part having been inflated during clearing.
Therefore when using the following key it is best to determine the presence or absence of an
oblique truncation using dry-mounted material.
Antennal funicle . This does not include the anellus (or false ring-joint of Timberlake, 19226:
168, 172), which may be present or absent but is almost always hidden by the pedicel and
invisible in dry-mounted material. In the Encyrtidae the anellus never bears setae, whereas the
funicle segments always bear setae (although sometimes very short). The relative length of the
setae to the diameter of the segments can be taken directly from the text-figures.
Eye. The measurements of length and breadth are the maximum and minimum diameters
respectively; the points from which the measurements are taken should be equidistant from the
objective of the microscope (i.e. both in focus simultaneously).
Frontovertex width. The measurements are taken either at the level of the anterior ocellus or at
the point where the fronto vertex is narrowest, as stated in text.
Head width. The maximum width of the head either in frontal view (as in Fig. 3) or side view (as
in Fig. 4), as stated in text.
Malar space. The minimum distance between eye and mouth margin. The measurement is taken
as for eye (above).
Malar sulcus. The sulcus joining the lower margin of the eye and mouth margin (see Figs 3, 4),
sometimes absent but usually indicated by a slight change of sculpture.
Mandibles. The dentition can vary as follows: without teeth (Fig. 218), with one long curved
tooth (Fig. 129), one tooth and a broad truncation (Figs 14, 121, 189, 229, 271), two teeth, two
teeth and a truncation (Figs 75, 122, 225, 347, 381), two teeth and a rudimentary third tooth,
three teeth (Figs 76, 123, 136, 144, 178, 221, 397, 435, 443) or four teeth (Figs 116, 188,293,294).
However, this is not always clear since the distinction between two teeth and a truncation and
three teeth is often not very great (see Figs 76, 123, 347). Similarly for the difference between
one tooth and a truncation and two teeth and a truncation (see Figs 74, 115, 319), between three
teeth and four teeth (see Fig. 188) and occasionally also between two teeth and a truncation and
four teeth.
OOL. The minimum distance between the eye margin and the nearest posterior ocellus (see Fig.
2).
POL. The minimum distance between the posterior ocelli (see Fig. 2).
Thorax (Figs 5-7)
Forewing (Fig. 5).
Filum spinosum: a series of peg-like setae on distal margin of linea calva which are clearly
stouter than adjacent setae.
Length of forewing: measured from most proximal part of costal cell to apex of wing.
Linea calva (or speculum of some authors): an oblique hairless line extending from just below
marginal and stigmal veins to posterior margin of forewing.
Marginal vein: measured from where the submarginal vein reaches the anterior margin of
wing (as shown in Fig. 5), or from where the anterior edge of the venation at the junction of the
submarginal vein is abruptly angled and not from the subapical hyaline break of the submarginal
vein.
Parastigma: a very slight to strong swelling of the apical one-third of the submarginal vein.
Postmarginal vein: measured as shown in Fig. 5, its apex usually indicated by a single,
relatively long, suberect seta.
OOL POL
INDO-PACIFIC ENCYRTIDAE
-flagellum
137
pedicel
ocelli
-^postmarginal vein
'*-. marginal vein
basal cell
filum spinosum
linea calva
Figs 1-5 1, generalized encyrtid $ antenna, left, outer aspect; 2, generalized encyrtid 9 head, dorsal
aspect; 3, generalized encyrtid $ head, frontal aspect; 4, generalized encyrtid $ head, aspect from left
side; 5, generalized encyrtid forewing, upper surface.
138 J- S. NOYES & M. HAYAT
Stigmal vein (or radial vein of some authors): measured as shown in Fig. 5. There are usually
four (sometimes fewer) circular sensillae at its apex. The relative position and number of these
sensillae are occasionally very useful in separating generic groups.
Uncus: beak-like process often arising from apex of stigmal vein.
Notaular lines (or parapsidal lines of some authors) (Fig. 6). These are occasionally difficult to
see in dry-mounted material unless viewed under correct light conditions.
Propodeum. The length is measured along the mid-line.
Scutellum. The length is measured along the mid-line; the breadth excludes the axillae.
Caster (Fig. 8)
Cerci (or pygostyles of some authors). The relative position is measured in dry-mounted
material; if it is measured in material which has been in alcohol or slide-mounted, the gaster may
be distended and the cerci will be positioned relatively nearer the apex of the gaster.
Gonostylus. The third valvula, or ovipositor sheath, as seen in slide-mounted material.
Hypopygium (or subgenital plate of some authors). The relative position of the apex is measured
in dry-mounted material. Care must be taken to take this measurement from specimens in which
the ovipositor has not dropped down into the laying position, particularly in the Encyrtinae.
Here the hypopygium is usually retracted during oviposition and thus a hypopygium which
normally reaches the apex of the gaster will often appear to reach only half to two-thirds of the
way along the gaster.
Last tergite (syntergum or epipygium of some authors). Its length is measured from its apex to
the centre of an imaginary line connecting the cereal plates.
Ovipositor. The length of the exserted part is measured from the apex of the last gastral tergite
(never hypopygium) in dry-mounted material. If material has been in alcohol the gaster may be
distorted and the ovipositor may appear to be relatively more exserted; in this case it would be
better to use the relative lengths of the exserted parts of the gonostyli (ovipositor sheaths).
Ovipositor sheath. The gonostylus as seen in dry-mounted material.
Abbreviations
AMNH American Museum of Natural History, New York, USA.
ANIC Australian National Insect Collection, Division of Entomology CSIRO, Canberra, Australia.
BMNH British Museum (Natural History), London, UK.
BPBM Bernice P. Bishop Museum, Honolulu, Hawaii.
CNC Canadian National Collection, Biosystematics Research Institute, Ottawa, Canada.
DSIR Division of Entomology, Department of Scientific and Industrial Research, Auckland, New
Zealand.
GC Gijwijt collection, c/o M. J. Giswijt, PO Box 4, 1243 ZG, 'S-Graveland, Netherlands.
HC Hayat collection, c/o M. Hayat, Department of Zoology, Aligarh Muslim University, Aligarh,
India.
HDOU Hope Museum, Oxford University, Oxford, England.
HNHM Hungarian Natural History Museum, Budapest, Hungary.
IPK Institute fur Pflanzenschutzforschung, Eberswalde, DDR.
MCSN Museo Civico di Storia Naturale, Genova, Italy.
PPRI Plant Protection Research Institute, Pretoria, South Africa.
QM Queensland Museum, Brisbane, Australia.
UCR University of California, Riverside, California, USA.
USNM National Museum of Natural History, Washington DC, USA.
RMNH Rijksmuseum van Natuurlijke Historic, Leiden, Netherlands.
SAM South Australian Museum, Adelaide, Australia.
ZI Zoological Institute, Academy of Sciences, Leningrad, USSR.
ZMCU Zoological Museum, Cambridge University, Cambridge, England.
INDO-PACIFIC ENCYRTIDAE
mesoscutum
139
pronotum
notaular line
axilla
tegula
hind
coxa
mid coxa metapleurum
mesopleurum
hypopygium
Figs 6-8 6, Homalotylusflaminius (Dalman) $ , thorax, dorsal aspect; 7, Charitopus sp. $ , thorax, aspect
from left side; 8, generalized encyrtid $ gaster, aspect from left side.
140 J. S. NOYES & M. HAYAT
Key to genera (females)
1 Tarsi four-segmented 45 (p. 143)
Tarsi five-segmented 2
2(1) Funicle with fewer than six segments 3
Funicle with at least six segments 4
3 (2) Funicle three- or four-segmented 46 (p. 143)
Funicle five-segmented 55 (p. 143)
4 (2) Forewing shortened, clearly not reaching apex of gaster 5
Forewing normal, at least very nearly reaching apex of gaster 6
5 (4) Hypopygium reaching or very nearly reaching apex of gaster (at least four-fifths
along gaster) 73 (p. 144)
Hypopygium not reaching more than two-thirds along gaster 85 (p. 146)
6 (4) Scutellum either with a group of coarse, long, dark setae arranged in a more or
less compact tuft or bundle (Fig. 47) , or with two or more scale-like setae (Figs
44,48) 95 (p. 148)
Scutellum without a distinct tuft or bundle of setae or scale-like setae 7
7 (6) Scape not more than three times as long as broad 8
Scape more than three times as long as broad 15
8(7) Flagellum broadened and flattened, at most only first funicle segment not
transverse (Figs51-54, 56, 57, 302) 104 (p. 150)
Flagellum not flattened, more or less cylindrical to broadly oval in cross-section,
or if appearing flattened then at least first two segments longer than broad ... 9
9 (8) Forewing infuscate or with a very distinct pattern of dark and pale setae and thus
appearing infuscate (excluding those species with a very indistinct suffusion of
yellow or pale brown or with a very small spot beneath marginal vein which
does not or hardly extends past apex of stigmal vein) 10
Forewing hyaline (including those species with a very indistinct suffusion of
yellow or pale brown, or with a small spot beneath marginal vein which does
not or hardly extends past apex of stigmal vein) 12
10 (9) Clava strongly apically obliquely truncate, the truncate part clearly longer than
remaining portion of ventral surface of clava (Figs 60, 62, 65, 67, 69, 239, 318);
pattern of forewing not composed of well-defined stripes and fuscous fasciae 120 (p. 154)
Clava more or less apically rounded or transversely truncate, or if sutures of
clava are oblique and clava thus appears to be obliquely truncate then either
truncate part is shorter than remaining portion of ventral surface of clava or
forewing has a strong pattern of well-defined stripes and fasciae 11
11(10) Hypopygium with apex not reaching more than two-thirds of way along gaster . 134 (p. 156)
Hypopygium reaching apex of gaster 143 (p. 158)
12 (9) Mesoscutum or scutellum or both at least partly yellow, orange or pale orange-
brown 159 (p. 160)
Mesoscutum and scutellum completely dark, not yellow, orange or pale brown 13
13 (12) Face with a pair of longitudinal membranous lines joined below anterior ocellus
by a short transverse membranous line (Fig. 105) (these occasionally obscure
in dry-mounted material because the head collapses inwards; best seen in
slide-mounted specimens) 182 (p. 164)
Face without membranous lines 14
14 (13) Hypopygium with apex not more than four-fifths along gaster, or if more then
exserted part of ovipositor is more than one-third as long as gaster 183 (p. 164)
Hypopygium with apex more or less reaching apex of gaster; ovipositor not or
hardly exserted 205 (p. 168)
15 (7) Malar space very short, not more than one-fifth as long as eye, eye very nearly
reaching base of mandible; body metallic green and often with distinct
punctate sculpture although this may be relatively shallow; notaular lines
absent PARABLASTOTHRIX (p. 314)
Malar space longer, at least one-quarter as long as eye, or if shorter then body
not metallic green, sculpture not punctate or notaular lines present and
complete 16
16 (15) All funicle segments longer than broad 17
INDO-PACIFIC ENCYRTIDAE 141
Not all funicle segments longer than broad, at least one segment quadrate or
transverse 27
17 (16) Forewing infuscate (excluding those species with only a pattern of dark and light
setae, or with an indistinct suffusion of yellow or pale brown, or with a small
spot beneath marginal vein which does not or hardly extends past apex of
stigmal vein) 18
Forewing hyaline (including those species with only a pattern of dark and light
setae, or with an indistinct suffusion of yellow or pale brown, or with a small
spot beneath marginal vein which does not or hardly extends past apex of
stigmal vein) 20
18 (17) Antennal toruli situated relatively high on head and close together so that they
are separated from mouth margin by at least one and one-half times the
minimum distance between them (Fig. 128) 222 (p. 172)
Antennal toruli separated from mouth margin by much less than one and
one-half times the minimum distance between them 19
19 (18) First funicle segment longer than pedicel 225 (p. 172)
First funicle segment not longer than pedicel 238 (p. 176)
20 (17) Either forewing with linea calva not interrupted on dorsal surface or filum
spinosum present or antennal toruli high on head, nearly twice their own
lengths from mouth margin 21
Forewing with linea calva interrupted or closed on dorsal surface of wing by
more than one line of setae and filum spinosum absent and antennal toruli not
more than their own lengths from mouth margin 23
21 (20) Body distinctly dorso-ventrally flattened; pronotum longitudinally divided in
middle (as in Fig. 38) ; ovipositor not or hardly exserted ; mandible bidentate 266 (p. 180)
Either body not dorso-ventrally flattened, or ovipositor exserted and exserted
part at least about half as long as gaster; mandible not bidentate; pronotum
entire 22
22 (21) Either notaular lines present or forewing with submarginal vein having a
strongly swollen parastigma (Figs 148, 150, 151); hypopygium always
reaching apex of gaster ; paratergites usually evident 23
Notaular lines absent; forewing with parastigma not or hardly swollen (Figs 132,
152-154, 156-158, 238) or if conspicuously swollen then hypopygium does not
reach more than halfway along gaster; hypopygium sometimes reaching apex
of gaster; paratergites almost always absent 24
23 (20,22) Notaular lines present in at least anterior part of mesoscutum; linea calva of
forewing not interrupted, although occasionally closed on dorsal surface of
wing; parastigma clearly swollen (Figs 148, 150, 151) 262 (p. 180)
Notaular lines completely absent; linea calva almost always interrupted or
widely closed on dorsal surface of wing; parastigma rarely swollen, usually not
or hardly wider than proximal part of submarginal vein (Figs 91, 95, 159,414) 266 (p. 180)
24 (22) Marginal vein of forewing punctiform or absent 278 (p. 182)
Marginal vein of forewing longer than broad 25
25 (24) Either exserted part of ovipositor at least one-third as long as gaster or
propodeum medially more than one-fifth as long as scutellum 290 (p. 186)
Neither ovipositor with exserted part as long as one-third length of gaster nor
propodeum medially longer than one-fifth length of scutellum 26
26(25) Either mesoscutum or scutellum (including axillae) at least partly orange,
yellow or orange-brown 307 (p. 188)
Both mesoscutum and scutellum (including axillae) dark, not partly orange,
yellow or orange-brown 317 (p. 190)
27 ( 16) Exserted part of ovipositor (measured from apex of last tergite of gaster to apex
of ovipositor) at least as long as one-third length of gaster 28
Ovipositor not exserted, or if exserted then exserted part not longer than
one-quarter length of gaster 29
28 (27) Hypopygium not extending more than three-quarters along gaster 344 (p. 196)
Hypopygium reaching or very nearly reaching apex of gaster 352 (p. 196)
29(27) Either mesoscutum, axillae or scutellum at least partly yellow, orange or
orange-brown 30
142 J. S. NOYES & M. HAY AT
Mesoscutum, axillae and scutellum completely dark, not partly yellow, orange
or orange-brown 32
30 (29) Either notaular lines present in at least anterior part of mesoscutum , or f orewing
infuscate (excluding those species with only a pattern of dark and light setae,
or with an indistinct suffusion of yellow or pale brown, or with a small spot
beneath marginal vein which does not or hardly extends past apex of stigmal
vein) 370 (p. 200)
Notaular lines completely absent; forewing hyaline (including those species with
a pattern of dark and light setae only, or with an indistinct suffusion of yellow
or pale brown, or with a small spot beneath marginal vein which does not or
hardly extends past apex of stigmal vein) 31
31(30) Head completely dark, not yellow, orange or orange-brown and usually metallic 391 (p. 204)
Head at least partly yellow, orange or orange-brown, not metallic 400 (p. 206)
32 (29) Submarginal vein of forewing with a subapical triangular expansion (usually
indicated by a single, long, semi-erect seta) (Figs 107, 109, 207) 415 (p. 208)
Submarginal vein of forewing without a subapical triangular expansion 33
33 (32) First funicle segment longer than broad 34
First funicle segment not longer than broad 40
34 (33) Mesoscutum with complete notaular lines (Fig. 6) HOMALOTYLUS (p. 287)
Mesoscutum without notaular lines 35
35 (34) Marginal vein of forewing punctiform or absent 418 (p. 208)
Marginal vein of forewing longer than broad 36
36 (35) Hind tibia foliaceously flattened and expanded, not more than two and one-half
times as long as broad (Fig. 213) NEOCLADIA (p. 306)
Hind tibia not expanded and flattened, or if slightly so then at least three times as
long as broad 37
37 (36) Linea calva completely obliterated on both dorsal and ventral surfaces of
forewing by short, dense setae so that forewing is densely and evenly hairy
from base to apex (Fig. 214) NATHISMUSIA (p. 302)
Forewing with linea calva not obliterated 38
38 (37) Forewing infuscate (excluding those species with an indistinct suffusion of
yellow or pale brown, or with a small spot beneath marginal vein which does
not or hardly extends past apex of stigmal vein) 39
Forewing hyaline (including those species with an indistinct suffusion of yellow
or pale brown, or with a small spot beneath marginal vein which does not or
hardly extends past apex of stigmal vein 434 (p. 212)
39 (38) First funicle segment at least as long as pedicel 457 (p. 214)
First funicle segment shorter than pedicel 464 (p. 214)
40 (33) Frontovertex with distinct piliferous punctures which give a thimble-like
appearance, if punctures shallow then generally separated by not more than
their own diameters 475 (p. 215)
Frontovertex without deep and distinct piliferous punctures, and not with
appearance of surface of a thimble 41
41 (40) Forewing infuscate (excluding those species with an indistinct suffusion of
yellow or pale brown, or with a small spot beneath marginal vein which does
not or hardly extends past apex of stigmal vein) 481 (p. 215)
Forewing hyaline (including those species with an indistinct suffusion of yellow
or pale brown, or with a small spot beneath marginal vein which does not or
hardly extends past apex of stigmal vein) 42
42 (41) Scutellum very convex with fine reticulate or reticulate-striate sculpture of a
matt or silky appearance; all funicle segments transverse except occasionally
the sixth (Figs390, 395) PARABLATTICIDA (p. 314)
Scutellum either not convex or without a reticulate-striate sculpture of silky
appearance; if appearing slightly convex and with silky appearance then only
first funicle segment is not longer than broad 43
43 (42) Marginal vein of forewing punctiform 490 (p. 216)
Marginal vein of forewing longer than broad 44
44 (43) Hypopygium more or less reaching apex of gaster 499 (p. 217)
INDO-PACIFIC ENCYRTIDAE 143
Hypopygium not reaching more than four-fifths along gaster 510 (p. 218)
45 (1) Antenna with two to four anelliform segments that are adpressed with clava,
clava large, at least as long as remainder of antenna (Fig. 9); forewing broad,
at most two and one-quarter times as long as broad, with marginal fringe
much shorter than maximum wing width (Fig. 10); mandible with a single
pointed tooth ARRHENOPHAGUS (p. 235)
Antenna with five or six funicle segments that are clearly separated from clava,
clava at most as long as funicle and pedicel combined (Fig. 13); forewing
narrow, not less than three and one-half times as long as wide, with marginal
fringe at least as long as wing width (Fig. 12); mandible with apex broadly
truncate or serrate (Fig. 14) ANTHEMUS (p. 233)
46 (3) Forewing hyaline 47
Forewing infuscate 52
47 (46) Funicle three-segmented 48
Funicle four-segmented 49
48 (47) Frontovertex with a transverse membranous line between anterior ocellus and
antennal toruli, this joined to antennal toruli, or nearly so, by longitudinal
membranous lines (Fig. 16); funicle segments strongly transverse and closely
adpressed together, clava solid, apically obliquely truncate and much longer
than pedicel and funicle together (Fig. 15) ARRHENOPHAGOIDEA (p. 235)
Frontovertex without any membranous lines; funicle segments clearly separated
and each quadrate or slightly longer than broad, clava three-segmented, not
obliquely truncate and slightly shorter than pedicel and funicle together
MARXELLA (p. 295)
49 (47) Funicle segments all longer than broad (Fig. 17); forewing with marginal vein
shorter than stigmal (Fig. 18) ; hypopygium reaching apex of gaster or beyond
CERCOBELUS (p. 247)
Not all funicle segments longer than broad, usually broader than long or
quadrate; forewing with marginal vein as long as or longer than stigmal;
hypopygium not extending to apex of gaster 50
50 (49) Clava two-segmented; mandibles with three acute teeth NASSAUIA (p. 302)
Clava three-segmented; mandibles with one or two teeth and a truncation or
four teeth 51
51 (50) First funicle segment longer than broad and at least a little longer than the fourth
COCCIDENCYRTUS (p. 253)
First funicle segment clearly shorter than fourth and transverse .... PLAGIOMERUS (p. 325)
52(46) Forewing more or less uniformly infuscate, without sharply delimited rays,
bands or spots; hypopygium extending to apex of gaster BRACHYPLATYCERUS (p. 243)
Forewing either with infuscate rays or bands , or infuscate with hyaline patches 53
53(52) All antennal segments flattened, clava obscurely two-segmented (Fig. 19);
scutellum without any apical lamelliform setae.
Forewing as in Fig. 20 SPANIOPTERUS (p. 338)
At most only scape flattened with flagellar segments cylindrical, clava three-
segmented ; apex of scutellum with at least one pair of lamelliform setae .... 54
54 (53) All funicle segments longer than broad HOMALOPODA (p. 287)
Not all funicle segments longer than broad, at least first two segments transverse
CAENOHOMALOPODA (p. 243)
55 (3) Antennal flagellum flattened; forewing with an infuscate band ANARHOPUS (p. 231)
Flagellum more or less cylindrical, not flattened; forewing hyaline or lightly
infuscate, without a distinct band 56
56 (55) Body dorso-ventrally flattened; pronotum longitudinally divided NEORHOPUS (p. 307)
Body robust, not dorso-ventrally flattened but if so then pronotum entire 57
57 (56) Wings shortened, not reaching apex of gaster; clava three-segmented 58
Either wings fully developed and reaching apex of gaster, or clava entire 59
58 (57) Body entirely yellow ZEALANDENCYRTUS (p. 350)
Body at least partly dark and metallic TETRACNEMOIDEA (p. 341)
59 (57) Forewing with area immediately below venation from proximal part of para-
stigma to apex of stigmal vein completely naked and continuous with the
144 J. S. NOYES & M. HAYAT
relatively wide linea calva which is conspicuously broader than length of
marginal vein (Fig. 22); mandible bidentate.
Cf antenna branched (Fig. 21) TETRACNEMOIDEA (p. 341)
Forewing with area immediately below distal one-third of venation with several
setae and not naked, linea calva not or hardly broader than length of marginal
vein; mandible with three teeth or one or two teeth and a truncation 60
60 (59) Head or thorax at least partly yellow or orange 61
Head and thorax dark, often shiny and metallic 67
61 (60) Clavasolid (Fig. 23) 62
Clava two- or three-segmented 63
62 (61) Body dorso-ventrally flattened; head prognathous; pronotum more than half as
longasmesoscutum(Fig.24) INDAPHYCUS (p. 289)
Body not dorso-ventrally flattened; head hypognathous; pronotum much shor-
ter than one-third length of mesoscutum ACEROPHAGUS (p. 220)
63 (61) Clava two-segmented (Fig. 28) PSEUDECTROMA (p. 329)
Clava three-segmented 64
64(63) Notaular lines present in anterior one-third of mesoscutum; ovipositor not
exserted ;hypopygium not quite reaching apex of gaster BEETHOVENA (p. 241)
Notaular lines absent; exserted part of ovipositor at least as long as one-fifth
length of gaster; hypopygium reaching apex of gaster 65
65 (64) Head and thorax clothed with conspicuous dark setae; scape not longer than
minimum width of frontovertex; antennal toruli separated from mouth
margin by about their own lengths; forewing with postmarginal vein about as
long as stigmal; mandible with two teeth and a truncation MOZARTELLA (p. 300)
Head and thorax clothed with pale or silvery white setae, or if setae dark then
scape is longer than minimum width of frontovertex, antennal toruli are
nearly at mouth margin being separated from it by much less than their own
lengths (Fig. 25) and forewing with postmarginal vein clearly shorter than
stigmal; mandible with three acute teeth 66
66 (65) Antenna unicolorous, yellow or orange ACEROPHAGUS (p. 220)
Clava at least partly white contrasting with brown or yellowish brown segments
offunicle(Fig. 26)
Forewing as in Fig. 27 PSEUDAPHYCUS (p. 328)
67 (60) Forewing with postmarginal vein at least about twice as long as stigmal
HOLCOTHORAX (p. 287)
Forewing with postmarginal vein not or hardly longer than stigmal 68
68(67) Clava transversely or obliquely truncate; notaular lines completely absent;
forewing with sensillae at apex of stigmal vein arranged symmetrically in a
square 69
Either clava apically rounded or notaular lines present; forewing with sensillae
at apex of stigmal vein arranged asymmetrically, not in a square 70
69 (68) Clava entire with apex strongly obliquely truncate. (Fig. 29)
Base of forewing as in Fig. 30 COPIDOSOMOPSIS (p. 258)
Clava three-segmented with apex more or less transversely truncate . RAFFAELLIA (p. 332)
70 (68) Notaular lines absent; exserted part of ovipositor at least one-fifth as long as
gaster 71
Notaular lines present; ovipositor not or hardly exserted 72
71 (70) Mandible with three acute teeth ; forewing with postmarginal vein a little shorter
than stigmal PARARHOPELLA (p. 318)
Mandible with one or two teeth and a truncation; forewing with post marginal
vein slightly longer than stigmal MESORHOPELLA (p. 297)
72 (70) Forewing with marginal vein more or less absent, venation not quite touching
anterior margin of wing, submarginal vein with parastigma not conspicuously
swollen (Fig. 31); scutellum always lustrous blue or green TRECHNITES (p. 345)
Forewing with venation touching anterior margin of wing and marginal vein
more or less quadrate, submarginal vein with parastigma conspicuously
swollen (Fig. 32); scutellum dull COCCIDAPHYCUS (p. 253)
73 (5) Propodeum medially at least one-third as long as scutellum (Fig. 33) 74
BRITISH
A review of the genera of Indo-Pacific Encyrtidae
(Hymenoptera: Chalcidoidea)
John S. Noyes
Department of Entomology, British Museum (Natural History), Cromwell Road, London
SW7 5BD
M. Hayat
Department of Zoology, Aligarh Muslim University, Aligarh, Uttar Pradesh 202 001, India
Contents
Synopsis ................................................................ 131
Introduction ............................................................ 131
Notes on generic review ................................................... 134
Notes on key ............................................... ............. 135
Notes on terms and measurements .......................................... 135
Abbreviations ........................................................... 138
Key to genera (females) ................................................... 140
Notes on genera ......................................................... 220
Incertae sedis ........................................................... 352
Systematic relationships of Indo-Pacific encyrtid genera ....................... 353
Host index ...................................................... ........ 355
Proposed new synonymies ................................................ 357
Proposed new combinations ............................................... 359
Proposed new status ...................................................... 365
Replacement names ...................................................... 365
Lectotype designations ................................................... 365
Acknowledgements ...................................................... 366
References ............................................................. 366
Index . ................................. 383
Synopsis
A key to females of the 263 described genera of Encyrtidae recognised from the Indo-Pacific region is
provided. Notes on each genus are included and give information on known world distribution, number of
described species, distribution of each genus within the area under review, a list of species known from the
region, references to original descriptions, redescriptions, revisions or other useful papers, biology, and
systematic placement of the genus. Lectotypes are designated for 44 species; 23 genera and 18 species are
described as new; one subtribe and one subspecies are raised to tribe and species level respectively; one
tribal, one subtribal, 107 generic and 41 specific synonymies, 358 combinations and three replacement
names for junior specific homonyms are newly proposed.
Introduction
The importance of the Hymenoptera Parasitica in biological control programmes is unquestion-
able . Clausen (1978) reviews a large amount of literature dealing with the introduction of natural
enemies to control weeds and pest species of arthropods. A brief scan through this review soon
reveals that the majority of insect species introduced to control pests are parasitic Hymenoptera,
and that the most important of these are the Chalcidoidea. Perhaps an indicator of the
importance of the Chalcidoidea in the field of biological control is Biocontrol News and
Bull. Br. Mus. not. Hist. (Ent.) 48 (3): 131-395 Issued 28 June 1984
132 J. S. NOYES & M. HAYAT
Information (published by Commonwealth Agricultural Bureaux, Slough, England), a review
of literature relevant to all forms of biological control. Of all the papers reviewed, no fewer than
16 per cent contain references to chalcids. Within the Chalcidoidea, the most important families
in this context are the Aphelinidae, Encyrtidae and Trichogrammatidae, species of which are
most commonly used to control lepidopterous and hemipterous pests. Of the seven major,
successful biological control projects listed by Bosch et al. (1982) for California, three have
utilised species of Encyrtidae as the controlling agent. That is not to say that species of
Encyrtidae are the main controlling agent for 40 per cent of all successful biological control
projects, but merely to illustrate that they are, economically, a very important group.
It is essential to be able to identify species accurately in order to convey information about
useful or potentially useful species. An important step facilitating the accurate identification of
species is a stable classification at the generic and possibly tribal level. Thus, the present review
has three aims. Firstly, to attempt to arrange the many poorly understood Australian species and
genera of Encyrtidae into some general pattern which agrees as closely as possible with
Trjapitzin's (1973fl,ft) classification of the group. Secondly, to bring together all relevant
taxonomic information available on the Encyrtidae of the Indo-Pacific region. Thirdly, to
facilitate the identification of material collected in this region.
The Indo-Pacific region is defined here as the area south of a line drawn from the north-
ernmost tip of Pakistan to the Hawaiian Islands (also north to Midway Island). This therefore
excludes Japan and Korea, but includes southern China, the Pacific islands, Australia and New
Zealand. Keys to the genera of the region have been published previously by Girault (19150) for
Australia, Beardsley (1976) for the Hawaiian Islands, Hayat etal. (1975), Shafee et al. (1975)
and Alam & Shafee (1982) for India. Unfortunately most of these keys are now obsolete or very
incomplete.
The fauna of the Indian subcontinent is probably the best known of any within the region,
except perhaps that of Australia. Even so, despite the work of earlier authors, e.g. Howard (in
Howard & Ashmead, 1896), Gahan (1914), Ayyar & Margabandhu (1934a,ft) and Mani (1935;
1939; 1941), only 30 genera and 50 species had been recorded from there by the middle of the
present century. Later work by other authors, e.g. Subba Rao (1957; 1967), Agarwal (1965),
Mani etal. (1973; 1974), Hayat etal. (1975), Shafee etal. (1975), added many more species and
genera. Several papers have since been published to clarify the systematic position of many
Indian genera and species, notably those of Subba Rao (1976) and Hayat (1979ft; 1981a,ft).
More recently Hayat & Subba Rao (1981) listed 117 genera and 276 species from the Indian
subcontinent.
In contrast, largely as a result of the work of A. A. Girault (1911-1941), the number of genera
and species 'known' from Australia is much greater. Girault alone described some 150 genera
and 347 species of Encyrtidae from that continent. Further species have been described by other
authors, e.g. Walker (1839), Howard (1898ft), Dodd (1917), Timberlake (1929), Compere
(1940) and Ferriere (1947). However, until recently, most of Girault's taxa have remained
unrecognised, mainly because of his inadequate descriptions and poor treatment of material,
and the inaccessibility of his type-material to taxonomists outside Australia. Fortunately the
work of E. C. Dahms at the Queensland Museum, Brisbane has now enabled a number of
specialists to study the Girault Australian type-material, e.g. Boucek (all families except
Encyrtidae, Aphelinidae and Mymaridae), De Bach & Rosen, Hayat (Aphelinidae), New
(Mymaridae), and Gordh & Dahms (Encyrtidae). The work of Gordh (UCR) & Dahms (QM)
overlaps with the present review since it includes detailed, illustrated, redescriptions of all
encyrtid genera described by Girault from Australia. Unfortunately it is not yet available but
should be published shortly after the present review. Therefore we are unable to include
comment on their opinions concerning these genera and many of the species included in them by
Girault. However, in discussion with both Gordh and Dahms it is apparent that there is a large
measure of agreement between us concerning the status of many of Girault's genera and the
placement of most species, but at the same time there is also some disagreement. The latter is
inevitable considering the state of many of Girault's types, but at least it may show where future
INDOPACIFIC ENCYRTIDAE
145
Figs 9-18 9-11, Arrhenophagus sp., (9) right antenna, outer aspect, $, (10) right forewing, upper
surface, $, (11) head, frontal aspect, $; 12-14, Anthemus maculatus Subba Rao, (12) right forewing,
upper surface, $ , (13) right antenna, outer aspect, $ (14) right mandible, $ ; 15, 16, Arrhenophagoidea
bicoloripes Girault, (15) right antenna, outer aspect, $ , (16) head, frontal aspect, $ ; 17, 18, Cercobdus
jugaeus (Walker) (extra-limital species), (17) left antenna, inner aspect, $, (18) apex of right forewing
venation, upper surface, $ .
146 J. S. NOYES & M. HAYAT
Propodeum medially not more than one-fifth as long as scutellum (Figs 34, 35) . 77
74 (73) Antenna with scape broadened and flattened, not more than three times as long
as broad 75
Antenna with scape not strongly flattened, at least five times as long as broad . . 76
75 (74) Clava solid ; scutellum concave with a line of scale-like setae at apex
COELASPIDIA (p. 225)
Clava three-segmented, scutellum flat or convex without an apical line of
scale-like setae XENANUSIA (p. 347)
76(74) Pronotum long, medially clearly much longer than mesoscutum, mandible
bidentate SCHILLERIELLA (p. 338)
Pronotum medially shorter than mesoscutum (Fig. 33); mandible with three
teeth SAKENCYRTUS (p. 336)
77 (73) Antenna with all segments broadened and flattened MIRA (p. 299)
Antenna with pedicel and flagellum more or less cylindrical, scape occasionally
broadened and flattened 78
78(77) All funicle segments longer than pedicel (Fig. 37); either funicle seven-
segmented and clava two-segmented, or flagellum not differentiated into
funicle and clava ANOMALICORNIA (p. 232)
Not all funicle segments longer than pedicel; funicle six-segmented and clava
two- or three-segmented 79
79 (78) Visible part of mesoscutum at least three times as broad as long (Fig. 34) or
mesoscutum completely hidden by pronotum 80
Visible part of mesoscutum not more than two and one-half times as broad as
long 81
80 (79) Wings moderately long and capable of meeting at apex of scutellum; fronto-
vertex at narrowest point not more than one and one-half times as broad as
length of scape; mandible with three teeth AUSTROCHOREIA (p. 237)
Wings very short, clearly not capable of meeting at mid-line; frontovertex at
narrowest point twice as wide as length of scape; mandible bidentate
NEODUSMETIA (p. 306)
81 (79) Antennal toruli very high on head, separated from mouth margin by more than
their own lengths; head and thorax covered with very conspicuous dark setae;
mandible with one or two teeth and a truncation HUNTERELLUS (p. 288)
Antennal toruli separated from mouth margin by less than their own lengths;
head and thorax not conspicuously hairy; mandible bidentate 82
82 (81) Body not dorso-ventrally flattened; pronotum entire (Fig. 35) 83
Body dorso-ventrally flattened; pronotum longitudinally divided in middle
(Fig. 38) 84
83 (82) Antennal flagellum with brown and white segments (Fig. 36); posterior margin
of eye straight or slightly convex CREMESINA (p. 260)
Antennal flagellum unicolorous, dark brown; posterior margin of eye concave
so that eye has a kidney-shaped appearance PARECTROMOIDELLA (p. 319)
84 (82) Eye larger, longer than malar space (Fig. 40) RHOPUS (p. 332)
Eye smaller, at least a little shorter than malar space (Fig. 39) . HAMUSENCYRTUS (p. 283)
85 (5) Antenna with all segments distinctly broadened and flattened (Fig. 41)
CERAPTEROCERUS (p. 245)
Antenna with pedicel and flagellum more or less cylindrical, scape occasionally
broadened and flattened 86
Figs 19-30 19, 20, Spaniopterus crucifer Gahan, (19) right antenna, outer aspect, 9 > (20) left forewing,
upper surface, $ ; 21, 22, Tetracnemoidea indica (Ayyar), (21) left antenna, outer aspect, cf , (22) base of
left forewing, upper surface, $; 23, Acerophagus solidus Hayat, left antenna, inner aspect, $; 24,
Indaphycus planus Hayat, pronotum, mesoscutum and scutellum, dorsal aspect, $; 25, Pseudaphycus
utilis Timberlake, head, frontal aspect, 9 ; 26, 27, Pseudaphycus orientalis Ferriere, (26) right antenna,
outer aspect, $, (27) base of right forewing, upper surface, $; 28, Pseudectroma sp., right antenna,
inner aspect (clava slightly collapsed), 9 ; 29, 30, Copidosomopsis nacoleiae (Eady), (29) right antenna,
outer aspect showing truncate sensory surface at apex of clava, $, (30) base of left forewing, upper
surface, $.
INDO-PACIFIC ENCYRTIDAE
147
20
148 J. S. NOYES & M. HAY AT
86 (85) Scutellum with a subapical group of dark coarse setae arranged in a more or less
compact bundle (as in Fig. 47) 87
Scutellum without such a group of setae 88
87 (86) Mesoscutum with a distinct transverse depression in its posterior one-third;
either mesoscutum with a more or less distinct bundle of setae in middle or
posterior margin or pronotum has a line of stiff black bristles . . DIVERSINERVUS (p. 265)
Mesoscutum without a transverse posterior depression; neither mesoscutum
with a median bundle of setae nor posterior margin of pronotum with a line of
stiff black bristles CHEILONEURUS (p. 249)
88 (86) Mesoscutum (including part hidden by pronotum) strongly transverse, at least
about three times as broad as long and entirely or almost entirely covered by
posterior margin of pronotum; mandible with three acute teeth
AUSTROCHOREIA (p. 237)
Mesoscutum (including part hidden by pronotum) not or hardly more than twice
as broad as long and only slightly covered by pronotum anteriorly, or if about
three times as broad as long then mesoscutum only slightly covered by
pronotum anteriorly and mandible with one or two teeth and a truncation ... 89
89 (88) Thorax entirely dark and metallic, not partly yellow or orange 90
Thorax at least partly yellow or orange 92
90 (89) Clava with a strong oblique apical truncation (as in Fig. 43) ; posterior margin of
mesoscutum more or less straight and not projecting over axillae so that when
thorax viewed from above the axillae more or less meet (Fig. 42); gaster
entirely dark; mandible with three acute teeth HYPERGONATOPUS (p. 288)
Clava apically more or less rounded (Fig. 45); posterior margin of mesoscutum
covering axillae in middle so that when thorax viewed from above the axillae
appear to be widely separated (Fig. 44); gaster often orange or yellow at base;
mandible usually with one or two teeth and a truncation, although occasion-
ally obscurely tridentate 91
91 (90) Forewing with at least apex infuscate XENOENCYRTUS (p. 348)
Forewing hyaline OOENCYRTUS (p. 309)
92 (89) Scutellum with a thin apical flange PARAPHAENODISCUS (p. 317)
Scutellum without a distinct apical flange 93
93 (92) Wing entirely hyaline ECTROMA (p. 268)
Wing infuscate 94
94 (93) Pronotum with a pair of distinct, sublateral, elongate white spots
*PROCHEILONEURUS Girault (p. 326)
Pronotum unicolorous, without a pair of sublateral white spots MICROTERYS (p. 299)
95 (6) Scutellum with two or more scale-like setae 96
Scutellum with a group of coarse, long, dark setae arranged in a more or less
compact bundle 98
96 (95) Apical one-third or so of scutellum with a few short, scale-like setae and with a
pair of slightly larger scale-like setae at apex (Fig. 46); forewing more or less
uniformly infuscate; head and thorax mostly yellow LAKSHAPHAGUS (p. 291)
Apex of scutellum with conspicuously longer, more distinctly scale-like setae
than remainder, these occasionally very large and up to 12 or more in number
(Fig. 48); forewing infuscate with well-defined hyaline areas; body wholly
dark and metallic 97
* Not to be contused with Prochiloneurus Silvestri (p. 327)
Figs 31-41 31, Trechnites flavipes (Mercet) (extra-limital species), base of right forewing, upper surface,
$ ; 32, Coccidaphycus sp. , base of right forewing, upper surface, $ ; 33, Sakencyrtus sp. , thorax, dorsal
aspect, 9; 34, Neodusmetia sangwani (Subba Rao), thorax, dorsal aspect, ; 35, 36, Cremesina spp.,
(35) thorax, dorsal aspect, $ , (36) right antenna, outer aspect, $ ; 37, Anomalicornia sp. , right antenna,
outer aspect, $ ; 38, 39, Hamusencyrtus mymaricoides (Compere, Subba Rao & Kaur), (38) pronotum,
mesoscutum and scutellum, dorsal aspect, 9 , (39) head, frontal aspect, $ ; 40, Rhopus sp. , head, frontal
aspect, $; 41, Cerapterocerus mirabilis Westwood (extra-limital species), right antenna, outer aspect,
INDO-PACIFIC ENCYRTIDAE
149
31
150
J. S. NOYES & M. HAY AT
97 (96) Apex of scutellum with about 10 to 14 long, slightly flattened, scale-like setae
arranged in a line RUSKINIANA (p. 336)
Apex of scutellum with at most two pairs, usually only with one, of slightly to
strongly broadened and flattened scale-like setae (Fig. 48) HABROLEPIS (p. 281)
98 (95) Mesoscutum with a group of coarse, long, dark setae arranged in a more or less
compact bundle and with a transverse depression in posterior one-third which
bears silvery white setae DIVERSINERVUS (p. 265)
Mesoscutum without such a bundle of setae, posterior one-third without a
transverse depression, although occasionally with silvery white setae 99
99 (98) Forewing with marginal vein at most only a little longer than broad, several
times shorter than either stigmal or postmarginal veins; mandible edentate
witharounded, sharp edge ENCYRTUS (p. 268)
Forewing with marginal vein at least nearly as long as stigmal; mandible with
three teeth or two teeth and a truncation 100
100 (99) Forewing hyaline 101
Forewing infuscate 102
101 (100) Forewing with marginal vein at least three times as long as stigmal, parastigma
strongly downcurved (Fig. 50) CHEILONEURUS (p. 249)
Forewing with marginal vein only slightly longer than stigmal, parastigma
normal (Fig. 49) ZAOMMA (p. 349)
102 (100) Forewing with a pair of interrupted hyaline fasciae distad of apex of venation,
marginal vein not longer than stigmal MICROTERYS (p. 299)
Forewing without hyaline fasciae distad of apex of venation, marginal vein at
least twice as long as stigmal 103
103 (102) Hypopygium extending to apex of gaster; ovipositor always strongly exserted,
the exserted part at least one-third as long as gaster
*PROCHILONEURUSSilvestri (p. 327)
Hypopygium not extending more than three-quarters along gaster; either
ovipositor not or hardly exserted, or if strongly so then hypopygium hardly
extends more than halfway along gaster CHEILONEURUS (p. 249)
104 (8) Forewing with a pattern of infuscate rays or bands 105
Forewing hyaline or more or less uniformly infuscate with one or two hyaline
spots or bands, not with infuscate rays or bands 109
105 (104) Forewing with one or two longitudinal infuscate rays (Fig. 303) .... COMPERIELLA (p. 256)
Forewing with one or two fuscous fasciae or with several fuscous lines radiating
from a longitudinal fuscous line in centre of wing between which are wedge-
shaped hyaline spots (Fig. 292) 106
106 (105) Hypopygium reaching apex of gaster; mandible bidentate 107
Hypopygium not reaching more than halfway along gaster; mandible with three
teeth 108
107 (106) Forewing with a central longitudinal fuscous line from which radiate several
fuscous lines between which are wedge-shaped triangular spots; scape more
or less rectangular in profile XENANUSIA (p. 347)
Forewing with fuscous fasciae; scape triangular in profile EPANUSIA (p. 271)
108 (106) Scape triangular in shape (Fig. 289); submarginal vein of forewing without a
subapical triangular expansion (Fig. 290) CERAPTEROCEROIDES (p. 245)
* Not to be confused with Procheiloneurus Girault (p. 326)
Figs 42-53 42, 43, Hypergonatopus hawaiiensis (Perkins), (42) pronotum, mesoscutum and scutellum,
dorsal aspect (from card-mounted specimen), $, (43) left antenna, outer aspect (from card-mounted
specimen), 9; 44, 45, Xenoencyrtus niger Riek, (44) pronotum, mesoscutum and scutellum, dorsal
aspect, $, (45) right antenna, outer aspect, $; 46, Lakshaphagus daulai (Shafee, Alam & Agarwal),
scutellum, dorsal aspect (from card-mounted specimen), $; 47, Cheiloneurus pyrillae Mani, scutellum,
dorsal aspect, 9; 48, Habrolepis rouxi Compere, scutellum, dorsal aspect, $; 49, Zaomma sp., base of
right forewing, upper surface, $; 50, Cheiloneurus sp., base of right forewing, upper surface (from
card-mounted specimen), $; 51, Eusemion cornigerum (Walker), right antenna, outer aspect, $; 52,
Anicetus integrellus Trjapitzin, left antenna, outer aspect, $ ; 53, Leurocerus hongkongensis Subba Rao,
right antenna, outer aspect, 9-
INDO-PACIFIC ENCYRTIDAE
151
152
J. S. NOYES & M. HAY AT
Scape more or less rectangular with dorsal and ventral margins subparallel;
submarginal vein of forewing with a subapical triangular expansion (Fig. 291)
CERAPTEROCERUS (p. 245)
109 (104) Hypopygium not reaching more than about two-thirds along gaster; mandible
with two teeth and a truncation, or three or four teeth 110
Hypopygium reaching apex of gaster; mandible with two, rarely three, teeth . . 114
110 (109) Forewing hyaline NEOCLADELLA (p. 305)
Forewing darkened Ill
111 (110) Body dark and metallic, not yellow or orange 112
At least head and thorax largely yellow or orange 113
112 (111) Forewing entirely infuscate, the infuscation gradually fading towards apex of
wing; clava entire (Fig. 53); marginal vein of forewing punctiform LEUROCERUS (p. 293)
Forewing with apex broadly hyaline; clava three-segmented (Fig. 51); marginal
vein of forewing more than twice as long as broad EUSEMION (p. 277)
113 (111) Scape tending to be subrectangular, the flattened part of upper edge more than
one-half as long as the straight part of the lower edge . . PARACERAPTROCERUS (p. 315)
Scape tending to be triangular, the flattened part of the upper edge less than half
as long as the straight part of the lower edge (Fig. 52) ANICETUS (p. 231)
114 (109) Forewing with postmarginal vein well developed, at most only about one-third
shorter than stigmal; pedicel usually longer and broader than first funicle
segment 115
Forewing with postmarginal vein very short or absent; pedicel narrower than
and at most about as long as first funicle segment 117
115 (114) Forewing with basal cell as densely and as evenly hairy as disc, linea calva closed
towards posterior margin, wing with a well-defined but irregular pattern;
thorax with punctate-reticulate sculpture and matt; facial carina dorsally with
two or three lines of very short, white squamous hairs CERAPTROCERELLA (p. 246)
Forewing with basal cell naked proximally, linea calva more or less open
posteriorly (Fig. 55), wing more or less evenly infuscate except in proximal
one-quarter where it is more or less hyaline; thorax with very shallow
sculpture and slightly to very shiny; facial carina without a distinct line of pale
setae dorsally 116
116(115) Forewing with proximal margin of linea calva with at least a few flattened
scale-like setae (Fig. 55); antennal flagellum in profile with subparallel sides
(Fig. 54) CHRYSOPLATYCERUS (p. 250)
Forewing with proximal margin of linea calva without any flattened scale-like
setae; antennal flagellum distinctly oval in profile (Fig. 56) ... NEOPLATYCERUS (p. 306)
117(114) Scutellum with a distinct, thin apical flange (Fig. 58); pedicel only slightly
shorter than first funicle segment, clava solid (Fig. 57) PRALEUROCERUS (p. 325)
Scutellum without a distinct apical flange; pedicel very small, much shorter than
first funicle segment, clava three-segmented 118
118 (117) Head prognathous and in frontal view elongate, nearly one-half longer than
broad MONSTRANUSIA (p. 300)
Head hypognathous and in frontal view about as long as broad 119
119 (118) First funicle segment at least three times as broad as pedicel which is triangularly
flattened, the distal segments narrowing but still at least about twice as wide as
pedicel; forewing with postmarginal vein very short, almost absent
CRYPTANUSIA (p. 262)
First funicle segment subequal in size to sixth, both much less than twice as
Figs 54-64 54, 55, Chrysoplatycerus splendens (Howard), (54) right antenna, outer aspect, $ (55) left
forewing, upper surface, $; 56, Neoplatycerus sp., left antenna, outer aspect (from card-mounted
specimen), $; 57, 58, Praleurocerus viridis (Agarwal), (57) right antenna, outer aspect, $, (58)
scutellum and propodeum, dorsal aspect, $ ; 59, 60, Proleurocerus fulgoridis Ferriere, (59) apex of right
forewing venation, upper surface, 9 (60) right antenna, inner aspect, $ ; 61 , Zozoros sinemarginissp.
n., base of right forewing, upper surface, 9; 62-64, Hambletonia pseudococcina Compere, (62) right
antenna, outer aspect, $ , (63) base of right forewing, upper surface, $ , (64) head, dorso-f rental aspect,
INDO-PACIFIC ENCYRTIDAE
153
154
J. S. NO YES & M. HAYAT
broad as pedicel which is subconical; forewing with postmarginal vein at least
about half as long as stigmal PARECTROMOIDELLA (p. 319)
120 (10) Forewing either with marginal vein absent, the postmarginal and stigmal veins
emitted from apex of submarginal with postmarginal vein not touching
margin of wing, or marginal vein punctiform (or occasionally slightly longer
than broad) and apex of stigmal vein joined to apex of postmarginal vein by a
distinct, often hyaline, hairless streak (Figs 59, 61, 63) 121
Forewing either with marginal vein distinctly longer than broad or with marginal
vein punctiform (or slightly longer than broad) and without distinct naked
streak from apex of postmarginal vein to apex of stigmal vein 125
121 (120) Clava shorter than funicle, about as long as preceding three funicle segments
together PENTELICUS (p. 322)
Clava at least as long as funicle, usually longer 122
122 (121) Pedicel with a few conspicuous long scale-like setae, clava large and oval
(Fig. 63); facial impression margined above by a sharp ridge (Fig. 64)
HAMBLETONIA (p. 282)
Pedicel with normal setae, clava not oval; facial impression at most with a
rounded edge 123
123 (122) Head smooth with very fine punctures; mandible bidentate LUTHERISCA (p. 294)
Head with deep , conspicuous piliferous punctures ; mandible with three teeth . 124
124 (123) Clava solid, funicle segments not less than twice as broad as long (Fig. 60); body
wholly dark and metallic, not partly yellow-brown PROLEUROCERUS (p. 328)
Clava three-segmented, funicle segments from slightly transverse to clearly
longer than broad (Fig. 65); body partly yellow-brown ZOZOROS (p. 350)
125 (120) Exserted part of ovipositor at least about one-third length of gaster
*PROCHILONEURUSS\\\estri (p. 327)
Ovipositor not or hardly exserted 126
126 (125) All funicle segments broader than long 127
Not all funicle segments broader than long, at least first funicle segment longer
than broad 132
127 (126) Hypopygium extending to apex of gaster; forewing with marginal vein shorter
than stigmal 128
Hypopygium not extending to apex of gaster; forewing with marginal vein at
least a little longer than stigmal 130
128 (127) Head and dorsum of thorax with fine punctate-reticulate sculpture and of matt
or velvety appearance; facial impression bordered above by a very strong,
almost straight transverse carina extending from gena to gena; pedicel
dorsally flattened and shiny CERAPTROCERELLA (p. 246)
Head and dorsum of thorax with shallow reticulate and shallow to moderately
deep piliferous punctures which often give a thimble-like appearance; face
without a strong transverse carina (although antennal scrobes may be very
sharply margined); pedicel not flattened dorsally and not shiny 129
129 (128) Frontovertex one-sixth to one-third head width, head with punctures descend-
ing at least some way between eye and facial impression; mandible bidentate
AENASIUS (p. 225)
Frontovertex less than one-sixth head width, head only with fine punctures
between eye and facial impression; mandible tridentate NEODISCODES (p. 306)
* Not to be confused with Procheiloneurus Girault (p. 326)
Figs 65-77 65, Zozoros sinemarginis sp. n., right antenna, outer aspect, 9 ; 66, Doddanusia sp. , base of
right forewing, upper surface, $f; 67, 68, Ovaloencyrtus fijiensis sp. n., (67) right antenna, outer aspect,
9 , (68) base of right forewing, upper surface, 9 ; 69-71 , Paratetralophidea sp. , (69) right antenna, outer
aspect, 9> (70) left forewing showing pattern and relative strength of infuscation, $, (71) head, frontal
aspect, $ ; 72, Epitetracnemus zetterstedtii (Westwood), head, aspect from left side, 9 ; 73, Paksimmond-
sius pakistanensis Ahmad & Ghani, base of right forewing, upper surface, 9; 74, Psyllaephagus
worcesteri (Girault), left mandible, 9; 75, Psyllaephagus dyari (Girault), right mandible, $; 76,
Aenasiella brachyscelidis Girault, right mandible, 9; 77, Lakshaphagus hautefeuilli (Mahdihassan),
apex of right forewing venation, upper surface, 9
INDO-PACIFIC ENCYRTIDAE
155
133
156 J. S. NOYES & M. HAYAT
130 (127) Thorax, excluding legs, entirely dark and metallic NEBLATTICIDA (p. 304)
Thorax, excluding legs, largely yellow or orange 131
131 (130) Forewing generally suffused pale brown without any hyaline areas, although
occasionally paler towards apex of wing, marginal vein less than twice as long
as stigmal, filum spinosum in posterior half of wing (Fig. 66) DODDANUSIA (p. 265)
Forewing with at least proximal one-third hyaline, distally strongly infuscate but
usually with some paler areas at apex of venation, on opposite side of wing
and at apex of wing, marginal vein at least twice as long as stigmal, filum
spinosum in anterior half of wing CHEILONEURUS (p. 249)
132 (126) Frontovertex very narrow, less than one-tenth head width; head and thorax with
fine punctate sculpture and silvery white recumbent hairs; funicle with some
white segments; forewing infuscate with a curved hyaline band distad of
venation, disc of forewing densely setose proximad of lineacalva .... COMPERIA (p. 256)
Frontovertex at least about one-seventh of head width; head and thorax smooth
or with shallow reticulate sculpture and brownish setae; forewing with a
fuscous band in middle, paler or hyaline in basal one-third and distad of
venation, disc of forewing proximad of linea calva with a large, bare area ....
133 (132) Clava about as long as funicle and apically pointed (Fig. 67); mid tibial spur
shorter than basal mid tarsal segment; infuscation of forewing weak (Fig. 68);
antennal scrobes long, much longer than toruli and meeting dorsally, not
delimited laterally by a sharp carina OVALOENCYRTUS (p. 310)
Clava clearly much shorter than funicle and, although strongly truncate, with
apex square (Fig. 69); mid tibial spur longer than basal mid tarsal segment;
infuscation of forewing strong (Fig. 70); antennal scrobes not longer than
toruli nor meeting dorsally, often delimited laterally by a sharp carina
(Fig. 71) PARATETRALOPHIDEA (p. 319)
134 (11) Costal cell of forewing abruptly narrowed at apex (Fig. 73); frontovertex with
deep piliferous punctures PAKSIMMONDSIUS (p. 312)
Costal cell of forewing not abruptly narrowed at apex but gradually tapered;
frontovertex without deep piliferous punctures 135
135 (134) Scutellum with a thin apical flange 136
Scutellum without a distinct apical flange 137
136 (135) Antennal clava white, longer than preceding three funicle segments combined
HESPERENCYRTUS (p. 286)
Clava not white, not longer than preceding three funicle segments combined
PARAPHAENODISCUS (p. 317)
137 (135) Basal cell of forewing with two separate infuscate areas, both areas clothed in
dark setae, one adjacent to base of wing and the other adjacent to linea calva,
the area between these appearing as a fascia of pale setae or completely
naked ; pronotum often with a pair of sublateral rectangular white spots
*PROCHEILONEURUS Girault (p. 326)
Basal cell of forewing otherwise and never with two areas of dark setae either
side of a hyaline or naked area ; pronotum never with a pair of sublateral white
spots
138(137) Body (excluding legs, antennae, wings and tegulae) at least partly yellow or
orange
Body (excluding legs, antennae, wings and tegulae) completely dark, not yellow
or orange, although occasionally with a narrow yellowish area between
metanotum and propodeum 140
139 (138) Forewing with submarginal vein with a subapical triangular expansion (Fig. 77) ,
wing usually uniformly infuscate; antennal scrobes sharply bordered above
and on sides LAKSHAPHAGUS (p. 291)
Forewing with submarginal vein without a subapical triangular expansion, wing
usually with transverse hyaline bands that may occasionally be interrupted;
antennal scrobes not deep and not sharply bordered MICROTERYS (p. 299)
140 (138) Head triangular in profile, strongly inflexed inwards at top of antennal scrobes
138
139
[ Not to be confused with Prochiloneurus Silvestri (p. 327)
INDO-PACIFIC ENCYRTIDAE
157
78
Figs 78-86 78, Eugahania ishiharai Tachikawa, base of right forewing, upper surface, $; 79, Parectro-
moidella lowelli (Girault), right forewing, $; 80, Cremesina sp., right forewing, $; 81, Tongyus nesus
sp. n. , base of right forewing, upper surface, 9 ; 82, Yasumatsuiola sp. , right forewing showing pattern of
infuscation, $; 83, Holanusomyia pulchripennis Girault, right forewing showing pattern of infuscation,
$ ; 84, 85, Gentakola trifasciata (Saraswat), (84) left forewing, 9 , (85) right antenna, outer aspect, 9 ; 86,
Anagyrietta sp., right forewing, 9-
158
J. S. NOYES & M. HAY AT
(Fig. 72) and with a distinct transverse line of silvery white setae across face at
this point and continuing below eyes EPITETRACNEMUS (p. 273)
Head in profile more or less gradually anteriorly rounded, not strongly inflexed
inwards at top of antennal scrobes and without a distinct transverse line of
silvery white setae 141
141 (140) Stigmal vein of forewing shorter than marginal vein ZOOENCYRTUS (p. 350)
Stigmal vein of forewing longer than marginal vein 142
142 (141) Mandible with one or two teeth and a truncation (Figs 74, 75); antenna usually
with all funicle segments longer than broad, although rarely all subquadrate
ortransverse PSYLLAEPHAGUS (p. 330)
Mandible with three acute teeth (Fig. 76); not all funicle segments longer than
broad AENASIELLA (p. 224)
143 (11) Costal cell of forewing strongly excised at apex (Fig. 78) EUGAHANIA (p. 276)
Costal cell of forewing not or hardly excised at apex 144
144 (143) All funicle segments longer than broad; mandible always bidentate 145
Not all funicle segments longer than broad; mandible occasionally bidentate,
but usually otherwise 151
145 (144) Body (excluding legs) wholly dark and with silvery white setae, those on
scutellum usually arranged in a distinct pattern PARANATHRIX (p. 317)
Body (excluding legs) at least partly yellow or red; setae on thorax not silvery
white, or if so then those on scutellum are evenly distributed and not arranged
in a distinct pattern 146
146 (145) Forewing with at least a broad fuscous band in middle one-third of wing but
usually more extensively infuscate (Figs 79, 80) and not with a pattern of dark
and pale setae 147
Either forewing less extensively infuscate, the infuscation limited to one or two
narrow fasciae or to basal one-third or to small areas below venation which do
not extend more than one-third across wing, or wing with a distinct pattern of
dark and pale setae 148
147 (146) Frontovertex relatively broad, at narrowest point only a little narrower than
length of scape CREMESINA (p. 260)
Frontovertex relatively narrow, at narrowest point less than half as wide as
length of scape PARECTROMOIDELLA (p. 319)
148 (146) Forewing with postmarginal vein longer than stigmal 149
Forewing with postmarginal vein not longer than stigmal 150
149 (148) Forewing with one or two distinct fuscous bands LEPTOMASTIDEA (p. 292)
Forewing with infuscation limited to longitudinal streaks adjacent to venation
GYRANUSOIDEA (p. 280)
150 (148) Forewing with a distinct infuscate area at base and a diffuse band from stigmal
vein across wing (Fig. 81) and not with a pattern of dark and pale setae,
remainder hyaline; flagellar segments clearly slightly flattened from side to
side TONGYUS (p. 343)
Forewing more or less generally suffused pale fuscous or with only longitudinal
infuscate streaks adjacent to venation or with a pattern of dark and pale setae;
flagellar segments cylindrical (N.B., if material has been dried from alcohol
the flagellar segments may have collapsed giving a flattened appearance)
ANAGYRUS (p. 229)
151 (144) Eyes much shorter than minimum width of frontovertex 152
Eyes not shorter than minimum width of frontovertex 153
Figs 87-98 87, 88, Alamella flava Agarwal, (87) head, frontal aspect, $, (88) apex of right forewing
venation, upper surface, 9 ; 89, 90, Philosindia longicornis sp. n. , (89) head, frontal aspect, $ , (90) apex
of right forewing venation, upper surface (discal setae omitted), 9; 91, Rhopus sp., right forewing, $;
92, Hamusencyrtus sp., right forewing, $; 93, Gyranusoidea phenacocci (Beardsley), apex of right
forewing venation, upper surface, 9 ; 94, Epidinocarsis californicus (Compere), head, frontal aspect, 9
95, Anagyrus swezeyi Timberlake, base of right forewing, upper surface, $; 96, Anagyrus antoninae
Timberlake, apex of right forewing venation, upper surface, $; 97, 98, Doliphoceras nigricans
(Perkins), (97) head, frontal aspect, $, (98) base of right forewing, upper surface, $.
INDO-PACIFIC ENCYRTIDAE
159
160 J. S. NOYES & M. HAY AT
152 (151) Body foliaceously flattened; head prognathous; antennal toruli at mouth mar-
gin; pronotum longitudinally divided in middle (as in Fig. 38) ... PLATYRHOPUS (p. 325)
Body not foliaceously flattened; head opisthognathous; antennal toruli separ-
ated from mouth margin by more than their own lengths; pronotum entire
HUNTERELLUS (p. 288)
153 (151) Exserted part of ovipositor at least about one-fifth length of gaster; notaular
lines usually present in anterior part of mesoscutum PSEUDOCOCCOBIUS (p. 329)
Ovipositor not, or hardly, exserted; notaular lines completely absent 154
154 (153) Forewing with marginal vein absent, stigmal vein arising directly from submar-
ginal vein before it reaches anterior margin of wing, costal cell very slightly
incised at apex; antennal scrobes bordered dorsally and laterally by a very
sharp carina;clava solid TROPIDOPHRYNE (p. 346)
Forewing with marginal vein at least a little longer than broad, costal cell not
incised at apex; antennal scrobes not bordered above or at sides by a sharp
carina; clava two- or three-segmented 155
155 (154) Clava two-segmented (Fig. 85); forewing with venation not reaching half way
along wing (Fig. 84); mandible with three teeth GENTAKOLA (p. 278)
Clava three-segmented; forewing with venation extending more than half way
along wing; mandible bidentate 156
156 (155) First funicle segment not longer than pedicel 157
First funicle segment longer than pedicel 158
157 (156) Forewing with a pattern of radiating darker setae interspersed with wedge-
shaped paler areas and hyaline fasciae (Fig. 86); legs more or less unicolorous
yellow ANAGYRIETTA (p. 228)
Forewing largely infuscate without radiating fuscous areas but with a transverse
hyaline band (occasionally apical one-third of forewing entirely hyaline) at
apex of venation (Fig. 79) ; legs at least partly strongly infuscate
PARECTROMOIDELLA (p. 319)
158 (156) Forewing with stigmal vein very long, nearly one-quarter length of venation
from origin of submarginal vein to apex of postmarginal vein; apex of costal
cell and submarginal vein distinct (Figs 83, 355) HOLANUSOMYIA (p. 286)
Forewing with stigmal vein less than one-eighth as long as combined lengths of
submarginal, marginal and postmarginal veins; apex of costal cell not easily
distinguishable (i.e. difficult to make out where submarginal vein ends and
marginal vein begins) (Fig. 82) YASUMATSUIOLA (p. 348)
159 (12) Antennal toruli more than their own lengths from mouth margin, their lower
margins not below the lower eye margin when head viewed from front (Figs
87, 89), or if slightly so then first funicle segment at least about twice as long as
pedicel 160
Antennal toruli much less than their own lengths from mouth margin, or if
more then their lower margins are clearly below lower eye margins when
head viewed from front and first funicle segment not or hardly longer than
pedicel 162
160 (159) Forewing with postmarginal vein at least as long as stigmal (Fig. 90); hypopy-
gium not reaching apex of gaster PHILOSINDIA (p. 323)
Forewing with postmarginal vein shorter than stigmal (Fig. 88); hypopygium
reaching apex of gaster 161
161 (160) Mandible bidentate; forewing with linea calva interrupted on dorsal surface of
Figs 99-109 99, Rhytidothorax Imarlatti Ashmead (extra-limital species), scutellum and propodeum,
dorsal aspect, $ ; 100, Coelopencyrtus mauiensis Timberlake, scutellum and propodeum, dorsal aspect,
9; 101, Neastymachus auraticorpus Girault, thorax, aspect from left side, $; 102, Psyllaephagus sp.,
thorax, aspect from left side, 9; 103, Erencyrtus dewitzii Mahdihassan, base of left forewing, upper
surface, 9; 104, Metaphycus helvolus (Compere), base of right forewing, upper surface, 9; 105, 106,
Avetianella sp., (105) head, frontal aspect, 9> (106) right antenna, outer aspect, 9; 107, Tyndarichus
sp., base of right forewing, upper surface, 9; 108, 109, Tyndaricopsis davatus (Eady), (108) right
antenna, outer aspect, 9 , (109) base of right forewing, upper surface, 9
INDO-PACIFIC ENCYRTIDAE
161
162
J. S. NOYES & M. HAY AT
wing by two or three lines of setae and also more or less closed near posterior
margin ALAMELLA (p. 227)
Mandible tridentate; forewing with linea calva uninterrupted (except perhaps
by one or two setae) and open posteriorly 162
162 (159, Hypopygium extending to apex of gaster 163
161)
Hypopygium not reaching more than two-thirds along gaster 177
163 (162) Exserted part of ovipositor at least one-fifth as long as gaster 164
Ovipositor not or hardly exserted 168
164(163) Mandible bidentate; stigmal vein of forewing without a distinct apical uncus
(Figs 95, 96, 98) ;notaular lines completely absent 165
Mandible tridentate; stigmal vein of forewing with a distinct apical uncus;
notaular lines often present in anterior part of mesoscutum 166
165 (164) Head and thorax with very fine punctate-reticulate or vermiculate sculpture
which gives it a silky or velvety appearance ANAGYRUS (p. 229)
Head and thorax with shallow reticulate sculpture and relatively shiny
DOLIPHOCERAS (p. 266)
166 (164) Forewing with linea calva not interrupted (except perhaps by one or two setae)
on dorsal surface of wing; notaular lines completely absent PARAPHYCUS (p. 317)
Either linea calva interrupted on dorsal surface of forewing by two or three lines
of setae, or notaular lines present in anterior part of mesoscutum 167
167 (166) Clava clearly shorter than funicle AENASIOIDEA (p. 225)
Clava at least as long as funicle PSEUDOCOCCOBIUS (p. 329)
168 (163) Notaular lines completely absent 169
Notaular lines present in anterior part of mesoscutum 177
169 (168) Body strongly dorso-ventrally flattened; pronotum longitudinally divided in
middle (Fig. 38) 170
Body not or hardly dorso-ventrally flattened; pronotum entire 171
170 (169) Forewing with linea calva poorly defined (Fig. 92); eyes smaller and not longer
than malar space (Fig. 39) HAMUSENCYRTUS (p. 283)
Forewing with well-defined linea calva (Figs 91, 414); eyes larger and longer
than malar space (Fig. 40) RHOPUS (p. 332)
171 (169) Forewing with linea calva interrupted on dorsal surface by at least two lines
of setae and filum spinosum absent (Figs 95, 98); mandible with two equal
teeth 172
Forewing with linea calva not interrupted on dorsal surface by more than two or
three setae and with filum spinosum present (Figs 103, 104, 248, 394);
mandible with one to three teeth , or if with two teeth then one is clearly longer
than the other 175
172 (171) Forewing with postmarginal vein at least one-quarter longer than stigmal
(Fig. 93) GYRANUSOIDEA (p. 280)
Forewing with postmarginal vein not or hardly longer than stigmal (Figs 95, 96,
98) 173
Figs 110-127 110, 111, Neodadella compressipes Girault, (110) antenna, $, (111) right mandible, $;
112, Ectopiognatha sp., right mandible, $; 113, GahaniellasaissetiaeTimberlakQ, head, frontal aspect,
$; 114, Thomsonisca pakistanensis (Ahmad), right antenna, outer aspect, $; 115, Epitetralophidea
bicinctipes Girault, right mandible, $ ; 116, Adelencyrtus moderatus (Howard), right mandible, $ ; 117,
Coccidencyrtus ochraceipes Gahan, base of right forewing, upper surface, $; 118, Coelopencyrtus
odyneri Timberlake, head, frontal aspect, $; 119, Coelopencyrtus kaalae (Ashmead), head, frontal
aspect, $; 120, Phauloencyrtus mirisimilis Girault, right antenna, outer aspect (from card-mounted
specimen), $; 121, Psyllaephagus burnsi (Girault), right mandible, $; 122, Psyllaephagus channingi
(Girault), left mandible, $; 123, Rhopalencyrtoidea purpureicorpus Girault, right mandible, $; 124,
Asitus phragmitis (Ferriere), pronotum, mesoscutum and scutellum, dorsal aspect, $; 125, Coccidocto-
nus trinidadensis Crawford (extra-limital species), gaster, dorsal aspect, 9; 126, Pentelicus sp., apex of
right forewing venation, upper surface, $ ; 127, Cerchysiella sp. , base of right forewing, upper surface,
9-
INDO-PACIFIC ENCYRTIDAE
163
164
J. S. NO YES & M. HAY AT
173 (172) Head and dorsum of thorax with very fine punctate-reticulate or vermiculate
sculpture of silky appearance ANAGYRUS (p. 229)
Head and mesoscutum with shallow reticulate sculpture and at least slightly
shiny 174
174 (173) Eye relatively small, shorter than minimum width of frontovertex (Fig. 97)
DOLIPHOCERAS (p. 266)
Eye larger, clearly longer than minimum width of frontovertex (Fig. 94)
EPIDINOCARSIS (p. 272)
175 (171) Mesoscutum and scutellum both strongly convex, both with striate-reticulate or
distinctly elongate reticulate sculpture, scutellum never smooth and shiny
PARABLATTICIDA (p. 314)
Scutellum flat, not strongly convex; mesoscutum moderately flat, neither
mesoscutum nor scutellum with elongate or striate-reticulae sculpture,
scutellum sometimes smooth and shiny 176
176 (175) Propodeum relatively long, medially at least about one-fifth as long as scutellum
and with some sculpture medially (Fig. 99); scutellum usually with an apical
carina (although often very fine); gonostyli always hidden, never visible;
mandible usually with one or two teeth, rarely with three . . RHYTIDOTHORAX (p. 333)
Propodeum very short and smooth, medially not more than one-eighth as long
as scutellum (Fig. 100) which is rounded apically and without a carina;
gonostyli only slightly exserted but visible externally; mandible large and
with three teeth COELOPENCYRTUS (p. 255)
177 (162, Clava two-segmented AENASOMYIELLA (p. 226)
168
Clava three-segmented 178
178 (177) Forewing with postmarginal vein longer than stigmal (Fig. 103) ERENCYRTUS (p. 274)
Forewing with postmarginal vein not longer than stigmal 179
179 (178) Mesoscutum and scutellum largely metallic green ZARHOPALOIDES (p. 349)
Neither mesoscutum nor scutellum even partly metallic green, occasionally
brown or darker but never metallic 180
180 (179) Forewing with linea calva interrupted or closed on dorsal surface of wing by at
least one line of setae (Fig. 104); notaular lines often present on mesoscutum
METAPHYCVS (p. 298)
Forewing with linea calva neither interrupted nor closed on dorsal surface of
wing; notaular lines absent 181
181 (180) Forewing with stigmal vein less than twice as long as marginal; mesopleurum
posteriorly enlarged so that when thorax is viewed from side it is more or less
touching basal segment of gaster and thus clearly separating metapleurum
and propodeum from hind coxa (Fig. 101) NEASTYMACHUS (p. 304)
Forewing with stigmal vein more than three times as long as marginal; meso-
pleurum more or less normal so that when thorax viewed from side meta-
pleurum together with propodeum at least narrowly in contact with hind coxa
and thus clearly separating it from basal segment of gaster (Fig. 102)
PSYLLAEPHAGUS (p. 330)
182 (13) Clava three segmented (Fig. 106) AVETIANELLA (p. 239)
Clava entire SZELENYIOLA (p. 339)
183 (14) Submarginal vein of forewing with a subapical triangular expansion (Figs 107,
109) 184
Figs 128-140 128-131, Carabuniasp., (128) head, frontal aspect, $, (129) right mandible, $, (130) base
of right forewing, upper surface, 9 > (131) left antenna, inner aspect, $ ; 132, Kataka mudigerensis sp. n. ,
base of right forewing, upper surface, $; 133, Cyrtocoryphes viridiceps Timberlake, apex of right
forewing venation, upper surface, $; 134-136, Ruanderoma sankarani sp. n., (134) right forewing
showing pattern of infuscation, $, (135) thorax, dorsal aspect, $, (136) right mandible, $; 137,
Parencyrtomyia niveiclava Girault, right antenna, outer aspect, $; 138, Trichomasthus sp., thorax,
aspect from left side, $ ; 139, Rhytidothorax sp. , thorax, aspect from left side, 9 ; 140, Copidosoma sp. ,
thorax, aspect from left side, $ .
INDO-PACIFIC ENCYRTIDAE
165
140
166
J. S. NOYES & M. HAY AT
Submarginal vein of forewing at most slightly broadened apically but without a
subapical triangular expansion (Figs 117,238,394) 185
184 (183) Clava three-segmented TYNDARICHUS (p. 346)
Clava entire (Fig. 108) TYNDARICOPSIS (p. 347)
185 (183) Clava entire AUSTRALANUSIA (p. 236)
Clava three-segmented 186
186 (185) Scape hardly longer than broad, much less than one and one-half times as long as
broad and only about one and one-half times as long as pedicel (Fig. 110);
antennal toruli a little more than twice their own lengths from mouth margin;
mandible with four teeth (Fig. Ill); gaster unicolorous, dark , not yellow
NEOCLADELLA (p. 305)
Scape more than one and one-half times as long as broad and at least about twice
as long as pedicel; antennal toruli not more than twice their own lengths from
mouth margin; mandibles with three teeth or one or two teeth and a
truncation, or if with four teeth (Fig. 112) then base of gaster is yellow
contrasting with the dark remainder 187
187 (186) Mesopleurum posteriorly enlarged so that it nearly touches base of gaster, when
thorax viewed from side it clearly separates propodeum and metapleurum
from hind coxa (as in Figs 101, 138, 177); base of gaster yellowish; mandible
with four teeth (Fig. 112) ECTOPIOGNATHA (p. 267)
Mesopleurum not so enlarged and clearly separated from base of gaster by
metapleurum together with propodeum which are at least narrowly in contact
with hind coxa (as in Figs 102, 139, 140), or if mesopleurum enlarged as in
alternate, then gaster is unicolorous and dark; mandible with three teeth or
one or two teeth and a truncation or rarely with four 188
188 (187) Forewing with postmarginal vein clearly much longer than stigmal AGENIASPIS (p. 226)
Forewing with postmarginal vein not or hardly longer than stigmal 189
189 (188) Antennal toruli situated relatively high on head, their lower margins level with,
or above, lower eye margins when head viewed from front (Fig. 113); eye not
distinctly hairy 190
Antennal toruli relatively lower, their lower margins clearly below the lowest
eye margins when head viewed from front; eye often very hairy 192
190 (189) Thorax dorsally convex; antennal scape not longer than malar space GAHANIELLA (p. 278)
Thorax dorsally fairly flat ; antennal scape at least a little longer than malar space 191
191 (190) Antennal clava with apex pointed, its dorsal margin strongly curved whilst its
ventral margin is more or less straight, first funicle segment clearly shorter
than pedicel and subsequent segments becoming broader and larger towards
apex of antenna; mandible with three sharp teeth MAYRIDIA (p. 295)
Antennal clava apically rounded and more or less cylindrical, funicle segments
subequal in length and usually also in breadth to pedicel and not distinctly
widening towards apex of antenna (Fig. 114); mandible with one or two teeth
and a truncation THOMSONISCA (p. 343)
192(189) Scutellum and mesoscutum flat, at least scutellum matt and not strongly
metallic, often with relatively deep reticulate sculpture; eye not distinctly
hairy 193
Scutellum and mesoscutum clearly convex, or if flat then either both are strongly
metallic or the eye is conspicuously hairy 195
193 (192) Mandible with four teeth (Fig. 116) ADELENCYRTUS (p. 223)
Figs 141-151 141 , Saprencyrtus casuarinae (Girault), right forewing showing pattern of infuscation (from
card-mounted specimen), $; 142, Copidosoma sp., apex of right forewing venation, upper surface, $;
143, 144, Tachinaephagus sp., (143) apex of left forewing venation, upper surface, $, (144) right
mandible, $ ; 145, Manicnemus indicus (Mani & Saraswat), right forewing showing pattern of infusca-
tion, 9; 146, Homalotylus sp., right antenna, outer aspect, $; 147, Mahencyrtus comara (Walker)
(extra-limital species), base of left forewing, upper surface, $; 148, Adektitopus gordhi sp. n., bas.e of
right forewing, upper surface, $; 149, Cheiloneurella sp., thorax, dorsal aspect, $; 150, Eotopus
beneficus (Shafee) , base of right forewing, upper surface, $ ; 151 , Paraclausenia herbicola Hayat, base of
right forewing, upper surface, $ .
INDO-PACIFIC ENCYRTIDAE
167
151
168
J. S. NOYES & M. HAY AT
Mandible with one or two teeth and a truncation (Fig. 115) 194
194 (193) Forewing with linea calva closed or interrupted on dorsal surface of wing
(Fig. 117) COCCIDENCYRTUS (p. 253)
Forewing with linea calva neither closed nor interrupted . . . EPITETRALOPHIDEA (p. 273)
195 (192) Both mesoscutum and scutellum very convex and dull, not shiny, with fine
striate-reticulate or punctate-reticulate sculpture PARABLATTICIDA (p. 314)
Mesoscutum and scutellum flat or only slightly convex and usually at least a little
shiny; sculpture shallow reticulate, or if punctate-reticulate then dorsum of
thorax distinctly metallic blue, green or purple 196
196 (195) Exserted part of ovipositor at least as long as one-quarter length of gaster 197
Ovipositor not or hardly exserted 200
197 (196) Exserted part of ovipositor slightly but distinctly downcurved; mandible with
two teeth and a truncation EPIBLATTICIDA (p. 272)
Exserted part of ovipositor straight; mandible with three acute teeth 198
198 (197) Exserted part of ovipositor at least about two-thirds as long as gaster
NEZARHOPALUS (p. 307)
Exserted part of ovipositor less than half as long as gaster 199
199 (198) Hypopygium extending past apex of gaster so that it is clearly visible in dorsal
view (Fig. 125) COCCIDOCTONUS (p. 254)
Hypopygium not extending past apex of gaster and not visible in dorsal view
TELETEREBRATUS (p. 341)
200 (196) Eye with conspicuous, long, dark setae 201
Eye more or less naked 202
201 (200) First funicle segment anelliform and clearly much shorter than second which
is subequal to the remaining funicle segments, all of which are slightly
transverse (Fig. 120); scutellum with longitudinally reticulate sculpture
which is clearly much deeper than the more regularly reticulate sculpture of
mesoscutum PHAULOENCYRTUS (p. 323)
First funicle segment not contrasting with remaining segments as in alternate,
the funicle segments usually enlarging distally; sculpture of scutellum not
deeper than that of mesoscutum, usually more shallow EXORISTOBIA (p. 277)
202 (200) Clava strongly obliquely truncate and clearly longer than funicle 203
Clava without a strong oblique truncation and shorter than funicle 204
203 (202) Head and mesoscutum with punctate-reticulate sculpture similar to that of
scutellum BAEOANUSIA (p. 241)
Head and mesoscutum with shallow irregular reticulate sculpture, almost
smooth and clearly much shallower than that on scutellum MESANUSIA (p. 296)
204 (202) Mandible with one or two teeth and a truncation (Figs 121, 122); forewing with
marginal vein punctiform or rarely longer than broad PSYLLAEPHAGUS (p. 330)
Mandible with three acute teeth (Fig. 76); forewing with marginal vein always
longer than broad AENASIELLA (p. 224)
205 (14) Body foliaceously dorso-ventrally flattened ; pronotum longitudinally divided in
middle (Fig. 124) 206
Body not or hardly dorso-ventrally flattened; pronotum entire 207
206 (205) Clava three-segmented; marginal fringe of forewing about one-eighth as long
as maximum width of wing PLATYRHOPUS (p. 325)
Clava entire; marginal fringe of forewing at most a little longer than one-fifth
maximum width of wing ASITUS (p. 236)
207 (205) Forewing with a naked line connecting apex of stigmal vein to apex of postmar-
ginal vein and anterior wing margin (Fig. 126) PENTELICUS (p. 322)
Setation at apex of forewing venation normal , naked line not present 208
208 (207) Either not all funicle segments longer than broad or forewing with postmarginal
vein longer than stigmal 209
All funicle segments longer than broad and forewing with postmarginal vein not
longer than stigmal 220
209 (208) Head, dorsum of thorax and mesopleurum with distinctive deep punctate
sculpture; forewing with postmarginal vein at least a little longer than stigmal;
scutellum never with an apical flange BLASTOTHRIX (p. 242)
INDO-PACIFIC ENCYRTIDAE
169
159
Figs 152-159 152, Ooencyrtus sp. , base of right forewing, upper surface, $ ; 153, Trichomasthus sp. , base
of right forewing, upper surface, $ ; 154, Hengata spinosa sp. n. , base of right forewing, upper surface,
$ ; 155, Cheiloneurella sp. , base of right forewing, upper surface, $ ; 156, Ethoris dahmsi sp. n. , base of
right forewing, upper surface, $; 157, Protyndarichoides sp., base of right forewing, upper surface, $;
158, Diasula glabriscutellum (Girault), base of left forewing, upper surface (from card-mounted
specimen), $ ; 159, Mashhoodia indica Shafee, left forewing showing pattern of light and dark setae, 9 .
170 J. S. NOYES & M. HAY AT
Head and at least mesoscutum and mesopleurum with shallow reticulate
sculpture and occasionally relatively deep piliferous punctures, but never,
except on scutellum, with punctate sculpture, or if so then scutellum has a
distinct apical flange; forewing with postmarginal vein usually not longer than
stigmal, although occasionally longer 210
210 (209) Frontovertex relatively narrow, at narrowest point not more than one-sixth
head width NEODISCODES (p. 306)
Frontovertex broader, at narrowest point at least one-quarter head width 211
211 (210) Apex of scutellum produced in a short thin flange; forewing with postmarginal
vein more than one and one-half times as long as stigmal; occipital margin
very sharp almost to base of mandible ERICYDNUS (p. 274)
Apex of scutellum without an apical flange ; forewing with postmarginal vein less
than one and one-half times as long as stigmal; occipital margin rounded or
sharp but not as extensively sharp as in alternate 212
212 (211) Dorsum of thorax strongly convex, mesoscutum and scutellum dull with at least
the scutellum and often mesoscutum with fine longitudinally striate sculpture;
forewing with postmarginal vein as long as or longer than stigmal
PARABLATTICIDA (p. 314)
Dorsum of thorax moderately flat, not strongly convex, neither mesoscutum nor
scutellum with longitudinally striate sculpture and often quite shiny; forewing
with postmarginal vein occasionally as long as or longer than stigmal but
usually a little shorter 213
213 (212) Eye relatively small and clearly not reaching occipital margin which is more or
less rounded, the greatest length of eye not more than minimum width of
frontovertex 214
Eye larger, at least slightly longer than minimum width of frontovertex and
more or less reaching occipital margin which is sharp 215
214 (213) Antennal toruli close to mouth margin, separated from it by less than half their
own lengths (Figs 442, 443); head prognathous, in profile more or less
gradually and evenly curved anteriorly and not triangular; mandible with two
or three sharp teeth (Fig. 443) ZAOMMOENCYRTUS (p. 349)
Antennal toruli more than their own lengths from mouth margin; head op-
isthognathous and in side view appearing triangular being acutely angled
inwards at top of antennal scrobes; mandible with one or two teeth and a
broad truncation HUNTERELLUS (p. 288)
215 (213) Forewing with filum spinosum directed towards junction of marginal and
submarginal veins and thus clearly converging with the line of setae on the
proximal margin of the linea calva (Fig. 127) CERCHYSIELLA (p. 246)
Forewing with filum spinosum absent or directed towards junction of stigmal
and marginal veins and thus more or less parallel with line of setae on
proximal margin of linea calva 216
216 (215) Propodeum relatively long, medially at least about one-fifth length of scutellum
and with some carinae (Fig. 99); scutellum usually with an apical carina
(although often very fine); gonostyli always hidden externally RHYTIDOTHORAX (p. 333)
Figs 160-175 160, l6l,Adektitopusgordhisp. n., (160) sculpture in centre of mesoscutum (area approx.
0-1 mm square), 9> (161) sculpture in centre of scutellum (area approx. 0-1 mm square), $; 162,
Anomalicornia sp., apex of right forewing venation, upper surface, $; 163, Leptomastidea aurantiaca
Mercet (extra-limital species), apex of right forewing venation, upper surface, $; 164, Leptomastix
nigrocoxalis Compere, apex of right forewing venation, upper surface, $; 165, Bacalusa fuscipennis
sp. n., base of right forewing, upper surface, $; 166, Apoleptomastix spoliata Kerrich, right antenna,
inner aspect, $; 167, Alamella flava Agarwal, right antenna, outer aspect, $; 168, Leptomastix
dactylopii Howard, left antenna, outer aspect, 9 ; 169, Anomalencyrtus longicornis Hayat & Verma, left
antenna, inner aspect, 9 ; 170, 171, Paratetracnemoidea malenotti (Mercet) (extra-limital species), (170)
right antenna, outer aspect, 9 , (171) apex of left forewing venation, upper surface, 9 ; 172, Heterococci-
doxenus schlechtendali (Mayr), apex of right forewing venation, upper surface, 9 '> 173, Bacalusa
tachikawai (Shafee, Alam & Agarwal), right antenna, outer aspect, 9 ; 174, Bacalusa fuscipennis sp. n. ,
right antenna, outer aspect, 9 ; 175, Doliphoceras nigricans (Perkins), right antenna, outer aspect, 9
INDO-PACIFIC ENCYRTIDAE
171
itgggK
172
J. S. NOYES & M. HAY AT
Propodeum medially very short, without any carinae medially and medially not
more than one-eighth as long as scutellum which is apically without a carina
(Fig. 100) ;gonostyli often clearly visible externally 217
217 (216) Postmarginal vein of forewing not longer than stigmal 218
Forewing with postmarginal vein longer than stigmal 219
218 (217) Mandible with three acute teeth (Figs 118, 119); tegula always dark; legs usually
extensively dark and never marked with pale yellow COELOPENCYRTUS (p. 255)
Mandible with one or two teeth and a truncation (Figs 121, 122); tegula and legs
often at least partly pale yellow PSYLLAEPHAGUS (p. 330)
219 (217) Mandible with three acute teeth (Fig. 123); exserted part of ovipositor at least
one-third as long as gaster RHOPALENCYRTOIDEA (p. 332)
Mandible bidentate; ovipositor not or hardly exserted 220
220 (208, Head and dorsum of thorax with fine punctate-reticulate or vermiculate sculp-
219) ture which gives these a silky appearance ANAGYRUS (p. 229)
At least head and mesoscutum with shallow reticulate sculpture which is not
silky in appearance and which is more distinctly shiny 221
221 (220) Frontovertex at narrowest point at least half head width; antennal flagel-
lum unicolorous; sculpture of scutellum almost same as that on head and
mesoscutum DOLIPHOCERAS (p. 266)
Frontovertex at narrowest point much less than half head width; antennal
flagellum usually with at least one white segment contrasting with dark brown
segments; scutellum with fine punctate-reticulate sculpture which contrasts
strongly with shallower sculpture of mesoscutum EPIDINOCARSIS (p. 272)
222 (18) Eye nearly touching base of mandible so that malar space is not more than
one-fifth length of eye; ovipositor distinctly exserted, the exserted part at least
about one-third as long as gaster HEXENCYRTUS (p. 286)
Eye clearly separated from base of mandible, malar space at least nearly
one-half as long as eye; ovipositor not exserted 223
223 (222) Occipital margin rounded ; forewing with postmarginal vein shorter than stigmal
(Fig. 132) KATAKA (p. 290)
Occipital margin sharp; forewing with postmarginal vein longer than stigmal
(Fig. 130) 224
224(223) Clava three-segmented; mandible broad and truncate, without teeth; eye
separated from occipital margin by at least about twice diameter of a facet
PRIONOMASTIX (p. 325)
Clava entire (Fig. 131); mandible with one long, acute tooth (Fig. 129); eye
separated from occipital margin by not more than diameter of a facet
CARABUNIA (p. 244)
225 (19) Forewing with linea calva interrupted or more or less closed on dorsal surface by
several lines of setae towards posterior margin 226
Forewing with linea calva neither interrupted nor closed on dorsal surface,
except perhaps by one or two setae 231
226 (225) Forewing with postmarginal vein very short or absent, much shorter than
stigmal (Fig. 133) 227
Figs 176-192 176, Cerchysius sp. , aspect from left side, $ ; 177, Ooencyrtus sp. , thorax, aspect from left
side, 9 ; 178, Paraenasomyia orro Girault, right mandible, 9 ; 179, Australia minuta Girault, antenna, $ ;
180, Paraschedius sp., apex of right forewing venation, upper surface (discal setae omitted), $; 181,
Ooencyrtus pacificus Waterston, apex of left forewing venation, upper surface, 9; 182, Ooencyrtus
batocerae Ferriere, apex of right forewing venation, upper surface, 9 ; 183, Copidosoma sp. , apex of left
forewing venation, upper surface, 9 ! 184, 185, Tachardiaephagus tachardiae Howard, (184) head frontal
aspect, 9, (185) head, dorsal aspect, 9; 186, Syrphophagushofferi(Hayat), base of left forewing, upper
surface, 9; 187, Ethoris dahmsi sp. n., apex of right forewing venation, upper surface, 9; 188,
Lemennaisia ambigua (Nees), left mandible, 9; 189, Coccidencyrtus bicolor (Girault), right mandible,
9; 190, Papuna nemis sp. n., apex of right forewing venation, upper surface (discal setae omitted), 9;
191 , Cowperia indica (Kerrich), apex of right forewing venation, upper surface, 9 ; 192, Echthrogonato-
pus nigricornis (Hayat), base of right forewing, upper surface, 9
INDO-PACIFIC ENCYRTIDAE
173
183
192
174
J. S. NOYES & M. HAY AT
Forewing with postmarginal vein nearly as long as or longer than stigmal (Figs
134, 164) 228
227 (226) Scutellum without dense white setae arranged in a more or less distinct pattern;
infuscation of forewing restricted to a broad band below apical half of
venation; head and dorsum of thorax metallic green or blue . . CYRTOCORYPHES (p. 263)
Scutellum with a pattern of dense silvery- white setae; forewing with infuscation
pale but more or less evenly distributed; head and thorax black and dull but
with some slight brassy reflections PARANATHRIX (p. 317)
228 (226) Notaular lines present in anterior part of mesoscutum (Fig. 135) ; mandible with
three acute teeth (Fig. 136) RUANDEROMA (p. 334)
Notaular lines absent; mandible with two acute teeth 229
229 (228) Forewing brown with hyaline bands or spots (similar to Fig. 135)
CALLIPTEROMA (p. 244)
Forewing hyaline with one or more pale brown longitudinal streaks or generally
suffused pale brown 230
230 (229) Forewing with one or more pale brown longitudinal streaks; antennal flagellum
unicolorous LEPTOMASTIX (p. 293)
Forewing generally suffused pale brown distad of bend of submarginal vein;
antennal flagellum with contrasting dark and pale segments ANAGYRUS (p. 229)
231 (225) Clava apically with a slight but distinct oblique truncation and outer suture
converging with inner (Fig. 137) 232
Clava apically transversely truncate or rounded so that the outer suture is more
or less parallel with inner 233
232 (231) Antennal flagellum unicolorous and brown; body dark and moderately to
strongly lustrous (latter especially on face); mandible with one tooth and a
broad truncation [Cerchysius] australiensis Ashmead (p. 352)
Clava white contrasting with the brown funicle; body orange or brown, not
lustrous; mandible with three acute teeth PARENCYRTOMYIA (p. 320)
233 (231) Mesopleurum not enlarged posteriorly and not in contact with basal segment of
gaster so that when thorax viewed from side metapleurum and propodeum
together are broadly (very rarely narrowly) in contact with hind coxa (Figs
139, 140); mandible with one, two or three sharp teeth, never with a
truncation; hypopygium often reaching apex of gaster 234
Mesopleurum enlarged posteriorly and touching or almost touching basal
segment of gaster so that when thorax viewed from side it separates meta-
pleurum and propodeum from hind coxa (Fig. 138) or these are only very
narrowly meeting; mandible with one or two teeth and a truncation; hypo-
pygium never extending more than three-quarters along gaster 237
234 (233) Apex of hypopygium not reaching more than about one-third along gaster;
forewing with a complete hyaline fascia distad of venation (Fig. 141)
SAPRENCYRTUS (p. 336)
Apex of hypopygium reaching to at least about two-thirds along gaster, often to
apex; forewing without a complete hyaline fascia distad of venation 235
235 (234) Antennal toruli separated from mouth margin by less than half their own
lengths; forewing with sensillae at apex of stigmal vein arranged symmetri-
cally in a square, uncus absent (Fig. 142) COPIDOSOMA (p. 258)
Antennal toruli separated from mouth margin by at least only a little less than
their own lengths; forewing with sensillae at apex of stigmal vein not arranged
in a square, uncus present (Fig. 143) 236
Figs 193-201 193, Copidosomyia ambiguous (Subba Rao), right antenna, outer aspect showing sensory
truncate surface at apex of clava, $; 194, Mashhoodiella echthromorpha Hayat, antenna, $; 195, 196,
Sakencyrtus sp. , (195) thorax, dorsal aspect, 9 , (196) right forewing, upper surface, 9 ; 197, Pseudococ-
cobius terryi (Fullaway), base of right forewing, upper surface, 9 ; 198, Homalotylus sp., base of right
forewing, upper surface, $; 199, Aphycus sp., base of right forewing, upper surface, 9; 200, 201,
Isodromus axillaris Timberlake, (200) right antenna, outer aspect, 9 , (201) base of right forewing, upper
surface, 9-
INDO-PACIFIC ENCYRTIDAE
175
176
J. S. NOYES & M. HAY AT
236 (235) Ovipositor at least slightly exserted so that gonostyli are externally visible;
mandible always with three sharp teeth (Fig. 144) TACHINAEPHAGUS (p. 340)
Ovipositor never exserted and gonostyli never visible externally; mandible
usually with one or two sharp teeth and only very rarely with three
RHYTIDOTHORAX (p. 333)
237 (233) Scutellum with punctate-reticulate sculpture which is conspicuously deeper than
the shallow reticulate sculpture of mesoscutum; infuscation of forewing pale
and inconspicuous and represented only by a subapical band TRICHOMASTHUS (p. 346)
Scutellum with shallow reticulate sculpture which is not deeper than that of
mesoscutum and apically distinctly shallower; infuscation of forewing more
extensive, usually covering apical two-thirds of wing but often with one or two
hyaline bands distad of venation MICROTERYS (p. 299)
238 (19) Notaular lines present in at least anterior one-third of mesoscutum 239
Notaular lines absent 243
239 (238) Notaular lines complete (Fig. 6); clava with an oblique apical truncation (Fig.
146) HOMALOTYLUS (p. 287)
Notaular lines not reaching more than half way across mesoscutum; clava
apically rounded 240
240 (239) Head and thorax orange-red, not metallic BACALUSA (p. 239)
Head and thorax at least partly metallic 241
241 (240) Head and thorax with several white , yellow and orange areas
ECHTHROBACCELLA (p. 267)
Head and thorax entirely dark and metallic without any pale areas 242
242 (241) Forewing with postmarginal vein longer than stigmal (Fig. 148) .... ADEKTITOPUS (p. 221)
Forewing with postmarginal vein shorter than stigmal (Fig. 145) MANICNEMUS (p. 294)
243 (238) Hypopygium reaching apex of gaster; ovipositor not exserted; mandible with
two acute teeth (possibly three in Ameniscocephalus) 244
Hypopygium not reaching more than four-fifths along gaster, or if so then
ovipositor is exserted and exserted part at least one-third as long as gaster;
mandible with one or two teeth and a truncation or three or four teeth 246
244 (243) Head lenticular, in side view about three times as long as wide; forewing evenly
infuscate without any hyaline areas AMENISCOCEPHALUS (p. 227)
Head not lenticular, in side view not or hardly more than twice as long as wide;
forewing not evenly infuscate and with hyaline areas 245
245 (244) Forewing with postmarginal vein at least as long as stigmal, infuscation usually
restricted to apex or to two narrow fasciae LEPTOMASTIDEA (p. 292)
Forewing with postmarginal vein clearly shorter than stigmal, infuscation at
least a wide band across wing from apical one-third of venation and often
another at apex (Fig. 79) PARECTROMOIDELLA (p. 319)
246 (243) Head and thorax (excluding legs and antennae) dark and metallic 247
Head and thorax (excluding legs and antennae) not completely dark and
metallic, at least partly pale 256
247 (246) Head triangular in profile , strongly inflexed at top of antennal scrobes (as in Fig.
72); mandible with four teeth (Fig. 116) ADELENCYRTUS (p. 223)
Head in profile more or less evenly rounded anteriorly, not strongly inflexed at
top of antennal scrobes; mandible with one or two teeth and a truncation or
three teeth . 248
Figs 202-212 202, 203, Agarwalencyrtus citri (Agarwal), (202) right antenna, outer aspect showing
sensory truncate surface on apex of clava, 9 (203) thorax, dorsal aspect showing distribution of setae
(left side) and sculpture (right side), $; 204, Copidosoma sp., right antenna, outer aspect showing
sensory truncate surface at apex of clava, $; 205, Proleuroceroides sp., left antenna, inner aspect
showing pattern of infuscation, 9; 206, 207, Parechthrodryinus davicornis Cameron, (206) scutellum
and propodeum, dorsal aspect, $, (207) apex of right forewing venation, upper surface, 9; 208, 209,
Astymachus japonicus Howard, (208) right antenna, outer aspect, $, (209) head, thorax and gaster,
dorsal aspect, 9 ; 210, Mahencyrtus comara (Walker) (extra-limital species), scutellum and propodeum,
dorsal aspect, 9; 211, 212, Taftia siassetiae Gahan, (211) left antenna, inner aspect, 9> (212) right
forewing, upper surface, 9
INDOPACIFIC ENCYRTIDAE
177
178
J. S. NOYES & M. HAY AT
248 (247) Postmarginal vein of forewing longer than stigmal ENCYRTOIDEA (p. 268)
Postmarginal vein of forewing not longer than stigmal 249
249 (248) Posterior margin of mesoscutum strongly projecting backwards so that when
thorax is in normal resting position it projects above axillae and separates
them by at least the length of the visible part of the axilla (Fig. 44)
XENOENCYRTUS (p. 348)
Posterior margin of mesoscutum hardly projecting above axillae so that they
appear to meet medially or almost so when thorax is in normal resting position
(asinFig.42) 250
250 (249) Forewing with sensillae at apex of stigmal vein arranged symmetrically in a
square, uncus absent (Fig. 142) COPIDOSOMA (p. 257)
Forewing with sensillae at apex of stigmal vein not arranged in a square, uncus
present and distinct (Figs 147, 148, 150-152) 251
251 (250) Forewing with stigmal vein longer than marginal 252
Forewing with stigmal vein not longer than marginal 253
252 (251) Mandible with one or two teeth and a truncation (Figs 121, 122); forewing with
marginal vein not more than twice as long as broad PSYLLAEPHAGUS (p. 330)
Mandible with three acute teeth; forewing with marginal vein at least three
times as long as broad CONCHYNJLLA (p. 256)
253 (25 1) Infuscation of forewing restricted to a median longitudinal wedge-shaped streak
from apex of wing to about level with apex of venation, submarginal vein with
parastigma slightly to strongly broadened, this broadening often triangular
and indicated by a single erect seta (as in Fig. 147) MAHENCYRTUS (p. 294)
Infuscation of forewing more extensive than in alternate, usually extending
from apex of submarginal vein to near apex of wing and enclosing one or two
hyaline areas, submarginal vein with parastigma not so distinctly expanded . 254
254 (253) Hypopygium reaching apex of gaster * PROCHILONEURUS Silvestri (p. 327)
Hypopygium not reaching more than two-thirds along gaster 255
255 (254) Gaster unicolorous, completely dark not white or yellow basally
NEABROLEPOIDEVS (p. 303)
Gaster with basal white or yellow ring contrasting with the dark remainder
MESOCALOCERINUS (p. 297)
256 (246) Clava obliquely truncate, the sutures strongly converging 257
Clava apically rounded , the sutures usually parallel or only slightly converging 258
257 (256) Hypopygium reaching apex of gaster; exserted part of ovipositor at least half as
long as gaster * PROCHILONEURUS Silvestri (p. 327)
Hypopygium not reaching more than two-thirds along gaster; ovipositor not or
hardly exserted MASHHOODIELLA (p. 295)
258 (256) Body almost entirely orange or yellow 259
At least dorsum of thorax brown or green 260
259 (258) Pronotum triangular and conspicuous in dorsal view, at least about as long as
mesoscutum (Fig. 149) CHEILONEURELLA (p. 248)
Pronotum strongly transverse and inconspicuous, not more than one-quarter as
long as mesoscutum PARASCHEDIUS (p. 318)
* Not to be confused with Procheiloneurus Girault (p. 326)
Figs 213-226 213, Neocladia sp., right hind tibia and tarsus, outer aspect, $; 214, Nathismusia
southwoodi sp. n., base of left forewing, upper surface, 9; 215, 216, Muluencyrtus nudipennis sp. n.,
(215) apex of right forewing venation, upper surface (from card-mounted specimen), $ , (216) right hind
tibia and tarsus, outer aspect (from card-mounted specimen), $ ; 217-219, Olypusa hirsuta sp. n., (217)
base of right forewing, upper surface, $ , (218) head, frontal aspect (from card-mounted specimen), $ ,
(219) left antenna, outer aspect (from card-mounted specimen), $; 220, Trichomasthus sp., right
antenna, outer aspect, $ ; 221 , 222, Saprencyrtus casuarinae (Girault), (221) left mandible, 9 , (222) base
of right forewing, upper surface (from card-mounted specimen), $; 223, Echthrogonatopus exitiosus
Perkins, right mandible, $ ; 224, Echthrogonatopus nigricornis (Hayat), right antenna, inner aspect, $;
225, Ovaloencyrtus fijiensis sp. n., left mandible, 9; 226, Hypergonatopus bifasciatus (Timberlake),
apex of left forewing venation, upper surface, $ .
INDO-PACIFIC ENCYRTIDAE
179
180
J. S. NO YES & M. HAY AT
260 (258) Forewing with marginal vein punctiform or nearly so, wing with a single large
fuscous blotch or broad fascia below marginal vein, hyaline fascia absent
OOENCYRTUS (p. 309)
Forewing with marginal vein at least twice as long as broad, infuscation of wing
more extensive than in alternate and often with at least one hyaline fascia
distad of apex of venation 261
261 (260) Forewing with a hyaline fascia at apex of venation MICROTERYS (p. 299)
Forewing without a hyaline fascia at apex of venation AUSTROMIRA (p. 238)
262 (23) Forewing with postmarginal vein not longer than stigmal (Fig. 150) 263
Forewing with postmarginal vein at least a little longer than stigmal (Figs 148,
151) 264
263 (262) Notaular lines complete; antenna mostly dark brown CHARITOPUS (p. 248)
Notaular lines not reaching more than half way across mesoscutum; antenna
completely yellowish orange EOTOPUS (p. 269)
264 (262) Scutellum with punctate-reticulate sculpture which is clearly much deeper than
that of mesoscutum (Figs 160, 161) ADEKTITOPUS (p. 221)
Scutellum with shallow reticulate sculpture which is not deeper than that of
mesoscutum and often quite smooth and shiny 265
265 (264) Sculpture of scutellum clearly more shallow than that of mesoscutum, almost
smooth; eyes overreaching occipital margin; fore wing relatively broad, much
less than two and one-half times as long as broad; mandible bidentate
CLAUSENIA (p. 251)
Sculpture of scutellum about same as that of mesoscutum ; eyes not overreaching
occipital margin; forewing at least slightly more than two and one-half times
as long as broad; mandible with three acute teeth PARACLAUSENIA (p. 316)
266 (21, 23) Forewing with a distinct pattern of dark and light setae (Fig. 159) . MASHHOODIA (p. 295)
Forewing without a pattern of dark and light setae 267
267 (266) Head and dorsum of thorax with very fine punctate or vermiculate sculpture
which gives a velvety or silky appearance, the sculpture of the mesoscutum
not conspicuously different from that on either scutellum or head 268
Head and dorsum of thorax with reticulate sculpture and not with fine punctate
or vermiculate sculpture and not with silky or velvety appearance, or if partly
so then at least mesoscutum has distinctly shallower and less fine sculpture
than either head or scutellum 269
268 (267) Forewing with postm arginal vein at least a little longer than stigmal
GYRANUSOIDEA (p. 280)
Forewing with postmarginal vein not longer than stigmal ANAGYRUS (p. 229)
269 (267) Scutellum with dense silvery white setae which are conspicuously more dense
than those on mesoscutum and often arranged in a distinct pattern; at least
some of the flagellar segments pale and contrasting with others which are dark 270
Scutellum with dark setae, or if with white setae then these are not conspicu-
ously deeper than on mesoscutum and are never arranged in a distinct
pattern; flagellum usually unicolorous although occasionally with contrasting
white and dark segments 271
270 (269) Forewing with postmarginal vein longer than stigmal; setae on scutellum not
arranged in a distinct pattern; scape cylindrical and not conspicuously
flattened, second segment of funicle white (Fig. 166) APOLEPTOMASTIX (p. 235)
Forewing with postmarginal vein not longer than stigmal; setae on scutellum
arranged in a distinct pattern; scape at least slightly broadened and flattened,
second segment of funicle dark, not white PARANATHRIX (p. 317)
Figs 227-238 227, Gahaniella saissetiae Timberlake, right antenna, outer aspect, $ ; 228-230, Pasulinia
gentha sp. n., (228) head, aspect from left side, $, (229) left mandible, $, (230) right antenna, outer
aspect, 9 ; 231 , 232, Protyndarichoides spp. , apex of right forewing venation, upper surface, 9 ; 233-234,
Protyndarichoides spp., right antenna, outer aspect, 9; 235, Cladiscodes sacchari Subba Rao, right
forewing, upper surface, 9, 236, Tachinaephagus sp., base of left forewing, upper surface, 9; 237,
Metaphaenodiscus aligarhensis Hayat, apex of left forewing venation, upper surface, 9; 238, Exoristo-
bia philippinensis Ashmead, base of left forewing, upper surface, 9
INDO-PACIFIC ENCYRTIDAE
181
237
182
J. S. NO YES & M. HAY AT
271 (269) Funicle seven-segmented, clava two-segmented (Figs 37, 167) 272
Funicle six-segmented, clava one- two- or three-segmented 273
272 (271) Forewing relatively broad, only a little more than twice as long as broad, stigmal
vein moderately long and slender, postmarginal vein relatively long and
distinct (Fig. 88) ALAMELLA (p. 227)
Forewing relatively slender, nearly three times as long as broad, stigmal vein
very short, subsessile, postmarginal vein more or less absent (Fig. 162)
ANOMALICORNIA (p. 232)
273 (271) Body distinctly dorso-ventrally flattened; pronotum longitudinally divided in
middle (as in Fig. 38) RHOPUS (p. 332)
Body more robust, not clearly flattened; pronotum entire 274
274 (273) Clava entire (Fig. 169) ANOMALENCYRTUS (p. 232)
Clava three-segmented 275
275 (274) Either postmarginal vein or forewing as long or longer than stigmal (Figs 163,
164) or forewing relatively long and narrow and more than three times as long
as broad 276
Postmarginal vein of forewing shorter than stigmal (Figs 98, 165) and forewing
not more than three times as long as broad 277
276 (275) First funicle segment not longer than pedicel; generally smaller species (length
less than 1-3 mm) LEPTOMASTIDEA (p. 292)
First funicle segment at least one and one-half times as long as pedicel (Fig. 168);
generally larger species (length greater than 1-4 mm) LEPTOMASTIX (p. 293)
277 (275) Flagellum widening towards clava so that clava is nearly twice as broad as first
funicle segment (Figs 173, 174); gaster white or pale orange at base, thorax
brown; ovipositor hardly as long as mid tibia BACALUSA (p. 239)
Flagellum not clearly widening towards clava so that clava is not or hardly wider
than first funicle segment (Fig. 175); gaster usually dark brown or if orange
then thorax is also partly orange; ovipositor clearly longer than mid tibia
DOLIPHOCERAS (p. 266)
278 (24) Forewing with postmarginal vein longer than stigmal PRIONOMASTIX (p. 325)
Forewing with postmarginal vein not longer than stigmal 279
279 (278) Forewing with marginal vein absent, stigmal vein arising from submarginal
before it reaches anterior margin of wing (Figs 171, 172); frontovertex usually
with distinct piliferous punctures giving it a thimble-like appearance 280
Forewing with marginal vein present; frontovertex without conspicuous pilifer-
ous punctures 281
280 (279) Forewing with marginal fringe absent; mesoscutum with deep piliferous punc-
tures giving it a thimble-like appearance; interantennal prominence normal;
clava with apex more or less rounded HETEROCOCCIDOXENUS (p. 286)
Forewing with marginal fringe present; mesoscutum with raised reticulate
sculpture, without deep piliferous punctures; international prominence
strongly produced into an acute tooth level with lowest margins of antennal
toruli; clava with a strongly obliquely truncate apex (Fig. 170)
PARATETRACNEMOIDEA (p. 318)
281 (279) Mesopleurum enlarged posteriorly so that it touches or nearly touches base of
gaster, thorax in side view with hind coxa more or less clearly separated from
metapleurum and propodeum by posterior margin of mesopleurum (Fig.
177) ; hypopygium not extending more than three-quarters along gaster 282
Figs 239-251 239, 240, Lutherisca sp., (239) left antenna, outer aspect, <j>, (240) apex of left forewing
venation, upper surface, $; 241, Copidosoma sp., left antenna, outer aspect showing truncate sensory
surface at apex of clava, $ ; 242, Coccidoxenoides peregrinus (Timberlake), base of left forewing, upper
surface, 9 ; 243, 244, Neocharitopus sp., (243) apex of right forewing venation, upper surface, $ , (244)
scutellum, dorsal aspect showing sculpture (left side) and distribution of setae (right side), $; 245,
Coagerus bouceki sp. n., right forewing, $; 246, Ooencyrtus sp., right antenna, outer aspect, $; 247,
Coelopencyrtus sp. , apex of right forewing venation, upper surface, $ ; 248, Coelopencyrtus sp. , base of
right forewing, upper surface, $; 249, Copidosoma sp., base of right forewing, upper surface, $; 250,
25 1 , Zaommoencyrtus spp . , apex of right forewing venation , upper surface , 9 .
INDO-PACIFIC ENCYRTIDAE
183
184
J. S. NO YES & M. HAYAT
Mesopleurum not so enlarged posteriorly, when thorax viewed from side the
hind coxa is more or less broadly in contact with metapleurum and pro-
podeum and is thus clearly separating mesopleurum from basal segment of
gaster (Figs 102, 140, 176), or if as in alternate then hypopygium extends to
apex of gaster 284
282 (281) Pedicel subtriangular, shorter than first funicle segment, clava not or hardly
longer than first funicle segment; antennal toruli with lower margins at level of
or above lower eye margins; frontovertex with relatively deep piliferous
punctures and with an almost thimble-like appearance BOTHRIOPHRYNE (p. 243)
Pedicel clearly much longer than broad and longer than first funicle segment;
antennal toruli with their lower margins clearly much below lower eye
margins; frontovertex with inconspicuous piliferous punctures 283
283 (282) Hind margin of mesoscutum more or less straight so that it does not project
above axillae medially and therefore the axillae appear to meet (as in Fig. 42);
mandible with three sharp teeth HELEGONATOPUS (p. 283)
Hind margin of mesoscutum distinctly projecting backwards above axillae so
that they appear to be quite widely separated when thorax viewed from above
when thorax in normal resting position (as in Fig. 44); mandible almost always
with one or two teeth and a truncation, although occasionally tridentate
OOENCYRTUS (p. 309)
284 (281) Head and thorax mostly orange with some fuscous markings, but not metallic . . 285
At least head and usually thorax completely dark and metallic, not orange .... 286
285 (284) Forewing with linea calva interrupted on dorsal surface by three or four lines of
setae; exserted part of ovipositor about as long as gaster AUSTRALAPHYCUS (p. 237)
Forewing with linea clava not interrupted, except perhaps by two or three setae;
exserted part of ovipositor less than half as long as gaster PARAPHYCUS (p. 317)
286 (284) Antennal toruli almost touching mouth margin, separated by not more than half
their own lengths; hypopygium extending to apex of gaster; mandible with
three acute teeth 287
Antennal toruli more than half their lengths from mouth margin; hypopygium
not reaching more than two-thirds along gaster, or if so then mandible with
one or two teeth and a truncation 288
287 (286) Forewing with sensillae at apex of stigmal vein symmetrical, arranged in a
square, apex of stigmal vein without a distinct uncus (Fig. 183) . . . COPIDOSOMA (p. 257)
Forewing with sensillae at apex of stigmal vein not arranged in a square and not
symmetrical, apex of stigmal vein with a distinct uncus PRIONOMITOIDES (p. 326)
288 (286) Ovipositor strongly exserted and with sheaths flattened from side to side and
slightly downcurved towards apex (Fig. 176) (most apparent along ventral
surface of sheaths) CERCHYSIUS (p. 247)
Ovipositor not exserted, or if so then sheaths are more or less cylindrical in
cross-section and are straight (ventral surface of sheaths straight or slightly
tapering upwards) 289
289 (288) Mandible with one or two teeth and a truncation (Figs 121, 122); mesoscutum
and scutellum usually green or blue-green and quite shiny, although oc-
casionally dull PSYLLAEPHAGUS (p. 330)
Mandible with three acute teeth; mesoscutum and scutellum dark with green
and blue reflections, not strongly metallic SYRPHOPHAGUS (p. 338)
Figs 252-266 252, Tassonia sp., apex of right forewing venation, upper surface, $; 253, 254, Haligra
concolor sp. n. , (253) right antenna, outer aspect showing truncate sensory surface at apex of clava, $ ,
(254) sculpture in centre of scutellum (area approx. 0-1 mm square), $; 255, Adektitopus sp., right
antenna, outer aspect, $; 256-263, Adektitopus gordhi sp. n., (256) right antenna, outer aspect, 9,
(257) apex of right forewing venation, upper surface, $?, (258) sculpture on frontovertex anterior to
anterior ocellus (area approx. 0-1 mm square), 9> (259) hypopygium, $, (260) right antenna, outer
aspect, d", (261) base of right forewing, upper surface, cf , (262) genitalia, cf j (263) digiti and associated
structures; 264, Anagyrietta sp. , left antenna, outer aspect, $ ; 265, Amicencyrtus obscurus Hayat, apex
of right forewing venation, upper surface, $; 266, Anagyrus swezeyi Timberlake, right antenna, outer
aspect, 9-
INDO-PACIFIC ENCYRTIDAE
185
252
254
: "^SiJs.^s*t_ "***^
186 J. S. NOYES & M. HAY AT
290 (25) Antennal toruli situated relatively high on head and close together so that their
distance from mouth margin is more than one and one-half times the
minimum distance between them HEXENCYRTUS (p. 286)
Distance of antennal toruli from mouth margin less than one and one-half times
the distance between them 291
291 (290) Clavaentire 292
Clava three-segemented 294
292 (291) Clava with a very strong oblique apical truncation which is much longer than
remainder of ventral surface of clava; forewing with sensillae at apex of
stigmal vein arranged symmetrically in a square, stigmal vein without a
distinct uncus (Fig. 183) COPIDOSOMA (p. 257)
Clava more or less apically rounded; forewing with sensillae at apex of stigmal
vein asymmetrical and not arranged in a square, stigmal vein with an apical
uncus 293
293 (292) Hypopygium not reaching four-fifths along gaster AUSTROENCYRTUS (p. 238)
Hypopygium reaching apex of gaster ASEIRBA (p. 235)
294 (291) Mandible bidentate HEMILEUCOCERUS (p. 284)
Mandible tridentate or with one or two teeth and a truncation 295
295 (294) Propodeum medially less than one-fifth as long as scutellum 296
Propodeum medially at least one-fifth as long as scutellum 303
296 (295) Dorsum of thorax at least partly orange or yellow XENOSTRYXIS (p. 348)
Dorsum of thorax dark brown, green or purple 297
297 (296) Forewing with sensillae at apex of stigmal vein arranged symmetrically in a
square; stigmal vein without a distinct apical uncus (Fig. 183) . . . COPIDOSOMA (p. 257)
Forewing with sensillae at apex of stigmal vein not arranged in a square,
asymmetrical; stigmal vein with a distinct apical uncus 298
298 (297) Antenna long and filiform, funicle segments increasing in length so that sixth is
about three times as long as broad, all segments of clava longer or hardly
broader than sixth funicle segment (Fig. 179) AUSTRALIA (p. 237)
Antenna not as in alternate; segments of clava only a little longer than broad,
quadrate or transverse 299
299 (298) Hypopygium reaching apex of gaster 300
Hypopygium not reaching apex of gaster 301
300 (299) Hypopygium extending past apex of last tergite so that it is plainly visible in
dorsal view (Fig. 125) COCCIDOCTONUS (p. 254)
Hypopygium not extending past apex of last tergite and not visible in dorsal view
RHOPALENCYRTOIDEA (p. 332)
301 (299) Forewing with stigmal vein not more than one and one-half times as long as
marginal (Fig. 186) SYRPHOPHAGUS (p. 338)
Forewing with stigmal vein at least twice as long as marginal 302
302 (301) Mandible with one or two teeth and a broad truncation (Figs 121, 122)
PSYLLAEPHAGUS (p. 330)
Mandible with three acute teeth (Fig. 178) PARAENASOMYIA (p. 316)
303 (295) Gaster with basal segment yellow or yellowish orange and contrasting with the
dark remainder PROTYNDARICHOIDES (p. 328)
Gaster unicolorous, dark and not paler basally 304
304(303) Ovipositor visible externally; scutellum with distinct reticulate sculpture; eye
with short inconspicuous hairs , each not longer than diameter of a facet 305
Figs 267-279 267, Anagyrus dactylopii (Howard), left antenna, inner aspect, $ ; 268, Anagyrus antoninae
Timberlake, right antenna, outer aspect, $; 269-271, Aphycomorpha araucariae Timberlake, (269)
apex of right forewing venation, upper surface, 9> (270) right antenna, outer aspect, $, (271) head,
frontal aspect, <j>; 272, 273, Aphycus spp., (272) right antenna, outer aspect, <j>, (273) left forewing,
upper surface, $; 274, 275, Asltus phragmitis (Ferriere), (274) apex of left forewing venation, upper
surface, $, (275) head, frontal aspect, $5 276, Astymachus japonicus Howard, apex of left forewing
venation, upper surface, $ ; 277, Avetianella sp. , apex of left forewing venation, upper surface, $ ; 278,
Austroencyrtus sp., right antenna, inner aspect, $; 279, Bacalusa fuscipennis sp. n., right forewing
showing pattern of infuscation, $ .
INDO-PACIFIC ENCYRTIDAE
187
188
J. S. NOYES & M. HAY AT
Either ovipositor not visible externally or scutellum more or less smooth and
shiny; eye distinctly hairy, each hair usually much longer than diameter of a
facet 306
305 (304) Forewing with submarginal vein with parastigma clearly broadened and forming
a weak triangular expansion , this indicated by a single erect seta (Fig. 147)
MAHENCYRTUS (p. 294)
Submarginal vein of forewing with parastigma not enlarged, not or hardly
wider than proximal part of submarginal vein ZOOENCYRTUS (p. 350)
306 (304) Gonostyli free and not fused with second valvifers (Fig. 430) and at least slightly
exserted and visible externally; scutellum always smooth and shiny
TACHINAEPHAGUS (p. 340)
Gonostyli fused with second valvifers (Fig. 415) and never visible externally;
scutellum usually distinctly sculptured but occasionally smooth and shiny
RHYTIDOTHORAX (p. 333)
307 (26) Mesoscutum with an inconspicuous median longitudinal ridge in posterior half
HENGATA (p. 284)
Mesoscutum without a median longitudinal ridge 308
308 (307) Antennal scrobes long, straight and deeply impressed, clearly reaching to at
least three-quarters distance from antennal toruli to anterior ocellus; inter-
antennal prominence dorsally very sharply margined and pointed and clearly
separated from frontovertex (Figs 184, 185) TACHARDIAEPHAGUS (p. 340)
Antennal scrobes relatively shallow, more or less semi-circular and not long,
reaching to at most a little more than half way between antennal toruli and
anterior ocellus; interantennal prominence not pointed dorsally and not
sharply margined, often confluent with frontovertex 309
309 (308) Pronotum triangular and conspicuous, in dorsal view about equal in length to
mesoscutum (Fig. 149) CHEILONEURELLA (p. 248)
Pronotum strongly transverse and inconspicuous, in dorsal view less than
one-third as long as mesoscutum 310
310 (309) Neither postmarginal vein of forewing longer than stigmal nor eye with dense
long setae (occasionally eye hairy, but hairs are not individually longer than a
facet); hypopygium not reaching apex of gaster; mandible with one or two
teeth and a truncation or rarely with three sharp teeth 311
Either postmarginal vein of forewing longer than stigmal or eye clothed in
conspicuous setae, each clearly much longer than diameter of a facet;
hypopygium often reaching apex of gaster; mandible with from one to three
sharp teeth 314
311 (310) Dorsum of thorax quite flat; hind leg yellow with one or two conspicuous dark
bands; forewing with stigmal vein short, subsessile (Fig. 180) . . . PARASCHEDIUS (p. 318)
Dorsum of thorax conspicuously convex; legs completely yellow or yellow-
orange without any conspicuous dark bands; forewing with stigmal vein
relatively long (Figs 181, 182) 312
312 (311) Gaster unicolorous, from yellowish brown to orange-brown NEASTYMACHUS (p. 304)
Gaster dark brown with a contrasting basal yellow band 313
313 (312) Mesopleurum enlarged and more or less touching basal segment of gaster so that
when thorax viewed from side it clearly separates hind coxa from meta-
pleurum and propodeum (Fig. 177); eye with very dense short, translucent
hairs . ! OOENCYRTUS (p. 309)
Figs 280-294 280-288, Bacalusa fuscipennis sp. n. , (280) sculpture on frontovertex anterior to anterior
ocellus (area approx. 0-1 mm square), $ , (281) sculpture in centre of mesoscutum (area approx. 0-1 mm
square), $, (282) sculpture in centre of scutellum (area approx. 0-1 mm square), $, (283) genitalia, $,
(284) hypopygium, $, (285) head, frontal aspect, d", (286) right antenna, outer aspect, C?, (287)
genitalia, cf, (288) digiti and apex of aedeagus; 289, 290, Cerapteroceroides sp., (289) right antenna,
outer aspect, $, (290) apex of left forewing venation, upper surface, $; 291, 292, Cerapterocerus sp.,
(291) apex of left forewing venation, upper surface, $, (292) left forewing showing pattern of
infuscation, $; 293, 294, Cercobelus jugaeus (Walker) (extra-limital species), (293) right mandible, $,
(294) head, frontal aspect, $.
INDO-PACIFIC ENCYRTIDAE
189
294
190
J. S. NO YES & M. HAY AT
Mesopleurum not so enlarged and not touching basal segment of gaster so that
when thorax viewed from side metapleurum together with propodeum are
narrowly in contact with hind coxa (as in Fig. 140) ; eye naked
DIAPHORENCYRTUS (p. 263)
314 (310) Mandible with three acute teeth and scutellum with deep reticulate sculpture . . 315
Either mandible with only one or two acute teeth or scutellum more or less
smooth and shiny 316
315 (314) Hypopygium not extending more than two-thirds along gaster; forewing with
postmarginal vein longer than stigmal (Figs 156, 187) ETHORIS (p. 275)
Hypopygium extending to at least four-fifths along gaster; forewing with
postmarginal vein rarely longer than stigmal 316
316(314, Ovipositor at least slightly exserted with sheaths flattened from side to side;
315) gonostyli free and at least about one-quarter as long as ovipositor (Fig. 430);
mandible with three acute teeth (Fig. 144) TACHINAEPHAGUS (p. 340)
Ovipositor not exserted, gonostyli not visible externally and not more than
one-fifth as long as ovipositor and fused to second valvifers (Fig. 415);
mandible with one or two teeth RHYTIDOTHORAX (p. 333)
317 (26) Scape shorter than minimum width of frontovertex 318
Scape not shorter than minimum width of frontovertex 320
318 (317) Gaster dark with basal orange or yellow ring which contrasts with dark re-
mainder PROTYNDARICHOIDES (p. 328)
Gaster unicolorous, dark, without pale basal ring 319
319 (318) Legs, including coxae, completely yellow; malar space more than two-thirds as
long as eye; fore wing with setae extending to base (Fig. 132) KATAKA (p. 290)
Legs with at least mid tibia and coxae brownish; malar space less than half as
long as eye; forewing with proximal part of basal cell naked KAKAOBURRA (p. 289)
320 (317) Gaster with basal orange or yellow ring; head in profile anteriorly more or less
evenly rounded; eyes not overreaching occipital margin which is sharp;
mesopleurum not enlarged so that when thorax viewed from side hind coxa is
in contact with metapleurum and propodeum (as in Figs 139, 140); mandible
with three acute teeth PROTYNDARICHOIDES (p. 328)
Gaster unicolorous, dark and usually slightly metallic, or if basal segment yellow
then either head in profile is triangular and abruptly inflexed at top of antennal
toruli (as in Fig. 72) and mandible with one tooth and a broad truncation or
eyes overreach occipital margin which is more or less rounded and meso-
pleurum posteriorly enlarged so that when thorax viewed from side it clearly
separates hind coxa from metapleurum and propodeum (Fig. 177) 321
321 (320) Scutellum with very elongate striate-reticulate sculpture; hypopygium not
extending more than four-fifths along gaster; head in profile anteriorly more
or less evenly rounded; mesopleurum not enlarged so that when thorax
viewed from side hind coxa touches metapleurum and propodeum (as in Figs
139, 140) 322
Scutellum without striate sculpture, or if appearing striate then either hypo-
pygium reaches apex of gaster or head in profile triangular and abruptly
inflexed at top of antennal scrobes (as in Fig. 72) or mesopleurum posteriorly
enlarged and more or less touching basal segment of gaster so that when
Figs 295-308 295, Cheiloneurella sp. , right antenna, outer aspect, $ ; 296-299, Coagerus bouceki sp. n. ,
(296) apex of right forewing venation, upper surface, $, (297) scutellum and propodeum showing
sculpture (left side) and distribution of setae (right side), $ , (298) right antenna, outer aspect, also inner
aspect of clava, $, (299) genitalia, right side, ventral aspect, $; 300, Coelopencyrtus odyneri Timber-
lake, right antenna, outer aspect, $; 301, Coelopencyrtus sp., right antenna, inner aspect, $; 302, 303,
Comperiella lemniscata Compere & Annecke, (302) right antenna, outer aspect, $, (303) left forewing
showing pattern of infuscation, 9 ; 304, Cremesina sp. , brachypterous species, right fore and hind wings,
$ ; 305-308, Cremesina aquilonaris sp. n. , (305) apex of right forewing venation, upper surface, 9 , (306)
right antenna, outer aspect, 9> (307) sculpture in centre of mesoscutum (area approx. 0-1 mm square),
$ , (308) genitalia, right side, ventral aspect, $ .
INDO-PACIFIC ENCYRTIDAE
191
192
J. S. NOYES & M. HAY AT
thorax viewed from side it clearly separates hind coxa from metapleurum and
propodeum (Fig. 177) 323
322(321) Both mesoscutum and scutellum with striate-reticulate sculpture, that on
scutellum very fine so that it is completely matt and not metallic; hypopygium
reaching to only a little more than halfway along gaster; mandible tridentate
NEGENIASPIDIUS (p. 305)
Mesoscutum with shallow reticulate sculpture which contrasts strongly with the
striate sculpture of scutellum; scutellum at least slightly metallic; hypopygium
reaching to about four-fifths along gaster; mandible with four teeth (Fig. 188)
although occasionally with only three LAMENNAISIA (p. 292)
323 (321) Hypopygium more or less reaching apex of gaster; forewing with postmarginal
vein not or hardly longer than stigmal 324
Hypopygium not extending more than three-quarters along gaster, or if so then
postmarginal vein of forewing is at least one-half longer than stigmal 327
324 (323) Forewing with filum spinosum directed towards junction of submarginal and
marginal veins and converging with setae on proximal margin of linea calva
(Fig. 127) CERCHYSIELLA (p. 246)
Forewing with filum spinosum absent or directed towards junction of marginal
and stigmal veins and subparallel to setae on proximal margin of linea calva
(Figs 152, 153, 158, 186, 238, 249; also as in Figs 134, 135, 139, 166) 325
325 (324) Eye not overreaching occipital margin, separated from occiput by a sharp
occipital margin; forewing with sensillae at apex of stigmal vein arranged
symmetrically in a square, uncus absent (Figs 142, 183) COPIDOSOMA (p. 257)
Eye overreaching occipital margin which is rounded at this point; forewing with
sensillae at apex of stigmal vein asymmetrical and not arranged in a square,
uncus clearly present 326
326 (325) Forewing with marginal vein not more than twice as long as broad and at least a
little shorter than stigmal, postmarginal vein a little shorter than stigmal
TRJAPITZINELLUS (p. 346)
Forewing with marginal vein at least about four times as long as broad and
longer than stigmal, postmarginal vein as long as or slightly longer than
stigmal PARECTROMOIDES (p. 320)
327 (323) Head in profile triangular, abruptly inflexed at top of antennal scrobes (as in
Fig. 72) 328
Head in profile more or less gradually and evenly anteriorly rounded and not
abruptly inflexed at top of antennal scrobes 330
328 (327) Mandible with four teeth or with one tooth and a truncation 329
Mandible with three acute teeth 330
329 (328) Mandible with four teeth (Fig. 116) ADELENCYRTUS (p. 223)
Mandible with one tooth and a broad truncation (Fig. 189) . . . COCCIDENCYRTUS (p. 253)
330 (327, Forewing with postmarginal vein at least one and one-half times as long as
328) stigmal 331
Forewing with postmarginal vein not or hardly longer than stigmal 332
331 (330) Mesopleurum posteriorly enlarged and more or less touching basal segment of
gaster so that when thorax viewed from side it clearly separates hind coxa
from metapleurum and propodeum (as in Fig. 177); mandible with three
acute teeth ENCYRTOIDEA (p. 268)
Mesopleurum not posteriorly enlarged so that when thorax viewed from side it is
clearly separated from basal segment of gaster by metapleurum and pro-
Figs 309-324 309-313, Cremesina aquilonaris sp. n., (309) hypopygium, <j>, (301) right antenna, outer
aspect, d", (311) genitalia, cf , (312) apex of genitalia, d", (313) base of right forewing, upper surface, d";
314, 315, Diasula glabriscutellum (Girault), (314) left mandible, $, (315) left antenna, outer aspect, $;
316, Cyrtocoryphes viridiceps Timberlake, right antenna, outer apsect, 9; 317-321, Doddanusia sp.,
(317) apex of right forewing venation, upper surface, $ , (318) right antenna, outer aspect, 9 , (319) left
mandible, $, (320) hypopygium, $, (321) genitalia, $; 322, 323, Ectopiognatha sp., (322) apex of left
forewing venation, upper surface, $?, (323) right antenna, outer aspect, $?; 324, Gahaniella saissetiae
Timberlake, apex of right forewing venation, upper surface, $ .
INDO-PACIFIC ENCYRTIDAE
193
324
194
J. S. NOYES & M. HAY AT
podeum which are touching hind coxa (Fig. 139); mandible with one or two
teeth RHYTIDOTHORAX (p. 333)
332 (330) Clavasolid ZAMENHOFELLA (p. 348)
Clava three-segmented 333
333 (332) Scutellum flat and occipital margin rounded MAYRIDIA (p. 295)
Scutellum convex, or if more or less flat then occipital margin sharp 334
334 (333) Eye with conspicuous dense, dark setae, each longer than diameter of a facet;
mesoscutum and scutellum clothed in dense recumbent dark setae so that
dorsum of thorax is distinctly hairy EXORISTOBIA (p. 277)
Eye with inconspicuous translucent setae, each not longer than diameter of a
facet; mesoscutum and scutellum not noticeably hairy 335
335 (334) Mandible tridentate 336
Mandible bidentate or with one or two teeth and a truncation 338
336 (335) Posterior margin of mesoscutum more or less straight, not projecting above
axillae medially so that when thorax is viewed from above the axillae meet
medially (as in Fig. 42) 337
Posterior margin of mesoscutum clearly projecting backwards above axillae so
that when thorax viewed from above and in normal resting position it broadly
separates axillae medially (as in Fig. 44) 338
337 (336) Ovipositor not or hardly exserted; forewing with postmarginal vein shorter than
stigmal HELEGONATOPUS (p. 283)
Either exserted part of ovipositor at least about one-fifth as long as gaster or
postmarginal vein of forewing as long as or longer than stigmal 340
338 (335, Forewing with marginal vein less than twice as long as broad 339
336)
Forewing with marginal vein at least twice as long as broad 340
339 (338) Mandible bidentate; mesoscutum and scutellum both with deep punctate-
reticulate sculpture and not shiny FULGORIDICIDA (p. 278)
Mandibles not bidentate; mesoscutum never with punctate-reticulate sculpture
and always slightly metallic, scutellum occasionally with punctate sculpture
OOENCYRTUS (p. 309)
340 (337, Mesopleurum posteriorly enlarged and more or less touching basal segment of
338) gaster so that when thorax viewed from side it clearly separates hind coxa
from metapleurum and propodeum (as in Figs 138, 177) 341
Mesopleurum not posteriorly enlarged so that when thorax viewed from side
hind coxa clearly touches metapleurum and propodeum thus separating
mesopleurum from basal segment of gaster (as in Figs 139, 140) 342
341 (340) Occipital margin very sharp, carinate; eye not reaching occiput; head and thorax
without pale setae; sculpture of scutellum often slightly deeper and finer than
that of mesoscutum, but never strongly so SYRPHOPHAGUS (p. 338)
Occipital margin rounded or sharp, but not carinate, or if carinate then
scutellum has moderate to strong punctate sculpture which is conspicuously
deeper than that of mesoscutum; eyes usually overreaching occipital margin;
head and thorax often with conspicuous pale setae TRICHOMASTHUS (p. 346)
342 (340) Stigmal vein of forewing about twice as long as marginal PARAENASOMYIA (p. 316)
Forewing with stigmal vein not more than one and one-half times as long as
marginal 343
343 (342) Propodeum medially not more than one-tenth as long as scutellum and devoid of
Figs 325-341 325-327, Eotopus beneficus (Shafee), (325) right antenna, outer aspect, <J> , (326) genitalia,
$, (327) genitalia, cf ; 328-331, Ethoris dahmsisp. n., (328) right antenna, outer aspect, <j>, (329) right
mandible, $, (330) genitalia, left side, ventral aspect, $, (331) head, frontal aspect, $; 332-339,
Gentakola trifasdata (Saraswat), (332) head, dorso-frontal aspect, $, (333) genitalia, left side, ventral
aspect, $ , (334) base of left forewing, upper surface, $ , (335) hypopygium, $ , (336) genitalia, cf , (337)
digiti and apex of aedeagus, (338) base of right forewing, upper surface, cf , (339) right antenna, outer
aspect, cf ; 340, 341, Haligra concolor sp. n., (340) sculpture in centre of mesoscutum (area approx.
0-1 mm square), $, (341) left mandible, $.
INDO-PACIFIC ENCYRTIDAE
195
341
196
J. S. NO YES & M. HAY AT
a median carina; scutellum moderately convex but not very shiny
SYRPHOPHAGUS (p. 338)
Propodeum medially more than one-sixth as long as scutellum and with a
shallow but distinct median carina; scutellum strongly convex and very shiny,
at least in its apical one-half DIASULA (p. 263)
344 (28) Forewing with marginal vein punctiform or only very slightly longer than broad 345
Marginal vein of forewing at least twice as long as broad 346
345 (344) Ovipositor sheaths strongly flattened from side to side and downcurved towards
apex (Fig. 176); mandible with three acute teeth CERCHYSIUS (p. 247)
Ovipositor sheaths more or less cylindrical and straight; mandible with one or
two teeth and a truncation PSYLLAEPHAGUS (p. 330)
346 (344) Forewing with postmarginal vein longer than stigmal (Fig. 190) PAPUNA (p. 312)
Forewing with postmarginal vein not longer than stigmal 347
347 (346) Forewing hyaline 348
Forewing infuscate 350
348 (347) Head and thorax mostly yellowish with a few dark markings XENOSTR YXIS (p. 348)
Head and thorax not yellowish, completely dark and more or less shiny 349
349 (348) Forewing with marginal vein at least about three times as long as stigmal (Fig.
192); mesoscutum clothed in moderately dense white setae; scutellum fairly
flat with deep reticulate sculpture contrasting with shallower sculpture of
mesoscutum ECHTHROGONATOPUS (p. 267)
Forewing with marginal vein at most only a little longer than stigmal, often
shorter (Fig. 186); mesoscutum with dark setae; scutellum convex and with
sculpture similar to that of mesoscutum SYRPHOPHAGUS (p. 338)
350 (347) Infuscation of forewing limited to a longitudinal wedge-shaped mark from
apex, submarginal vein with parastigma slightly enlarged into an indistinct
triangular expansion indicated by a single erect seta (Fig. 147) . . MAHENCYRTUS (p. 294)
Infuscation of forewing quite extensive and usually forming a distinct pattern,
parastigma not or hardly swollen 351
351 (350) Head in profile triangular and abruptly inflexed at top of antennal scrobes (as in
Fig. 72); clava never obliquely truncate and with sutures subparallel; meso-
scutum and scutellum never orange or yellowish, always dark and metallic
ADELENCYRTUS (p. 223)
Head in profile anteriorly more or less evenly curved, although occasionally
slightly triangular but not strongly so; clava usually with an oblique apical
truncation, or at least with sutures oblique and converging; mesoscutum or
scutellum often partly orange or yellowish CHEILONEURUS (p. 249)
352 (28) Forewing with marginal vein absent (Fig. 191) COWPERIA (p. 259)
Forewing with marginal vein present, although sometimes short 353
353 (352) Clava with a strong oblique apical truncation, the truncate surface about as long
or longer than remainder of ventral surface (Figs 146, 193), if clava three-
segmented then sutures strongly converge 354
Clava apically more or less rounded or if with a slight oblique truncation then
truncate surface clearly much shorter than remainder of ventral surface of
clava and sutures , if present , subparallel or occasionally slightly converging . 358
354 (353) Notaular lines complete (Fig. 6) HOMALOTYLUS (p. 287)
Notaular lines absent . 355
Figs 342-357 342-345, Haligra concolor sp. n., (342) apex of left forewing venation, upper surface, $,
(343) hypopygium, $, (344) base of right forewing, upper surface, $, (345) genitalia, $; 346,
Hamusencyrtus mymaricoides (Compere, Subba Rao & Kaur), left antenna, inner aspect, $; 347-354,
Hengata spinosa sp. n., (347) left mandible, $, (348) right antenna, outer aspect, $, (349) apex of right
forewing venation, upper surface, $ , (350) hypopygium, $ , (351) genitalia, left side, ventral aspect, $ ,
(352) right antenna, outer aspect, Q", (353) genitalia, cT, (354) digiti and associated structures; 355, 356,
Holanusomyia pulchripennis Girault, (355) apex of right forewing venation and linea calva, upper
surface, 9, (356) left antenna, inner aspect, 9; 357, Indaphycus planus Hayat, base of right forewing,
upper surf ace, $.
INDO-PACIFIC ENCYRTIDAE
197
342
357
198 J. S. NOYES & M. HAY AT
355 (354) Clava entire (Fig. 193); eyes clearly separated from occipital margin by at least
the diameter of an ocellus COPIDOSOMYIA (p. 259)
Clava three-segmented ; eyes reaching or very nearly reaching occipital margin 356
356 (355) Forewing with marginal vein not longer than stigmal, wing completely hyaline
except for a small inconspicuous cloud below marginal vein
PENTACLADOCERUS (p. 322)
Forewing with marginal vein at least three times as long as stigmal, forewing
more strongly infuscate 357
357 (356) Mesoscutum with a few scattered dark setae; dorsum of thorax completely dark
and metallic, at least apical one-third of scutellum shiny TINEOPHOCTONUS (p. 343)
Mesoscutum with moderate to very dense white setae; dorsum of thorax usually
at least partly orange or yellow, although occasionally completely dark;
scutellum, except extreme apex, with fine reticulate sculpture giving it a matt
appearance *PROCHILONEURUSSi\vestri (p. 327)
358 (353) Forewing infuscate with a distinct dark pattern, or with a fuscous spot in centre
of wing 359
Forewing hyaline or generally suffused very pale brown, without a distinct
pattern 363
359 (358) Forewing with postmarginal vein longer than stigmal; mesoscutum with deep
piliferous punctures giving it a thimble-like appearance BORROWELLA (p. 242)
Forewing with postmarginal vein not longer than stigmal; mesoscutum without
deep piliferous punctures 360
360 (359) Forewing with infuscation restricted to a large pale fuscous spot below marginal
vein, marginal vein punctiform (Fig. 197); notaular lines reaching to about
one-third way across mesoscutum PSEUDOCOCCOBIUS (p. 329)
Forewing with infuscation more distinct and extensive than in alternate, mar-
ginal vein usually at least a little longer than broad; notaular lines almost
always absent 361
361 (360) Head and thorax never partly yellowish or orange, always completely dark and
metallic; mandible bidentate TETRACNEMUS (p. 342)
Head and thorax at least partly yellowish or orange ; mandible tridentate 362
362 (36 1 ) Forewing with marginal vein at least five times as long as broad
*PROCHILONEURUSSi\\estri (p. 327)
Forewing with marginal vein less than twice as long as broad (Fig. 199) . . APHYCUS (p. 234)
363 (358) Body largely yellow or orange, not metallic; notaular lines often present in
anterior part of mesoscutum 364
Body dark reddish, reddish brown or darker and often metallic; notaular lines
absent 365
364 (363) Mandible with three acute teeth; notaular lines usually absent APHYCUS (p. 234)
Mandible bidentate; notaular lines always present in anterior part of meso-
scutum EPISTENOTERYS (p. 273)
365 (363) Dorsum of thorax at least partly reddish or reddish brown; scutellum moder-
ately smooth and shiny; propodeum medially at least one-sixth length of
scutellum and distinctly sculptured TACHINAEPHAGUS (p. 340)
Dorsum of thorax completely dark and metallic; scutellum with distinct
although sometimes shallow reticulate sculpture; propodeum medially not
more than one-eighth as long as scutellum and not sculptured 366
366 (365) Hypopygium extending past apex of last tergite so that it is visible when gaster
viewed from above (Fig. 125) 367
Hypopygium not extending past apex of last tergite 368
367 (366) Ovipositor sheaths slightly but distinctly curving downwards towards apex;
mandible with two teeth and a truncation; forewing never with postmarginal
vein longer than stigmal EPIBLATTICIDA (p. 272)
Ovipositor sheaths more or less straight and not curving downwards towards
apex; mandible with three teeth; forewing with postmarginal vein often
longer than stigmal COCCIDOCTONUS (p. 254)
* Not to be confused with Procheiloneurus Girault (p. 326)
INDO-PACIFIC ENCYRTIDAE
199
Figs 358-370 358-367, Kataka mudigerensis sp. n., (358) head, frontal aspect, <j>, (359) right antenna,
outer aspect, $, (360) left mandible, $, (361) thorax, aspect from left side, $, (362) apex of right
forewing venation, upper surf ace, 9,(363)hypopygium, 9, (364) genitalia, right side, ventral aspect, 9,
(365) genitalia, cf, (366) digiti and apex of aedeagus, (367) right antenna, outer apsect, cf; 368,
Lakshaphagus hautefeuilli (Mahdihassan), right antenna, outer aspect, 9; 369, 370, Manicnemus
indicus (Mani & Saraswat), (369) right antenna, outer aspect, $, (370) scutellum and propodeum
showing sculpture of scutellum, $ .
200 J. S. NOYES & M. HAY AT
368 (366) First funicle segment anelliform and clearly much smaller than the second which
is subquadrate; mandible with one tooth and a broad truncation BACHIANA (p. 241)
First funicle segment subequal in size to second and both clearly transverse;
mandible with three sharp teeth 369
369 (368) Forewing with postmarginal vein longer than stigmal . . . RHOPALENCYRTOIDEA (p. 332)
Forewing with postmarginal vein not longer than stigmal TELETEREBRATUS (p. 341)
370 (30) Scutellum extremely convex and dome-like and separated from axillae by deep
grooves; antennal flagellum strongly compressed from side to side, clava
entire ANANUSIA (p. 231)
Scutellum not or hardly convex and never dome-like, not separated from axillae
by deep grooves; antennal flagellum usually cylindrical to oval in cross-
section, but if flattened then clava three-segmented 371
371 (370) Either first funicle segment longer than pedicel or forewing with postmarginal
vein longer than stigmal; notaular lines absent 372
Neither first funicle segment longer than pedicel nor postmarginal vein of
forewing longer than stigmal, or if postmarginal vein a little longer than
stigmal then notaular lines present 374
372 (371) Eye clearly reaching occipital margin; posterior ocellus distinctly nearer to eye
than to occipital margin; mandible with three teeth PARENCYRTOMYIA (p. 320)
Eye separated from occipital margin by at least about the diameter of a facet;
posterior ocellus at least as near to occipital margin as to eye; mandible
truncate without teeth 373
373 (372) Forewing with postmarginal vein longer than stigmal; frontovertex distinctly
broader than half head width PRIONOMASTIX (p. 325)
Forewing with postmarginal vein not longer than stigmal; frontovertex less
than half head width ENCYRTUS (p. 268)
374 (371) Hypopygium not reaching more than four-fifths along gaster; cerci situated in
basal half of gaster 375
Hypopygium reaching or almost reaching apex of gaster; cerci often situated in
apical half of gaster 385
375 (374) Either forewing with linea calva interrupted or closed on dorsal surface by at
least one line of setae in posterior one-third (Fig. 104), or posterior margin of
pronotum whitish or yellowish and contrasting with orange or darker colour
of remainder of pronotum or mesoscutum 376
Forewing with linea calva not interrupted and open posteriorly; posterior
margin of pronotum translucent or concolorous with mesoscutum 378
376 (375) Mesoscutum and scutellum largely metallic green ZARHOPALOIDES (p. 349)
Neither mesoscutum nor scutellum metallic, usually yellow, orange or dark
brown 377
377 (376) Mandible with three acute teeth METAPHYCUS (p. 298)
Mandible with one tooth and a broad truncation (as in Fig. 189) APHYCOPSIS (p. 233)
378 (375) Apex of clava obliquely truncate with truncate surface longer than adjacent
surface on same side of clava and sutures oblique and converging (Fig. 194);
forewing with marginal vein not longer than stigmal MASHHOODIELLA (p. 295)
Apex of clava rounded, sutures more or less parallel, or if truncate then either
truncate surface shorter than adjacent surface on same side of clava or
marginal vein of forewing at least twice as long as stigmal 379
Figs 371-384 371-373, Muluencyrtus nudipennis sp. n., (371) left antenna, outer aspect (from card-
mounted specimen), $, (372) head, frontal aspect (from card-mounted specimen), $, (373) right
forewing, upper surface (from card-mounted specimen), $; 374-376, Nathismusia southwoodi sp. n.,
(374) left antenna, outer aspect (from card-mounted specimen), 9, (375) head, frontal aspect (from
card-mounted specimen), $, (376) apex of left forewing venation, upper surface, $; 377, 378,
Neodusmetiasangwani (Subba Rao), (377) right antenna, outer aspect, $ , (378) head, frontal aspect, $ ;
379, Olypusa hirsuta sp. n., left antenna, outer aspect, cf ; 380, Neocharitopus sp., right antenna, outer
aspect, $; 381, Ooencyrtus sp., left mandible, $; 382-384, Ovaloencyrtus fijiensis sp. n., (382) apex of
right forewing venation, upper surface (discal setae omitted), $ , (383) genitalia, $ , (384) hypopygium,
9-
INDO-PACIFIC ENCYRTIDAE
201
382
384
202
J. S. NOYES & M. HAY AT
379 (378) Inner margins of eyes clearly converging below anterior ocellus; mesopleurum
not posteriorly enlarged so that when gaster viewed from side metapleurum
and propodeum are at least narrowly touching hind coxa (as in Fig. 139)
CHEILONEUROMYIA (p. 249)
Inner margins of eyes not converging below anterior ocellus; mesopleurum
often posteriorly enlarged and more or less touching basal segment of gaster
so that when thorax viewed from side it clearly separates hind coxa from
metapleurum and propodeum (as in Figs 138, 177) 380
380 (379) Scutellum with a very thin, apical flange which projects above median part of
propodeum PARAPHAENODISCUS (p. 317)
Scutellum without an apical flange 381
381 (380) Forewing with marginal vein punctiform or nearly so OOENCYRTUS (p. 309)
Forewing with marginal vein at least about three times as long as broad 382
382 (381) Forewing with marginal vein less than one and one-half times as long as stigmal;
thorax never strongly metallic 383
Forewing with marginal vein at least twice as long as stigmal; thorax often partly
lustrous and metallic 384
383 (382) Posterior margin of mesoscutum projecting above axillae so that axillae appear
to be separated (as in Fig. 44); scutellum flat, smooth and shiny; clava and
apical funicle segments white and contrasting with dark proximal segments
LEEFMANSIA (p. 292)
Posterior margin of mesoscutum not projecting above axillae, axillae more or
less meeting in middle, separated only by a short carina; scutellum usually
convex or if flat then with distinct sculpture; clava dark and concolorous with
proximal funicle segments MICROTERYS (p. 299)
384 (382) Basal cell of forewing with two areas of dark setae either side of a naked area or
fascia of pale translucent setae *PROCHEILONEURUSGirault (p. 326)
Basal cell of forewing with only one area of dark setae, this adjacent to linea
calva, proximal of this either naked or a small patch of pale setae
CHEILONEURUS (p. 249)
385(374) Forewing with marginal vein at least twice as long as stigmal (Fig. 196);
propodeum medially at least half as long as scutellum (Fig. 195) SAKENCYRTUS (p. 336)
Forewing with marginal vein not longer than stigmal, or if slightly so then
propodeum medially much less than half as long as scutellum 386
386 (385) Notaular lines absent; paratergites present (at least represented by a membra-
nousstrip) PARECTROMOIDELLA (p. 319)
Notaular lines present in at least anterior one-third of mesoscutum; paratergites
absent 387
387 (386) Notaular lines complete 388
Notaular lines not reaching more than halfway across mesoscutum 389
388 (387) Forewing with stigmal vein straight, not abruptly bent immediately below
marginal vein and thus forming an angle of about 45, linea calva clearly open
towards posterior margin of wing (Fig. 199) APHYCUS (p. 234)
Forewing with stigmal vein abruptly bent below marginal vein and thus running
nearly parallel to anterior margin of wing and forming an angle of clearly less
than 30 , linea calva closed towards posterior margin of wing by at least two
lines of setae on dorsal surface (Fig. 198) HOMALOTYLUS (p. 287)
389 (387) Clava solid with a strong oblique truncation (Fig. 200); forewing with stigmal
vein arising from submarginal vein before it reaches anterior wing margin,
linea calva broadening towards posterior wing margin and very clearly open
(Fig. 201) ISODROMUS (p. 289)
Clava two- or three-segmented with apex rounded; forewing with stigmal vein
arising from marginal vein at anterior wing margin, linea calva with sides
subparallel and more or less closed near posterior margin by setae on dorsal
surface of wing (Fig. 197) 390
390 (389) Clava two-segmented; mandible bidentate EPISTENOTERYS (p. 273)
Clava three-segmented; mandible tridentate PSEUDOCOCCOBIUS (p. 329)
* Not to be confused with Prochiloneurus Silvestri (p. 327)
INDO-PACIFIC ENCYRTIDAE
203
Figs 385-397 385-389, Papuna nemis sp. n., (385) right antenna, outer aspect, <j>, (386) base of right
forewing, upper surface, $, (387) head, frontal aspect, $, (388) right mandible, J, (389) genitalia, left
side, ventral aspect, $; 390-395, Parablatticida spp., (390) left antenna, outer aspect, $, (391) left
antenna, outer aspect, cf, (392) left mandible, $, (393) right antenna, outer aspect, $, (394) base of
right forewing, upper surface, $ , (395) left antenna, outer aspect, $ ; 396, 397, Paraclausenia herbicola
Hayat, (396) right antenna, outer aspect, $, (397) right mandible, $.
204 J. S. NOYES & M. HAY AT
391 (31) Mesopleurum posteriorly enlarged and more or less touching basal segment of
gaster so that when thorax viewed from side it clearly separates hind coxa
from metapleurum and propodeum (as in Figs 138, 177) 392
Mesopleurum not enlarged so that when thorax viewed from side hind coxa
touches metapleurum and propodeum thus separating mesopleurum from
gaster (as in Figs 139, 140) 394
392 (391) Antennal scrobes narrow, elongate and deeply impressed, separated from
anterior ocellus by only a little more than its own diameter; interantennal
prominence very long and dorsally sharply delimited, pointed and clearly
separate from frontovertex (Figs 184, 185) TACHARDIAEPHAGUS (p. 340)
Antennal scrobes relatively shallow, not long and more or less semicircular and
dorsally separated from anterior ocellus by at least twice its diameter;
interantennal prominence dorsally more or less rounded or confluent with
frontovertex and not sharply delimited 393
393 (392) Clava very large with a strong oblique apical truncation and at least twice as long
as funicle (Fig. 202); mesoscutum and scutellum with striate-reticulate sculp-
ture (Fig. 203) AGARWALENCYRTUS (p. 226)
Clava smaller, usually shorter than funicle although occasionally a little longer,
but never twice as long, usually more or less apically rounded, although
occasionally with a short oblique truncation (Fig. 246); mesoscutum and
scutellum without striate-reticulate sculpture OOENCYRTUS (p. 309)
394 (391) Clava entire (Fig. 204) COPIDOSOMA (p. 257)
Clava two- or three-segmented 395
395 (394) Body strongly flattened; pronotum longitudinally divided in middle (as in Fig.
38); mandible bidentate with two equal teeth RHOPUS (p. 332)
Body not flattened ; pronotum entire ; mandible not bidentate , or if so then teeth
are unequal in length 396
396 (395) Antennal scrobes extending more than half way between toruli and anterior
ocellus, their upper limit not semicircular; forewing with postmarginal vein
longer than stigmal (Fig. 103) ERENCYRTUS(p.274)
Antennal scrobes more or less semicircular and only occasionally reaching more
than half way between toruli and anterior ocellus, but if so then postmarginal
vein of forewing not longer than stigmal 397
397(396) Gaster dark with basal segment yellow or orange; ovipositor, although not
exserted, curved upwards PROTYNDARICHOIDES (p. 328)
Gaster unicolorous, dark, without paler basal segment; ovipositor straight or
curved downwards 398
398 (397) Eye clothed in short translucent setae, each not longer than diameter of a facet;
hypopygium not extending more than two-thirds along gaster; mandible with
two teeth and a truncation; forewing with postmarginal vein not longer than
stigmal ; propodeum medially not more than one-fifth as long as scutellum
SYRPHOPHAGUS (p. 338)
Eye clothed in long, occasionally very dense, setae, each clearly longer than a
facet; hypopygium almost always reaching apex of gaster or nearly so;
mandible with from one to three sharp teeth, never with a truncation;
forewing with postmarginal vein sometimes longer than stigmal; propodeum
often medially more than one-fifth as long as scutellum 399
Figs 398-411 398, 399, Paraschedius spp., (398) right antenna, outer aspect, $, (399) right forewing,
upper surface, $; 400, Parectromoidella lowelli (Girault), apex of right forewing venation, upper
surface, $; 401-406, Pasulinia gentha sp. n., (401) right forewing, upper surface, 9, (402) sculpture in
centre of mesoscutum (area approx. 0-1 mm square), $, (403) sculpture in centre of scutellum (area
approx. 0-1 mm square), $, (404) hypopygium, $, (405) mid tibia and tarsus, $, (406) genitalia, left
side, ventral aspect, $; 407-410, Philosindia longicornis sp. n., (407) right antenna, outer aspect, $,
(408) genitalia, ventral aspect, left side, $, (409) sculpture in centre of mesoscutum (area approx. 0-1
mm square), $, (410) sculpture in centre of scutellum (area approx. 0-1 mm square), $; 411,
Praleurocerus viridis (Agarwal), apex of right forewing venation, upper surface, $ .
INDO-PACIFIC ENCYRTIDAE
205
206 J. S. NOYES & M. HAY AT
399 (398) Ovipositor at least slightly exserted and with sheaths flattened from side to side ;
gonostyli free and at least one-quarter length of ovipositor (Fig. 430);
mandible with three acute teeth (Fig. 144) TACHINAEPHAGUS (p. 340)
Ovipositor not exserted and not visible externally; gonostyli fused to second
valvifers (Fig. 415) and not more than one-quarter as long as ovipositor;
mandible with one or two teeth RHYTIDOTHORAX (p. 333)
400(31) Clava apically obliquely truncate and entire or three-segmented, if three-
segmented then outer suture strongly oblique and converging with inner (Figs
205, 21 1) 401
Clava apically rounded and two- or three-segmented, sutures more or less
parallel 403
401 (400) Clava three-segmented (Fig. 205) PROLEUROCEROIDES (p. 327)
Clava entire (Fig. 211) 402
402 (401) Scutellum strongly convex and separated from axillae by deep grooves; head
and dorsum of thorax with extremely dense, very short, appressed setae;
forewing with marginal vein longer than broad ANANUSIA (p. 231)
Scutellum moderately convex and separated from axillae by normal sutures;
head and dorsum of thorax with sparse , moderately long erect setae ; forewing
with marginal vein punctiform (Fig. 212) TAFTIA (p. 341)
403 (400) Body distinctly dorso-ventrally flattened; clava frequently two-segmented 404
Body not dorso-ventrally flattened; clava always three-segmented 405
404 (403) Exserted part of ovipositor about one-quarter length of gaster with sheaths dark
brown and contrasting with remainder of body which is yellow; pronotum
clearly visible, entire and triangular in dorsal view and longer than meso-
scutum (Fig. 209); head sub-opisthognathous; mandible tridentate; clava
two-segmented (Fig. 208) ASTYMACHUS (p. 236)
Ovipositor not exserted; pronotum not clearly visible, obscured by head in
dorsal view and longitudinally divided in middle, not triangular in shape (as in
Fig. 38) and shorter than mesoscutum; head prognathous; mandible biden-
tate; clava frequently three-segmented RHOPUS (p. 332)
405 (403) Forewing with linea calva interrupted in its posterior one-third by at least two
lines of setae on dorsal surface of wing (Figs 95, 104), or closed at this point by
several lines of setae (Fig. 159) 406
Forewing with linea calva not interrupted, or if closed then by not more than one
line of setae near posterior margin of wing 408
406 (405) Forewing with linea calva interrupted in its posterior half (Fig. 104); mandible
tridentate; hypopygium not reaching apex of gaster METAPHYCUS (p. 298)
Forewing with linea calva more or less entirely closed in its posterior one-half by
setae on dorsal surface of wing; mandible bidentate; hypopygium reaching
apex of gaster 407
407 (406) Forewing with a distinct pattern of dark and pale setae, stigmal vein long, more
than one-quarter length of submarginal vein, marginal vein not quite reaching
anterior margin of wing (Fig. 159) MASHHOODIA (p. 295)
Forewing without a distinct pattern of dark and pale setae, stigmal vein less than
one-quarter as long as submarginal vein, marginal vein confluent with an-
terior margin of forewing (Fig. 95) ANAGYRUS (p. 229)
408 (405) Pronotum triangular in dorsal view, about twice as broad as long and at least
about two-thirds as long as mesoscutum (Fig. 149) CHEILONEURELLA (p. 248)
Pronotum very transverse, in dorsal view at least about five times as broad as
long and not more than about half as long as mesoscutum 409
409 (408) Antennal scrobes deeply impressed and more or less sharply margined laterally;
interantennal prominence long, reaching more than half way between
antennal toruli and anterior ocellus, sharp at its apex and not confluent with
frontovertex (Figs 184, 185) TACHARDIAEPHAGUS (p. 340)
Antennal scrobes shallow to moderately impressed with lateral margins
rounded and not well defined; interantennal prominence short, not reaching
half way between antennal toruli and anterior ocellus and rounded at its apex,
or if longer then confluent with frontovertex 410
INDO-PACIFIC ENCYRTIDAE
207
Figs 412-424 412-414, Rhopus spp., (412) right antenna, outer aspect, $, (413) left antenna, outer
aspect, $, (414) base of left forewing, upper surface, 9; 415, Rhytidothomx sp., genitalia, left side,
ventral aspect, $; 416, Proleuroceroides sp., base of right forewing, upper surface, 9; 417-421,
Ruanderoma sankarani sp. nov., (417) genitalia, left side, ventral aspect, $, (418) genitalia, cf , (419)
digiti and apex of aedeagus, (420) right antenna, outer aspect, $ , (421) right antenna, outer aspect, cf ;
422, Sakencyrtus sp., right antenna, outer aspect, $; 423, 424, Sakencyrtus sp., (423) right antenna,
outer aspect, $ , (424) right forewing, upper surface, $ .
208
J. S. NOYES & M. HAY AT
410 (409) Eye and frontovertex with very conspicuous dark setae which are subequal in
length APHYCOMORPHA (p. 233)
Eye more or less naked, if setae on eye clearly visible then they are distinctly
shorter than those on frontovertex 411
411 (410) Body entirely yellow, except for a black spot in centre of pronotum; fronto-
vertex and dorsum of thorax clothed in conspicuous black setae
PSYLLAPHYCUS (p. 332)
Body generally orange or darker; frontovertex clothed in relatively incon-
spicuous setae , dorsum of thorax occasionally with conspicuous setae 412
412 (411) Dorsum of thorax and mesopleura largely metallic green ZARHOPALOIDES (p. 349)
If dorsum of thorax and mesopleura with dark areas then these usually dark
brown, but never metallic green 413
413 (412) Mesopleurum brown and distinctly darker than dorsum of thorax; mandible
with three teeth [Mesanusia] speciosa Girault (p. 353)
Mesopleurum not darker than dorsum of thorax; mandible with two teeth and a
truncation 414
414 (413) Scape longer than minimum width of frontovertex NEASTYMACHUS (p. 304)
Scape shorter than minimum width of frontovertex APHYCOMORPHA (p. 233)
415 (32) Clava entire (Fig. 108) TYNDARICOPSIS (p. 347)
Clava three-segmented 416
416 (415) Clava much broader than funicle, strongly obliquely truncate and at least as long
as funicle; scutellum (excluding axillae) at least a little longer than broad and
convex TYNDARICHUS (p. 346)
Clava apically rounded or occasionally obliquely truncate and shorter than
funicle, or if obliquely truncate and longer than funicle then scutellum convex
but at least a little broader than long 417
417 (416) Propodeum medially at least one-fifth as long as scutellum and usually with
some irregular carinae present medially (Fig. 210); occipital margin rounded;
triangular expansion of submarginal vein of forewing weak (Fig. 147),
forewing often with an elongate wedge-shaped pale fuscous mark from apex;
top of antennal toruli at least a little above lower eye margins . . . MAHENCYRTUS (p. 294)
Propodeum medially much less than one-sixth as long as scutellum and without
carinae (Fig. 206); occipital margin more or less sharp; forewing with triangu-
lar expansion of submarginal vein distinct (Fig. 207), forewing completely
hyaline ; top of antennal toruli below level of lower eye margins
PARECHTHRODRYINUS (p. 319)
418 (35) Either forewing with marginal vein absent (Figs 172, 191) or head with distinct
piliferous punctures which are usually not separated by more than their own
diameters giving it a thimble-like appearance 419
Forewing with marginal vein present ; head without deep piliferous punctures . 426
419(418) Marginal fringe of forewing absent; clava apically rounded with sutures
parallel HETEROCOCCIDOXENUS (p. 286)
Forewing with marginal fringe present; clava usually obliquely truncate and
with sutures oblique and converging 420
420(419) Clava weakly obliquely truncate; first funicle segment almost always at least
slightly longer than pedicel, rarely a little shorter 421
Clava strongly obliquely truncate; first funicle segment always distinctly shorter
than pedicel 423
421 (420) Forewing with postmarginal vein not longer than stigmal (Fig. 191) COWPERIA (p. 259)
Figs 425-439 425, 426, Sakencyrtus sp. , (425) right antenna outer aspect, 9 , (426) left forewing, $ ; 427,
Saprencyrtus casuarinae (Girault), left antenna, outer aspect (from card-mounted specimen), $; 428,
Tachinaephagus sp., right antenna, outer aspect, $; 429, Thomsonisca pakistanensis (Ahmad), apex of
left forewing venation, $; 430, Tachinaephagus sp., genitalia, left side, ventral aspect, 9; 431, 432,
Teleterebratus perversus Compere & Zinna, (431) right antenna, outer aspect, 9, (432) right mandible,
9 ; 433-439, Tongyus nesus sp. n. , (433) right antenna, outer aspect, 9 , (434) genitalia left side, 9 , (435)
hypopygium, 9> (436) sculpture of frontovertex anterior to anterior ocellus (area approx. 0-1 mm
square), 9 , (437) right antenna, outer aspect, cf , (438) genitalia, cf , (439) digiti and apex of aedeagus.
INDO-PACIFIC ENCYRTIDAE
209
439
210
J. S. NO YES & M. HAYAT
Forewing with postmarginal vein clearly longer than stigmal 422
422 (421) Clava entire PARACLADELLA (p. 315)
Clava three-segmented ANAGYRODES (p. 228)
423 (420) Frontovertex narrow , at narrowest point not more than one-quarter head width ;
antennal scrobes very deep and dorsally sharply margined; mesopleurum
enlarged posteriorly and more or less touching basal segment of gaster so that
when thorax viewed from side the hind coxa is clearly separated from
metapleurum and propodeum by hind margin of mesopleurum (as in Fig. 177)
AMIRA (p. 228)
Frontovertex broad, at narrowest point at least one-third head width; antennal
scrobes very shallow or absent and not sharply margined; mesopleurum not
posteriorly enlarged and clearly separated from first segment of gaster so that
when thorax is viewed from side the hind coxa is broadly in contact with
metapleurum and propodeum (as in Fig. 140) 424
424 (423) Scutellum broad and flat, clearly much broader than long AMICENCYRTUS (p. 227)
Scutellum distinctly convex, at least as long as broad, usually longer 425
425 (424) Mesoscutum and scutellum with very deep distinct piliferous punctures which
are more or less touching each other and give a thimble-like appearance;
propodeum relatively long medially, at least about one-quarter as long as
scutellum BOTHRIOTHORAX (p. 243)
Mesoscutum with shallow, very indistinct piliferous punctures, scutellum with
raised, closely meshed, reticulate sculpture; propodeum very short, not more
than one-tenth as long as scutellum MENISCOCEPHALUS (p. 296)
426 (418) Forewing with postmarginal vein as long as or longer than stigmal 427
Forewing with postmarginal vein shorter than stigmal 430
427 (426) Clava strongly obliquely truncate and much broader than funicle (Fig. 211); face
with a strong transverse carina from below eyes and across top of antennal
scrobes TAFTIA (p. 341)
Clava more or less apically rounded and not or hardly broader than funicle; face
without a strong transverse carina 428
428 (427) Forewing with postmarginal vein clearly much longer than stigmal 429
Forewing with postmarginal vein about as long as stigmal 430
429 (428) Clava entire; mandible long with two very short apical teeth PARACLADELLA (p. 315)
Clava three-segmented; mandible short and broad with one very small tooth and
a broad truncation PRIONOMASTIX (p. 325)
430 (426, Posterior margin of mesoscutum projecting slightly backwards above axillae so
428) that when thorax is in normal resting position and viewed from above, the
axillae appear to be broadly separated (as in Fig. 44); mesopleurum enlarged
posteriorly and more or less touching basal segment of gaster so that when
thorax viewed from side it clearly separates metapleurum and propodeum
from hind coxa (Fig. 177) 431
Posterior margin of mesoscutum almost straight, not projecting above axillae
and so axillae appear to meet medially (as in Fig. 42); mesopleurum not
posteriorly enlarged and not touching basal segment of gaster so that when
thorax viewed from side the hind coxa touches metapleurum and propodeum
(asinFigs 139, 140) 432
43 1 (430) Mesoscutum and scutellum with coarse punctate-reticulate sculpture ; mandible
bidentate FULGORIDICIDA (p. 278)
Mesoscutum and occasionally also scutellum with shallow reticulate sculpture;
mandible with one or two teeth and a truncation or obscurely tridentate
OOENCYRTUS (p. 309)
432 (430) Hypopygium more or less reaching apex of gaster; occipital margin sharp behind
eyes; forewing with sensillae at apex of stigmal vein arranged symmetrically in
a square, uncus absent (Figs. 142, 183); mandible with three acute teeth
COPIDOSOMA (p. 257)
Hypopygium not usually extending more than two-thirds along gaster, although
very occasionally nearly reaching apex; occipital margin often more or less
rounded, particularly behind eyes; forewing with sensillae at apex of stigmal
INDO-PACIFIC ENCYRTIDAE
211
448
Figs 440-450 440, Xenostryxis sp. , right antenna, outer aspect, $ ; 441 , Yasumatsuiola sp. , right antenna,
outer aspect, $; 442-444, Zaommoencyrtus spp., (442) head, frontal aspect, $, (443) head, frontal
aspect, 9> (444), right antenna, outer aspect, $?; 445-450, Zozoros sinemarginis sp. n., (445) head,
frontal aspect, $?, (446) genitalia, right side, ventral aspect, $, (447) sculpture in centre of mesoscutum
(area approx. 0-1 mm square), $ , (448) sculpture in centre of scutellum (area approx. 0-1 mm square),
$, (449) hypopygium, $, (450) propodeum, dorsal aspect showing sculpture, $.
212
J. S. NO YES & M. HAYAT
vein arranged asymmetrically and not in a square, uncus present; mandible
with one or two teeth and a truncation, or rarely with three teeth 433
433 (432) Mandible with one or two teeth and a truncation (Figs 121-122) PSYLLAEPHAGUS (p. 330)
Mandible with three acute teeth (Fig. 178) PARAENASOMYIA (p. 316)
434 (38) Antennal toruli situated high on head, their lowest margins not or hardly below
level of lowest eye margins when head viewed from front (Fig. 113) 435
Antennal toruli lower on head, their lowest margins well below level of lowest
eye margins when head viewed from front 436
435 (434) Scape not or hardly longer than malar space; pedicel and funicle segments
subequal in size and shape (Fig. 227); posterior ocellus about equidistant from
eye and occipital margins, the latter sharp GAHANIELLA (p. 278)
Scape at least twice as long as malar space; pedicel and funicle segments not
subequal in size and shape, some at least distinctly narrower or shorter than
others; posterior ocellus much closer to eye margin than to occipital margin,
the latter more or less rounded MAYRIDIA (p. 295)
436 (434) Mesopleurum posteriorly enlarged so that it more or less touches basal segment
of gaster and when thorax viewed from side it clearly separates hind coxa from
metapleurum and propodeum (Fig. 177) 437
Mesopleurum not so conspicuously enlarged, so that when thorax viewed from
side hind coxa more or less broadly touches metapleurum and propodeum
and together they clearly separate mesopleurum from basal segment of gaster
(asinFigs. 139, 140) 439
437 (436) Mandible bidentate; vertex, mesoscutum and scutellum with coarse punctate-
reticulate sculpture FULGORIDICIDA (p. 278)
Mandible with one or two teeth and a truncation; at least mesoscutum with
relatively shallow sculpture 438
438 (437) Forewing with marginal vein less than twice as long as broad (Fig. 152)
OOENCYRTUS (p. 309)
Forewing with marginal vein more than twice as long as broad (Fig. 153)
TRICHOMASTHUS (p. 346)
439 (436) Head triangular in profile and strongly inflexed at top of antennal scrobes (Figs
72, 228); mandible with four teeth or one small tooth and a very broad
truncation (Figs 116, 229); mesoscutum with dark setae; hypopygium with
apex not more than two-thirds along gaster 440
Head in profile more or less anteriorly evenly rounded, not strongly inflexed at
top of antennal scrobes and not triangular, or if slightly so then either
mandible has three acute teeth and the mesoscutum is clothed in dense white
setae or the hypopygium reaches or very nearly reaches apex of gaster 441
440 (439) Clava more or less apically rounded, not truncate and with sutures more or less
parallel; gaster unicolorous and metallic; mandible with four teeth (Fig. 116)
ADELENCYRTUS (p. 223)
Clava strongly obliquely truncate with outer suture oblique (Fig. 230); gaster
largely yellow or orange; mandible with one small tooth and a very broad
truncation (Fig. 229) PASULINIA (p. 320)
441 (439) Head with very distinct, deep piliferous punctures, these usually shiny-
bottomed and contrasting with the relatively dull areas between, each
puncture giving rise to a broadened silvery white seta MENISCOCEPHALUS (p. 296)
Either head without deep piliferous punctures, or if punctures deep and
conspicuous then they give rise to dark setae and not broadened silvery white
setae 422
442 (441) Base of gaster at least partly yellow or orange; clava two- or three-segmented
and not obliquely truncate 443
Gaster unicolorous and dark, or if with base yellow then clava entire and
strongly obliquely truncate 444
443 (442) Forewing with stigmal vein long, at least a little longer than marginal; clava
apically rounded; mandible with two teeth and a truncation
DIAPHORENCYRTUS (p. 263)
Forewing with stigmal vein short or subsessile, distinctly shorter than marginal
INDO-PACIFIC ENCYRTIDAE
213
(Figs 231, 232); clava apically rounded or transversely truncate (Figs 233,
234); mandible with three acute teeth PROTYNDARICHOIDES (p. 328)
444(442) Hypopygium reaching or very nearly reaching apex of gaster, or if only
extending about three-quarters along gaster then clava apically rounded;
clava three-segmented 445
Hypopygium not extending more than two-thirds along gaster, or if slightly
more then clava either apically obliquely truncate or solid or two-segmented 451
445 (444) Forewing with filum spinosum directed towards junction of submarginal and
marginal veins and thus distinctly converging with setae on proximal margin
of lineacalva (Fig. 127) CERCHYSIELLA (p. 246)
Forewing with filum spinosum more or less directed towards junction of stigmal
and marginal veins and thus subparallel to setae on proximal margin of linea
calva (Figs 139, 236, 238) 436
446 (445) Head and thorax with deep, fine, punctate sculpture; forewing with postmar-
ginal vein longer than stigmal BLASTOTHRIX (p. 242)
Sculpture of head and thorax shallow, although scutellum occasionally has deep
reticulate sculpture, of if head and thorax with punctate sculpture then
postmarginal vein of forewing not longer than stigmal 447
447 (446) Eye clearly overreaching occipital margin PARECTROMOIDES (p. 320)
Eye clearly separated from occiput by occipital margin which is more or less
sharp at this point 448
448 (447) Eye more or less naked ; head and thorax bright metallic green or blue-green
PSYLLAEPHAGUS (p. 330)
Eye distinctly hairy; head and thorax usually dull but occasionally metallic 449
449 (448) Propodeum medially not more than one-sixth length of scutellum; mandible
with two teeth and a truncation, three teeth or four teeth EXORISTOBIA (p. 277)
Propodeum medially at least one-fifth as long as scutellum; mandible with from
one to three sharp teeth 450
450 (449) Mandible with three acute teeth (Fig. 144); ovipositor usually slightly exserted
with sheaths flattened from side to side TACHINAEPHAGUS (p. 340)
Mandible with one or two teeth; ovipositor always hidden and, together with
gonostyli, not visible externally RHYTIDOTHORAX (p. 333)
451(444) Forewing with postmarginal vein longer than stigmal ... AUSTROENCYRTOIDEA (p. 238)
Forewing with postmarginal vein not longer than stigmal 452
452 (451) Clava solid or two-segmented 453
Clava three-segmented 454
453 (452) Mandible with three acute teeth; eye clearly separated from occiput by sharp
occipital margin; forewing with sensillae at apex of stigmal vein arranged
symmetrically in a square, uncus absent (Figs 142, 183) COPIDOSOMA (p. 257)
Mandible with one or two teeth and a truncation; eye overreaching occipital
margin; forewing with sensillae at apex of stigmal vein not arranged in a
square, uncus present ISODROMOIDES (p. 289)
454 (452) Forewing with marginal vein at least three times as long as broad (Figs 186, 192);
clava either transversely or obliquely truncate 455
Forewing with marginal vein only slightly longer than broad, clava apically
rounded 456
455 (454) Clava with a strong oblique truncation (Fig. 224) ; scutellum fairly flat with fine ,
deep, reticulate sculpture which gives it a matt appearance; forewing with
marginal vein at least five times as long as broad (Fig. 192); mandible with
three acute teeth (Fig. 223) ECHTHROGONATOPUS (p. 267)
Clava with only a slightly oblique, transverse, apical truncation; scutellum
slightly convex with shallow, reticulate sculpture and at least slightly shiny;
forewing with marginal vein not more than four times as long as broad
(Fig. 186); mandible with one or two teeth and a truncation . . . SYRPHOPHAGUS (p. 353)
456 (454) Mandible with three acute teeth (Fig . 178) ; thorax dull purple-brown with green
and coppery reflections PARAENASOMYIA (p. 316)
Mandible with one tooth and a truncation (or occasionally with two teeth and a
truncation) (Figs 121, 122); thorax metallic green PSYLLAEPHAGUS (p. 330)
214
J. S. NOYES & M. HAY AT
457 (39) Cereal plates in apical half of gaster 458
Cereal plates in basal half of gaster 460
458(457) Forewing with postmarginal vein shorter than stigmal (Fig. 215); hind tibia
strongly oblique at apex (Fig. 216) MULUENCYRTUS (p. 300)
Forewing with postmarginal vein clearly longer than stigmal; hind tibia not
strongly oblique at apex 459
459 (458) Hypopygium reaching or very nearly reaching apex of gaster ; mandible with one
long, sickle-shaped tooth (or possibly two very short apical teeth giving the
mandible a unidentate appearance) PARALEPTOMASTIX (p. 316)
Hypopygium not reaching more than halfway along gaster; mandible with three
apical teeth EUCOMOMORPHELLA (p. 276)
460 (457) Costal cell of forewing with setae evenly distributed over its dorsal surface
(Fig. 217); clava entire and apically rounded (Fig. 219); mandible edentate
(Fig. 218) OLYPUSA (p. 307)
Forewing with setae on dorsal surface of costal cell restricted to a single line in
apical half only; clava two- or three-segmented; mandible with two or three
teeth 461
461 (460) Clava apically rounded; eye not overreaching occipital margin 462
Clava with a strong oblique apical truncation; eye reaching occiput 463
462 (461) Forewing with a complete hyaline fascia distal to venation, marginal vein clearly
longer than stigmal (Figs 141, 222); mandible with three acute teeth (Fig. 221)
SAPRENCYRTUS (p. 336)
Forewing without a complete hyaline fascia distal to venation, marginal vein
shorter than stigmal; mandible with one or two teeth and a truncation
(Figs 121, 122) PSYLLAEPHAGUS (p. 330)
463 (461) Mesopleurum not posteriorly enlarged and not touching basal segment of gaster
so that when thorax viewed from side hind coxa more or less broadly touches
metapleurum and propodeum (as in Fig. 140); dorsum of thorax dull and not
strongly metallic; clava white contrasting with dark funicle segments (Fig.
137); mandible tridentate PARENCYRTOMYIA (p. 320)
Mesopleurum posteriorly enlarged and more or less touching basal segment of
gaster so that when thorax viewed from side it clearly separates hind coxa
from metapleurum and propodeum (as in Fig. 177); dorsum of thorax strongly
shining purple or blue; flagellum unicolours and dark; mandible with one or
two teeth and a truncation (Fig. 71) PARATETRALOPHIDEA (p. 319)
464 (39) Forewing with postmarginal vein much longer than stigmal 465
Forewing with postmarginal vein not longer than stigmal 466
465 (464) Propodeum medially not longer than one-fifth length of scutellum; eye more or
less naked, setae clearly shorter than diameter of a facet; infuscation of
forewing diffuse and not forming a distinct pattern ENCYRTOIDEA (p. 268)
Propodeum medially more than one-fifth as long as scutellum; eye with numer-
ous short setae, each longer than diameter of a facet; infuscation of forewing
moderately strong and forming a distinct pattern BORROWELLA (p. 242)
466 (464) Mesopleurum with anterior half more or less smooth and shiny and posterior
half with distinct, clearly delimited shiny buttomed punctures
PARASTENOTERYS (p. 318)
Mesopleurum with similar sculpture in both anterior and posterior halves,
usually entirely smooth or nearly so 467
467 (466) Head and dorsum of thorax with conspicuous piliferous punctures of thimble-
like appearance RHYTIDOTHORAX (p. 333)
Head and dorsum of thorax with indistinct, shallow piliferous punctures and not
of thimble-like appearance 468
468 (467) Mesopleurum enlarged posteriorly so that it more or less touches basal segment
of gaster so that when thorax viewed from side it separates hind coxa from
metapleurum and propodeum (as in Figs 138, 177) 469
Mesopleurum not so posteriorly enlarged and not touching basal segment of
gaster so that when thorax viewed from side the hind coxa is more or less
broadly touching metapleurum and propodeum (as in Figs 139, 140) 471
INDO-PACIFIC ENCYRTIDAE
215
469 (468) Clava not obliquely truncate, truncate surface shorter than ventral surface of
clava(Fig. 220) TRICHOMASTHUS (p. 346)
Clava very strongly obliquely truncate, the truncate surface much longer than
remainder of ventral surface of clava (Figs 67, 224) 470
470 (469) Scutellum smooth and very shiny; fore wing with marginal vein not or hardly
longer than stigmal (Fig. 68); mandible with two teeth and a very broad
truncation (Fig. 225) OVALOENCYRTUS (p. 310)
Scutellum with deep reticulate sculpture; marginal vein of forewing more than
twice as long as stigmal (Fig. 192); mandible tridentate (Fig. 223)
ECHTHROGONATOPUS (p. 267)
471 (468) Forewing with marginal vein relatively short, not more than twice as long as
stigmal 472
Forewing with marginal vein longer, at least three times as long as stigmal 474
472 (471) Clava entire and obliquely truncate at apex ISODROMOIDES (p. 289)
Clava three-segmented and not obliquely truncate (although dorsal surface
occasionally more strongly curved than ventral) 473
473 (472) Clava with dorsal surface clearly more strongly curved than ventral surface;
antennal toruli separated from mouth margin by much more than their own
lengths MAYRIDIA (p. 295)
Clava with dorsal surface similarly curved to ventral surface; antennal toruli
separated from mouth margin by not more than their own lengths . . AENASIELLA (p. 224)
474 (471) Scutellum more or less flat with punctate-reticulate sculpture and matt; fore-
wing with apex of stigmal vein separated from anterior wing margin by less
than the maximum depth of uncus CHEILONEURUS (p. 249)
Scutellum clearly convex and with shallow reticulate sculpture and at least
slightly shiny; forewing with apex of stigmal vein separated from anterior
wing margin by more than maximum depth of uncus (Fig. 226)
HYPERGONATOPVS (p. 288)
475 (40) Forewing with marginal vein absent AMIRA (p. 228)
Forewing with marginal vein present 476
476 (475) Hypopygium not reaching more than two-thirds along gaster; propodeal spira-
cle surrounded by dense white setae which continue along sides of propodeum
and metapleurum to hind coxa which is also clothed in dense white setae
MENISCOCEPHALUS (p. 296)
Hypopygium more or less reaching apex of gaster; propodeal spiracle not
surrounded by dense white setae, or if so then these do not continue down
sides of propodeum and metapleurum to hind coxa 477
477 (476) Forewing with filum spinosum directed towards junction of submarginal and
marginal veins so that it clearly converges with line of setae on proximal
margin of lineacalva (Fig. 127) CERCHYSIELLA (p. 246)
Forewing with filum spinosum absent, margins of linea calva more or less
parallel (Fig. 235) 478
478 (477) Frontovertex at narrowest point about one-quarter head width 479
Frontovertex at narrowest point at least one-third head width, usually much
wider 480
479 (478) Forewing with postmarginal vein longer than stigmal BLEPYRUS (p. 242)
Forewing with postmarginal vein not longer than stigmal AENASIUS (p. 225)
480 (478) Forewing with marginal vein long, at least half as long as submarginal and at
least three times as long as either the short stigmal or postmarginal veins,
anterior margin of wing not incised at apex of costal cell (Fig. 237)
METAPHAENODISCUS (p. 297)
Forewing with marginal vein short, not more than half as long as either
postmarginal or stigmal veins and less than one-tenth as long as submarginal
vein, anterior margin of wing incised at apex or costal cell (Fig. 235)
CLADISCODES (p. 251)
481 (41) Head in profile more or less evenly rounded anteriorly, not triangular; occipi-
tal margin sharp; mandible with two or three acute teeth 482
Head in profile triangular, face strongly inflexed at top of antennal scrobes (as in
216
J. S. NOYES & M. HAYAT
Fig. 72); occipital margin more or less rounded; mandible with one or two
teeth and a truncation or four teeth 488
482 (481) Clava with a strong oblique truncation (Figs 43, 239,241) 483
Clava apically rounded 486
483 (482) Hypopygium reaching apex of gaster; forewing with marginal vein absent
(Fig. 240) ; frontovertex at narrowest point broader than length of scape
LUTHERISCA (p. 294)
Hypopygium not reaching more than two-thirds along gaster; forewing with
marginal vein present; frontovertex at narrowest point narrower than length
of scape 484
484 (483) Forewing with marginal vein not more than twice as long as broad, sensillae
at apex of stigmal vein arranged symmetrically in a square, uncus absent
(Figs 142, 183, 249) COPIDOSOMA (p. 257)
Forewing with marginal vein at least four times as long as broad, sensillae at
apex of stigmal vein not arranged symmetrically and not in a square, uncus
present (Fig. 226) 485
485 (484) Clava about as long as pedicel and funicle together and darker than funicle, all
funicle segments clearly transverse; mesoscutum with sparse white setae;
scutellum fairly flat and with distinctly deeper reticulate sculpture than that on
mesoscutum; mandible with two teeth and a truncation BAEOANUSIA (p. 241)
Clava not longer than funicle and concolorous; funicle segments subquadrate;
mesoscutum with dark setae; scutellum fairly convex and not with deeper
sculpture than mesoscutum; mandible tridentate HYPERGONATOPUS (p. 288)
486 (482) Basal segment of gaster at least dorsally white or yellow and contrasting with the
dark remainder of gaster 487
Gaster unicolorous, dark and shiny, not partly white or yellow 488
487 (486) Infuscation of forewing restricted to a subapical fuscous streak (Fig. 245) ; setae
on apical funicle segments normal; mandible tridentate COAGERUS (p. 251)
Infuscation of forewing more extensive with at least a broad complete fuscous
fascia from apex of venation; apical funicle segments occasionally with
flattened scale-like setae; mandible usually with one or two teeth and a
truncation, rarely with three teeth CHEILONEURUS (p. 249)
488 (481, Forewing strongly infuscate from apical one-third of submarginal vein to apex
486) and enclosing at least three hyaline spots ; head with a strong transverse line of
dense silvery white setae below eyes and across face below angle efface
EPITETRACNEMUS (p. 273)
Forewing with infuscation often rather weak and at most with only two hyaline
spots in infuscate area; transverse line of setae across face absent or with setae
very sparse 489
489 (488) Forewing with postmarginal vein clearly much longer than either marginal or
stigmal veins, stigmal vein not or hardly shorter than marginal; mandible with
two teeth and a truncation, never with four teeth ADELENCYRTOIDES (p. 223)
Forewing with postmarginal vein not or hardly longer than stigmal or marginal
veins, stigmal vein usually much shorter than marginal; mandible with two
teeth and a truncation or four teeth ADELENCYRTUS (p. 223)
490(43) Forewing with venation not reaching anterior margin of wing (Fig. 240);
antennal toruli separated from mouth margin by about their own lengths;
clava large and with a strong oblique truncation which is twice as long as
remainder of ventral surface of clava (Fig. 239); mandible bidentate
LUTHERISCA (p. 294)
Forewing with venation reaching anterior margin; antennal toruli less than their
own lengths from mouth margin ; clava usually with apex more or less rounded
or transversely truncate, although occasionally with an oblique truncation,
but this is only rarely longer than remainder of ventral surface of clava
(Neanagyrus spp.); mandible tridentate or with one or two teeth and a
truncation 491
491 (490) Forewing with postmarginal vein longer than stigmal ENCYRTOIDEA (p. 268)
Forewing with postmarginal vein not longer than stigmal 492
INDO-PACIFIC ENCYRTIDAE
217
492 (491) First funicle segment anelliform, the remainder subequal and subquadrate;
forewing completely naked immediately below marginal vein at top of linea
calva OVIDOENCYRTUS (p. 312)
First funicle segment only a little smaller than second, all funicle segments
usually gradually enlarging distally; forewing with at least a few setae im-
mediately below marginal vein at top of linea calva 493
493 (492) Mandible with one or two teeth and a truncation; apex of hypopygium usually
not reaching more than halfway along gaster, although occasionally reaching
apex 494
Mandible with three sharp teeth; hypopygium reaching more than half way
along gaster and usually to more than five-sixths 496
494 (493) Mesopleurum posteriorly enlarged and more or less touching basal segment of
gaster so that when thorax viewed from side it clearly separates hind coxa
from metapleurum and propodeum (Fig. 177); posterior margin of meso-
scutum projecting backwards above axillae so that when thorax in resting
position the axillae appear to be broadly separated (as in Fig. 44)
OOENCYRTUS (p. 309)
Mesopleurum not posteriorly enlarged and not touching basal segment of
gaster, in side view hind coxa more or less broadly touches metapleurum and
propodeum (as in Figs 139, 140); median portion of posterior margin of
mesoscutum not projecting above axillae so that with thorax in resting
position axillae appear to meet (as in Fig. 42) 495
495 (494) Clava with a very strong oblique truncation, the surface of which is clearly
longer than remainder of ventral surf ace of clava NEANAGYRUS (p. 303)
Clava with apex more or less rounded PSYLLAEPHAGUS (p. 330)
496(493) Antennal toruli separated from mouth margin by at least nearly their own
lengths; forewing with sensillae at apex of stigmal vein not arranged in a
square, uncus present (Figs 247, 248) 497
Antennal toruli separated from mouth margin by less than half their own
lengths; forewing with sensillae at apex of stigmal vein arranged symmetri-
cally in a square, uncus absent (Figs 142, 183, 249) 498
497 (496) Head and thorax bright metallic green with punctate-reticulate sculpture; scape
clearly much shorter than minimum width of frontovertex; hypopygium not
reaching more than three-quarters along gaster PARACHALCERINYS (p. 315)
Head and thorax often metallic but never bright green, with shallow reticulate
sculpture; scape at least about as long as minimum width of frontovertex;
hypopygium reaching apex of gaster COELOPENCYRTUS (p. 255)
498 (496) Antennal flagellum not unicolorous, consisting of at least a few white segments
contrasting with dark remainder, clava apically transversely truncate
PARALITOMASTIX (p. 316)
Antennal flagellum unicolorous; clava usually with rounded apex, although
occasionally apically truncate COPIDOSOMA (p. 257)
499(44) Scutellum strongly convex and separated from axillae by deep grooves; face
with strong transverse carina above antennal toruli ANANUSIA (p. 231)
Scutellum flat to moderately convex and separated from axillae by normal
sutures; face without a strong transverse carina above antennal toruli 500
500 (499) Forewing with postmarginal vein at least a little longer than stigmal (Figs 151,
243) 501
Forewing with postmarginal vein not longer than stigmal 503
501 (500) Notaular lines absent; clava longer than funicle; frontovertex not more than
one-third head width EURYRHOPALUS (p. 277)
Notaular lines present in anterior part of mesoscutum; clava shorter than
funicle; frontovertex at least one-third head width 502
502 (501) Forewing with marginal vein longer than stigmal (Fig. 151); malar space less
than half as long as eye; sculpture of scutellum more or less same as that on
mesoscutum, shallow and reticulate; mandible tridentate PARACLAUSENIA (p. 316)
Forewing with marginal vein not longer than stigmal (Fig. 243); malar space
longer than half length of eye; scutellum with deep longitudinally elongate
218
J. S. NO YES & M. HAYAT
reticulate sculpture (Fig. 244) which contrasts strongly with the reticulate
sculpture of mesoscutum; mandible bidentate NEOCHARITOPUS (p. 305)
503 (500) Forewing with parastigma clearly swollen (Fig. 242); mandible with two teeth;
scutellum distinctly sculptured throughout COCCIDOXENOIDES (p. 255)
Forewing with parastigma not swollen, or if slightly swollen than mandible has
three teeth and scutellum has at least apical half smooth and shiny and devoid
of sculpture 504
504 (503) Face largely green with a contrasting yellow pattern ZARHOPALOIDES (p. 349)
Head either completely yellow or darker and without a contrasting pattern .... 505
505 (504) Scutellum with at least shallow sculpture throughout and without lateral,
longitudinal grooves behind axillae; forewing with filum spinosum directed
towards junction of marginal and stigmal veins and thus not converging with
setae of proximal margin of linea calva (Figs 248, 249) 506
Scutellum with at least apical half smooth and shiny and devoid of sculpture and
also with a lateral, longitudinal groove behind each axilla; forewing with
filum spinosum, if distinct, directed towards junction of submarginal and
marginal veins and thus converging with setae of proximal margin of linea
calva (Figs 127, 250, 251) 509
506 (505) Hypopygium extending past apex of last tergite and thus clearly visible in dorsal
view (similar to Fig. 125); ovipositor slightly exserted and apically slightly
downcurved EPIBLATTICIDA (p. 272)
Hypopygium not quite or only just reaching apex of gaster and not visible in
dorsal view; ovipositor not exserted, or if so then apex is not downcurved . . . 507
507 (506) Clavasolid COPIDOSOMA (p. 257)
Clava three-segmented 508
508 (507) Frontovertex very narrow, not more than one-quarter head width; mouth
opening small, less than twice as wide as frontovertex or less than half head
width ;clava with a distinct oblique apical truncation TRJAPITZINELLUS (p. 346)
Frontovertex at least one-third as wide as head, or if narrower then mouth
opening much wider than frontovertex or at least about half as wide as head;
clava only seldom obliquely truncate COELOPENCYRTUS (p. 255)
509 (505) Eye relatively small and clearly not reaching occipital margin which is more or
less rounded; eye length never more than minimum width of frontovertex
ZAOMMOENCYRTUS (p. 349)
Eye larger and more or less reaching occipital margin which is sharp; length of
eye greater than minimum width of frontovertex CERCHYSIELLA (p. 246)
510 (44) Clava solid with apex rounded AUSTRALANUSIA (p. 236)
Either clava three-segmented or if solid then apex strongly obliquely truncate . 511
511 (510) Mesopleurum posteriorly enlarged and more or less touching basal segment of
gaster so that when thorax viewed from side it clearly separates hind coxa
from metapleurum and propodeum (Fig. 177) 512
Mesopleurum not so enlarged and not touching basal segment of gaster so that
when thorax viewed from side hind coxa more or less broadly touches
metapleurum and propodeum (as in Figs 139, 140) 515
512 (511) Forewing with postmarginal vein clearly longer than either stigmal or marginal
veins ENCYRTOIDEA (p. 268)
Forewing with either marginal or stigmal vein longer than postmarginal vein . . 513
513 (511) Forewing with marginal vein not more than twice as long as broad; posterior
margin of mesoscutum projecting slightly backwards above axillae so that
when thorax in resting position axillae appear to be broadly separated (as in
Fig. 44) 514
Forewing with marginal vein more than twice as long as broad; posterior margin
of mesoscutum not projecting above axillae so that axillae more or less
touching medially when thorax is in resting position (as in Fig. 42) 515
514(513) Clava very large, much wider than funicle and with a very strong oblique
truncation, longer than pedicel and funicle together (Fig. 202)
AGARWALENCYRTUS (p. 226)
Clava relatively smaller, rarely longer than funicle and apex generally more or
INDO-PACIFIC ENCYRTIDAE
219
less rounded, never strongly obliquely truncate (Fig. 246) OOENCYRTUS (p. 309)
515 (511, Occipital margin more or less rounded, or if appearing sharp then either head
513) triangular in profile, the face strongly inflexed at top of toruli (as in Fig. 72) or
forewing with postmarginal vein much more than one-sixth longer than
stigmal 516
Occipital margin sharp; head in profile anteriorly more or less evenly rounded;
forewing with postmarginal vein not more than one-sixth longer than stigmal 520
516 (515) Mandible with three acute teeth; scutellum slightly convex and at least about as
long as broad ACHALCERINYS (p. 220)
Mandible with four teeth or with one or two teeth and a truncation; scutellum
usually flat and distinctly broader than long 517
517 (516) Either forewing with postmarginal vein longer than stigmal or mandible with
fourteeth 518
Forewing with postmarginal vein not longer than stigmal; mandible with one or
two teeth and a truncation 519
518 (517) Forewing with postmarginal vein clearly much longer than either marginal or
stigmal veins, stigmal vein not or hardly shorter than marginal; mandible with
two teeth and a truncation ADELENCYRTOIDES (p. 223)
Postmarginal vein of forewing not or hardly longer than either stigmal or
marginal veins, stigmal vein usually much shorter than marginal; mandible
with four teeth (Fig. 116) ADELENCYRTVS (p. 223)
519 (517) Forewing with linea calva interrupted on dorsal surface by at least two lines of
setae COCCIDENCYRTUS (p. 253)
Forewing with linea calva neither interrupted nor closed on dorsal surface
EPITETRALOPHIDEA (p. 273)
520 (515) Forewing with a distinct hairless streak from apex of postmarginal vein to apex
of stigmal vein and extending slightly into disc (Fig. 252) TASSONIA (p. 341)
Forewing without a hairless streak at apex of venation 521
521 (520) Mandible with one or two teeth and a truncation 522
Mandible with three or four sharp teeth, if tridentate then uppermost tooth may
be short 527
522 (521) Clava with a strong oblique apical truncation, the truncate surface much longer
than remainder of ventral surface of clava 523
Clava without an oblique apical truncation or if so then truncate surface is much
shorter than remainder of ventral surface of clava 524
523 (522) Forewing with marginal vein at least about four times as long as broad and
clearly much longer than stigmal BAEOANUSIA (p. 241)
Forewing with marginal vein only a little longer than broad and less than half as
long as stigmal NEANAGYRUS (p. 303)
524 (522) Setae on eye short and inconspicuous, translucent and not or hardly longer than
diameter of a facet 525
Eye clothed in very long, conspicuous, dark setae, each at least twice as long as
diameter of a facet 527
525 (524) Forewing with stigmal vein at least nearly three times as long as marginal; head
and thorax generally bright metallic green or blue-green PSYLLAEPHAGUS (p. 330)
Forewing with stigmal vein less than twice as long as marginal; head and thorax
dark purple-brown with green or brassy reflections, occasionally mesoscutum
bright metallic green, but head and scutellum never strongly shiny 526
526 (525) Metapleurum clothed with distinct white setae extending to base of hind coxa
which is clothed in moderately dense conspicuous setae; base of gaster yellow
contrasting with remainder which is dark and shiny DIAPHORENCYRTUS (p. 263)
Metapleurum without any conspicuous white setae; hind coxae with only a few
sparse inconspicuous setae; gaster completely dark and shiny without a basal
yellow band SYRPHOPHAGUS (p. 338)
527 (521, Mesoscutum metallic with contrasting white setae; scutellum more or less flat
524) with fine reticulate sculpture giving it a matt appearance which strongly
contrasts with metallic colour of mesoscutum; clava strongly obliquely trun-
cate (Fig. 224) ECHTHROGONATOPUS (p. 267)
220
J. S. NOYES & M. HAY AT
Mesoscutum with dark setae; scutellum at least slightly convex although occa-
sionally with fine reticulate sculpture which gives it a matt appearance; clava
usually more or less rounded, although occasionally with a strong oblique
truncation 528
528 (527) Eye naked or with very short inconspicuous, translucent setae which are each
shorter than the diameter of a facet 529
Eye clothed in dense conspicuous, pale or dark setae, each at least as long as
diameter of a facet 531
529 (528) Forewing with sensillae at apex of stigmal vein arranged symmetrically in a
square, uncus absent (Figs 142, 183, 249); clava usually solid and with an
oblique apical truncation (Fig. 241), although occasionally rounded
COPIDOSOMA (p. 257)
Forewing with sensillae at apex of stigmal vein not arranged in a square, uncus
present; clava three-segmented and with apex rounded 530
530 (529) Scutellum with deep reticulate sculpture contrasting with shallow sculpture of
mesoscutum, latter dull purple and not shiny, scutellum green; mandible with
a long middle tooth and very short inner and outer teeth almost appearing
unidentate; ovipositor not visible externally RHYTIDOTHORAX (p. 333)
Scutellum with similar sculpture to mesoscutum, both very shiny and metallic;
mandible with three short subequal teeth; ovipositor slightly but distinctly
exserted STENOTEROPSIS (p. 338)
531 (528) Scutellum with very shallow reticulate sculpture, almost smooth EXORISTOBIA (p. 277)
Scutellum with fine striate-reticulate sculpture (Fig. 254) 532
532 (531) All funicle segments transverse (Fig. 253) HALIGRA (p. 281)
Only the first funicle segment not longer than broad, remainder each longer
than broad LAMENNAISIA (p. 292)
Notes on genera
ACEROPHAGUS Smith
(Key couplets: 62, 66. Fig. 23)
Acerophagus Smith, 1880: 83. Type-species: Acerophagus coccois Smith, by monotypy.
Rhopoideus Howard, 18986: 235. Type-species: Rhopoideus citrinus Howard, by monotypy.
DISTRIBUTION AND SPECIES. Sixteen species, New World; three species from review area: coccois
Smith; Rosen (1969: 57) (Hawaiian Is. ),solidusHayat(l981b: 13) (India) and texanus (Howard;
Rosen, 1969: 63) (Hawaiian Is.).
REFERENCES. World revision: Rosen (1969); see also Beardsley (1976).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Pseudectroma bryanti Girault may also run here in the key since it is possible that the
clava is entire and not two-segmented. We are retaining it in Pseudectroma pending examination
of freshly collected material.
Acerophagus is very close to Pseudaphycus, Pseudectroma, Indaphycus (tribe Aphycini,
subtribe Aphycina) and possibly also Mozartella, from all of which it can be separated using the
characters given in the key. It is possibly closest to Pseudectroma and can be most reliably
separated from this genus by the relatively more narrow frontovertex (see comments under
Pseudectroma).
ACHALCERINYS Girault
(Key couplet: 5 16)
Achalcerinys Girault, 1915a: 98. Type-species: Achalcerinys triclavata Girault, by original designation.
Echthrobacomyia Girault, 1920d: 142. Type-species: Echthrobacomyia niveipes Girault, by monotypy.
Syn. n.
INDO-PACIFIC ENCYRTIDAE 221
DISTRIBUTION AND SPECIES. Four species, possibly all synonymous, Neotropics (BMNH), Old
World including Afrotropical region; all four species from review area: gorodkovi (Myartseva,
1983: 66) (comb. n. from Parasyrpophagus) (India), lindus (Mercet, 1921: 271) (comb. n. from
Parasyrpophagus) (India), niveipes (Girault, 1920d: 142) (comb. n. from Echthrobacomyia)
(Australia) and triclavata Girault (1915a: 98) (Australia), also further undetermined material
from S. China and Vietnam to Papua New Guinea and Fiji (BPBM).
BIOLOGY. Unknown.
COMMENTS. The holotypes of Achalcerinys triclavata (QM) and Echthrobacomyia niveipes (QM)
have been examined and certainly are congeneric. They may also be conspecific but there are
some slight differences in setation and sculpture of the thorax and in the relative length of the
postmarginal vein of the forewing. Achalcerinys lindus (Mercet) may also be synonymous.
Achalcerinys gorodkovi (Myartseva) from Europe and India has dark hind femora, but probably
it is merely a colour form of lindus since there does not appear to be any consistent morphologi-
cal difference.
The genus can probably be placed best in the tribe Cheiloneurini. It is superficially similar to
Mahencyrtus but differs in lacking a strongly expanded parastigma, shorter propodeum and
generally much paler legs (the legs of Achalcerinys are completely yellow or with the hind
femora dark, those of Mahencyrtus are usually more extensively darkened). The head also has a
characteristic groove connecting the occipital foramen to the centre of the occipital margin
behind the ocelli.
ADEKTITOPUSgen. n.
(Key couplets: 242, 264. Figs 148, 160, 161, 255-263)
Type-species: Adektitopus gordhisp. n. Gender: masculine.
$. Head. In facial view a little broader than long, in profile about twice as long as broad and anteriorly
gradually curved. Eye with posterior margin very slightly concave, about one-half longer than broad and
with numerous fairly conspicuous translucent setae and more or less overreaching occipital margin which is
sharp only behind ocelli. Malar space distinctly to a little less than one-third length of eye and with malar
sulcus present. Frontovertex about one-third head width; ocelli approximately forming a right angle,
posterior ocellus separated from occipital margin by its own to twice its diameter and from eye margin by a
little less than twice its diameter. Antennal scrobes very shallow, meeting or not meeting dorsally and
nearly reaching halfway to anterior ocellus from antennal toruli; antennal torulus separated from mouth
margin by slightly less than its own length and from other torulus by slightly less than to about its own
length, its dorsal margin about level with ventral margins of eyes; clypeal margin broadly excised below
toruli. Antennal scape cylindrical, much longer than minimum width of frontovertex, about nine times as
long as broad, pedicel conical, about as long as or slightly longer than the funicle segments which are all
longer than broad, cylindrical and gradually widening distally; clava three-segmented about half as long as
funicle and with apex more or less rounded, outer suture very oblique and strongly converging with inner
suture ventrally so that they nearly meet (Figs 255, 256); longitudinal sensillae on all but first one or two
flagellar segments. Frontovertex with moderately deep, raised reticulate sculpture becoming more
squamiform on lower parts efface and on interantennal prominence; frontovertex clothed in sparse, short,
translucent setae, occasionally piliferous punctures large and giving the frontovertex a thimble-like
appearance. Mandible with three teeth, the uppermost tooth shortest and blunt, maxillary palpus
four-segmented, labial palpus three-segmented.
Thorax. In side view moderately deep with metapleurum and propodeum narrowly in contact with hind
coxa, dorsally with both mesoscutum and scutellum convex, almost flat. In dorsal view posterior margin of
pronotum quite concave; visible part of mesoscutum about twice as broad as long, with notaular lines
present (although in dry-mounted material rather obscure) and reaching about half way across mesoscu-
tum; axillae virtually meeting but appearing separate because posterior margin of mesoscutum projects a
little backwards medially; scutellum about as broad as long and about as long as mesoscutum, with apex
blunt; propodeum medially about one-quarter to one-fifth as long as scutellum. Mesoscutum with shallow
to fairly deep, raised squamiform-reticulate sculpture, scutellum with distinctly deeper, raised, more or
less fine vermiculate-reticulate sculpture; propodeum medially quite smooth; mesopleurum almost smooth
222 J. S. NOYES & M. HAY AT
but with shallow, raised, reticulate sculpture; dorsum of thorax with numerous, moderately long,
translucent, recumbent setae. Forewing hyaline with a faint hint of yellowish or faintly infuscate, wing two
and one-half to nearly three times as long as broad; linea calva not interrupted but more or less closed near
posterior margin of wing, filum spinosum absent; submarginal vein with an apical hyaline break, with
parastigma clearly swollen, much broader than proximal two-thirds of vein, marginal vein about seven to
nine times as long as broad, a little longer than postmarginal which is distinctly longer than stigmal; costal
cell about 16 times as long as broad and with a single line of setae dorsally in distal half. Hindwing hyaline,
about three-quarters as long as forewing, five times as long as broad, with marginal fringe about one-third
maximum wing width. Mid tibial spur about as long as basal mid tarsal segment.
Caster. About as long as thorax, cereal plates in anterior half, paratergites present but membranous, last
tergite about two-thirds to three-quarters as long as mid tibia, hypopygium reaching apex of gaster,
ovipositor slightly exserted and about as long as to one-third longer than mid tibia, gonostyli more or less
free and about one-fifth as long as ovipositor.
Cf . Differs from female as follows.
Head. Eye not quite reaching occiput, occipital margin more or less acute. Malar space about half as long
as eye; frontovertex nearly two-thirds head width; ocelli relatively larger, posterior ocellus separated from
occipital margin by about half its diameter and from eye margin by about its diameter; antennal scrobes
reaching more than halfway from toruli to anterior ocellus; antennal torulus separated from mouth margin
by slightly more than its own length and from other torulus by slightly less, its ventral margin slightly below
the ventral margins of the eyes; antennal scape a little shorter than minimum width of frontovertex,
cylindrical, about five times as long as broad; pedicel conical, slightly longer than broad, clearly several
times shorter than any of the funicle segments which are at least three times as long as broad, setae on
flagellum slightly longer than diameter of segments; clava in card-mounted specimens appearing to be
entire, but in slide-mounted specimens two-segmented, although suture is incomplete.
Thorax. In side view metapleurum and propodeum broadly in contact with hind coxa; forewing slightly
broader, about two and one-half times as long as broad; linea calva open; marginal vein about seven times
as long as broad, slightly shorter than postmarginal, both clearly longer than stigmal; costal cell a little
more than 20 times as long as broad.
Gaster. Shorter than thorax; genitalia with hooks on the digiti, aedeagus about one-third as long as mid
tibia or one-third longer than mid tibial spur.
COMMENTS. The genus belongs to the tribe Charitopidini (Tetracneminae) and can be separated
from other genera on the following combination of characters: incomplete notaular lines,
sculpture of thorax , lightly infuscate fore wings , postmarginal vein longer than stigmal , relatively
well-advanced cereal plates and long last tergite, and the unbranched antenna in the male.
The type-species of the genus is named in honour of Dr Gordon Gordh (UCR).
Adektitopusgordhisp. n.
(Figs 148, 160, 161,256-263)
9 Length: 1-24-1-56 mm (holotype, 1-56 mm).
Colour. Head black with faint greenish or bluish metallic sheen, antenna with scape honey yellow with
apex slightly darker, pedicel and flagellum from entirely pale brownish yellow to entirely dark brown, the
basal segments sometimes paler, clava dark brown; pronotum, axilla and tegula black with a slight purple
sheen, mesoscutum dark metallic blue with some purple reflections, scutellum basally dark purple,
gradually becoming blue and then green towards the apex; metanotum more or less orange-brown,
propodeum and mesopleurum strongly dark metallic purple; fore coxa dark brown with purple reflections,
mid coxa basally dark brown, remainder of legs and apex of mid coxa honey yellow; forewing mostly
hyaline but with faint yellowish suffusion distal to parastigma, occasionally a faint longitudinal fuscous
streak in apical one-half of wing; gaster mostly orange-yellow, the last tergite to a greater or lesser extent
dark brown; visible part of gonostyli dark brown.
Head. In profile very slightly less than twice as long as broad; posterior ocellus nearly one and one-half
times its own length from occipital margin; antennal scrobes not meeting dorsally; antennal toruli
separated from each other by about three-quarters their own lengths; sculpture of frontovertex as in Fig.
258. Relative measurements (holotype): head width (frontal view) 84, head length 75, minimum width of
frontovertex 15, malar space 18, eye length 59, eye width 41, POL 14, OOL 2, scape length 49, other
proportions of antenna as in Fig. 256.
INDO-PACIFIC ENCYRTIDAE 223
Thorax. Mesoscutum with shallow, raised, squamiform-reticulate sculpture (Fig. 160); scutellum with
much deeper vermiculate-reticulate sculpture (Fig. 161). Relative measurements of forewing (holotype):
length 73, width 25, other proportions as in Figs 148, 257; of hindwing: length 57, width 11.
Caster. Relative lengths (paratype): ovipositor 60, gonostylus 11, last tergite 37, [mid tibia 45].
Hypopygium as in Fig. 259.
Cf Length: 0-87-1-03 mm.
Colour. Head blackish, weakly metallic green, antennal scape testaceous yellow, darker apically;
pedicel and flagellum dark brown, apex of pedicel slightly paler; mesoscutum dark brown, weakly metallic
green or bluish with some purple reflections; scutellum metallic green; propodeum and mesopleurum dark
brown with a purplish sheen; metapleurum slightly yellowish in colour; legs as for female except fore and
mid coxae largely yellow; gaster mostly dark brown but ventrally and basally yellowish; forewing very
faintly infumate from level with parastigma to apex.
Head. Frontovertex with moderately deep, raised, reticulate sculpture, this becoming shallower and
more longitudinally elongate on lower parts of face, particularly genae. Relative measurements (para-
type): head width (facial view) 53, head length 47, minimum frontovertex width 30, malar space 13, eye
length 28, eye width 20, POL 11, OOL 7, scape length 17, other proportions of antenna as in Fig. 260.
Thorax. Mesoscutum with shallow, raised, reticulate sculpture; scutellum with distinctly deeper
(although shallower than in female), raised, reticulate sculpture. Relative measurements (paratype):
forewing length 141, forewing width 55, other proportions as in Fig. 261.
Gaster. Relative lengths (paratype): aedeagus 34, [mid tibial spur 26]. Genitalia as in Figs 262, 263.
DISTRIBUTION. India.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $, India: Tamil Nadu, Mudumalai Animal Sanctuary, 23-24.X.1979 (/. S. Noyes) (BMNH).
Paratypes. India: 7 9 , 3 cf , same data as holotype; 1 $ , Maharashtra, Elephanta (caves near Bombay),
28.X.1979 (M. Hayat). (BMNH, HC, USNM, ZI, PPRI.)
COMMENTS. A further four species, all from India, can be separated by the female coloration and
relative lengths of the antennal segments, general body colour, especially legs and gaster,
relative lengths of malar space, length of head in side view, distance of ocelli from occipital
margin, whether antennal scrobes meet dorsally, the sculpture of the frontovertex, the relative
distance separating the antennal toruli, sculpture of dorsum of thorax, strength of forewing
infuscation, relative lengths of ovipositor and last gastral tergite to mid tibia; in the male they
may be distinguished by leg coloration and sculpture of the head and thorax.
ADELENCYRTOIDESTachikavsa & Valentine
(Key couplets: 489, 5 18)
Adelencyrtoldes Tachikawa & Valentine, 1969ft: 548. Type-species: Adelencyrtoides novae zealandiae
Tachikawa & Valentine, by original designation.
DISTRIBUTION AND SPECIES. One species novaezealandiae Tachikawa & Valentine (1969ft: 548)
(New Zealand, Chatham I.); possibly also several other species from New Zealand (BMNH,
DSIR).
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. This genus will be dealt with in more detail in a paper in preparation on the New
Zealand Encyrtidae. It may contain several other species but the generic status of most of these
is not yet certain and many may be attributable to other, as yet, undescribed genera.
The genus is placed in the tribe Habrolepidini, subtribe Habrolepidina and can be separated
from other related genera using the key provided by Tachikawa & Valentine (19696).
ADELENCYRTUS Ashmead
(Key couplets: 193, 247, 329, 351, 440, 489, 518. Fig. 116)
Adelencyrtus Ashmead, 1900ft: 401. Type-species: Encyrtus chionaspidis Howard, by original designation.
224 J- S. NOYES & M. HAY AT
Epiencyrtoides Girault, 1915a: 108. Type-species: Epiencyrtoides quadridentatus Girault, by original
designation.
Rotrencyrtus Risbec, 1958: 39. Type-species: Rotrencyrtus depressus Risbec, by monotypy.
DISTRIBUTION AND SPECIES. Twenty-five species, cosmopolitan; 19 from review area: aulacaspidis
(Brethes; Mercet, 1921: 294) (New Zealand), axillaris (Girault, 19150: 108) (Australia),
bifasciatus (Ishii; Tachikawa, 1963: 163) (India, Bangladesh, Taiwan, Hawaiian Is.), bimacula-
tus Alam; Hayat etal. (1975: 85) (India), chionaspidis (Howard; Compere & Annecke, 1961: 52)
(Sri Lanka), clavatus Hayat, Alam & Agarwal (1975: 83) (India), coxalis Hayat, Alam &
Agarwal (1975: 78) (India) , funicularis Hayat, Alam & Agarwal (1975: 80) (India), longiclava-
tus Hayat, Alam & Agarwal (1975: 84) (India), mayurai (Subba Rao, 1957: 380) (comb. n. from
Anabrolepis} (India), minutus (Girault, 19150: 177) (comb. n. from Epitetralophidea) (Austra-
lia), moderatus (Howard; Noyes, 1979: 144) (Pakistan, India), oceanicus (Doutt, 1951: 501)
(comb. n. from Anabrolepis) (Caroline Is., Mariana Is.), odonaspidis Fullaway (19130: 27)
(Hawaiian Is.), quadridentatus (Girault, 19150: 108) (Australia), quadriguttus (Girault; Hayat,
1978: 33) (comb. n. from Epitetracnemus) (India), quinquedentatus (Girault, 19290: 3) (comb. n.
from Epiencyrtoides) (Australia), shafeei Hayat, Alam & Agarwal (1975: 84) (India), simmond-
si Compere (19476: 281) (Australia), also several other unidentified species from throughout the
region (BMNH, BPBM, CNC, AMNH).
REFERENCES. Compere & Annecke (1961: 49-58); review of Indian species: Hayat etal. (1975:
76-87).
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. Encyrtus solidus Howard, described from the male sex only, has been incorrectly
placed in Adelencyrtus (Schmiedeknecht, 1909: 253). The holotype cf (USNM) has been
examined, but its generic placement remains uncertain.
Two or three undescribed species from the area extending from Borneo and the Philippines to
the New Hebrides appear to form a distinct group. This group is characterised by each species
being relatively much larger in size (at least about 1-5 mm long), having all antennal segments at
least about as long as broad and more deeply infuscate forewings. We do not consider these
characters to be sufficient for separate generic status.
The genus belongs to the tribe Habrolepidini, subtribe Habrolepidina (Encyrtinae) and is
very closely related to Epitetracnemus. The two genera are not at all easy to distinguish and it is
our view that further study will show that they should be considered synonymous. However, for
the present we are retaining the two as distinct genera, but only on the basis of the presence or
absence of a line of silvery setae across the face and the pattern and strength of infuscation of the
forewings (see key). We do not consider that the mandibles are reliable in separating the two
genera since the difference between a quadridentate mandible and one with two teeth and a
truncation is not very great (see Tachikawa, 1963: fig. 70). Head shape also does not appear to
be a good character since this can vary significantly. The males of the Habrolepidini all have a
two-segmented funicle and a long unsegmented clava and are extremely difficult to separate. A
key to females of some of the genera included in this subtribe is also given by Tachikawa &
Valentine (19696).
AENASIELLA Girault
(Key couplets: 142, 204, 473. Fig. 76)
Aenasiella Girault, 1914o: 33. Type-species: Aenasiella brachyscelidis Girault, by original designation.
DISTRIBUTION AND SPECIES. Seven species, all Australian: apiomorphae Girault (19150: 80),
australia Girault (19176: 35), brachyscelidis Girault (19140: 33), eucalypti (Dodd, 1917: 354)
(comb. n. from Coccidencyrtus), lunulata (Girault, 19150: 140) (comb. n. from Coccidoxenus) ,
ovi Girault (19250: 2) and Sidney i (Girault, 19266: 59) (comb. n. from Encyrtoidea).
BIOLOGY. Parasites of Apiomorpha galls (Homoptera: Eriococcidae) on Eucalyptus. Also
INDO-PACIFIC ENCYRTIDAE 225
recorded as a parasite of eggs of a longicorn beetle (Coleoptera) on Eucalyptus but this is
possibly incorrect.
COMMENTS. The holotype of Coccidencyrtus eucalypti Dodd has not been examined but the
description of that species indicates that it must be closely related to brachyscelidis.
The genus is very close to Psyllaephagus (tribe Trechnitini, subtribe Metaprionomitina) and
can virtually only be separated reliably from this genus by having three teeth on the mandible
instead of two teeth and a truncation, although the difference here is not always very distinct
(compare Figs 75, 76). The marginal vein of the fore wing is always clearly longer than broad,
whereas in Psyllaephagus it is almost always more or less quadrate.
AENASIOIDEA Girault
(Key couplet: 167)
Aenasioidea Girault, 1911: 171. Type-species: Aenasioidea latiscapus Girault, by original designation.
DISTRIBUTION AND SPECIES. Eleven species, Holarctic, Afrotropical; only one species included
here: aligerhini (Girault, 1932a: 5) (comb. n. from Aphycus) (Australia), also one undetermined
species reared from Ceroplastes ceriferus (Fabricius) from the Philippines (USNM).
REFERENCES. Timberlake (1916: 579-585), Tachikawa (1963: 194-195).
BIOLOGY. Parasites of Kermesidae and Coccidae (Homoptera).
COMMENTS. The genus is extremely close to Metaphycus (tribe Aphycini, subtribe Paraphycina)
and differs in having the hypopygium reaching the apex of the gaster, the usually relatively long
funicle segments and characteristic shape of the ventral margin of the scape.
AENASIUS Walker
(Key couplets: 129, 479)
Aenasius Walker, 1846: 181. Type-species: Encyrtus hyettus Walker, by original designation.
Pseudanasius Hayat, Alam & Agarwal, 1975: 21. Type-species: Pseudanasius clavus Hayat, Alam &
Agarwal, by original designation.
DISTRIBUTION AND SPECIES. Twenty-nine species, mainly New World but also Afrotropical; only
one species from this area: advena Compere; Kerrich (1967: 207) (Pakistan, India, Bangladesh,
Malaysia, Philippines, Solomon Is., New Caledonia, Samoa, Fiji, Loyalty Is., Hawaiian Is.),
also one undescribed species from India (BMNH).
REFERENCE. Revision: Kerrich (1967).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The undescribed species from India is intermediate between Neodiscodes and
Aenasius, but we place it in the present genus because of the relatively broader frontovertex
(about one-quarter head width).
Trjapitzin (1973) places Aenasius in the subtribe Aenasiina of the tribe Rhinoencyrtini which
must be incorrect, since Aenasius (and thus the Aenasiina) belongs in the Tetracneminae whilst
Paratetracnemoidea (=Rhinoencyrtus) must belong in the Encyrtinae (see comments under
Paratetracnemoidea). The subtribe Aenasiina should now be given tribal status, i.e. Aenasiini
Kerrich, 1967 (stat. n.) since the oldest name previously applied to the group, Tetralophideina
Erdos & Novicky, 1955 is based on a misidentification of the genus Tetralophidea Howard. The
tribe Aenasiini thus contains the following genera: Aenasius, Blepyrus Chalcaspis, Euryrhopa-
lus, Metaphaenodiscus , Monodiscodes Hoffer and Neodiscodes (Aenasiini = Neodiscodini
Trjapitzin, 1973 syn. n.). Aenasius is nearest to Neodiscodes and Chalcaspis and can best be
separated from these genera using Kerrich's key (1967: 188-190), although it is our opinion that
further study will show that these three genera should be considered synonymous.
226 J. S. NO YES & M. HAY AT
AENASOMYIELLA Girault
(Key couplet: 177)
Aenasomyiella Girault, 1915a: 93. Type-species: Aenasomyiella coleridgei Girault, by original desig-
nation.
Zaomommoencyrtus Girault, 1917g: 143. Type-species: Zaomommoencyrtus poeta Girault, by original
designation. Syn. n.
DISTRIBUTION AND SPECIES. Three species, all Australian: cervidncta Girault (1922e: 151),
coleridgei Girault (19150: 93) and poeta (Girault, 1917g: 143) (comb. n. from Zaomommoen-
cyrtus).
BIOLOGY. Doubtfully reared from a psyllid (Homoptera, Psyllidae) nymph under bark of
Eucalyptus.
COMMENTS. The three species included here are very close but can be separated from each other
by the relative position, shape and size of the purple spot on the scape: at apex only -poeta;
restricted to basal half or so - coleridgei; extending from base along ventral margin nearly to
apex of scape - cervidncta.
The genus is very close to Metaphycus (tribe Aphycini, subtribe Paraphycina) and can be
separated by the two-segmented clava, relatively shorter scape, i.e. not or hardly longer than the
malar space, uninterrupted linea calva and characteristic purplish spot on outer surface of scape.
AGAR WALENCYRTUS Hayat
(Key couplets: 393, 514. Figs 202, 203)
Agarwalencyrtus Hayat, 1981ft: 15. Type-species: Coccidencyrtus citri Agarwal, by original designation.
DISTRIBUTION AND SPECIES. One species, Afrotropical, Oriental and Australasian: citri
(Agarwal; Hayat, 19816: 15) (India, Bangladesh, Hong Kong, Java and Solomon Is.), probably
a second species from Taiwan (BPBM) which differs from citri in the relative position of the
ocelli and proportions of the antennal segments.
BIOLOGY. Recorded as a parasite of Planococcus citri (Risso) (Homoptera, Pseudococcidae) by
Agarwal (1965) but some material (BMNH) reported as being reared from Pipunculidae
(Diptera).
COMMENTS. The type-species appears to vary quite considerably in colour, some specimens
being almost entirely reddish orange whilst others are almost entirely black. This variation in
colour does not appear to be related to distribution.
The genus is probably closely related to Ooencyrtus (tribe Microteryini, subtribe Ooen-
cyrtina) and can be easily separated by the relatively large, obliquely truncate clava and strongly
transverse funicle segments.
AGEMASPIS Dahlbom
(Key couplet: 188)
Ageniaspis Dahlbom, 1857: 293. Type-species: Encyrtus fuscicollis Dalman, by designation of Ashmead
(1904c: 303).
Leuroceroides Girault, 1915a: 114. Type-species: Leuroceroides niger Girault, by original designation.
Syn. n.
Microrhopus Girault, 19326: 1. Type-species: Microrhopus striatithorax Girault, by monotypy. Syn. n.
DISTRIBUTION AND SPECIES. Nine species, cosmopolitan; three from review area: dtricola
Loginovskaya (1983: 610) (Vietnam), nigra (Girault, 19150: 114) (comb. n. from Leuroceroides)
(Australia) and striatithorax (Girault, 19326: 1) (comb. n. from Microrhopus), also further
undetermined specimens from Papua New Guinea (BPBM).
BIOLOGY. Polyembryonic parasites of larvae of Yponomeutidae (Lepidoptera).
INDO-PACIFIC ENCYRTIDAE 227
COMMENTS. Ageniaspis nigra and striatithorax are extremely close and may eventually prove to
be synonymous when freshly collected material can be carefully compared with the types of the
two species.
In his description of Microrhopus striatithorax, Girault did not state that the genus Microrho-
pus was being described as new. It is here taken as an available name since the species epithet is
valid (under Article llg(ii) of the Code of Zoological Nomenclature) and the generic name is not
unavailable for reasons of homonymy.
Trjapitzin (19736) places the genus in the subtribe Ageniaspidiina, tribe Copidosomatini.
Where their biology is known, all included species are polyembryonic parasites of Lepidoptera.
However, further study may show that the Ageniaspidiina are in fact not as closely related to the
Copidosomatina as their biology suggests. This is indicated by the difference in fore wing
venation (notably the arrangement of the sensillae at the apex of the stigmal vein, and the long
postmarginal vein) and structure of the gaster (notably the ovipositor). They may in fact be more
closely related to the Microteryini. The subtribe Ageniaspidiina contains three other genera,
including Holcothorax. Ageniaspis can be separated from Holcothorax by having a six-
segmented funicle (Holcothorax has a five-segmented funicle). (See also comments under
Holcothorax.)
ALAMELLA Agarwal
(Key couplets: 161, 272. Figs 87-88, 167)
Alamella Agarwal, 1966: 74. Type-species: Alamellaflava Agarwal, by original designation.
DISTRIBUTION AND SPECIES. Two species, Afrotropical and Oriental; one from review area:/7va
Agarwal (1966: 77) (India, Pakistan), also one probably undescribed species from Taiwan
(BPBM).
REFERENCES. Annecke (1969: 453-457), Hayat & Verma (1980), Hayat (19816: 16-17).
BIOLOGY. Parasites of Pseudococcidae (Homoptera); erroneously recorded from Eriococcidae
(Homoptera).
COMMENTS. The genus most probably belongs near Anagyrus (tribe Anagyrini, subtribe
Anagyrina) but can easily be separated from this and related genera by the very distinct structure
of the antenna (Fig. 167).
AMENISCOCEPHALUS Girault
(Key couplet: 244)
Ameniscocephalus Girault, I9l5a: 167. Type-species: Ameniscocephalus meniscocephalus Girault, by
original designation.
DISTRIBUTION AND SPECIES. One species, Australian only: meniscocephalus Girault (19156: 167).
BIOLOGY. Unknown, but probably parasitic on Pseudococcidae (Homoptera).
COMMENTS. Almost certainly related to Metaphaenodiscus (tribe Aenasiini - see comments
under Aenasius) but can easily be distinguished on body colour and venation. Metaphaenodiscus
is dark and metallic with a relatively short stigmal vein and distinct postmarginal vein, whilst
Ameniscocephalus has a relatively long stigmal vein (subequal to marginal) and the postmar-
ginal vein absent. The venation is very similar to that found in genera near Paraphaenodiscus .
AMICENCYRTUS Hayat
(Key couplet: 424. Fig. 265)
Amicencyrtus Hayat, 19816: 16. Type-species: Amicencyrtus obscurus Hayat, by original designation.
DISTRIBUTION AND SPECIES. Afrotropical, Oriental, Australasian, one described species: obscu-
228 J. S. NOYES & M. HAY AT
rus Hayat (19816: 17) (India), also undetermined material from Hong Kong, Malaysia, Brunei,
Sulawesi, Philippines, Java and Australia (BMNH, BPBM).
BIOLOGY. Unknown.
COMMENTS. Closer examination of the undetermined material may show that it all belongs to
obscurus. However, there is at least one further undescribed species from the Afrotropical
region (BMNH).
The genus is close to Cowperia (tribe Bothriothoracini, subtribe Aminellina) from which it
can be separated principally by the much flatter thoracic dorsum, particularly the scutellum (that
of Cowperia is quite convex). Other characters for separating this genus from Cowperia are
given by Hayat (19816: 17).
AMIRA Girault
(Key couplets: 423, 475)
Amira Girault, 1913c: 93. Type-species: Amirafabrei Girault, by original designation.
Bregmencyrtus Annecke, 1974: 369. Type-species: Eucomys durantae Risbec, by original designation.
DISTRIBUTION AND SPECIES. Three species, Afrotropical, Oriental, Australasian; two from
review area: fabrei Girault; Noyes (1977: 49) (India, Australia) and tarsata (Ashmead; 19056:
403) (comb. n. from Howardiella) (Philippines), also undetermined material from India,
Borneo and Solomon Is. (BMNH, BMBM).
REFERENCE. Revision: Noyes (1977).
BIOLOGY. Parasites of the eggs of spiders (Araneida).
COMMENTS. The genus has been placed in a separate tribe in the Encyrtinae (Amirini) by
Trjapitzin (19736) but closer examination of material belonging to Amira, and its biology,
indicate that it is very closely related to Ooencyrtus (Microteryini, Ooencyrtina). If this
suggested affinity proves to be correct then Amirini Girault, 1913 will have precedence as the
valid tribal name over Microteryini Hoffer, 1955. However, the generic relationships within the
Encyrtinae are so poorly understood at present that we feel a formal synonymy of these tribal
names is premature and probably unnecessary at the present time.
ANAGYRIETTA Ferriere
(Key couplet: 157. Figs 86, 264)
Anagyrietta Ferriere, 1955: 121. Type-species: Anagyrietta pantherina Ferriere, by original designation.
DISTRIBUTION AND SPECIES. Palaearctic and Oriental, one species known, but not from review
area; one undescribed species from India (BMNH).
BIOLOGY. The type-species has been reared from Spinococcus calluneti (Lindinger) (Homop-
tera, Pseudococcidae) on Calluna vulgaris.
COMMENTS. The Indian species differs from the type-species in having the linea calva completely
closed (in pantherina it is interrupted anteriorly and then closed near posterior margin of wing),
the areas of the forewing where the dark setae are situated not infuscate as in pantherina, notauli
completely absent, filum spinosum present (apparently absent in pantherina, although we have
been unable to examine a slide-mounted forewing). The presence of the filum spinosum is very
rare in tetracnemine encyrtids and in particular the Anagyrini. We do not believe that these
differences require separate generic status for the Indian species.
ANAGYRODES Girault
(Key couplet: 422)
Anagyrodes Girault, 1915a: 155. Type-species: Anagyrodes maximus Girault, by original designation.
INDO-PACIFIC ENCYRTIDAE 229
DISTRIBUTION AND SPECIES. Oriental and Australasian, seven species: baethei Girault (19226:
103) (Australia), del (Girault, 19226: 100) (comb. n. from Paradadella) (Australia), giganteus
Girault (19150: 156) (Australia), maximus Girault (1915a: 155) (Australia), odacon (Walker,
18386: 476) (comb. n. from Encyrtus) (Australia), perkinsi (Subba Rao, 1971: 212) (comb. n.
from Neocladia) (Australia) and punctaticeps Girault (19286: 449) (Philippines), also undeter-
mined material from India, Papua New Guinea and Borneo (BMNH, BPBM).
BIOLOGY. The unidentified Indian material has been reared from Batmchomorphus indicus
(Lethierry) nymphs (Homoptera, Cicadellidae).
COMMENTS. The single extant female syntype of Encyrtus odacon Walker (BMNH) is here
designated LECTOTYPE. This species is very close to del and perkinsi.
This genus is related to those placed in the tribes Encyrtini, Eugahaniini, Prionomasticini,
Neocladiini and Aethognathini by Trjapitzin (19736). It is probably closest to Eugahania and
can easily be separated from this genus in that it lacks the incision at the apex of the costal cell of
the fore wing. It can be separated from other related genera by the combination of the
three-segmented clava, long sickle-shaped mandibular tooth, more or less absent marginal vein
of fore wing and hypopygium extending to the apex of the gaster or nearly so. Future study may
show that these five tribes should be considered synonymous.
ANAGYR US Howard
(Key couplets: 150, 165, 173, 220, 230, 268, 407. Figs 95, 96, 266-268)
Anagyrus Howard in Howard & Ashmead, 1896: 638. Type-species: Anagyrus greeni Howard, by
monotypy.
Heterarthrellus Howard, 18986: 239. Type-species: Heterarthrellus australiensis Howard, by monotypy.
Paranusia Brethes, 1913: 102. Type-species Paranusia bifasciata Brethes, by monotypy.
Philoponectroma Brethes, 1913: 104. Type-species: Philoponectroma pectinatum Brethes, by original
designation.
Gyranusia Brethes, 1920: 137. Type-species: Gyranusia porteri Brethes, by monotypy.
Gyranusa Mercet, 1921: 123. Type-species: Gyranusa matritensis Mercet, by original designation.
Protanagyrus Blanchard, 1940: 115. Type-species: Protanagyrus aciculatus Blanchard, by monotypy.
Xiphomastix De Santis, 1972: 45. Type-species: Xiphomastix bellator De Santis, by original designation.
DISTRIBUTION AND SPECIES. About 125 species, cosmopolitan; 58 from review area: adamsoni
Timberlake (1941: 227) (Tahiti), agraensis Saraswat in Saraswat & Mukerjee (1975: 41) (India),
alami Hayat (19700: 112) (India), aligarhensis Agarwal (1965: 52) (India), almoriensis Shafee,
Alam & Agarwal (1975: 13) (India), amoenus Compere (1939: 12) (India), ananaitis Gahan
(1949: 357) (Hawaiian Is.), antoninae Timberlake; Beardsley (1969: 291) (Hawaiian Is.),
australiensis (Howard, 18986: 239) (Australia), bellus (Girault, 19216: 190) (comb. n. from
Dinocarsis} (Australia), citri Agarwal (1965: 48) (India), comperei Subba Rao & Rai (1970: 91)
(India), cooki (Girault, 19196: 57) (comb. n. from Dinocarsis) (Java), dactylopii (Howard,
18986: 242) (India, Hong Kong, Hawaiian Is.), darevskii (Trjapitzin, 1965: 310) (comb. n. from
Doliphoceras) (Indonesia), diversicornis Mercet (1921: 134) (India), fasciiscapus (Girault,
19326: 1) (comb. n. from Dinocarsis) ( Australia), ferus (nom. n. ioiflavus Shafee, 1974: 325 nee
Ishii, 1928) (India) , flaviceps Timberlake (1941: 221) (Marquesas Is.),flavidus Shafee Alam &
Agarwal (1975: 20) (India), flavimesopleurum (Girault, 1917g: 137) (comb. n. from Dinocarsis)
( Australia), foersteri (Girault, 19150: 145) (comb. n. from Epidinocarsis) ( Australia), fusciven-
tris (Girault, 19150: 144) (Australia), greeni Howard in Howard & Ashmead (1896: 639) (Sri
Lanka), hipocoon Trjapitzin (1965: 317) (Indonesia), indicus (Subba Rao, 1967: 1) (India),
inopus (nom. n. for indicus Shafee, Alam & Agarwal, 1975: 13) (India, Mariana Is.), kivuensis
Compere (1939: 11) (India) , laeviceps Perkins (1910: 654) (Hawaiian Is.), ///0dm' Ferriere (1937:
317) (Philippines), lineatipes (Girault, 19196: 57) (comb. n. from Dinocarsis) (Java), lon-
gipennis Shafee, Alam & Agarwal (1975: 16) (India), longiventris Hayat (19790: 173) (India),
major Perkins; Beardsley (1969: 289) (Hawaian Is.), mirus (Girault, 19150: 143) (comb. n. from
Epidinocarsis} (Australia), mumfordi Timberlake (1941: 222) (Marquesas Is.), nigricornis
230 J. S. NOYES & M. HAY AT
Timberlake (19196: 197) (Hawaiian Is.), nigricorpus Shafee, Alam & Agarwal (1975: 11)
(India), nigriflagellum (Girault, 1915: 145) (comb. n. from Epidinocarsis} (Australia), nigror-
adiculatus Subba Rao & Rai (1970: 94) (India), orbitalis Timberlake (1941: 220) (Marquesas
Is.), pseudococci (Girault, 1915: 185) (Pakistan, India), punctulatus Agarwal (1965: 50)
(India), qadrii (Hayat, Alam & Agarwal, 1975: 12) (comb. n. from Leptanusid) (India),
saccharicola Timberlake (1932: 159) (India, Taiwan, Thailand, Malaysia, Philippines, Fiji,
Hawaiian Is.), saipanensis Doutt (1952: 399) (Mariana Is.), sawadai Ishii (1928: 88) (India,
Taiwan), scutomaculatus Agarwal (1965: 49) (India), shahidi Hayat (1979a: 177) (India) , similis
(Girault. 1915: 145) (comb. n. from Epidinocarsis) (Australia), spica (Girault, 19216: 191)
(comb. n. from Dinocarsis) (Australia), subalbipes Ishii (1928: 90) (S. China), subflaviceps
(Girault, 1915a: 143) (comb. n. from Epidinocarsis) (Australia) , subproximus (Silvestri, 19156:
346) (Pakistan), swezeyi Timberlake (19196: 199) (Hawaiian Is., India), tibimaculatus Agarwal
(1965: 50) (India), varithorax (Girault, 1923d: 2) (comb. n. from Leptomastix) (Australia),
xanthogaster Perkins (1910: 653) (Hawaiian Is.), also much unidentified material from
throughout the region (BMNH, BPBM, CNC, UCR, HC).
REFERENCES. Beardsley (1969), Shafee etal. (1975: 9-21), Hayat (19790).
BIOLOGY. Parasites of Pseudococcidae (Homoptera) and Coccinellidae (Coleoptera) from
Australia whose larvae produce a waxy secretion, e.g. Telsimia sp.
COMMENTS. We have examined a specimen determined as Mashhoodia flava by Shafee and
believe that it belongs in Anagyrus. We have not examined material of Doliphoceras darevskii,
but from the description it would seem to be better placed in Anagyrus.
Anagyrus has been placed in the tribe Anagyrini, subtribe Anagyrina by Trjapitzin (1973a).
During our study of the species belonging to genera of this subtribe we have had some measure
of difficulty assigning many of the species to genera as they are understood at the present time.
Kerrich (1982) has summarised the characters previously used by workers to separate the genera
but we have found that the single characters or combinations of characters used by him are
largely unreliable, probably because his study was based on only a relatively small number of
species belonging to this group. In particular, we have had difficulty in separating A nagyrus from
Doliphoceras and Gyranusoidea, and Epidinocarsis from Doliphoceras, largely because many of
the species have unusual combinations of characters, e.g. a species which could be placed in
Doliphoceras with sculpture typical of Anagyrus or a species which could be placed in Anagyrus
with an elongate postmarginal vein as in Gyranusoidea. Our study has not been sufficiently
detailed to allow us to reach any satisfactory conclusions with regard to the possible natural
grouping of species of this subtribe. We do believe however, that many of the genera included in
this group are not necessary and it is very probable that many will be considered synonymous
when a more detailed study, on a world-wide basis, is undertaken. Any new genera or generic
synonymy proposed at this point could prove to be premature and almost certainly would lead to
a good deal of confusion and resentment amongst biological control workers. Therefore we have
found it necessary to use simple, convenient characters for separating these genera in the key
and, although we do not think that these characters alone will reflect the natural grouping of
species, it does allow most of the well-known described species to run in the key to genera where
they are placed by most workers at present. Anagyrus is here separated from Doliphoceras
almost solely on sculpture since we find scape coloration, shape of flagellar segments and body
shape all totally unreliable. Gyranusoidea is separated from Anagyrus by the postmarginal vein
of the forewing being at least one-quarter longer than the stigmal, whereas in Anagyrus it is not
or hardly longer. We have not found that sculpture or shape of the scape is totally reliable.
Unfortunately this has led to one relatively well-known species being transferred from Anagyrus
to Gyranusoidea, i.e. mirzai Agarwal. Anagyrus and Epidinocarsis are separated entirely on the
sculpture of the head and dorsum of thorax as no other characters were found to be reliable.
Other genera belonging to this group were separated on characters given in the key.
INDO-PACIFIC ENCYRTIDAE 231
ANANUSIA Girault
(Key couplets: 370, 402, 499)
Paranusia Girault, 1913e: 97. Type-species: Paranusia longiscapus Girault, by original designation.
[Homonym of Paranusia Brethes, 1913.]
Ananusia Girault, 1917g: 155. [Replacement name for Paranusia Girault.]
Myrmencyrtus Gordh & Trjapitzin, 1979ft: 107. Type-species: Myrmencyrtus australis Gordh &
Trjapitzin, by original designation. Syn. n.
DISTRIBUTION AND SPECIES. Two species, both Australian: australis (Gordh & Trjapitzin, 19796:
107) (comb. n. from Myrmencyrtus) and longiscapus (Girault, 1913e: 98).
BIOLOGY. Associated with nests of ants (Hymenoptera, Formicidae) and probably parasitic on
mealybugs within the nests (Homoptera, Pseudococcidae).
COMMENTS. We have not seen material of Myrmencyrtus australis, but it is clear from the
description that it is congeneric with, if not conspecific with, Ananusia longiscapus.
The genus is very close to Taftia (Chrysoplatycerini) and can easily be separated from it by the
extremely deep sutures between the scutellum and the axillae and by the strongly flattened
flagellum (in Taftia it is more or less cylindrical). A key to separate the genera of the
Chrysoplatycerini is given by Gordh & Trjapitzin (19796).
ANARHOPUS Timberlake
(Key couplet: 55)
Anarhopus Timberlake, 1929: 15. Type-species: Anarhopus sydneyensis Timberlake, by original des-
ignation.
DISTRIBUTION AND SPECIES. One species, Australasia and New World: sydneyensis Timberlake
(1929: 18) (= Arhopoideus semiargenteus Girault, 19296: 314 syn. n.) (Australia, New Zealand,
Hawaiian Is.).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. We have not examined the holotype of A. semiargenteus (SAM), but since Girault
himself, in an unpublished manuscript, synonymised this species with Anarhopus sydneyensis,
and the description agrees with sydneyensis, we have no hesitation in synonymising the two
species.
Placed in the tribe Tetracnemini, subtribe Arhopoideina which also includes Tetracnemoidea
and Zealandencyrtus from which it can be separated using the characters given in the key.
Tetracnemoidea and Anarhopus are extremely close and some of the species of Tetracnemoidea
found in New Zealand appear to be more or less intermediate. A more detailed reevaluation of
characters, taking these species into consideration, may eventually show that the two genera
should be considered synonymous.
ANICETUS Howard
(Key couplet: 113. Fig. 52)
Anicetus Howard in Howard & Ashmead, 1896: 639. Type-species: Anicetus ceylonensis Howard, by
monotypy.
Asteropaeus Howard, 1898ft: 231. Type-species: Asteropaeus primus Howard, by monotypy.
Habrolepopterygis Girault, 1915a: 86. Type-species: Habrolepopterygis felix Girault, by original des-
ignation.
Krishnieriella Mani, 1935: 421. Type-species: Krishnieriella ceroplastodis Mani, by original designation.
DISTRIBUTION AND SPECIES. Twenty-seven species, cosmopolitan except for more northerly
latitudes (40+); 16 species from review area: aligarhensis Hayat, Alam & Agarwal (1975: 34)
(India), angustus Hayat, Alam & Agarwal (1975: 35) (India), annulatus Timberlake; Annecke
232 J. S. NO YES & M. HAY AT
(1967: 110) (China, Australia, Hawaiian Is.), ashmeadi Hayat, Alam & Agarwal (1975: 33)
(India), beneficus Ishii & Yasumatsu; Tachikawa (1963: 126) (India), ceylonensis Howard:
Annecke (1967: 108) (India, Sri Lanka), chinensis Girault; Annecke (1967: 128), (China),
communis Annecke (1967: 121) (Australia), deltoideus Annecke (1967: 118) (India, China,
Borneo), dodonia Ferriere; Annecke (1967: 120) (Pakistan, India) , felix Girault (1915: 86),
(Australia), howardi Hayat, Alam & Agarwal (1975: 36) (India), integrellus Trjapitzin;
Annecke (1967: 129) (Pakistan, India), mirabilis (Girault; Annecke, 19716: 258) (Australia),
stylatus Subba Rao (1977: 16) (India) andyasumatsui Subba Rao (1965: 73) (India), also further
undetermined material from throughout the region (BMNH, BPBM, HC).
REFERENCES. Revision: Annecke (1967: 105-130); also Hayat etal. (1975: 30-38).
BIOLOGY. Parasites of Coccidae (Homoptera).
COMMENTS. Placed in the tribe Cerapterocerini; Anicetus and other genera of this tribe can be
separated using the present key and the more detailed one provided by Annecke (1967:
100-101).
ANOMALENCYRTUS Hayat & Verma
(Key couplet: 274. Fig. 169)
Anomalencyrtus Hayat & Verma, 1980: 341. Type-species: Anomalencyrtus longicornis Hayat & Verma,
by original designation.
DISTRIBUTION AND SPECIES. Afrotropical, Oriental, one described species only: longicornis
Hayat & Verma (1980: 344) (India).
BIOLOGY. Unknown.
COMMENTS. Material from the Afrotropical region (Zimbabwe - BMNH) almost certainly
belongs to the type-species, but differs in having the metanotum, propodeum and mesopleurum
extensively dark brown, whereas in the Indian specimens these parts are yellowish.
The genus can best be placed in the tribe Anagyrini, subtribe Anagyrina and differs from all
other genera of the subtribe by the peculiar structure of the antenna, notably the long,
unsegmented clava.
ANOMALICORNIA Mercet
(Key couplets: 78, 272. Figs 37, 162)
Anomalicornia Mercet, 1921: 85. Type-species: Anomalicornia tenuicornis Mercet, by original des-
ignation.
DISTRIBUTION AND SPECIES. Palaearctic. Afrotropical, only one species recognised: tenuicornis
Mercet (1921: 85; also 19226: 294 as ruschkai) (India), also undetermined material from Java
and Australia (BMNH, CNC).
BIOLOGY. A parasite of Pseudococcidae (Homoptera).
COMMENTS. More than one species is known to us since material from Cameroun (BMNH)
represents an undescribed species. We are not certain that all of the material from India or
Australia is actually tenuicornis because there are some differences in the relative lengths of the
funicle segments.
Trjapitzin (1973) places this genus in a separate tribe, the Anomalicorniini, but we believe
that it could be accommodated in the Anagyrini, possibly as a separate subtribe. The unique
structure of the antenna and forewing venation (in fully winged forms) should serve to
distinguish this genus from others included in the Anagyrini.
INDO-PACIFIC ENCYRTIDAE 233
ANTHEMUS Howard
(Key couplet: 45. Figs 12-14)
Anthemus Howard in Howard & Ashmead, 1896: 643. Type-species: Anthemus chionaspidis Howard, by
monotypy.
Hexalis Bakkendorf, 1939: 84. Type-species: Hexalis funicular is Bakkendorf, by monotypy.
DISTRIBUTION AND SPECIES. Ten species, Old World; four from review area: chionaspidis Howard
in Howard & Ashmead (1896: 643) (Sri Lanka), inconspicuus Doutt (1966: 226) (Pakistan), hilli
Dodd (1917: 352, as var. of chionaspidis) (Australia) and maculatus Subba Rao (1976: 685)
(Pakistan).
REFERENCE. Key to world species: Subba Rao (1976: 685).
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. The types of Anthemus emersoni Girault (19206: 98) and A. nigriceps Girault
(1934a: 2) have been examined (QM). Both belong to the family Mymaridae: A. emersoni to a
genus very probably near to Parallelaptera Enock, and A. nigriceps may be a species oiArescon
Walker.
The genus is the sole representative of the tribe Anthemini (Encyrtinae).
APHYCOMORPHA Timberlake
(Key couplets: 410, 414. Figs 269-271)
Aphycomorpha Timberlake, 19196: 225. Type-species: Aphycomorpha araucariae Timberlake, by
original designation.
DISTRIBUTION AND SPECIES. Two species, Neotropics, Pacific and New Zealand: araucariae
Timberlake (19196: 227) (Hawaiian Is.) and aspidioti Tachikawa & Valentine (19690: 535)
(New Zealand and offshore islands).
BIOLOGY. Parasites of Diaspididae and Eriococcidae (Homoptera).
COMMENTS. Aphycomorpha aspidioti (key couplet 414) may be incorrectly placed in this genus
and may be closer to Aphycopsis since the mesopleurum is not as strongly enlarged as in
araucariae (see below). It differs from Aphycopsis australiensis in biology and in lacking
notaular lines on the mesoscutum; however, the latter are very obscure in australiensis and this
species would run to Aphycomorpha (couplet 414) in the key if the notaular lines are
overlooked.
The genus has been placed in the tribe Aphycini by Trjapitzin (19736) but we believe that
almost certainly it would be better placed in the Microteryini. This is indicated by the shape of
the mandible (two teeth and a broad truncation) and the enlarged mesopleurum (as in
Ooencyrtus and Trichomasthus , although in aspidioti it is not so strongly enlarged) which
appears to be typical of most genera which belong in the Microteryini. It is probably closest to
Aphycopsis (see comments under Aphycopsis) and can be separated from most other genera of
the tribe by its resemblance to some species of Metaphycus and Aphycus.
APHYCOPSIS Timberlake
(Key couplet: 377)
Aphycopsis Timberlake, 1916: 585. Type-species: Aphycus australiensis Howard, by original designation.
DISTRIBUTION AND SPECIES. One species, Australia only: australiensis (Howard; Timberlake,
1916:586).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. We have examined the two female syntypes of Aphycus australiensis Howard
234 J. S. NOYES & M. HAY AT
(USNM). One has the head still attached to the body but partially eaten away at the vertex and
an antenna and forewing mounted on a slide. The other has the head removed and mounted
separately on a slide. The head has been dissected and crushed in typical Girault fashion but is
otherwise complete. The latter specimen is here designated as LECTOTYPE and has been so
labelled.
Mesanusia speciosa Girault (p. 353) may also belong to this genus.
The genus bears a superficial resemblance to species ofMetaphycus but differs in the mandible
having a single small tooth and a broad truncation (as in Fig. 189) whereas all species of
Metaphycus have tridentate mandibles. The structure of the mandible and general habitus
suggest that the genus is related to Aphycomorpha and Mozartella which very probably belong
in the Microteryini. It differs from Aphycomorpha in having the hind coxa narrowly in contact
with the propodeum in side view (see comments under Aphycomorpha), forewing with a
punctiform marginal vein, and notaular lines present (although very obscure) in the anterior
part of the mesoscutum. In Aphycomorpha the marginal vein is at least about twice as long as
broad and the notaular lines are absent. It differs from Mozartella in having a six-segmented
funicle and notaular lines present (Mozartella has a five-segmented funicle and notaular lines
absent).
APHYCUS Mayr
(Key couplets: 362, 364, 388. Figs 199, 272-273)
Aphycus Mayr, 1876: 695. Type-species: Encyrtus apicalis Dalman, by designation of Ashmead
(1900ft: 383).
Aphycoideus Williams, 1916: 153. Type-species: Aphycoideus io Williams, by monotypy.
Waterstonia Mercet, 1917c: 268. Type-species: Waterston ia prima Mercet, by original designation.
Euaphycus Mercet, 1921: 197. Type-species: Encyrtus hederaceus Westwood, by original designation. (As
subgenus of Aphycus . )
Aphycaspis Hoffer, 1954: 170. Type-species: Aphycus snoflaki Hoffer, by original designation. (As
subgenus of Aphycus.)
DISTRIBUTION AND SPECIES. Twenty-seven species, cosmopolitan except Neotropics; four
from review area: coccidiphagus Girault (1917g: 134) (Australia), nassaui Girault (19320: 4)
(Australia), parisoti Girault (1936: 1) (Australia) and rubescens (Compere & Annecke,
1961: 41) (Taiwan), also several undetermined species from Pakistan, India and Australia
(BMNH, BPBM).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. One of the Indian species placed here has complete notaular lines and appears to be
very close to snoflaki Hoffer which is the type-species of subgenus Aphycaspis. We have
considered the possibility that Aphycaspis should be raised to generic status because of the
complete notaular lines, but believe that it would be best to defer this until a more detailed study
of related genera can be undertaken e.g. Cirrhencyrtus Timberlake and Echthroplexiella
Mercet.
Aphycus is placed in the tribe Aphycini by Trjapitzin (19736) and almost certainly the tribe
should also include Echthroplexiella (type-genus of subtribe Echthroplexiellina). (Trjapitzin
(19736) places the subtribe Echthroplexiellina in the tribe Miraini which must be incorrect since
Mira (type-genus of the Miraini) belongs in the Tetracneminae near the tribe Charitopidini (see
also comments under Mira).) It is very probable that further study will show that Echthro-
plexiella and Aphycus are very close and may even be synonymous since most of the characters
used to separate these two genera are unreliable. We also believe that the status of the tribe
Homalotylini should be reconsidered because it is very close to the Aphycini and often difficult
to separate, even at a subtribal level. The presence or absence of notaular lines is not reliable
since many species of Aphycus, e.g. hederaceus (Westwood), have notaular lines present in the
extreme anterior part of the mesoscutum. These are visible on examination of well-cleared
specimens using a phase contrast microscope.
INDO-PACIFIC ENCYRTIDAE 235
APOLEPTOMASTIX Kerrich
(Key couplet: 270. Fig. 166)
Apoleptomastix Kerrich, 1982: 416. Type-species: Apoleptomastix spoliata Kerrich, by original des-
ignation.
DISTRIBUTION AND SPECIES. Six species, Oriental, Afrotropical and Australasian; five from
review area: bicoloricornis (Girault; Kerrich, 1982: 427) ( Australia), poonensis (Mani & Kaul;
Kerrich, 1982: 420) (India), rufipleurus Kerrich (1982: 421) (India), rufiscapus Kerrich (1982:
422) (India) and spoliata Kerrich (1982: 424) (Pakistan, India), also at least two further species
from India, Bangladesh, Cambodia, Laos, Thailand, Vietnam, China and Hong Kong (BMNH,
BPBM, GC).
BIOLOGY. Parasites of Pseudoccoidae (Homoptera).
COMMENTS. A. bicoloricornis and spoliata are extremely close and may be synonymous.
The genus can be placed in the tribe Anagyrini, subtribe Anagyrina (Tetracneminae) and can
be separated from related genera by the characters given in the key and also by using the
characters listed by Kerrich (1982).
ARRHENOPHAGOIDEA Girault
(Key couplet: 48. Figs 15, 16)
Arrhenophagoidea Girault, 1915a: 73. Type-species: Arrhenophagoidea coloripes Girault, by original
designation.
DISTRIBUTION AND SPECIES. Four species, Neotropical, Afrotropical and Australasian; one from
review area: coloripes Girault; Annecke & Prinsloo (1974: 41) (Australia, New Zealand).
REFERENCE. Revision: Annecke & Prinsloo (1974).
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. Placed in the tribe Psyllechthrini (Encyrtinae).
ARRHENOPHAGUS Aurivillius
(Key couplet: 45. Figs 9-11)
Arrhenophagus Aurivillius, 1888: 144. Type-species: Arrhenophagus chionaspidis Aurivillius, by
monotypy.
Mymariella Risbec, 1951: 402. Type-species: Mymariella parlatoriae Risbec, by monotypy.
DISTRIBUTION AND SPECIES. Two species, New World, Palaearctic, Afrotropical, Oriental and
Pacific; both from review area: albitibiae Girault; Annecke & Prinsloo (1974: 38) (Sri Lanka,
Hong Kong, China, Hawaiian Is.) and chionaspidis Aurivillius; Annecke & Prinsloo (1974: 36)
(India, Sri Lanka, New Zealand), also one undetermined species from Samoa (BMNH).
REFERENCE. Revision: Annecke & Prinsloo (1974).
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. The only included genus in the tribe Arrhenophagini (Encyrtinae).
ASEIRBA Cameron
(Key couplet: 293)
Aseirba Cameron, 1884: 127. Type-species: Aseirba caudata Cameron, by monotypy.
DISTRIBUTION AND SPECIES. One, Neotropical; one undescribed species from Sarawak (BPBM).
236 J. S. NOYES & M. HAY AT
BIOLOGY. Unknown.
REFERENCE. Noyes (1980: 179).
COMMENTS. The species from Sarawak differs from caudata in having tridentate mandibles,
although on one side the third, inner tooth is rather obscure. Having taken into consideration
other characters and the size and shape of the mandible we do not consider that this difference is
generic. Aseirba is very close to Austroencyrtus and more or less differs only in having the
hypopygium reaching the apex of the gaster, the marginal vein of the forewing relatively shorter,
not or hardly longer than broad, whereas in Austroencyrtus the hypopygium does not reach
more than two-thirds along the gaster and the marginal vein is clearly several times longer than
broad. The undescribed Neotropical species previously placed in Aseirba (Noyes, 1980: 179) can
be better placed in Austroencyrtus.
Placement of the genus according to Trjapitzin's (19736) classification is difficult but it may be
related to the Bothriothoracini, possibly in an as yet undefined group of genera which would also
include Austroencyrtus and Hemileucocerus . It can be separated from these genera by the
characters given in the key.
(Key couplet: 206. Figs 124, 274-275)
Asitus Erdos, 1955: 47. Type-species: Asitus ciliatus Erdos, by original designation.
Ferriereus Ghesquiere, 1956: 698. Type-species: Xanthoencyrtus phragmitis Ferriere, by original des-
ignation.
DISTRIBUTION AND SPECIES. One, Palaearctic and Oriental: phragmitis (Ferriere, 1955: 13)
(Pakistan).
BIOLOGY. Parasitic on mealybugs associated with Phragmites (Homoptera, Pseudococcidae).
COMMENTS. The genus belongs to the subtribe Rhopina (tribe Anagyrini, Tetracneminae) and
can be separated from other related genera of the subtribe by having an extremely dorso-
ventrally flattened body and a solid clava.
ASTYMACHUS Howard
(Key couplet: 402. Figs 208, 209, 276)
Astymachus Howard, 1898ft: 238. Type-species: Astymachus japonicus Howard, by monotypy.
DISTRIBUTION AND SPECIES. Two, Palaearctic, Oriental; one from review area: japonicus
Howard; Tachikawa (1963: 160) (India, Pakistan, Malaysia).
BIOLOGY. Reared from Aclerdidae (Homoptera) on sugarcane. Also reported from Pseudococ-
cidae (Homoptera) on sugarcane which is possibly erroneous.
COMMENTS. Placed in the tribe Astymachini by Trjapitzin (19736) as the sole included genus. It is
quite possibly related to genera in the tribe Aphycini.
AUSTRALANUSIA Girault
(Key couplets: 185, 5 10)
Australanusia Girault, 1922a: 47. Type-species: Australanusia pilosithorax Girault, by monotypy.
DISTRIBUTION AND SPECIES. Two species, both Australian: pilosithorax Girault (1922a: 47) and
tarsalis Girault (1923d: 2).
BIOLOGY. Unknown.
INDO-PACIFIC ENCYRTIDAE 237
COMMENTS. The two included species are extremely close and may be synonymous; they appear
to differ only in the characters given by Girault (1923d: 2).
The genus probably belongs to the tribe Microteryini and should be distinguishable from
other genera placed here by Trjaptzin by the combination of the solid clava, transverse funicle
segments, very conspicuous hairs on the eyes and dorsum of thorax, and the setae in basal cell of
forewing being about as dense of those in centre of wing.
AUSTRALAPHYCUS Girault
(Key couplet: 285)
Australaphycus Girault, 1923c: 143. Type-species: Australaphycus albioviductus Girault, by monotypy.
DISTRIBUTION AND SPECIES. Australia, one species: albioviductus Girault (1923c: 143).
BIOLOGY. Unknown.
COMMENTS. Girault, in his description, states 'ovipositor free', which we take to mean that the
hypopygium does not extend to near the apex of the gaster. Examination of the holotype (QM)
of albioviductus shows that the hypopygium appears to reach at least to the apex of the gaster, or
perhaps very slightly beyond. However, since the specimen is badly mounted on a microscope
slide that may be misleading. The genus appears to be close to Aenasioidea (tribe Aphycini,
subtribe Paraphycina) and may prove to be synonymous when fresh material is collected.
A USTRALIA Girault
(Key couplet: 298. Fig. 179)
Australia Girault, 1928a: 3. Type-species: Australia minuta Girault, by monotypy.
DISTRIBUTION AND SPECIES. One species, Australia only: minuta Girault (1928a: 3).
BIOLOGY. Unknown.
COMMENTS. This genus possibly also includes Parachalcerinys coccidoxenoides Girault from
which it can be separated by the relative lengths of the antennal segments.
Australia almost certainly belongs in the tribe Aphycini (Encyrtinae) and can be separated
from other genera placed here by the metallic green body, wing venation and conformation of
the antennae. Most other genera of the tribe have non-metallic bodies.
AUSTROCHOREIA Girault
(Key couplets: 80, 88)
Chinchilla Girault, 1928a: 1. Type-species: Chinchilla keatsi Girault, by monotypy. [Homonym of
Chinchilla Bennett, 1829.] Syn. n.
Austrochoreia Girault, 1929a: 3. Type-species: Austrochoreia latiscutum Girault, by monotypy.
Chinchillisca Ghesquiere, 1946: 369. [Replacement name for Chinchilla Girault.] Syn. n.
DISTRIBUTION AND SPECIES. Two described species, both Australian: keatsi (Girault, 1928a: 1)
(comb. n. from Chinchilla) and latiscutum Girault (1929a: 3), also several other species from
Australia and New Zealand (BMNH, DSIR).
BIOLOGY. Unknown.
COMMENTS. Taken in isolation, the two described species appear to exhibit enough morphologi-
cal differences to warrant retaining the genera they were described in as distinct. Austrochoreia
latiscutum has a very transverse scutellum without a distinct flange apically or laterally and the
mesoscutum is slightly exposed posteriorly, whilst keatsi has a much longer, more rounded
scutellum with a clear flange apically and laterally under which the wing partly fits and the
mesoscutum is completely hidden by the pronotum. Other differences, e.g. body size, colour
238 J. S. NO YES & M. HAY AT
and relative proportions of the antennal segments, could largely be taken as specific. However, a
study of the other material available has shown that the differences in the scutellum and
pronotum length are inconsistent and therefore we propose that the two genera be synonymised.
The genus is best placed in the tribe discodini (Encyrtinae) and can be distinguished from
other included genera by the elongate pronotum, which largely covers the mesoscutum, the lack
of notaular lines and the abbreviated wings.
AUSTROENCYRTOIDEA Girault
(Key couplet: 451)
Austroencyrtoidea Girault, I922d: 206. Type-species: Austroencyrtoidea leichhardti Girault, by monotypy.
DISTRIBUTION AND SPECIES. One species, Australia only: leichhardti Girault (1922d: 206).
BIOLOGY. Unknown.
COMMENTS. The genus may belong to the subtribe Syrphophagina (tribe Microteryini, En-
cyrtinae) and can be separated from other genera placed there by the combination of the solid
clava, elongate postmarginal vein of the forewing and strongly tridentate mandibles. All other
genera included in this subtribe have a two- or three-segmented clava, although some oc-
casionally have an elongate postmarginal vein and tridentate mandibles.
AUSTROENCYRTUS Girault
(Key couplet: 293. Fig. 278)
Austroencyrtus Girault, 1923c: 141. Type-species: Austroencyrtus annulicornis Girault, by monotypy.
DISTRIBUTION AND SPECIES. Neotropics and Australasia; two species from review area: annuli-
cornis Girault (1923c: 141) (Australia) and guamensis (Fullaway, 1946: 208) (comb. n. from
Cerchysius) (Mariana Is.), also at least three further undescribed species from Papua New
Guinea and New Hebrides (BMNH, BPBM).
BIOLOGY. Associated with Cerambycidae and other beetles (Coleoptera) in rotting logs.
COMMENTS. The genus is near Aseirba and Hemileucocerus (see comments under Aseirba) . The
holotype of Cerchysius guamensis has been examined (USNM) and belongs to the present
genus.
AUSTROMIRA Girault
(Key couplet: 261)
Austromira Girault, 19246: 3. Type-species: Austromira muironi Girault, by monotypy.
DISTRIBUTION AND SPECIES. Australia only, one species: muironi Girault (19246: 3) and possibly
one other from Australia (BMNH).
BIOLOGY. Unknown.
COMMENTS. Almost certainly related to Cheiloneurus (Encyrtinae, tribe Cheiloneurini) and
distinguished from other related genera by the lack of an apical tuft of setae on the scutellum,
more or less evenly infuscate forewings (except apex which is hyaline), pattern of setae on the
forewing (a central area just distal to linea calva where the setae are distinctly more sparse than
proximal or distal to it, setae in basal cell more or less extending to base of forewing) and
coloration and structure of antennae (apical funicle segments and clava white, contrasting with
basal segments, and flagellum cylindrical, not conspicuously broadening apically).
INDOPACIFIC ENCYRTIDAE 239
A VETIANELLA Trjapitzin
(Key couplet: 182. Figs 105, 106, 277)
A vetianella Trjapitzin, 1968: 97. Type-species: AvetianellacapnodiobiaTrjapitzin, by original designation.
DISTRIBUTION AND SPECIES. Two described species, Holarctic, also Neotropics and Oriental; one
undescribed species from India (BMNH).
BIOLOGY. Parasites of eggs of Cerambycidae and Scolytidae (Coleoptera).
COMMENTS. Placed in the tribe Microteryini, subtribe Oobiina (Encyrtinae). Trjapitzin (1977)
provides a key to the genera of this subtribe.
BACALUSA gen. n.
(Key couplets: 240, 277. Figs 165, 173, 174, 279-288)
Type-species: Bacalusa fuscipennis sp. n. Gender: feminine.
$ . Head. In facial view about as broad as long, or clearly broader than long, in profile about one-half longer
than wide and anteriorly more or less gradually and evenly curved. Eye with posterior margin straight,
about one-half longer than broad, with a few short inconspicuous setae and reaching occipital margin which
is sharp. Malar space a little less than half as long as eye, with malar sulcur present. Frontovertex slightly
less than half width of head; ocelli forming a right angle, posterior ocellus separated from occipital margin
by less than to about its own diameter, and from eye margin by its own diameter or a little more. Antennal
scrobes moderately deep, but not strongly margined, more or less meeting dorsally and reaching a little
more than halfway to anterior ocellus from toruli; antennal torulus separated from mouth margin by about
its own length and from other torulus by a little less than one and a half times its own length, its dorsal
margin slightly below the ventral level of the eyes. Antennal scape more or less cylindrical, about as long as
maximum width of frontovertex, about six times as long as broad, pedicel conical and distinctly longer than
any of the funicle segments which are all clearly longer than broad, cylindrical and slightly broadening
distally, funicle six-segmented, clava three-segmented, about half to two-thirds as long as, and slightly
wider than, the funicle, with apex more or less rounded and sutures parallel; longitudinal sensillae on all
but the first two or three flagellar segments. Frontovertex with very fine, raised, rugose to rugose-reticulate
sculpture which may give it a silky appearance, more irregular and elongate on cheeks and more
squamiform-reticulate on inter-antennal prominence, numerous short translucent or white recumbent
setae on frontovertex. Mandible narrow with two acute apical teeth, maxillary palpus three-segmented,
labial palpus two-segmented.
Thorax. In side view moderately deep and dorsally quite flat with metapleurum and propodeum broadly
in contact with hind coxa. Pronotum in dorsal view with hind margin gradually curved and moderately
concave; visible part of mesoscutum about twice as broad as long, with notaular lines absent or present in
anterior half; axillae meeting; scutellum a little broader than long, slightly longer than visible part of
mesoscutum, with apex rounded; propodeum short medially, not more than about one-seventh length of
scutellum. Mesoscutum with fine, raised, squamiform-reticulate sculpture, scutellum with similar but
rather more longitudinally elongate sculpture, both mesoscutum and scutellum often having an almost silky
appearance, propodeum with or without shallow, raised sculpture medially, mesopleurum with shallow
raised reticulate sculpture; dorsum of thorax with numerous short, appressed, translucent setae. Forewing
hyaline or with a distinct fuscous pattern, wing from about two and one half to a little less than three times
as long as broad, linea calva interrupted or closed by about three lines of setae, filum spinosum absent,
submarginal vein with an apical hyaline break, marginal vein about twice as long as broad, about as long as
or longer than postmarginal and slightly shorter than stigmal, costal cell relatively narrow, over 18 times as
long as broad and with a single line of setae dorsally in distal half. Hindwing hyaline, about five to six and
one-half times as long as broad with marginal fringe about two-thirds maximum wing width . Mid tibial spur
a little shorter than basal mid tarsal segment.
Caster. About as long as thorax, cereal plates in anterior half, paratergites present, last tergite from only
a little longer than half length of to as long as mid tibia, hypopygium reaching apex of gaster, ovipositor
very slightly exserted and about two-thirds length of mid tibia, gonostyli fused to second valvifers and
about one-sixth length of ovipositor.
0". Differs from female as follows.
Head. Malar space at least about two-thirds length of eye; frontovertex clearly broader than half head
240 J. S. NOYES & M. HAY AT
width, posterior ocellus separated from occipital margin by less than its own diameter and from eye margin
by a little less than twice its diameter; antennal scrobes absent; antennal torulus separated from mouth
margin by about twice its own length and from other torulus by about its own length, its ventral margin
slightly below to well above the ventral margins of the eyes; antennal scape about as long as to distinctly
shorter than width of frontovertex, pedicel short, a little longer than broad and about half as long as any of
the funicle segments which are at least about twice as long as broad; longest setae on funicle about four
times as long as maximum width of segments, longitudinal sensillae on all flagellar segments but the first
one or two, scale-like sensillae in distal half of sixth funicle segment or proximal half of clava. Sculpture of
head similar to that of female but less silky in appearance.
Thorax. Similar to that of female, except if infuscation of forewing present then less strong than in
corresponding females and sculpture of dorsum of thorax less fine and lacking silky appearance.
Caster. Similar to female except cereal plates of distal half of gaster and genitalia: digiti about
one-quarter to one-eighth length of aedeagus which in turn is a little less than half as long as mid tibia or
about twice as long as mid tibial spur.
COMMENTS. The genus belongs to the tribe Anagyrini, subtribe Anagyrina (Tetracneminae). It
can be distinguished from other members of the subtribe by the conformation of the antenna,
the occasional presence of notaular lines on the mesoscutum and infuscation of the forewing.
Bacalusa fuscipennis sp. n.
(Figs 165, 174, 279-288)
<j>. Length: approx. 0-78-0-81 mm (holotype, 0-81 mm).
Colour. Head orange, antenna yellow with basal half of scape and apex of clava slightly dusky, thorax
and gaster dark orange, distinctly dusky in centre and anterior margin of mesoscutum, apex of tegula,
along midline of scutellum, sides of propodeum immediately above hind coxa and on gaster immediately
distad of cereal plates, legs yellow; infuscation of forewing as in Fig. 279.
Head. Frontovertex above scrobes with very fine, raised, transversely rugose-reticulate sculpture of
moderately silky appearance (Fig. 280), lower parts of face and interantennal prominence with more
squamiform-reticulate sculpture; posterior ocellus separated from occipital margin by a little more than
half its own length and from eye margin by about one and one-half times its own length. Relative
measurements (holotype): head width (facial view) 55, head length 55, minimum frontovertex width 26,
malar space 17, eye length 34, eye width 23, POL 8, OOL 6, scape length 30, other proportions of antenna
as in Fig. 174.
Thorax. Notaular lines present and reaching slightly more than halfway across mesoscutum; mesoscu-
tum with fine, raised, squamiform-reticulate sculpture (Fig. 281), that on scutellum similar but laterally
more longitudinally elongate (Fig. 282), both mesoscutum and scutellum distinctly less silky in appearance
than frontovertex and with only a few appressed, translucent setae. Relative measurements of forewing
(holotype): length 129, width 42, other proportions as in Figs 165, 279; of hindwing: length 100, width 15.
Gaster. Relative lengths (paratype): ovipositor 68, gonostylus approx. 10, last tergite 60, [mid tibia 105].
Ovipositor Fig. 283, hypopygium Fig. 284.
Cf . Length: approx. 0.75 mm.
Similar to.female except following. Coloration slightly darker and infuscation of forewing distinctly
paler. Antennal toruli with their lower margins clearly a little above ventral margins of eyes (Fig. 285),
otherwise differs from female as in generic description. Relative measurements (paratype): minimum
frontovertex width 36, head width 63, scape length 35, proportions of antenna as in Fig. 286; aedeagus
length 31, length mid tibial spur 14, genitalia as in Figs 287, 288.
DISTRIBUTION. India.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $, India: Tamil Nadu, 3 km E. Manjaler Dam, 15-18.X.1979 (J. S. Noyes) (BMNH).
Paratypes. India: 1 $, 1 Cf , same data as holotype (BMNH).
COMMENTS. We consider that Doliphoceras tachikawai Shafee, Alam & Agarwal (1975: 26) also
belongs to this genus (comb. n.). It differs from fuscipennis in coloration, hyaline forewings and
lack of notaular lines. There is also a possible third species which appears to be distinguishable
INDO-PACIFIC ENCYRTIDAE 241
from fuscipennis in having the mesoscutum and scutellum more silky in appearance and more
dense, white, appressed setae, the subapical fuscous marking of the fore wing more oblique and
the body coloration of the female generally more reddish or orange. This third species has been
found in India and Zimbabwe (BMNH), but may only be a form of fuscipennis .
BACHIANA Girault
(Key couplet: 368)
Bachiana Girault, 1940: 149. Type-species: Bachiana curiosa Girault, by monotypy.
DISTRIBUTION AND SPECIES. One species, Australia only: curiosa Girault (1940: 149).
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. Placement of the genus is difficult because the type-material is in poor condition.
However, the mandible (one tooth and a truncation) and wing venation suggest that it could be
placed in the Microteryini (Encyrtinae). It is possibly related to the genera placed in the subtribe
Syrphophagina and can be easily separated from these by having an anelliform first funicle
segment, interrupted linea calva and by the shape of the mandible. Girault's unpublished
manuscript (QM) states that the clava is entire, but the only clava located on either of the two
slides containing syntypes appears to be two-segmented. There appear to be no intact female
antennae on either of these slides. (See also comments under Coccidoctonus .)
BAEOANUSIA Girault
(Key couplets: 203, 485, 523)
Baeoanusia Girault, 19150: 163. Type-species: Baeoanusia magnidava Girault, by original designation.
DISTRIBUTION AND SPECIES. Australia only, three species: albifunide Girault (19320: 3), magni-
dava Girault (19150: 164) and persimilis Girault (19150: 164).
BIOLOGY. Unknown. However, albifunide (which is misplaced in this genus, see below) has been
reared from the eggs oiParopsis sp. (Coleoptera, Chrysomelidae) (Riek, 1962c).
COMMENTS. Baeoanusia albifunide is misplaced in this genus, appearing to be intermediate
between Mesanusia and Baeoanusia and, in some respects, has some resemblance to a large
species of Zaomma. Almost certainly a new genus is required to accommodate it. However,
since the species can be reasonably well placed in Baeoanusia, we feel that a new genus is
unnecessary at the present time, at least until fresh material becomes available and the
relationships between these genera can be studied in more detail.
The genus belongs to the tribe Cheiloneurini (Encyrtinae) and is closest to Neblattidda from
which it can be separated by having hyaline wings, whilst Neblattidda has infuscate wings. A
more detailed study of fresh material may indicate that the two genera should be considered
synonymous. Baeoanusia can be separated from other members of the tribe by having finely
punctate sculpture on the head and dorsum of the thorax (as in Blastothrix) and a large antennal
clava, and lacking an apical tuft of setae on the scutellum.
BEETHOVENA Girault
(Key couplet: 64)
Beethovena Girault, 1932a: 3. Type-species: Beethovena longifasciata Girault, my monotypy.
DISTRIBUTION AND SPECIES. One species, Australia only: longifasdata Girault (19320: 2).
BIOLOGY. Unknown.
COMMENTS. The genus is very close to Metaphycus (Aphycini, subtribe Paraphycina) from which
it can be separated by having a five-segmented funicle, whilst Metaphycus has a six-segmented
funicle.
242 J. S. NO YES & M. HAY AT
BLASTOTHRIX Mayr
(Key couplets: 209, 446)
Blastothrix Mayr, 1876: 697. Type-species: Encyrtus sericeus Dalman, by designation of Ashmead (1900ft:
389).
DISTRIBUTION AND SPECIES. Twenty-seven species, Holarctic, Afrotropical, Oriental; three
species from review area: britannica Girault; Sugonjaev (1964: 382) (Pakistan), sericea (Dal-
man; Sugonjaev, 1964: 381) (Pakistan, India) and siddiqii (Bhatnagar, 1952: 167) (comb. n.
from Encyrtus) (India), also one undetermined species from India (BMNH).
REFERENCE. Revision of Palaearctic species: Sugonjaev (1964).
BIOLOGY. Parasites of Coccidae and possibly also Kermesidae (Homoptera).
COMMENTS. We have not seen the holotype of Encyrtus siddiqii but from the description of the
species and the host record we feel certain that it belongs in Blastothrix.
Trjapitzin (19736) places the genus in the tribe Aphycini (subtribe Blastothrichina) which we
believe may be incorrect since it appears to be closely related to Psyllaephagus. The latter is
placed in the tribe Trechnitini, subtribe Metaprionomitina. The subtribe Blastothrichina should
probably be transferred from the Aphycini to the Trechnitini, but we do not formally propose
this since it is beyond the scope of the present work. Blastothrix is recognisable by the metallic
green or blue-green colour, deep puntcate sculpture of the head and thorax, the mandible
having one tooth and a broad truncation, and the fore wing with a marginal vein at least three
times as long as broad.
BLEPYRUS Howard
(Key couplet: 479)
Blepyrus Howard, 1898ft: 233. Type-species: Blepyrus mexicanus Howard, by designation of Ashmead
(1900ft: 373).
Coccophoctonus Ashmead, 1900ft: 375. Type-species: Coccophoctonus dactylopii Ashmead, by original
designation.
DISTRIBUTION AND SPECIES. Three species, circumtropical; one from review area: insularis
(Cameron; Kerrich, 1967: 226) found throughout the area except New Zealand.
REFERENCE. Revision: Kerrich (1967: 225-228).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The genus is a member of the tribe Aenasiini (see comments under Aenasius). It can
be separated from other related genera by the characters given in the key, notably the stigmal
vein of the forewing being clearly shorter than the postmarginal and the relatively wide
fronto vertex. Kerrich (1967: 188-190) also provides a key to most of the genera of the tribe.
BORROWELLA Girault
(Key couplets: 359, 465)
Borrowella Girault, 1923ft: 99. Type-species: Borrowella bioculata Girault, by monotypy.
DISTRIBUTION AND SPECIES. Australia only; two species: bioculata Girault (19236: 99)
(= Borrowella consobrina Girault, 1923d: 2 syn. n.) and punctatinotum Girault (19236:
100); possibly also one further species near punctatinotum but with the ovipositor less exserted.
BIOLOGY. Unknown.
COMMENTS. Borrowella consobrina appears to be a colour form of bioculata and the two are
therefore regarded as synonymous.
The genus probably belongs in the tribe Bothriothoracini. It can be separated from other
INDO-PACIFIC ENCYRTlDAE 243
members of the tribe principally by the darkened forewing and the postmarginal vein being
longer than the stigmal.
BOTHRIOPHR YNE Compere
(Key couplet: 282)
Bothriophryne Compere, 1937: 45. Type-species: Bothriophryne ceroplastae Compere, by original de-
signation.
DISTRIBUTION AND SPECIES. Seven species, Afrotropical; none from review area but two
undescribed species from India (Agarwal etal., 1980: 30).
REFERENCE. Prinsloo & Annecke (19786: 323-325).
BIOLOGY. Parasites of Coccidae (Homoptera).
COMMENTS. The genus is close to Trichomasthus (Microteryini, subtribe Microteryina) and is
most easily separated by having the antennal toruli relatively high on the head, their lowest
margins at or above the level of the lowest eye margins (in Trichomasthus they are well below),
and the marginal vein of forewing punctiform (in Trichomasthus it is generally much longer than
broad).
BOTHRIOTHORAX Ratzeburg
(Key couplet: 425)
Bothriothorax Ratzeburg, 1844: 209. Type-species: Bothriothorax aliens teinii Ratzeburg, by monotypy.
Trimorphocerus Dahlbom, 1857: 292. Type-species: Bothriothorax altensteinii Ratzeburg, by designation
of Gahan & Pagan (1923: 149).
DISTRIBUTION AND SPECIES. Thirty species, Holarctic and Oriental; two undetermined species
from Taiwan and India (BMNH, BPBM).
REFERENCE. Peck (1963: 375).
BIOLOGY. Parasites of larvae of Syrphidae (Diptera).
COMMENTS. Placed in the tribe Bothriothoracini, subtribe Bothriothoracina (Encyrtinae).
BRACHYPLATYCERUS De Santis
(Key couplet: 52)
Brachyplatycerus De Santis, 1972: 49. Type-species: Brachyplatycerus minutum De Santis, by original
designation.
DISTRIBUTION AND SPECIES. One species, Neotropical; also one species reported from India
(Hayat & Subba Rao, 1981).
BIOLOGY. Unknown.
COMMENTS. We have been unable to locate the material referred to by Hayat & Subba Rao
(1981: 109) and presumably either the material was misidentified or has been lost.
The genus is related to Pentelicus (see comments under Pentelicus), differing in the number of
funicle segments (see key).
CAENOHOMALOPODA Tachikawa
(Key couplet: 54)
Caenohomalopoda Tachikawa, 1979a: 169. Type-species: Pseudhomalopoda shikokuensis Tachikawa, by
original designation.
DISTRIBUTION AND SPECIES. Four species, eastern Palaearctic, Oriental, Australasian; two from
244 J. S. NOYES & M. HAY AT
review area: guamensis (Fullaway; Tachikawa, 19790: 169) (Mariana Is., Hawaiian Is.) and
nagaii (Tachikawa, 1978: 65) (Indonesia), also undetermined material from India, Taiwan,
Indonesia, Brunei, Philippines and Australia (BMNH, BPBM).
REFERENCE. Key to species: Tachikawa et al. (1981).
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. In addition to the characters given by Tachikawa (19790) to separate this genus from
Pseudhomalopoda Girault we have found that the shape of the pronotum is important. In
Caenohomalopoda the posterior margin of the pronotum is almost straight, whereas in
Pseudhomalopoda it is strongly concave and medially incised.
The genus belongs in the tribe Habrolepidini, subtribe Habrolepidina (Encyrtinae) and can
be separated from most other genera included here by the characters given in the key or by the
key to genera of the subtribe provided by Tachikawa (19790).
CALLIPTEROMA Motschulsky
(Key couplet: 229)
Callipteroma Motschulsky, 1863: 35. Type-species: Callipteroma quinqueguttata Motschulsky, by desig-
nation of Ashmead (19006: 402).
Calocerinella Girault, 1913d: 46. Type-species: Calocerinella trifasdata Girault, by original designation.
Vosleria Timberlake, 1926: 1. Type-species: Vosleria signata Timberlake, by original designation. Syn. n.
DISTRIBUTION AND SPECIES. Five species, old World; three from review area: australia (Girault;
Noyes, 1978: 543) (= Vosleria signata Timberlake, 1926: 3 syn. n.) (Australia), sexguttata
Motschulsky; Noyes (1978: 546, 548) (Pakistan to Australia) and testacea Motschulsky; Noyes
(1978: 549) (Pakistan to Australia).
REFERENCE. Revision: Noyes (1978).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The holotype of Vosleria signata has not been examined, but it is clear from
Timberlake's detailed description that it is the same as australia.
We regard sexguttata and quinqueguttata as synonymous. This synonymy was first proposed by
Boucek (19770: 70) and has been confirmed by the examination of much fresh material collected
in southern India and other parts of the region.
The genus is placed in the tribe Anagyrini, subtribe Leptomastideina (Tetracneminae) by
Trjapitzin (19730), although the Leptomastideina is here considered synonymous with the
Anagyrina (see comments under Leptomastided) . The forewing venation suggests that Callipter-
oma may possibly be better placed in the Dinocarsini (or Dinocarsina if reduced to subtribal
status within the Anagyrini).
CARABUNIA Waterston
(Key couplet: 224. Figs 128-131)
Carabunia Waterston, 1928a: 249. Type-species: Carabunia myersi Waterston, by original designation.
Elijahia Girault, 1928a: 1. Type-species: Elijahia poeta Girault, by monotypy. Syn. n.
Schillerana Girault, 1932a: 6. Type-species: Schillerana dilatata Girault, by monotypy. Syn. n.
DISTRIBUTION AND SPECIES. Seven species, Neotropical, Oriental and Australasian; four from
review area: dilatata (Girault, 19320: 6) (comb. n. from Schillerana) (Australia) , longimarginalis
Subba Rao (1973: 486) (India, Malaysia), orientalis Subba Rao (1971: 211) (India, Bangladesh,
Thailand) and poeta (Girault, 19280: 1) (comb. n. from Elijahia) (Australia), also unidentified
material from India to Philippines, Papua New Guinea and Solomon Is. (BMNH, BPBM, CNC,
GC).
IN DO-PACIFIC ENCYRTIDAE 245
REFERENCE. Part revision: Subba Rao (1973).
BIOLOGY. Parasites of nymphs of Cercopidae (Homoptera).
COMMENTS. The types of dilatata and poeta have been examined (QM) and are congeneric with
the two above species described in Carabunia by Subba Rao. The Oriental and Australasian
species of the genus (which could be called thepoeta-group) differ from the Neotropical species
in having a much more elongate postmarginal vein (that in myersi is very obscure, or perhaps
absent, shortly distal to the apex of the stigmal vein); we do not consider this to be a generic
difference.
The genus is placed in the tribe Neocladiini (Encyrtinae) by Trjapitzin (1973b) but we believe
that this tribe is too narrowly defined (see comments under Anagyrodes). It can be separated
from related genera by the combination of the mandible having one long sickle-shaped tooth,
solid clava, long marginal and postmarginal veins of forewing, lightly to moderately infuscate
fore wing and the hypopygium more or less extending to apex of gaster.
CERAPTEROCEROIDES Ashmead
(Key couplet: 108. Figs 289, 290)
Cerapteroceroides Ashmead, 1904b: 156. Type-species: Cerapteroceroides japonicus Ashmead, by
monotypy.
Metacerapteroceruslshii, 1928: 151. Type-species: Cerapterocerus fortunatuslshii, by original designation.
DISTRIBUTION AND SPECIES. Three species, Oriental and eastern Palaearctic; two species
from review area: japonicus Ashmead; Tachikawa (1963: 148) (Pakistan) and similis (Ishii;
Tachikawa, 1963: 150) (India); also undetermined material from Sri Lanka, India, Taiwan,
Indonesia and Sarawak (BMNH, BPBM, UCR).
REFERENCE. Revision: Tachikawa (1963: 142-151).
BIOLOGY. Hyperparasitic on various Homoptera (Psyllidae, Aphididae, Coccidae, Pseudococci-
dae, Diaspididae) via other Encyrtidae and Aphelinidae (Hymenoptera).
COMMENTS. Placed in the tribe Cerapterocerini (Encyrtinae). It can be separated from other
closely related genera by the characters provided in the key and by the key given by Annecke
(1967: 100-101).
CERAPTEROCERUS Westwood
(Key couplets: 85, 108. Figs 41, 291, 292)
Cerapterocerus Westwood, 18336: 495. Type-species: Cerapterocerus mirabilis Westwood, by monotypy.
Jurinia Costa, 1839: 115. Type-species: Jurinia platicera Costa, by designation of Boucek (1970: 86).
Telegraphus Ratzeburg, 1848: 152. Type-species: Telegraphus maculipennis Ratzeburg, by monotypy.
DISTRIBUTION AND SPECIES. Eight species, Holarctic, Afrotropical, Oriental, Australasian; four
species from review area: australia Girault (I9lle: 97) (Australia), emersoni Girault (19150:
102) (Australia), subapterus Girault (19220: 48) (Australia) and virens Agarwal (1963: 398)
(India), also further undetermined material from India, China, Hong Kong, Singapore,
Malaysia, Sarawak and Sulawesi (BMNH, BPBM, USNM).
BIOLOGY. Hyperparasites of Coccidae (Homoptera) via other Encyrtidae.
COMMENTS. The holotype of australia appears to be lost but it may be possible to recognise the
species from Girault's description when freshly collected material becomes available. However,
until that time the name should be considered a nomen dubium.
Placed in the tribe Cerapterocerini (Encyrtinae). It can be separated from other closely
related genera by the characters given in the key and also by the key provided by Annecke (1967:
100-101).
246 J. S. NOYES & M. HAY AT
CERAPTROCERELLA Girault
(Key couplets: 115, 128)
Ceraptrocerella Girault, 1918: 1. Type-species: Ceraptrocerella apus Girault, by original designation.
Austrotropidia Kerrich, 1978: 143. Type-species: Tropidophryne flandersi Compere, by original desig-
nation. Syn. n.
DISTRIBUTION AND SPECIES. Australia only; one species: apus Girault (1918: 1) (= Tropidophryne
flandersi Compere; Kerrich, 1978: 143 syn. n.).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. We have not seen the holotype of flandersi but a specimen determined as this by
Kerrich (who has examined the holotype) is the same species as one compared with the syntypes
of apus.
The genus belongs in the tribe Chrysoplatycerini, which also contains Chrysoplatycerus,
Hambletonia, Neoplatycerus and Tropidophryne. It can be separated from these genera using
the characters given in the key, or by using the key provided by Kerrich (1978: 113-114).
CERCHYSIELLA Girault
(Key couplets: 215, 324, 445, 447, 509. Fig. 127)
Aratus Howard, 1897: 155. Type-species: Aratus scutellatus Howard, by monotypy. [Homonym of Aratus
Milne-Edwards, 1853.] Syn. n.
Cerchysiella Girault, 1914ft: 60. Type-species: Cerchysiella nigrella Girault, by monotypy.
Zeteticontus Silvestri, 1915ft: 343. Type-species: Zeteticontus abilis Silvestri, by original designation.
Syn. n.
Mirrencyrtus Girault, 1915a: 115. Type-species: Mirrencyrtus glabriscutellum Girault, by original desig-
nation. Syn. n.
Ericydnella Girault, 1915a: 169. Type-species: Ericydnella ashmeadi Girault, by original designation.
Syn. n.
Aratiscus Ghesquiere, 1946: 368. [Replacement name for Aratus Howard.] Syn. n.
Prolitomastix Hoffer, 1954: 173. Type-species: Prolitomastix vestonicensis Hoffer, by original designation.
Syn. n.
DISTRIBUTION AND SPECIES. Eighteen species, cosmopolitan; eight from review area: abilis
(Silvestri, 19156: 345) (comb. n. from Zeteticontus) (Pakistan), glabriscutellum (Girault, 19150:
115) (comb. n. from Mirrencyrtus) (= Mimencyrtus [sic] arboris Girault, 19230: 47 syn. n.)
(Australia), kamathi (Mani & Saraswat in Mani et al., 1974: 84) (comb. n. from Prionomitus)
(India), nigra Girault (19150: 82) (Australia), nigrella Girault (19146: 60) (=Ericydnella
ashmeadi Girault, 19150: 169 syn. n.) ( Australia), perkinsi (Timberlake, 1924: 402) (comb. n.
from Zeteticontus}, umbilicata Girault (19150: 83) (Australia) and utilis (Noyes, 1982: 457)
(comb. n. from Zeteticontus) (Hawaiian Is.), also further undetermined material from through-
out the region except New Zealand (BMNH, BPBM, CNC, AMNH, USNM, UCR, HC).
REFERENCE. Revision: Subba Rao (1972).
BIOLOGY. Parasites of larvae of Nitidulidae, Erotylidae and Silvanidae (Coleoptera) and
apparently also Trypetidae (Diptera) (Tachikawa, 1981: 100).
COMMENTS. Girault (19150: 82) published the formal description of the genus Cerchysiella
stating that the type-species was nigra described on the same page. However, the generic name
had been validated one year previously with the publication of the description of Cerchysiella
nigrella Girault. The holotype of Cerchysiella nigrella has been examined (QM) and is
congeneric with Zeteticontus abilis Silvestri, the type-species of Zetiticontus . Unfortunately
therefore the name Cerchysiella has precedence over Zeteticontus. We do not feel that a
submission to the International Commission on Zoological Nomenclature to ask for suppression
of Cerchysiella in favour of Zeteticontus is necessary since the name Zeteticontus has been
IN DO-PACIFIC ENCYRTIDAE 247
relatively little used in the literature. In addition to the above, therefore, we propose that the
following extra-limital species all be transferred to Cerchysiella (all comb, n.): amurensis
Khlopunov (from Zeteticontus) , centennalis Erdos (from Zeteticontus) , insularis Howard (from
Bothriothorax) , laevigata De Santis (fromAratiscus), laeviscutum Thomson (from Microterys),
planiscutellum Mercet (from Zeteticontus), punctiscutellum Subba Rao (from Zeteticontus),
scutellata Howard (from Aratus) , takenakai Tachikawa (from Zeteticontus) andxanthopus Masi
(from Zeteticontus) .
The genus is placed in the tribe Bothriothoracini, subtribe Coenocercina. It can be separated
from related genera by the characteristic arrangement of the pegs constituting the filum
spinosum (Fig. 127), and from related genera (Pentacladocerus and Zaommoencyrtus) by the
characters given in the key.
CERCHYSIUS Westwood
(Key couplets: 288, 345. Fig. 176)
Cerchysius Westwood, 1832: 128. Type-species: Encyrtus urocerus Dalman, by designation of Westwood
(1840: 73).
DISTRIBUTION AND SPECIES. Eleven species, cosmopolitan; five from review area: australiensis
Ashmead (1900a: 342) (Australia), australis (Girault, 19146: 59) (comb. n. from Copidosomd)
(= Cerchysius australis Girault, 1915: 85) (Australia), hispidiscutum Girault (1915a: 83)
(Australia), laticeps Kerrich (1954: 372) (India, Malaysia) and robustus Girault (1915a: 84)
(Australia), also further material, which may include several undescribed species, from India,
Malaysia, Philippines and Australia (BMNH, BPBM, HC).
REFERENCE. Partial world revision: Kerrich (1954).
BIOLOGY. Parasites of Chamaemyiidae (Diptera).
COMMENTS. Girault (1917e: 96) synonymised Copidosoma australis with Cerchysius australis. It
is clear from a comparison of the two descriptions that he inadvertently described the same
specimen as a new species in two different genera in separate publications.
The genus is placed in the Microteryini, subtribe Pseudencyrtina (Encyrtinae) by Trjapitzin
(19736). However, it may be related to the subtribe Metaprionomitina of the Trechnitini since it
can be very difficult to separate from some Australian species of Psyllaephagus which also have a
long exserted ovipositor (see key).
CERCOBELUS Walker
(Key couplet: 49. Figs 17, 18, 293, 294)
Cercobelus Walker, 1842: vi. Type-species: Encyrtus jugaeus Walker, by monotypy.
DISTRIBUTION AND SPECIES. One described species, Europe; at least one undescribed species
from Afrotropical region (BMNH) and from review area: India, Sarawak and Australia
(BMNH, BPBM, QM, ANIC).
REFERENCE. Kryger (1951: 99-103).
BIOLOGY. Parasites of nymphs of Psyllidae (Homoptera).
COMMENTS. The genus is commonly attributed to Walker and dated 1840. However, Graham
(1969) threw some doubt on the authorship of the genus, saying that the plate on which
Cercobelus jugaeus was figured was actually drawn by Haliday. It is quite possible that Haliday
drew the figures and that Walker wrote the legends to the plates. Whichever is the truth of the
matter we shall probably never know and therefore we retain Walker as the author of the genus.
Almost certainly the date commonly attributed is incorrect since the legends to the figures were
published along with the index to volume 1 of the Entomologist and it is highly unlikely that the
248 J. S. NOYES & M. HAY AT
index was published before the final part of this volume which was published in 1842. Therefore
we have no hesitation in dating the genus 1842 and not 1840.
The genus is the sole representative of the tribe Cercobelini (Encyrtinae). Trjapitzin (19736)
states that the mandible is tridentate which is incorrect since a fourth tooth is present (Fig. 293).
The structure of the gaster is very unusual in the Encyrtidae since it is highly telescopic (see
Kryger, 1951: 101-102) and is probably adapted for its particular mode of oviposition.
CHARITOPUS Forster
(Key couplet: 263. Fig. 7)
Charitopus Forster, 1856: 31. Type-species: Charitopus fulviventrls Forster, by designation of Forster
(1860: 112).
Leptorhopala Motschulsky, 1863: 60. Type-species: Leptorhopala cuprifrons Motschulsky, by monotypy.
Eupelmomorpha Girault, 1915a: 43. Type-species: Eupelmomorpha quadricolor Girault, by designation
of Gahan & Pagan (1923: 60). Syn. n.
Diversicornia Mercet, 1916c: 371. Type-species: Diversicornia pinicola Mercet, by original designation.
DISTRIBUTION AND SPECIES. Thirteen species, Palaearctic, Afrotropical, Oriental, Australasian;
seven species from review area: apicatus (Mani & Saraswat in Mani et al., 1974: 79) (India),
bicolor (Girault, 1915: 44) (comb. n. from Eupelmomorpha) ( Australia), fulviventris Forster;
Trjapitzin (1969a: 675) (India), cuprifrons (Motschulsky; Trjapitzin, 19646: 242) (Sri Lanka),
panchgania (Mani & Saraswat in Mani etal., 1974: 81) (India), quadricolor (Girault, 1915<z: 43)
(comb. n. from Eupelmomorpha) (Australia) and tricolor (Girault, 19150: 43) (comb. n. from
Eupelmomorpha) (= Eupelmomorpha hawthornei Girault, 1915a: 44 syn. n.), also undeter-
mined material from Sulawesi and Bangladesh (BMNH).
REFERENCES. Keys to species: Trjapitzin (19690: 675) and Hoffer (1980: 388).
BIOLOGY. Unknown, but almost certainly parasites of Pseudococcidae (Homoptera).
COMMENTS. There appears to be some considerable variation in colour within some species and it
is probable that many of the above species are synonymous since they are separated largely on
colour differences, e.g. bicolor, quadricolor and tricolor.
The genus is placed in the tribe Charitopidini (Tetracneminae) which probably contains some
of the most primitive encyrtids known. They are characterised by the very long marginal vein of
the forewing, well-developed notaular lines and short last gastral tergite so that the cereal plates
are situated near the apex of the gaster. Most genera have membranous areas surrounding the
mid coxae which allow the mid legs to be flexed forwards, particularly when dead. This is also
characteristic of the Tanaostigmatidae and some Eupelmidae.
CHEILONEURELLA Girault
(Key couplets: 259, 309, 408. Figs 149, 155, 295)
Cheiloneurella Girault, 1915a: 177. Type-species: Cheiloneurella binotativentris Girault, by original
designation.
DISTRIBUTION AND SPECIES. Only one described species: binotativentris Girault (19150: 177)
(Australia), but also other material, containing at least one undescribed species, from India,
Thailand, Hong Kong, Malaysia, Indonesia and Philippines (BMNH, BPBM).
BIOLOGY. Unknown.
COMMENTS. The genus very probably belongs in the tribe Cheiloneurini (Encyrtinae) and can be
separated from other genera of the tribe by having a very long pronotum which is triangular in
dorsal view (Fig. 149) and not covered by the head.
INDO-PACIFIC ENCYRTIDAE 249
CHEILONEUROMYIA Girault
(Key couplet: 379)
Cheiloneuromyia Girault, 19150: 178. Type-species: Cheiloneuromyia simpliciscutellum Girault, by
original designation.
DISTRIBUTION AND SPECIES. Three species, Oriental and Australasian: javensis Girault (1916c:
480) (Indonesia, Hawaiian Is.), planchoniae (Howard in Howard & Ashmead, 1896: 637)
(comb. n. from Encyrtus) (Sri Lanka) and simpliciscutellum Girault (19150: 178) (Australia),
also some undetermined material from India and Solomon Is. (BMNH, BPBM).
BIOLOGY. Parasites of Coccidae (Homoptera).
COMMENTS. The genus most probably belongs in the tribe Cheiloneurini (Encyrtinae).
CHEILONEURUS Westwood
(Key couplets: 87, 101, 103, 131, 351, 384, 474, 487. Figs 47, 50)
Cheiloneurus Westwood, 1833a: 343. Type-species: Encyrtus elegans Dalman, by monotypy.
Chrysopophagus Ashmead, 1894: 245. Type-species: Chrysopophagus compressicornis Ashmead, by
monotypy.
Blatticida Ashmead, 1904c: 305. Type-species: Blatticida pulchra Ashmead, by original designation.
Saronotwn Perkins, 1906: 259. Type-species: Saronotum australiae Perkins, by designation of Gahan &
Pagan (1923: 130).
Cristatothorax Girault, 1911: 169. Type-species: Cristatothorax pulcher Girault, by original designation.
Eusemionella Girault, 1915a: 78. Type-species: Eusemionella cristata Girault, by original designation.
Syn. n.
Chrysopophagoid.es Girault, 1915a: 90. Type-species: Chrysopophagoides westwoodi Girault, by mono-
typy. Syn. n.
Paracheiloneurus Girault, 1915a: 119. Type-species: Cheiloneurus perpulcher Girault, by original designa-
tion (as subgenus of Cheiloneurus). Syn. n.
Epicheiloneurus Girault, 1915a: 173. Type-species: Epicheiloneurus albicoxa Girault, by original desig-
nation. Syn. n.
Eusemionopsis Girault, 1918: 3. Type-species: Eusemionopsis centaurus Girault, by original designation.
Syn. n.
Metacheiloneurus Hoffer, 1957: 336. Type-species: Metacheiloneurus moestus Hoffer, by monotypy.
DISTRIBUTION AND SPECIES. Well over 100 species, cosmopolitan; 45 from review area: albifuni-
culus Hayat, Alam & Agarwal (1975: 48) (India), axillaris Hayat, Alam & Agarwal (1975: 55)
(India), australiae (Perkins, 1906: 260) (Australia), bangalorensis (Subba Rao, 1957: 382)
(India), basin Hayat, Alam & Agarwal (1975: 53) (India), beerwahi (Girault, 19256: 100)
(comb. n. from Epicheiloneurus) (Australia), burnsi (Girault, 19266: 69) (comb. n. from
Eusemionella} (Australia), cheles (Walker, 1839: 37) (comb. n. from Encyrtus) (Australia),
chlorodryini Perkins (1906: 261) (= Cheiloneurus dubius Girault, 19150: 88 syn. n.) (Australia),
chrysopae Fullaway (1946: 207) (Mariana ls.),cinctiventris (Girault, 19296: 311) (comb. n. from
Epicheiloneurus) (Australia), cristatus (Girault, 19150: 78) (comb. n. from Eusemionella)
(Australia), cupreicollis (Ashmead; Noyes, 1979: 50) (Australia), diversicolor Hayat, Alam &
Agarwal (1975: 134) (India), dumasi Girault (19320: 2) (Australia), flaccus (Walker, =
americanus Perkins, 1906: 260) (Hawaiian Is.), gonatopodis Perkins (1906: 261) (Australia),
hemipterus (Girault, 19200: 48) (comb. n. from Eusemionella) (Australia), hugoi (Girault,
1915a: 156) (comb. n. from Cristatothorax) (= Cristathorax nobilis Girault, 19226: 100 syn. n.)
(Australia), javanus Perkins (1912: 17) (Java, Brunei), javensis Girault (19170: 3) (Java),
kerrichi Hayat, Alam & Agarwal (1975: 127) (India), latifrons Hayat, Alam & Agarwal (1975:
108) (India), latiscapus (Girault, 1916c: 481) (comb. n. from Cristatothorax) (Malaysia, Java),
longicornis Hayat, Alam & Agarwal (1975: 120) (India), malayensis Noyes & Chua (1977: 544)
(Malaysia), margiscutellum (Girault, 1917g: 141) (comb. n. from Bavanusia [sic]) (Australia),
mazzinini (Girault, 19150: 103) (comb. n. from Chrysopophagus) (Australia), nepalensis Khan
250 J. S. NO YES & M. HAY AT
& Agarwal (1978: 23) (Nepal), nigricornis Hayat, Alam & Agarwal (1975: 122) (India),
novimandibularis (Girault, 19150: 158) (comb. n. from Cristatothorax) (= Cristatothorax
mandibularis Girault, 19150: 157 syn. n., = Cristatothorax mackayensis Girault, 19150: 158 syn.
n., = Cristatothorax sublimis Girault, 19296: 314 syn. n., = Cristatothorax partipes Girault,
19320: 3 syn. n.) (Australia), noxius Compere (1925: 302) (Hawaiian Is.), pasteuri (Girault,
19150: 159) (comb. n. from Cristatothorax) (= Cristatothorax bidentimaxillae Girault, 19150:
157 syn. n., = Cristatothorax vinculum Girault, 19150: 159 syn. n., = Epicheiloneurus albicoxa
Girault, 19150: 177 syn. n., = Cristatothorax bidentimaxillae poeta Girault, 19320: 3 syn. n.)
(Australia), perpulcher Girault (19150: 88) (Australia), purpureicinctus (Girault, 19150: 104)
(comb. n. from Chrysopophagus) (= Eusemionopsis centaurus Girault, 1918: 3 syn. n., =
Chrysopophagus variocelli Girault, 19240: 2 syn. n.) (Australia), purpureiventris Girault
(19150: 87) (Australia), pyrillae Mani (1939: 73) (India), quadricolor (Girault 19150: 157)
(Pakistan, India, Australia), V0r0 (Girault, 19220: 42) (comb. n. from Eusemionelld) (Austra-
lia), regis (Girault, 19320: 3) (comb. n. from Cristatothorax) (Australia), saissetiae Noyes &
Chua (1977: 541) (Malaysia), seminigridavus Girault (19150: 88) (Australia), unicolor Mercet
(19220: 155) (Java), viridiscutum (Girault, 19150: 158) (comb. n. from Cristatothorax) (Austra-
lia), westwoodi (Girault, 19150: 90) (comb. n. from Chrysopophagoides) (Australia) and
yasumatsui Trjapitzin (19716: 123) (India), also probably many other species amongst material
from throughout the region (BMNH, BPBM, DSIR, QM, ANIC, CNC, UCR, HC).
REFERENCES. Key to Palaearctic species: Trjapitzin (19716: 123-125), key to Indian species:
Hayat etal. (1975: 45-47), Khan & Agarwal (1978: 21).
BIOLOGY. Parasites of Dryinidae and chalcids (Hymenoptera), mainly Aphelinidae and Encyrti-
dae, parasitic on other insects, notably Homoptera (Auchenorrhyncha, also Coccidae, Pseudo-
coccidae, etc.) and also predatory Diptera, e.g. Drosophilidae.
COMMENTS. The single extant female syntype of Encyrtus cheles Walker (BMNH) is here
designated LECTOTYPE. It belongs to the same species-group as novimandibularis, but is in
poor condition, lacking both fore wings and most of the antennae.
We have not examined the holotype of Cheiloneurus rufescens Motschulsky (1863: 53), but
according to Z. Boucek (pers. comm.) it belongs to the family Eulophidae.
The genus is placed in the tribe Cheiloneurini (Encyrtinae). It appears to be a very large and
diverse genus whose limits are uncertain. Generally speaking, it is characterised by the
arrangement of the setae in the basal cell of the forewing, by the wing venation (relatively long
marginal and short stigmal and postmarginal veins), normally infuscate forewing, the usual
presence of an apical tuft of setae on the scutellum , and the hypopygium never reaching the apex
of the gaster . We have included here in Cheiloneurus two unusual species, one being cinctiventris
which has the unusual character of the basal cell of the forewing being almost entirely setose and
the other, an undescribed species from Papua New Guinea (BPBM), which has the forewing
entirely hyaline and an unusually long marginal vein (Fig. 50). It is possible that once this
difficult complex of genera (which includes Tobiasia Trjapitzin, Neabrolepoideus, Baeoanusia,
Neblatticida and Mesocalocerinus) is studied in more detail a number of them will be considered
synonymous with Cheiloneurus.
CHR YSOPLA TYCERUS Ashmead
(Key couplet: 116. Figs 54, 55)
Rileya Howard in Smith, 1888: 80. Type-species: Rileya splendens Howard, by monotypy. [Homonym of
Rileya Ashmead, 1888.]
Chrysoplatycerus Ashmead, 1889: 38. [Replacement name for Rileya Howard.]
Encyrtolophus De Santis, 1972: 49. Type-species: Encyrtolophus flavicollis De Santis, by original
designation.
Paraplatycerus Hall, 1974: 19. Type-species: Paraplatycerus citriculus Hall, by original designation.
Metaplatycerus Gordh & Trjapitzin in Trjapitzin & Gordh, 1978a: 384. Type-species: Chrysoplatycerus
ferrisi Timberlake, by original designation.
INDO-PACIFIC ENCYRTIDAE 251
DISTRIBUTION AND SPECIES. Four species, New World, Afrotropical; one species in review area:
splendens (Howard; Kerrich, 1978: 140) (Hawaiian Is.).
REFERENCE. Revision: Kerrich (1978: 136-142).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Chrysoplatycerini, subtribe Chrysoplatycerina (Tetracneminae).
CLADISCODES Subba Rao
(Key couplet: 480. Fig. 235)
Cladiscodes Subba Rao, 1977: 18. Type-species: Cladiscodes sacchari Subba Rao, by original designation.
DISTRIBUTION AND SPECIES. One species: sacchari Subba Rao (1977: 19) (India), also undeter-
mined material from Laos, Vietnam and Australia (BMNH, BPBM).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The genus is related to Monodiscodes and Metaphaenodiscus (tribe Aenasiini, see
comments under Aenasius) and is characterised by the costal cell of the forewing being abruptly
incised at its apex and by the distinct venation and shape of the wing (Fig. 235).
CLAUSENIA Ishii
(Key couplet: 265)
Clausenia Ishii, 1923: 98. Type-species: Clausenia purpurea Ishii, by original designation.
DISTRIBUTION AND SPECIES. Eleven species, Afrotropical, Palaearctic, Oriental and Pacific; two
from review area: lacca (Agarwal, 1962: 278) (India) and purpurea Ishii (Kerrich, 1967: 182)
(S. China, Taiwan, Hawaiian Is.).
REFERENCE. Revision: Kerrich (1967: 181-188).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Ericydnini by Trjapitzin (1973) but probably would be better
placed in the Charitopidini.
COAGERUSgtn. n.
(Key couplet: 487. Figs 245, 296-299)
Type-species: Coagerus boucekisp. n. Gender: feminine.
9 . Head. In facial view about as long as broad and in profile clearly less than twice as long as broad and
anteriorly more or less gradually and evenly curved, but most strongly at top of antennal scrobes. Eye with
posterior margin very slightly concave, almost straight, about one-third longer than broad, almost naked
with some sparse short setae each not longer than half the diameter of a facet, eye just reaching occipital
margin which is sharp. Malar space a little more than half length of eye and with sulcus absent.
Frontovertex about one-fifth head width, ocelli forming an angle of about 45, posterior ocellus a little less
than half its own major diameter from occipital margin and touching eye margin. Antennal scrobes
shallow, meeting dorsally and not quite reaching half way between toruli and anterior ocellus; antennal
torulus separated from mouth margin by about half its length and from other torulus by about two to three
times its own length, its upper margin at least about its own length below lowest eye margin, clypeal margin
very shallowly produced below toruli. Antennal scape about twice as long as minimum width of
frontovertex and slightly flattened and broadened, slightly more than three times as long as broad, pedicel
slightly less than half as long as scape, cylindrical and clearly longer than any of the funicle segments the
first four of which are distinctly transverse and the fifth and sixth subquadrate; clava three-segmented with
an oblique apical truncation, the outer suture strongly converging with the inner, the truncated surface
with two rows of 'tubular' setae and also a few scattered on other surfaces of apical segment of clava, these
only visible on slide material examined at higher magnifications (x250+); longitudinal sensillae on fifth
252 J. S. NOYES & M. HAY AT
and sixth funicle segment and clava only. Frontovertex with shallow, regular, hexagonal, raised, reticulate
sculpture, this becoming irregular and more longitudinally elongate on lower parts of face; setae very
sparse, inconspicuous, brown and short, not present on frontovertex below anterior ocellus except along
eye margin. Mandible with three equal acute apical teeth; maxillary and labial palpi not visible in
slide-mounted material available.
Thorax. In side view with mesoscutum and scutellum distinctly convex and with metapleurum and
propodeum together quite broadly in contact with hind coxa. In dorsal view with posterior margin of
pronotum moderately concave; visible part of mesoscutum about twice as broad as long, its posterior
margin slightly convex medially; axillae touching; scutellum a little broader than long, its apex rounded;
propodeum medially not more than about one-tenth length of scutellum. Mesoscutum with very shallow,
raised, squamiform-reticulate sculpture, that on axillae similar but a little deeper; scutellum with deep,
fine, raised, longitudinally striate-reticulate sculpture (Fig. 297) clearly a lot deeper than sculpture of
mesoscutum, apical one-fifth and extreme sides smooth and polished; dorsum of thorax with fairly
numerous, moderately long, brown setae. Forewing more or less hyaline, but with a short fuscous streak in
disc beyond venation and slight clouding below marginal vein, wing nearly three times as long as broad;
linea calva not interrupted and open; film spinosum present; submarginal vein with parastigma not
conspicuously thickened, with an apical hyaline break; costal cell about 13 times as long as broad with only
two or three setae dorsally near apex; marginal vein about four or five times as long as broad, about twice as
long as stigmal which is clearly longer than postmarginal, stigmal vein with three apical sensillae arranged
in a line. Hindwing hyaline, about two-thirds as long as forewing, about four to five times as long as broad,
marginal fringe about one-third wing width. Mid tibial spur about as long as or a little shorter than basal
mid tarsal segment.
Caster. Slightly longer than thorax with ovipositor slightly exserted, cereal plates in basal half,
hypopygium reaching to about half way along gaster, last tergite slightly shorter than mid tibia; ovipositor
about one-third longer than mid tibia, gonostyli free and about one-third to two-fifths as long as ovipositor.
Cf . Unknown.
COMMENTS. At first glance this new genus bears a striking superficial resemblance to Para-
leurocerus Girault but is easily separated by the three-segmented clava (in Paraleurocerus it is
entire), postmarginal vein of forewing shorter than stigmal (in Paraleurocerus it is clearly
longer) and infuscate forewings. However, the basic type of wing venation, strongly tridentate
mandible and structure of ovipositor suggest that it has some affinity with the group of genera to
which Paraleurocerus belongs, i.e. tribe Copidosomatini, subtribe Ageniaspidiina, but can be
separated from all other genera included in this subtribe by the postmarginal vein of the
forewing being shorter than the stigmal.
The type-species of the genus is named in honour of Dr Z. Boucek.
Coagerus bouceki sp. n.
(Figs 245, 296-299)
$. Length (excluding ovipositor): 0-67-0-97 mm (holotype, 0-97 mm).
Colour. Head black dorsally with dull greenish and brassy reflections, around mouth and antennal toruli
slightly purplish ; antennal torulus , basal half or so of scape and pedicel dark brown , remainder of antenna
yellow, the apex of clava indistinctly fuscous; pronotum purplish brown, mesoscutum shining metallic
green, along anterior and posterior margins a little purplish; tegula brown; scutellum matt, black, apical
one-fifth or so and extreme sides polished and metallic green; mesopleurum purplish brown, slightly shiny
with some brassy, green and bluish reflections; propodeum dark purple-brown laterally on outer face with
distinct bluish hue; legs white to yellow with apical one-third of mid femur, extreme base and a narrow
sub-basal band on mid tibia and extreme apex of hind femur dark brown; forewing as in Fig. 245; gaster
with venter and basal area dorsally yellow; apex dorsally continuing along sides to base dark purplish
brown; exserted part of gonostyli dark brown, apices yellowish.
Head. Relative measurements (holotype): head length 51, head width (facial view) 51, head width (side
view) 29, minimum frontovertex width 10-5, malar space 22, eye length 35, eye width 28, POL 6, OOL
0-25, scape length 23, scape width 6, other proportions of antenna Fig. 298. Smaller specimens tend to have
the eyes a little smaller and thus the frontovertex correspondingly wider.
Thorax. Sculpture of scutellum Fig. 297. Relative measurements (holotype): forewing length 128,
INDO-PACIFIC ENCYRTIDAE 253
forewing width 45, other proportions of forewing as in Figs 245, 296; hindwing length 90, hindwing width
19.
Caster. Relative lengths (paratype): last tergite 53, ovipositor 85, gonostyli 26, [mid tibia 62]; genitalia
Fig. 299.
Cf . Unknown.
BIOLOGY. Unknown.
DISTRIBUTION. India.
MATERIAL EXAMINED
Holotype $, India: Tamil Nadu, Coimbatore, 25.ix.-l.x.l979 (J. S. Noyes) (BMNH).
Paratypes. India: 5 $, same data as holotype; 1 $, Karnataka, Bangalore, 3.xi.l979 (Z. Boucek)
(BMNH).
COCCIDAPHYCUS Blanchard
(Key couplet: 72. Fig. 32)
Coccidaphycus Blanchard, 1940: 110. Type-species: Coccidaphycus nigricans Blanchard, by original
designation.
DISTRIBUTION AND SPECIES. One described species, Neotropical; one undescribed species from
Sarawak (BMNH).
BIOLOGY. Parasites of Coccidae (Homoptera).
COMMENTS. The genus is closely related to Trechnites (tribe Trechnitini, subtribe Trechnitina),
differing in biology (Trechnites spp. are parasites of nymphs of Psyllidae) and in the characters
given in the key.
COCCIDENCYRTUS Ashmead
(Key couplets: 194, 329, 519. Figs 117, 189)
Coccidencyrtus Ashmead, 19006: 383. Type-species: Encyrtus ensifer Howard, by original designation.
Encyrtomyia Girault, 19150: 131. Type-species: Encyrtomyia albiflagellum Girault, by original desig-
nation. Syn. n.
Omphalencyrtus Girault, 1915a: 169. Type-species: Omphalencyrtus wallacei Girault, by original desig-
nation. Syn. n.
Coccidencyrtoides Blanchard, 1940: 107. Type-species: Coccidencyrtoides annulipes Blanchard, by de-
signation of De Santis (1967: 161).
Neoadelencyrtus Hayat, Alam & Agarwal, 1975: 72. Type-species: Neoadelencyrtus mandibularis Hayat,
Alam & Agarwal, by original designation. Syn n.
DISTRIBUTION AND SPECIES. Thirty-three species, cosmopolitan; nine species from review area:
albiflagellum (Girault, 1915a: 131) (comb. n. from Encyrtomyia) (Australia), albitarsis (Girault,
1915a: 132) (comb. n. from Encyrtomyia) (Australia), auricornis (Girault, 19240: 2) (comb. n.
from Epitetracnemus) (Australia), australis (Girault, 1915: 132) (comb. n. from Encyrtomyia),
(Australia) bicolor (Girault, 1915: 141) (comb. n. from Coccidoxenus) (Australia), mandibu-
laris (Hayat, Alam & Agarwal, 1975: 74) (comb. n. from Neoadelencyrtus) (India), ochraceipes
Gahan (1927a: 18) (Hawaiian Is.), secundus (Girault, 19150: 131) (comb. n. from Encyrtomyia)
(Australia) and wallacei (Girault, 1915: 169) (comb. n. from Omphalencyrtus) (Australia).
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. The type-species of Neoadelencyrtus differs from the other species of Cocciden-
cyrtus only in the number of segments in the maxillary and labial palpi: mandibularis has
four-segmented maxillary and three-segmented labial palpi whilst all species of Coccidencyrtus
that have been examined have one segment fewer in each. We do not think that this difference
warrants separate generic status.
254 J. S. NOYES & M. HAY AT
Encyrtomyia and Omphalencyrtus are rather more problematic. The type-species of
Omphalencyrtus has the funicle distinctly four-segmented, whilst that of Encyrtomyia has the
first segment with two partial sutures so that in slide-mounted material it could be taken as
one-segmented (i.e. four-segmented funicle) and in dry-mounted material it appears three-
segmented (i.e. six-segmented funicle). Examination of the extant types of albitarsis and
secundus (which may be synonymous with albiflagellum} also shows this to be the case, whilst in
australis the funicle is definitely four-segmented. Several other specimens with partial segmen-
tation of the first funicle segment have also been examined. The occurrence of an apparent
partial fusion of the first three funicle segments in some specimens, or species, is therefore not
uncommon. Taking this into consideration and the fact that the first funicle segment of both
australis and wallacei is longer than those following and is about as long as that which might be
expected if the first three segments became fused, we have no hesitation in regarding
Omphalencyrtus and Encyrtomyia as synonymous with Coccidencyrtus .
The genus is placed in the tribe Habrolepidini (Encyrtinae) by Trjaptizin & Gordh (19786).
COCCIDOCTONUS Crawford
(Key couplets: 199, 300, 367. Fig. 125)
Coccidoctonus Crawford, 1912: 167. Type-species: Coccidoctonus trinidadensis Crawford, by original
designation.
Quaylea Timberlake, 19196: 214. Type-species: Cerchysius whittieri Girault, by original designation.
Cerchysiopsis Girault, 19226: 108. Type-species: Cerchysiopsis lowelli Girault, by monotypy. Syn. n.
DISTRIBUTION AND SPECIES. Seven described species, New World, Australasia, Pacific; six
species from review area: dubius (Girault, 1915: 102) (comb. n. from Rhopalencyrtoided)
(= Rhopalencyrtoidea cinctifemur Girault, 1925: 2 syn. n., = Paraenasomyia liszti Girault,
19326: 1 syn. n.) (Australia), lowelli (Girault, 19226: 108) (comb. n. from Cerchysiopsis)
(Australia), oviductus (Girault, 1915a: 85) (comb. n. from Cerchysius) (Australia), psyllae
(Riek, 19626: 189) (comb. n. from Echthroplexis) (Australia), terebratus (Hayat, Alam &
Agarwal, 1975: 69) (comb. n. from Echthroplexis) (India) and whittieri (Girault; = aliena
Timberlake, 19196: 216) (Hawaiian Is. , New Zealand), also several other species from Australia
(BMNH,QM,ANIC).
BIOLOGY. Hyperparasites of Coccidae, Pseudococcidae and Psyllidae (Homoptera) via Pter-
omalidae and other Encyrtidae (Hymenoptera).
COMMENTS. Girault (19326) incorrectly proposed liszti as a replacement name for Rhopalencyr-
toidea dubia Girault, 1915, believing that it was preoccupied by Paraenasomyia dubia Girault,
1923 when he presumably transferred this species to Paraenasomyia. This is obviously incorrect
and thus we revert to the original name. Paraenasomyia dubia Girault, 1923 is placed here in
Psyllaephagus . It is possible that Nezarhopalus caudatus also belongs in Coccidoctonus.
This genus is closely related to Syrphophagus (tribe Microteryini, subtribe Syrphophagina).
The subtribe to which these genera belong is a very difficult, complex group whose genera are
very difficult to define. We have separated them in the key by the use of the following simple
characters in order to retain most of the generic names as valid until a more detailed study of the
group can be undertaken. Two of the genera in this group have the hypopygium extending past
the apex of the last tergite, i.e. Coccidoctonus and Epiblatticida, whilst in the others it does not
extend past the apex of the last tergite. These two genera can be separated from each other by
the characters given in the key. Two of the remaining genera have the hypopygium more or less
reaching the apex of the gaster, one has the postmarginal vein of the forewing longer than the
stigmal (Rhopalencyrtoidea), whilst in the other it is not longer than the stigmal ( Teleterebratus] .
The remaining genera, Bachiana and Syrphophagus, have the hypopygium not reaching further
than four-fifths along the gaster. Bachiana has the clava two-segmented (or possibly entire)
whilst that of Syrphophagus is always three-segmented.
INDO-PACIFIC ENCYRTIDAE 255
COCCIDOXENOIDES Girault
(Key couplet: 503. Fig. 242)
Coccidoxenoides Girault, 1915a: 173. Type-species: Coccidoxenoides perminutus Girault, by original
designation.
Pauridia Timberlake, 1919ft: 206. Type-species: Pauridia peregrina Timberlake, by original designation.
Syn. n.
DISTRIBUTION AND SPECIES. Two species, New World, Afrotropical, Oriental, Australasian and
Pacific; both found in review area and possibly synonymous: peregrinus (Timberlake, 19196:
208) (comb. n. from Pauridia) (Pakistan, India, Java, Philippines, Hawaiian Is.) and perminutus
Girault (1915a: 173) (= Fulgoridicida babindae Girault, 1922a: 47 syn. n.) (Australia), also
material from the Cook Is. and New Caledonia (BMNH, BPBM, DSIR).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The holotype of Coccidoxenoides perminutus has been examined (QM). It is
congeneric with and possibly conspecific with Pauridia peregrina. Although Pauridia is a fairly
well-known name we prefer to follow the rules of zoological nomenclature and use the older
name, Coccidoxenoides, for the genus. We therefore do not think that it is necessary to apply to
the International Commission for Zoological Nomenclature to ask for suppression of Cocci-
doxenoides in favour of Pauridia.
The genus is placed in the Pauridiini (Tetracneminae).
COELASPIDIA Timberlake
(Key couplet: 75)
Coelaspidia Timberlake, 1923: 326. Type-species: Coelaspidia osborni Timberlake, by original desig-
nation.
DISTRIBUTION AND SPECIES. One described species, Neotropics and Hawaiian Is.: osborni
Timberlake (1923: 330).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Acroaspidiini (Tetracneminae).
COELOPENCYRTUS Timberlake
(Key couplets: 176, 218, 497, 508. Figs 100, 118, 119, 247, 248, 300, 301)
Coelopencyrtus Timberlake, 1919ft: 218. Type-species: Coelopencyrtus odyneri Timberlake, by original
designation.
Nesencyrtus Timberlake, 1919ft: 223. Type-species: Adelencyrtus kaalae Ashmead, by original desig-
nation.
Epaenasomyia Girault, 1919ft: 53. Type-species: Epaenasomyia pallidiceps Girault, by original desig-
nation. [Homonym of Epaenasomyia Girault, 1917.] Syn. n.
Giraultella Gahan & Pagan, 1923: 66. [Replacement name for Epaenasomyia Girault, 1919.] Syn. n.
Batrachencyrtus Jansson, 1957: 71. Type-species: Batrachencyrtus calidii Jansson, by monotypy.
Lymanera Szelenyi, 1972a: 125. Type-species: Lymanera crassicornis Szelenyi, by original designation.
Syn. n.
DISTRIBUTION AND SPECIES. Twenty-five species, cosmopolitan; 10 species from review area:
asperithorax (Rayment, 1949: 253) (comb. n. from Aphycus) (Australia), kaalae (Ashmead;
Timberlake, 1922a: 139) (Hawaiian Is.), krishnamurtii (Mahdihassan, 1957: 182) (comb. n.
from Giraultella) (India), mauiensis Timberlake (19220: 137) (Hawaiian Is.), odyneri Timber-
lake (19196: 221) (Hawaiian Is.), orbi Timberlake (1920: 422) (Hawaiian Is.), pallidiceps
(Girault, 19196: 53) (comb. n. from Epaenasomyia) (Java), sexramosus (Timberlake, 1922a:
141) (Hawaiian Is.), swezeyi Timberlake (19196: 222) (Hawaiian Is.) and xylocopae (Girault,
256 J. S. NOYES & M. HAYAT
19 196: 54) (comb. n. from Epaenasomyid) (Java), also further undetermined material from
India to Papua New Guinea (BMNH, BPBM, RMNH).
REFERENCE. Review of Hawaiian species: Timberlake (1922a: 135-142).
BIOLOGY. Polyembryonic parasites of larvae of Xylocopidae, Apidae and Hylaeidae (Hymen-
optera).
COMMENTS. The syntypes of Epaenasomyia pallidiceps in the Queensland Museum have been
examined. They are morphologically very close to species of Codopencyrtus (although differing
in colour) and therefore we propose that the two genera be considered synonymous.
The holytpe of Lymanera crassicornis has been examined (HNHM) and is a typical species of
Codopencyrtus.
There are several specimens determined as Giraultella krishnamurthi in the collections of the
USNM. They are almost certainly syntypes and belong to the genus Coelopencyrtus .
Zarhopaloides cinctithorax (Girault, 1939a: 20) and Anagyrus saintpierrei Girault (1913e:
112) may both be aberrant species belonging in this genus.
The genus is placed in the tribe Copidosomatini, subtribe Coelopencyrtina (Encyrtinae) by
Trjapitzin (19736).
COMPERIA Gomes
(Key couplet: 132)
Comperia Gomes, 1942: 41. Type-species: Dicarnosis merceti Compere, by original designation.
DISTRIBUTION AND SPECIES. Seven species, New World, Afrotropical; one species found in review
area: merceti (Compere, 1938: 317) (India, Hawaiian Is.), also some undetermined material of
at least one further species from Samoa and Australia (BMNH, BPBM).
BIOLOGY. Parasites of cockroach oothecae (Orthoptera, Blattodea).
COMMENTS. Placed in the tribe Comperiini by Trjaptzin (19736), but possibly should be
considered as a subtribe of the Microteryini.
COMPERIELLA Howard
(Key couplet: 105. Figs 302, 303)
Comperiella Howard, 1906: 121. Type-species: Comperiella bifasciata Howard, by monotypy.
Pseudanusia Girault, 1915a: 155. Type-species: Pseudanusia pia Girault, by original designation.
Habrolepistia Mercet,. 1921: 668. Type-species: Habrolepistia cerapterocera Mercet, by original des-
ignation.
DISTRIBUTION AND SPECIES. Eight species, cosmopolitan; six from review area: aspidiotiphaga
Subba Rao (1966: 137) (Pakistan, India), bifasciata Howard (= cerapterocera Mercet, 1921:
669) (Pakistan, China, Hawaiian Is.), indica Ayyar (1934: 219) (India), lemniscata Compere &
Annecke (1961: 32) (Pakistan, India, Hong Kong), pia (Girault; Sands & Snowball, 1980: 41)
(Australia) and unifasciata Ishii; Compere (1926: 49) (India).
REFERENCE. Review of species: Hayat (1977: 249).
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. Placed in the tribe Habrolepidini, subtribe Comperiellina (Encyrtinae).
CONCHYNILLA Girault
(Key couplet: 252)
Conchynilla Girault, 1923c: 148. Type-species: Conchy nilla fuscipennis Girault, by monotypy.
DISTRIBUTION AND SPECIES. One species, Australia only: fuscipennis Girault (1923c: 148).
INDO-PACIFIC ENCYRTIDAE 257
BIOLOGY. Unknown.
COMMENTS. Girault states in his unpublished manuscript (QM), under Cerchysiella fuscipennis ,
that the spelling of the generic name Conchynilla is incorrect and that it should have been
published as Cerchysiella. Thus when stating 'As genotype but . . .', he must have actually been
referring to Cerchysiella nigra (which he had designated as type-species of the genus). However,
Article 32a(ii) of the International Code of Zoological Nomenclature states that if there is no
clear evidence of mis-spelling in the original publication then the spelling must stand. In this case
there is clear evidence of an error by Girault, but not that the spelling of the generic name is
incorrect. Therefore, since the generic name Conchynilla is not unavailable for reasons of
homonymy and the species epithet is valid (under Article llg(g) of the Code) and fuscipennis
does not belong to any other genus known to us, we here regard the genus and generic name as
valid.
The genus is probably related to the subtribe Syrphophagina (Microteryini), but can be
distinguished from all other included genera by the distinctly infuscate fore wings and very long,
sharply tridentate mandible.
COPIDOSOMA Ratzeburg
(Key couplets: 235, 250, 287, 292, 297, 325, 394, 432, 453, 484, 498, 507, 529. Figs 140, 142, 183,
204,241,249)
Copidosoma Ratzeburg, 1844: 157. Type-species: Copidosoma boucheanum Ratzeburg, by monotypy.
Litomastix Thomson, 1876: 171 . Type-species: Encyrtus chalconotus Dalman, by designation of Ashmead
(1900: 363).
Pentacnemus Howard, 1892: 366. Type-species: Pentacnemus bucculatricis Howard, by monotypy. Syn. n.
Parapsilophrys Howard, 1898ft: 232. Type-species: Parapsilophrys gelechiae Howard, by monotypy.
Berecyntus Howard, 18986: 237. Type-species: Berecyntus bakeri Howard, by monotypy.
Pseudencyrtella Girault, I9l3e: 113. Type-species: Pseudencyrtella fasciata Girault, by original designa-
tion. Syn. n.
Zaomencyrtus Girault, 1915a: 107. Type-species: Zaomencyrtus lepidopterophagus Girault, by original
designation. Syn. n.
Paracaenocercus Girault, 1915a: 116. Type-species: Paracaenocercus perseverans Girault, by monotypy.
Syn. n.
Paracopidosomopsis Girault, 1916a: 49. Type-species: Berecyntus floridanus Ashmead, by original
designation.
Verdunia Mercet, 1917ft: 203. Type-species: Verdunia gloriosa Mercet, by original designation.
Litomastiellus Mercet, 1921: 443. Type-species: Litomastix claviger Mercet by designation of Peck in
Muesebeck et al. (1951: 481).
Limastotix Mercet, 1921: 443. Type-species: Litomastix hispanicus Mercet, by monotypy.
Angeliconana Girault, 1922e: 150. Type-species: Angeliconana eja Girault, by monotypy. Syn. n.
Parasteropaeus Girault, 1923a: 50. Type-species: Par aster op aeus lotae Girault, by monotypy. Syn. n.
Neocopidosoma Ishii, 1923: 101. Type-species: Neocopidosoma komabae Ishii, by monotypy.
Mesocopidosomyiia Girault, 1925ft: 93. Type-species: Mesocopidosomyiia variventris Girault, by mono-
typy. Syn. n.
Mesencyrtus Timberlake, 1941: 228. Type-species: Mesencyrtus insularis Timberlake, by original desig-
nation. Syn. n.
Berecyntiscus Ghesquiere, 1946: 368. [Unnecessary replacement name for Berecyntus Howard.]
Arrenoclavus Doutt, 1948: 145. Type-species: Copidosoma koehleri Blanchard, by original designation.
DISTRIBUTION AND SPECIES. About 150 species, cosmopolitan; 25 species from review area:
aeripes (Girault, 19326: 1) (comb. n. from Zaomencyrtus) (Australia), australia Girault (1917g:
133) (Australia), australicum Girault (1917g: 133) (Australia), australis Girault (1917g: 133)
(Australia), compressiventris Girault (1915a: 112) (Australia), daccaensis (Mani, 1941: 28)
(comb. n. from Litomastix) (Bangladesh), desantisi Annecke & Mynhardt (1974: 32) (Austra-
lia), fasciatum (Girault, 1913e: 113) (comb. n. from Pseudencyrtella) (Australia), insularis
(Timberlake, 1941: 230) (comb. n. from Mesencyrtus) (Marquesas ls.),javae (Girault, 1917a: 5)
258 J. S. NOYES & M. HAY AT
(comb. n. from Paracopidosomopsis] (India, 3a\a),javensis (Girault, 1919b: 56) (comb. n. from
Copidosomopsis) (Java), koehleri Blanchard; Annecke & Mynhardt (1974: 32) (India), lepidop-
terophagus (Girault, 19150: 107) (comb. n. from Zaomencyrtus) (Australia), longiartus
(Girault, 19320: 1) (comb. n. from Liothorax) (Australia), lotae (Girault, 19230: 50) (comb. n.
from Par aster opaeus) (Australia), lucetius (Walker, 1839: 36) (comb. n. from Encyrtus)
(Australia), maculatum (Ishii; Tachikawa, 1963: 199) (Australia, New Zealand), manilae
(Ashmead, 19040: 14) (comb. n. from Coccidencyrtus) (Philippines) , parkeri (Girault, 19320: 2)
(comb. n. from Helegonatopus) ( Australia), perseverans (Girault, 19150: 116) (comb. n. from
Paracaenocercus) (= Angeliconana eja Girault, 1922e: 150 syn. n.) (Australia), salacon
(Walker, 1839: 37) (comb. n. from Encyrtus) (Australia), shakespearei Girault (1923d: 2)
(Australia), truncatellum (Dalman; = aestivalis Mercet, 1921: 447) (Hawaiian Is.), variventris
(Girault, 19256: 94) (comb. n. from Mesocopidosomyiia) (Australia) and walshi (Mercet,
19220: 154) (comb. n. from Litomastix) (Java), also much undetermined material from
throughout the region, probably containing many undescribed species (BMNH, BPBM, QM,
ANIC, CNC, UCNM, HC, GC).
BIOLOGY. Polyembryonic parasites of larvae of Lepidoptera.
COMMENTS. The single extant male syntype of Encyrtus salacon Walker (BMNH) is here
designated LECTOTYPE.
There are two male syntypes of Encyrtus lucetius Walker in the BMNH. One of them is here
designated LECTOTYPE and has been labelled as such.
There has been much discussion concerning the maintaining of Copidosoma and Litomastix as
two separate genera. The majority of workers in North America have taken the view that they
should be considered synonymous, but most workers in Europe have regarded them as distinct.
Certainly the type-species of the respective genera are very different and in some regions of the
world (e.g. Europe) the genera can be separated easily and with confidence. However, it has
been our experience, whilst examining material from throughout the world, that the two genera
are impossible to separate. The usual combination of characters for separating them (obliquely
truncate solid clava, hypopygium not extending to the apex of the gaster for Litomastix and
apically rounded or transversely truncate solid or three-segmented clava, hypopygium extend-
ing to the apex of the gaster for Copidosoma} are not at all reliable. Even by using other
characters, e.g. whether the ovipositor is exserted, relative length of marginal vein of forewing,
sculpture, general body shape, etc., we have not been able to separate the species into these two
recognised genera with any degree of certainty. For example, a species in North America has a
well-exserted ovipositor and hypopygium reaching the apex of the gaster but with all other
characters typical of Litomastix; a species from India has the antenna typical of Copidosoma but
the rest of the body like Litomastix, and so on. With this in mind we are following the majority of
the North American workers in considering the two genera as synonymous. This decision is
further enhanced by the fact that where their biology is known, all species are polyembryonic
parasites of Lepidoptera, and all species have a characteristic square arrangement of the
sensillae at the apex of the stigmal vein of the forewing and the uncus absent (Figs 142, 183). This
latter character is not known to us in any other encyrtid group except some members of the tribes
Dinocarsini and Anagyrini (Tetracneminae) and Rhinoencyrtini (Encyrtinae).
Copidosoma is placed in the tribe Copidosomatini, subtribe Copidosomatina (Encyrtinae).
COPIDOSOMOPSIS Girault
(Key couplet: 69. Figs 29, 30)
Copidosomopsis Girault, 1915a: 94. Type-species: Copidosomopsis perminutus Girault, by monotypy.
Pseudolitomastix Eady, 1960a: 667. Type-species: Pseudolitomastix nacoleiae Eady, by original desig-
nation. [Homonym of Pseudolitomastix Risbec, 1954.] Syn. n.
Pentalitomastix Eady, 19606: 173. [Replacement name for Pseudolitomastix Eady.] Syn. n.
DISTRIBUTION AND SPECIES. Four species, Neotropical, Palaearctic, Australasian; two from
INDO-PACIFIC ENCYRTIDAE 259
review area: nacoleiae (Eady, 19600: 667) (comb. n. from Pseudolitomastix) (India, Singapore,
Malaysia, Indonesia, Papua New Guinea) and perminutus Girault (19150: 94) (Australia), also
undetermined material from Papua New Guinea and Australia (BMNH, AMNH).
BIOLOGY. Polyembryonic parasites of larvae of Pyralidae and Tortricidae (Lepidoptera).
COMMENTS. The holotype of Copidosomopsis perminutus (QM) has the body mounted on a card
and the head, one fore wing and antennae mounted on a slide. Girault did not mention in his
original description how many segments the funicle of Copidosomopsis consisted of, except by
inference when he compared it with Copidosomyia, which has a six-segmented funicle. Also in
his unpublished manuscript (QM) he states that the funicle is six-segmented. However,
examination of the parts on the slide reveals the following present: five funicle segments and a
pedicel, four funicle segments plus clava, four funicle segments plus pedicel and scape and two
funicle segments plus a clava. This adds up to 15 funicle, two pedicel, one scape and two clava
segments i.e. parts of at least three antennae present. In no case is there a complete, intact
funicle with six segments. It is almost certain that Girault drew up his description from the parts
on this slide and assumed that the funicle was six-segmented. However, amongst material
collected recently by Boucek is a specimen which agrees more or less exactly with the parts of the
holotype of perminutus. This specimen has only five funicle segments. Since the biology of
perminutus is the same as nacoleiae (both species have been reared from pyralid larvae) , we feel
certain that the funicle of this species is only five-segmented and that Girault was erroneous in
believing it to be six-segmented. The wing venation, hypopygium and other morphological
characters of perminutus are very much the same as those of nacoleiae, therefore we have
no hesitation in synonymising Pentalitomastix with Copidosomopsis. Thus the following
extra-limital species are also transferred to Copidosomopsis from Pentalitomastix: arenicola
Trjapitzin, bohemicus Hoffer and plethoricus Caltagirone (all comb. n.).
The genus is very near to Copidosoma (tribe Copidosomatini, subtribe Copidosomatina) and
can be separated from it by having a five-segmented funicle. It is also very close to Raffaellia,
from which it can be separated using the characters given in the key.
COPIDOSOMYIA Girault
(Key couplet: 355. Fig. 193)
Copidosomyia Girault, 1915a: 99. Type-species: Copidosomyia cinctiventris Girault, by original desig-
nation.
Acridencyrtus Subba Rao, 1979: 144. Type-species: Acridencyrtus ambiguous Subba Rao, by original
designation. Syn. n.
Neochrysopophilus Tachikawa, 1979ft: 175. Type-species: Neochrysopophilus bhimolpornae Tachikawa,
by original designation. Syn. n.
DISTRIBUTION AND SPECIES. Three species, Oriental and Australasian: ambiguous (Subba Rao,
1979: 145) (comb. n. from Acridencyrtus) (India, Bangladesh), bhimolpornae (Tachikawa,
19796: 174) (comb. n. from Neochrysopophilus) (Thailand) and cinctiventris Girault (1915a: 99)
(Australia), also undetermined material from Hong Kong (BPBM).
BIOLOGY. Parasites of Chrysopidae (Neuroptera). Records of this genus having been reared
from Pseudococcidae (Homoptera) are probably erroneous.
COMMENTS. The genus is very close to Homalotylus (tribe Homalotylini, subtribe Homalotylina)
but can be separated by the remarkable shape of the head (see Tachikawa, 19796) and lack of
notaular lines on the mesoscutum.
COWPERIA Girault
(Key couplets: 352, 421. Fig. 191)
Cowperia Girault, v. 1919a: 167. Type-species: Cowperia punctata Girault, by monotypy.
Aminellus Masi, ix. 1919: 286. Type-species: Aminellus niger Masi, by monotypy. Syn. n.
260 J. S. NOYES & M. HAY AT
DISTRIBUTION AND SPECIES. Four species, Palaearctic, Oriental, Australasian; three from review
area: indica (Kerrich, 1963: 362) (comb. n. from Aminellus) (India, Sri Lanka), punctata Girault
(19190: 167) (Singapore) and sumatraensis (Kerrich, 1963: 363) (comb. n. from Aminellus),
also further undetermined material, including at least one undescribed species, from India and
Sri Lanka to Borneo (BMNH, BPBM, UCR, USNM).
REFERENCE. Revision: Kerrich (1963).
BIOLOGY. Parasites of coccinellid (Coleoptera) larvae which are predaceous on Pseudococcidae
(Homoptera).
COMMENTS. The single extant syntype female of Cowperia punctata Girault is here designated
LECTOTYPE (BMNH). It is very close to Cowperia indica but differs in having a slightly flatter
scutellum which is more conspicuously carinate laterally and relatively more transverse funicle
segments.
Placed in the tribe Bothriothoracini, subtribe Aminellina by Trjapitzin (19736). It can be
separated from the other included genus, Amicencyrtus , by the more distinctly convex scutellum
(see also Hayat, 19816:17).
CREMESINA gen. n.
(Key couplets: 83, 147. Figs 35, 36, 80, 304, 313)
Type-species: Cremesina aquilonaris sp. n. Gender: feminine.
9 . Head. In facial view a little broader than long and in profile about twice as long as broad and anteriorly
more or less gradually and evenly curved. Eye with posterior margin straight, about two-thirds longer than
broad and with numerous short setae and reaching or nearly reaching occipital margin which is sharp.
Malar space about one-third length of eye, with sulcus absent or present. Frontovertex slightly less than
half head width; ocelli forming a slightly acute to slightly obtuse angle, posterior ocellus separated from
occipital margin by a little less than its own major diameter and from eye margin by about its own major
diameter. Antennal scrobes shallow, not meeting dorsally and reaching about half way from antennal
toruli to anterior ocellus; antennal torulus separated from mouth margin by about half its length and from
other torulus by about its own length, its dorsal margin about level with ventral eye margin; clypeal margin
broadly excised between toruli. Antennal scape broadened and flattened about two to two and one-half
times as long as broad and a little longer than mimimum width of frontovertex, pedicel conical, a little
longer than any of the funicle segments (except perhaps the first) which are cylindrical and all clearly longer
than broad, the first a little longer and narrower than the sixth, funicle six-segmented; clava three-
segmented, a little less than half as long as funicle and with apex more or less rounded, the sutures more or
less parallel, the outer suture slightly oblique; longitudinal sensillae on all flagellar segments. Frontovertex
with very fine, transversely rugose sculpture of silky appearance and clothed with fairly dense short white
setae. Mandible narrow with two apical teeth, maxillary palpus four-segmented, labial palpus probably
two- or possibly three-segmented.
Thorax. In side view moderately deep with metapleurum and propodeum broadly in contact with hind
coxa, dorsally with mesoscutum and scutellum flat. In dorsal view (Fig. 35) pronotum with posterior
margin slightly concave; visible part of mesoscutum a little more than twice as broad as long with notaular
lines absent; axillae meeting, scutellum a little broader than long and about as long as to one-third longer
than mesoscutum and with apex acute; propodeum medially not more than about one-tenth as long as
scutellum. Dorsum of thorax with similar sculpture to frontovertex, silky in appearance and covered with
numerous short, appressed setae. Macropterous species (Fig. 80) with forewing centrally strongly
infuscate, occasionally with only the basal and apical quarters hyaline, wing a little over two and one-half
times as long as broad, linea calva interrupted just below middle and closed near posterior margin of wing,
filum spinosum present, submarginal vein with an apical hyaline break, marginal vein about three times as
long as broad, clearly longer than the short postmarginal and about as long as the stigmal; costal cell about
20 times as long as broad and with a single line of setae dorsally in distal half. Hindwing hyaline and about
three-quarters as long as forewing and a little over four times as long as broad, with marginal fringe about
one-quarter as long as maximum wing width. Brachypterous species (Fig. 304) with forewing reaching
about halfway along gaster, the apex truncate, about four times as long as broad, venation nearly reaching
apex, linea calva absent; hindwing about two-thirds length of forewing, about eight times as long as broad,
INDO-PACIFIC ENCYRTIDAE 261
gradually tapering towards apex and with venation reaching apex. Mid tibial spur nearly as long as basal
mid tarsal segment.
Caster. Slightly longer than thorax, cereal plates in anterior half, paratergites present, last tergite a little
longer to one-half longer than mid tibia, hypopygium reaching apex of gaster, ovipositor not exserted and
about one-quarter to one-third longer than mid tibia, gonostyli fused to second valvifers and about
one-fifth to one-sixth length of ovipositor.
Cf . Differs from female as follows. Eye a little smaller, about one-half longer than broad and a little
separated from occipital margin; malar space about two-fifths to one-half length of eye; malar sulcus
present. Frontovertex a little more than half head width; posterior ocellus separated from eye margin by
about its own major diameter to much more than its own major diameter, antennal scrobes more or less
meeting dorsally; antennal torulus separated from mouth margin by a little less than its own length and
from other torulus by less than to about its own length, its ventral margin only a little below or about level
with ventral margin of eye; antennal scape stout, about four times as long as broad and about one-third
shorter than minimum width of frontovertex, pedicel conical, subquadrate, a little longer than broad and
clearly shorter than any funicle segments which are all longer than broad, setae on flagellum at least about
twice as long as maximum diameter of segments, longitudinal sensillae on all flagellar segments, scale-like
sensillae on sixth funicle segment and base of clava. Forewing entirely hyaline and about two and one-half
times as long as broad, filum spinosum absent; hindwing about five times as long as broad. Gaster about as
long as thorax; genitalia with digiti about one-tenth to one-twentieth as long as aedeagus and without
hooks, aedeagus about one-half as long as mid tibia or twice as long as mid tibial spur.
COMMENTS. The genus belongs to the Anagyrini, subtribe Anagyrina (Tetracneminae) and is
probably closest toAnagyrus. It can be separated from this genus by the pattern of infuscation of
the fore wing and the presence of the filum spinosum in the fully winged species. The latter is
apparently very rare in the Tetracneminae.
Cremesinaaquilonarissp. n.
(Figs 305-313)
<j>. Length: 1-02-1-59 mm (holotype, 1-44 mm).
Colour. Head and thorax generally reddish; antenna with radicle dark brown, scape dark brown and on
outer face with a short white stripe along its ventral margin to about one-quarter along margin, dorsal
margin white, apical quarter white with extreme apex dark brown, inner face similar but white stripe along
dorsal margin wider and brown areas in centre often pale yellowish-brown; pedicel with basal half dark
brown and apical half white, funicle segments two to five white, other flagellar segments dark brown (Fig.
306), occasionally fifth funicle segment also dark brown or segments two to five pale dusky brownish white;
tegula white with apex dark brown; occasionally posterior margin of propodeum laterally dark; gaster
completely reddish as in thorax but usually mixed with dark brownish to a lesser or greater extent; legs
including fore and mid coxae yellowish white, hind coxa usually reddish mixed with brownish, all femora
occasionally slightly dusky as well as fore tibia and bases of mid and hind tibiae outwardly, foretarsus
testaceous yellow, mid and hind pretarsi dark brown; forewing infuscate (as in Fig. 80) except for a distinct
hyaline break immediately distal to infuscate area.
Head. Setae on eyes generally dark and conspicuous, particularly in larger specimens, in smaller
specimens they may be pale, short and inconspicuous; malar sulcus absent; ocelli forming a distinctly
obtuse angle; antennal toruli separated from each other by very slightly more than their own lengths.
Relative measurements (holotype): head width (facial view) 74, head length 65, minimum frontovertex
width 32, malar space 13, eye length 51, eye width 30, POL 16, OOL 6, scape length 35, other proportions
of antenna as in Fig. 306.
Thorax. Scutellum a little longer to nearly one-third longer than mesoscutum, forewing fully developed.
Relative measurements of forewing (holotype): length 68, width 27, other proportions as in Fig. 305; of
hindwing: length 50, width 12. Sculpture of mesoscutum Fig. 307.
Gaster. Relative lengths (paratype): ovipositor 52, gonostylus approx. 9, last tergite 47, [mid tibia 38].
Ovipositor as in Fig. 308, hypopygium as in Fig. 309.
Cf . Length: 0-67-1-05 mm.
Differs from female as follows. Colour. Head, thorax and gaster generally dark brown and orange or
yellowish orange along margins of eyes and face below top of antennal scrobes, slightly dusky on
interantennal prominence and dark brown on lower part of gena near base of mandible; legs and tegula
262 J. S. NOYES & M. HAY AT
more or less as for female except hind coxa almost totally dark brown; sides of thorax mixed with orange,
particularly around perimeter of mesopleurum; antenna with scape dusky white basally , with a broad dark
brown median band and apical one-third or so more or less yellowish brown, pedicel and flagellum
yellowish with base of pedicel darker mixed with brown.
Head. Setae on eyes generally less conspicuous than in female; ventral margin of antennal torulus
slightly below lower eye margin, toruli separated by about their own lengths, scale-like sensillae present on
clava. Relative measurements (paratype): head width (facial view) 60, head length 53, minimum
frontovertex width 33, malar space 13, eye length 33, eye width 22, POL 15, OOL 7, scape length 26,
proportions of antenna as in Fig. 310.
Thorax. Base of forewing and venation as in Fig. 313.
Gaster. Relative lengths (paratype): aeadeagus 48, [mid tibial spur 19]; genitalia as in Figs 311, 312.
DISTRIBUTION. India.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype , India: Uttar Pradesh, Aligarh, 8-10.xi.1979 (/. 5. Noyes) (BMNH).
Paratypes. India: 11 9, 7 Cf , same date as holotype; 1$, Uttar Pradesh, Aligarh, 24. i. 1978 (M. Hayat);
1$, Uttar Pradesh, Aligarh, lO.x.1979 (M. Verma); I $ , Uttar Pradesh, Aligarh, 13.xii.1979 (M. Hayat &
M. Verma); 1 , Uttar Pradesh, Aligarh, 10.viii.1980 (M. Hayat); 1 $, Uttar Pradesh, Aligarh, S.iii. 1981
(M. Hayat); 21 $, 1 cT, Delhi, IARI area, x.1979 (Z. Boudek); 11 $, 10 cT, Uttar Pradesh, Dehra Dun,
x.1979 (Z. Boucek) (BMNH, HC, USNM, UCR, ZI, PPRI).
COMMENTS. A further three species from India and one from Cook Is. , the latter species differing
in having brachypterous hyaline fore wings. The species can be separated in the female by
general coloration, the relative distance between the antennal toruli, the relative lengths of the
funicle segments, the relative width of the frontovertex, the angle formed by the ocelli, the
presence or absence of a malar sulcus, the extent of the infuscation and the relative lengths of the
fore wings and the relative length of the scutellum to the mesoscutum; in the male they can be
separated by the relative distance that separates the antennal toruli, their position in relation to
the lower eye margin, the relative width of the frontovertex; the presence or absence of
scale-like sensillae on the clava and the relative length of the digiti of the genitalia.
CRYPTANUSIA Girault
(Key couplet: 119)
Cryptanusia Girault, 1917/: 14. Type-species: Cryptanusia albiclava Girault, by original designation.
Anusoidea Girault, 1926c: 128. Type-species: Anusoidea aureiscutellum Girault, by monotypy.
DISTRIBUTION AND SPECIES. Seven species, all Australasian: albiclava Girault; Gordh & Trjapit-
zin (1981: 15) (Java), aureiscutellum (Girault, 1926c: 128) (Australia), comperei (Timberlake,
1929: 11) (Australia), gigantea (Girault, 1917g: 138) (comb. n. from Xenanusia) (Australia),
luzonica (Gordh, 1974: 203) (Philippines) , phoonae (Tachikawa, 1968: 117) (Singapore) and
varia (Girault, 19276: 310) (Australia).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. It is quite possible that aureiscutellum, comperei, gigantea and varia are synonymous
since they differ only in colour and slightly in the arrangement of the setae proximal to the linea
calva. A series of specimens recently collected in Australia (BMNH) exhibits a large degree of
variation in these characters.
The genus is placed in the tribe Anagyrini, subtribe Anusiina (Tetracneminae) by Gordh &
Trjapitzin (1981: 17). In our view the group to which this genus belongs (including Cyrtocory-
phes, Parectromoidella, Epanusia and Xenanusia) might be better placed within the Dinocarsini
(or Dinocarsina if it is regarded as a subtribe of the Anagyrini). Cryptanusia can be separated
from related genera (see above) by the characters given in the key.
INDO-PACIFIC ENCYRTIDAE 263
CYRTOCOR YPHES Timberlake
(Key couplet: 227. Figs 133, 316)
Cyrtocoryphes Timberlake, 1926: 5. Type-species: Cyrtocoryphes viridiceps Timberlake, by monotypy.
DISTRIBUTION AND SPECIES. One species, Fiji only: viridiceps Timberlake (1926: 8).
BIOLOGY. Unknown.
COMMENTS. The genus is close to Parectromoidella (see also comments under Cryptanusia) from
which it can be separated by the characters given in the key.
DIAPHORENCYRTUS Hayat
(Key couplets: 313, 443, 526)
Diaphorencyrtus Hayat, 1981a: 18. Type-species: Aphidencyrtus aligarhensis Shafee, Alam & Agarwal, by
original designation.
DISTRIBUTION AND SPECIES. Three species, all Oriental and possibly synonymous: aligarhensis
(Shafee, Alam & Agarwal, 1975: 91) (India) , diaphorinae (Lin & Tao, 1979: 117) (comb. n. from
Psyllaephagus) (Taiwan) and diaphorinae (Myartseva & Trjapitzin, 1978: 793) (comb. n. from
Aphidencyrtus) (Vietnam), also one undescribed species from Hong Kong (BMNH).
BIOLOGY. Parasites of nymphs of Psyllidae (Homoptera).
COMMENTS. The genus is most probably best placed in the tribe Microteryini, subtribe
Syrphophagina (Encyrtinae) and can be separated from related genera (see comments under
Coccidoctonus} by the characters given in the key.
DIASULA gen. n.
(Key couplet: 343. Figs 158, 314, 315)
Type-species: Liothorax glabriscutellum Girault. Gender: feminine.
9 . Head. In frontal view slightly wider than long and in side view a little less than twice as long as broad and
more or less gradually and evenly curved anteriorly but a little more strongly so above top of antennal
scrobes. Eye almost naked, with sparse, very inconspicuous pale setae, each clearly shorter than the
diameter of a facet; posterior margin of eye very slightly concave, eye about one-third longer than broad
and reaching occipital margin which is sharply carinate, particularly behind ocelli. Malar space slightly
longer than one-third length of eye, with sulcus present; mouth opening relatively broad, about two-thirds
as wide as head. Frontovertex about one-third head width; ocelli more or less forming a right angle,
relatively large, the posterior ones very close to eye margin but separated from occipital margin by about
one and one-half times their own major diameters. Antennal scrobes fairly shallow and short, only about as
long as toruli, meeting dorsally, reaching to about one-third way from toruli to anterior ocellus; antennal
torulus separated from mouth margin by about its own length and from other torulus by slightly less than its
own length, its dorsal margin about half its length above ventral level of eyes; clypeus broadly but shallowly
concave, naked along mouth margin. Antennal scape clearly longer than width of frontovertex, subcylin-
drical, about five to six times as long as broad; pedicel conical, about two-fifths length of scape, subequal in
length or a little longer than any of the funicle segments, all of which are cylindrical and at least slightly
longer than broad; clava three-segmented, apically rounded with sutures parallel and about twice as long as
any funicle segment and not or hardly broader; longitudinal sensillae on all flagellar segments; longest
setae a little longer than diameter of segments. Frontovertex fairly smooth and polished behind ocelli;
between ocelli with very shallow, raised, reticulate sculpture, below this with very shallow, raised,
transverse, squamiform-reticulate sculpture, this becoming more longitudinally elongate between scrobes
and eyes and on lower parts of face; setae on head sparse, dark and not very conspicuous although each a
little longer than the diameter of an ocellus. Mandible broad with three acute teeth; maxillary palpus
four-segmented; labial palpus three-segmented.
Thorax. In side view fairly deep with mesoscutum only a little convex, but scutellum fairly strongly so;
metapleurum together with propodeum only narrowly in contact with hind coxa. Pronotum in dorsal view
264 J. S. NOYES & M. HAY AT
more or less triangular with its posterior margin slightly concave; visible part of mesoscutum about one and
one-half times as broad as long, with posterior margin slightly convex, notaular lines absent; axillae
meeting; scutellum very convex, clearly longer than broad with its apex narrow and rounded; propodeum
medially quite long, but not more than one-sixth as long as scutellum. Mesoscutum and scutellum with very
shallow, raised, squamiform-reticulate sculpture, sculpture of axillae similar but finer, anterior one-third
or so of scutellum with similar sculpture to mesoscutum but shallower, gradually becoming more shallow
posteriorly so that apical half of scutellum is almost completely smooth and polished; mesopleurum
smooth; propodeum smooth save for a very shallow incomplete carina medially; mesoscutum with a few
scattered, fairly long, dark setae; scutellum with about two dozen long conspicuous setae including two
pairs of long erect setae subapically. Forewing hyaline, about two and one-half times as long as broad; linea
calva neither interrupted nor closed; basal cell sparsely hairy; filum spinosum present; submarginal vein
with an apical hyaline break, parastigma not swollen; costal cell about 12 to 13 times as long as broad, with
only a few setae dorsally in its apical half or so; marginal vein about four to five times as long as broad,
about one and one-half times as long as stigmal which is subequal in length to postmarginal vein; venation
yellowish. Hindwing hyaline, about three-quarters as long as broad, with marginal fringe about one-
seventh as long as width of hindwing. Mid tibial spur about as long as basal mid tarsal segment.
Caster. Clearly longer than thorax and apically acute; cereal plates in basal one-third; ovipositor a little
exserted, exserted part less than one-tenth length of gaster; hypopygium reaching from about one-third to
nearly three-quarters along gaster; last tergite about as long as mid tibia.
Cf. Unknown.
COMMENTS. Diasula is possibly related to Helegonatopus (Encyrtinae, Chalceryini) since the
mandible has three acute teeth, the scutellum is convex and the wing venation is yellow.
However, it can be easily separated from this and related genera by the very sharp occipital
margin, long marginal vein and very shiny scutellum.
Diasula glabriscutellum (Girault) comb. n.
(Figs 158, 314, 315)
Liothorox glabriscutellum Girault, 1932a: 1. LECTOTYPE <j>, AUSTRALIA (QM), here designated
[examined].
$. Length: 2.08-2.22 mm.
Colour. Head and thorax metallic green with some purple reflections, particularly between ocelli and
occipital margin and on lower parts efface, sides of thorax orange-brown to dark purplish brown; antennal
pedicel and flagellum dark brown, scape, palpi and legs, excluding mid coxae, pale yellow, mid coxa dark
brown; wings hyaline, venation yellow; gaster towards base ventrally metallic green, remainder of dorsum
shining purple, ovipositor sheaths dark brown.
Head. Relative measurements (Australian specimen): head length 84, head width (frontal view) 89,
head width (side view) 51, minimum frontovertex width 31, POL 14, OOL 1-5, malar space 21, eye length
59, eye width 45, scape length 47, other proportions of antenna as in Fig. 315, mandible as in Fig. 314.
Thorax. Base of forewing as in Fig. 158. Relative measurements (Australian specimen): forewing length
278, forewing width 108, hindwing length 198, hindwing width 53.
Gaster. Hypopygium reaching about three-quarters along gaster. Relative lengths (Australian speci-
men): last tergite 110, [mid tibia 111].
Cf . Unknown
DISTRIBUTION. Australia, Philippines.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Lectotype $, Australia: Queensland, Nelson (A. P. Dodd).
Australia: 1 $, Queensland, 15 km SE. of Nambour, 6.xi.l976 (Z. Boucek} (compared with lectotype)
(BMNH). Philippines, 1 $ , Mt Montalban, Rizal Wa-Wa Dam, 150-200 m, 23.iii. 1965 (L. M. Torrevillas)
(BPBM).
COMMENTS. The single extant syntype of Liothorax glabriscutellum Girault (1932a: 1) in the
Queensland Museum has the body mounted on a card and labelled 'Liothorax glabri-
IN DO-PACIFIC ENCYRTIDAE 265
scutellum Gir. $ type'; the head and right forewing are on a separate slide labelled 'Liothorax
glabriscutellum Girault 9 Type'. It is here designated lectotype. The Australian specimen was
compared with the lectotype by one of us (JSN) during a visit to Brisbane in 1980.
Two other species are provisionally placed in this genus: Diasula semiargentipes (Girault,
19150: 105)(comb. n. from Parasyrpophagus) (Australia) and Diasula homeri (Girault, 1935: 3)
(comb. b. from Parasyrpophagus) (Australia), and should run here in the key.
DIVERSINERVUS Silverstri
(Key couplets: 87, 98)
Diversinervus Silvestri, 1915a: 301. Type-species: Diversinervus elegans Silvestri, by original designation.
Cheiloneuroides Girault, 1915a: 96. Type-species: Cheiloneuroides bicristatus Girault, by original des-
ignation.
DISTRIBUTION AND SPECIES. Eleven species, cosmopolitan; five from review area: cervantesi
(Girault, 1933: 4) (Australia. Malaysia, Samoa), elegans Silvestri (19150: 304) (India, Australia,
Fiji, Hawaiian Is.),intermedius Hayat, Alam & Agarwal (1975: 43) (India) , ma dgaoensis Hayat,
Alam & Agarwal (1975: 41) (India) and paradisicus (Motschulsky, 1863: 52) (Sri Lanka), also
undetermined material from New Caledonia (BPBM).
REFERENCE. Key to world species: Hayat et al. (1975: 39-41); Rosen & Alon (1983).
BIOLOGY. Parasites of Coccidae (Homoptera).
COMMENTS. The only species not included in the key by Hayat et al. is cervantesi but this has been
included by Rosen & Alon. It can be easily recognised since it is the only brachypterous species
known in the genus.
The genus is placed in the tribe Cheiloneurini (Encyrtinae) by Trjapitzin (19736), but the
forewing venation suggests a strong link with some genera of the Cerapterocerini, e.g. Anicetus.
DODDANUSIA gen. n.
(Key couplet: 131. Figs 66, 317-321)
Type-species: Anusia viridiflava Dodd. Gender: feminine
9- Head. In facial view nearly one-third broader than long, in side view a little less than twice as long as
broad and more or less evenly curved anteriorly except below top of antennal scrobes where it is almost
straight. Eye with fairly conspicuous translucent setae, each a little longer than the diameter of a facet,
posterior margin of eye straight, eye only very slightly longer than broad, reaching occipital margin which is
rounded, but not strongly so. Malar space about one-half to three-fifths length of eye, with sulcus absent.
Frontovertex less than one-quarter as wide as head; ocelli forming an acute angle of about 45-70, the
posterior ones clearly closer to eye margin than to occipital margin, separated from the latter by about their
own diameters. Antennal scrobes broadly semi-circular, meeting dorsally and more or less sharply
margined dorsally, reaching about one-third way from antennal toruli to anterior ocellus; antennal torulus
separated from mouth margin by slightly more than its own length and from other torulus by about one and
one-half times its own length, its dorsal margin clearly below the ventral level of the eyes; clypeus broadly
and shallowly emarginate. Antennal scape (Fig. 318) much longer than minimum width of frontovertex
and distinctly broadened and flattened, about twice as long as broad, pedicel conical, about one-quarter
length of scape and clearly longer than any of the funicle segments; funicle six-segmented, cylindrical,
clearly broadening distally; clava two- or three-segmented, with a strong oblique truncation, nearly as long
as funicle; longitudinal sensillae on fifth and sixth funicle segments and clava; longest setae clearly shorter
than diameter of first funicle segment, funicle segments and basal segment of clava with setae flattened and
scale-like. Frontovertex near ocelli with shallow to fairly deep polygonal reticulate sculpture, above
scrobes similar but transversely elongate or almost entirely smooth and shiny; between scrobes and eyes
moderately deep, polygonally reticulate sculpture, this becoming a little shallower and more longitudinally
elongate on genae; setae on frontovertex translucent or dark, about as long as the diameter of an ocellus.
266 J. S. NOYES & M. HAY AT
Mandible with one tooth and a broad truncation (Fig. 319); maxillary palpus four-segmented, labial palpus
three-segmented.
Thorax. In side view moderately robust but dorsally very flat; the mesopleurum enlarged and more or
less touching basal segment of gaster and thus clearly separating the hind coxa from the metapleurum and
propodeum. In dorsal view posterior margin of propodeum broadly and shallowly concave; visible part of
mesoscutum about twice as broad as long with notaular lines absent, its posterior margin almost straight;
axillae meeting; scutellum flat, triangular and slightly broader than long; propodeum medially a little less
than one-fifth length of scutellum. Dorsum of thorax with shallow, raised, squamiform-reticulate sculp-
ture, mesopleurum with raised reticulate sculpture of fine mesh medially and longer mesh posteriorly,
anteriorly rather more irregular and longitudinally elongate; propodeum with shallow, raised irregular
sculpture; setae on dorsum of thorax dark, sparse and of moderate length. Forewing generally suffused
pale brown and convex dorsally (as in Discodes), about twice as long as broad; linea calva not interrupted
but closed on dorsal surface by one or two lines of setae near posterior margin of wing; filum spinosum
present but in posterior half of wing (Fig. 66); submarginal vein with an indistinct apical hyaline break,
parastigma not swollen; costal cell about 10 or 11 times as long as broad, with one or two lines of setae
dorsally along its length; marginal vein about four to five times as long as broad, about three times as long as
postmarginal and a little longer than, to about same length as stigmal (Fig. 317); setae on dorsal surface of
wing fairly inconspicuous and short. Hindwing slightly longer than three-quarters length of fore wing,
about three times as long as broad, with marginal fringe about one-sixth as long as width of wing. Mid tibial
spur a little shorter than basal mid tarsal segment.
Gaster. A little shorter than thorax with hypopygium extending to about three-quarters along venter;
ovipositor not exserted; last tergite a little shorter than mid tibia; paratergites absent; ovipositor (Fig. 321)
about as long as mid tibia; gonostyli free, about one-sixth as long as ovipositor; hypopygium as in Fig. 320.
Cf . Unknown.
COMMENTS. This genus should probably be placed in the Microteryini (Encyrtinae), subtribe
Microteryina and can be separated easily from all other related genera by the strongly obliquely
truncate antennal clava, scale-like setae on the flagellum, and presence of a filum spinosum in
posterior half of forewing.
Doddanusia viridiffava (Dodd) comb. n.
Dodd's original (1924) description is probably sufficient to recognise this species. It can be
separated from a second species from the mainland of Australia: Queensland (BMNH) by
having the antennal scrobes partly metallic green, the frontovertex at its narrowest point about
one-fifth head width, malar space about one-half length of an eye, ocelli forming an angle of
about 45 and antennal scape more clearly triangular in shape. The species from the Australian
mainland has the scrobes non-metallic, frontovertex about one-quarter head width, ocelli in an
angle of about 65-70, malar space about three-fifths as long as an eye and antennal scape
subrectangular in shape (Fig. 318). Figs 66, 317-321 are of this second undescribed species.
DISTRIBUTION. Australia (Queensland and Norfolk Is.).
BIOLOGY. Unknown.
DOLIPHOCERAS Mercet
(Key couplets: 165, 174, 221, 277. Figs 97, 98, 175)
Doliphoceros Mercet, 1921: 91. Type-species: Pholidoceros integralis Mercet, by original designation.
Rhopomorphus Ghesquiere, 1958: 25. Type-species: Rhopomorphus varleyellus Ghesquiere, by original
designation.
DISTRIBUTION AND SPECIES. Twelve species, Palaearctic, Afrotropical, Oriental, Australasian
and Pacific; five from review area: fraternus (Perkins, 1910: 653)(comb. n. from Anagyrus)
(Hawaiian Is.), gracilis Hayat (19700: 114) (India), nigricans (Perkins, 1910: 653) (Hawaiian
Is.), punctifrons (Timberlake, 1941: 219) (comb. n. from Anagyrus) (Marquesas Is.) and
INDO-PACIFIC ENCYRTIDAE 267
tantaleus (Perkins, 1910: 654) (comb. n. from Anagyrus} (Hawaiian Is.), also undetermined
material from Nepal, Hong Kong, Fiji and Australia (BMNH, BPBM).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The genus is placed in the subtribe Rhopina of the tribe Anagyrini (Tetracenemi-
nae) by Trjapitzin (1973a), which is, without doubt, incorrect. Doliphoceras is extremely close
to Anagyrus (subtribe Anagyrina) and very probably should be considered synonymous, but for
the present we are maintaining the two genera as distinct (see comments under Anagyrus).
ECHTHROBACCELLA Girault
(Key couplet: 241)
Echthrobaccella Girault, 19150: 113. Type-species: Echthrobaccella argentinotata Girault, by original
designation.
DISTRIBUTION AND SPECIES. One species, Australia only: argentinotata Girault (19150: 113).
BIOLOGY. Unknown.
COMMENTS. Related to Cheiloneurus (Encyrtinae, tribe Cheiloneurini) from which it can be
separated by the apparent presence of notaular lines in the extreme anterior part of the
mesoscutum, dorsum of thorax with fine punctate sculpture of silky appearance and forewing
having dense setae throughout basal cell, submarginal vein not bent downwards subapically, a
characteristic infuscate pattern, and the distinctive coloration of the thorax (see Girault, 1915:
113).
ECHTHROGONA TOPUS Perkins
(Key couplet: 349, 455, 470, 527. Figs 192, 223, 224)
Echthrogonatopus Perkins, 1906: 256. Type-species: Echthrogonatopus exitiosus Perkins, by designation
of Gahan & Pagan (1923: 48).
DISTRIBUTION AND SPECIES. Four species, Nearctic, Afrotropical, Oriental, Australasian; three
species from review area: exitiosus Perkins (1906: 256) (Malaysia, Philippines, Fiji, Australia),
nigricornis (Hayat, 1980: 644) (India) andparvus (Hayat, 1980: 643) (India), also undetermined
material, including at least one undescribed species, from Bangladesh, Hong Kong, Philippines
and Samoa (BMNH, BPBM, GC).
BIOLOGY. Hyperparasites of leafhoppers (Homoptera, Auchenorrhyncha) via Dryinidae
(Hymenoptera).
COMMENTS. The genus is best placed in the tribe Cheiloneurini (Encyrtinae) and can be
separated from its nearest relatives, Zaomma, by the lack of an apical scutellar brush (see also
Hayat, 1980: 642-643), and Hypergonatopus by the flat dull scutellum and hyaline forewings
(the scutellum of Hypergonatopus is convex and at least a little shiny and the forewings are
darkened).
ECTOPIOGNATHA Perkins
(Key couplet: 187. Figs 112, 322, 323)
Ectopiognatha Perkins, 1906:254. Type-species: Ectopiognatha minor Perkins, by designation of Gahan &
Pagan (1923: 49).
DISTRIBUTION AND SPECIES. Two species, both Australian and probably synonymous: major
Perkins (1906: 255) and minor Perkins (1906: 255), also one undetermined specimen, lacking
antennae, from Irian Jaya (BPBM).
BIOLOGY. Parasites of eggs of Flatidae and Eurybrachidae (Homoptera).
268 J. S. NO YES & M. HAYAT
COMMENTS. Placement of the genus according to Trjapitzin's (1973) classification is difficult, but
it most probably belongs in the Microteryini.
ECTROMA Westwood
(Key couplet: 93)
Ectroma Westwood, 1833a: 344. Type-species: Ectroma fulvescens Westwood, by monotypy.
Metallon Walker, 1848: 219. Type-species: Metallon acacallis Walker, by monotypy.
Pezobius Forster, 1860: 129. Type-species: Pezobius polychromus Forster, by monotypy.
DISTRIBUTION AND SPECIES. Twelve described species, Neotropical, Palaearctic, Afrotropical;
none found in review area but one undescribed species from India (BMNH).
BIOLOGY. Unknown.
COMMENTS. Dalla Torre (1898: 238) lists Ectroma dunense Six (1876) as originating from Batavia
in Asia (Indonesia), but it is not included here since this species was described from the Batavia
peninsula in the Netherlands.
The mandibles of Ectroma are tridentate, although they were erroneously stated to be
bidentate by Noyes (1980: 114).
The genus has been placed in the tribe Miraini, subtribe Mayridiina (Encyrtinae) by
Trjapitzin (19736). This is obviously incorrect since Mira (and thus the tribe Miraini) belongs in
the Tetracneminae whereas Ectroma and its relatives belong in the Encyrtinae. We feel sure that
Ectroma can be accommodated in the Cheiloneurini, and is probably close to Cheiloneurus . It
can be difficult to separate from brachypterous forms of Cheiloneurus, particularly if the latter
lacks the usual subapical, scutellar brush (see key).
ENCYRTOIDEA Girault
(Key couplets: 248, 331, 465, 491, 512)
Encyrtoidea Girault, 1923c: 146. Type-species: Encyrtoidea punctatifrons Girault, by monotypy.
DISTRIBUTION AND SPECIES. Two species, Australia only: compressifemur (Girault, \923e: 5)
(comb. n. from Nezarhopalus) and punctatifrons Girault (1923c: 146), also two further species
from Australia (BMNH).
BIOLOGY. Unknown.
COMMENTS. Encyrtoidea compressifemur may be incorrectly placed in this genus since the
mandibles are very different from those of punctatifrons . The latter species has all three teeth
very nearly equal in size, whereas in compressifemur the lowest tooth is very much larger than
either of the two upper teeth.
The genus is very difficult to place according to Trjapitzin's (19736) classification of the
Encyrtinae. The venation and mandibles suggest an affinity with Borrowella (which is here
provisionally placed in the Bothriothoracini), but in general appearance it is not unsimilar to
some genera of the Microteryini e.g. Ooencyrtus and Syrphophagus.
ENCYRTUS Latreitte
(Key couplets: 99, 373)
Encyrtus Latreille , 1809: 31. Type-species: Chrysis infida Rossi, by designation of Latreille (1810: 436).
Eucomys Forster, 1856: 32. Type-species: Encyrtus swederiDalman, by original designation.
Comys Forster, 1856: 144. [Unnecessary replacement name for Eucomys Forster.]
Howardia Dalla Torre, 1897: 86. Type-species: Bothriothorax peckhami Ashmead, by original desig-
nation. [Homonym of Howardia Berlese & Leonardi, 1896.]
Howardiella Dalla Torre, 1898: 228. [Replacement name for Howardia Dalla Torre.]
INDO-PACIFIC ENCYRTIDAE 269
Allorhopoideus Brethes, 1916: 425. Type-species: Allorhopoideus mirabilis Brethes, by original des-
ignation.
Prorhopoideus Brethes, 1921: 80. Type-species: Prorhopoideus baezi Brethes, by original designation.
DISTRIBUTION AND SPECIES. About 90 species currently in the genus Encyrtus but only about 40
are correctly placed, cosmopolitan; six species from review area: albidus Hayat (19706: 61)
(India), argenticoxa (Girault, 1915: 129) (comb. n. from Eucomys) (= Eucomys hibisci
Girault, 1915a: 128 syn. n., = Eucomys aurantifasciata Girault, 19150: 129 syn. n., = Eucomys
argentiscapus Girault, 1915: 130 syn. n.) (Australia), infelix (Embleton, 1902: 223) (Fiji, New
Zealand, Hawaiian Is.), lecaniomm (Mayr, 1876: 740) (India, Philippines, New Zealand,
Hawaiian Is.), proserpinensis (Girault, 1915a: 130) (comb. n. from Eucomys) ( Eucomys
hortensis Girault, 1915a: 130 syn. n.) (Australia) and saissetiae (Yasumatsu & Yoshimura, 1945:
33) (comb. n. from Eucomys) (Mariana Is.), also much undetermined material from throughout
the region (BMNH, BPBM, DSIR, CNC, USNM).
REFERENCE. Review of Holarctic species: Sugonjaev & Gordh (1981); key to some species:
Hayat (1970b: 59).
BIOLOGY. Parasites of Coccidae (Homoptera).
COMMENTS. We have not seen the holotype of E. corvinus Motschulsky (1863: 55), but according
to Boucek (pers. comm.) it belongs to the subfamily Telenominae (Proctotrupoidea,
Scelionidae).
Placed in the tribe Encyrtini as the sole included genus (Trjapitzin, 19736). We feel that
Trjapitzin's definition of this tribe is probably too narrow and that it should also include those
genera of the tribes Eugahaniini, Prionomasticini, Neocladiini and Aethognathini. However,
further more detailed study of this group is desirable before this tribal synonymy can be
proposed formally.
EOTOPUSgen. n.
(Key couplet: 263. Figs 150, 325-327)
Type-species: Ericydnus beneficus Shafee. Gender: masculine.
$ . Head. In facial view clearly broader than long and in profile about two-thirds longer than broad and
anteriorly more or less gradually and evenly curved. Eye with posterior margin slightly concave, about
one-half longer than broad, with dense fairly conspicuous translucent setae and clearly overreaching
occipital margin which is more or less rounded. Malar space about one-quarter eye length, with malar
sulcus present. Frontovertex about one-third head width; ocelli in a very slightly acute angle, posterior
ocellus separated from occipital margin by about twice its diameter and from eye margin by about its own
diameter. Antennal scrobes shallow, meeting dorsally and clearly reaching more than halfway to anterior
ocellus from antennal toruli; antennal torulus separated from mouth margin by a little less than its own
diameter and separated from other torulus by nearly one and one-half times its length, its middle being
about level with lower eye margins; clypeal margin broadly excised between toruli. Antennal scape much
longer than minimum width of frontovertex, cylindrical, slightly wider near base, about five or six times as
long as broad, pedicel conical, at least slightly longer than any funicle segment, all of which are longer than
wide and slightly widening distally; clava three-segmented, about half as long as funicle, with apex more or
less rounded, its sutures almost parallel; longitudinal sensillae on all but the first two flagellar segments.
Frontovertex with squamiform-reticulate sculpture, deepest in front of anterior ocellus, fairly shallow
behind ocelli and towards lower parts efface where it becomes more longitudinally elongate; frontovertex
clothed in sparse, rather inconspicuous, moderately long translucent setae. Mandible tridentate, the upper
tooth short and blunt; maxillary palpus three-segmented, labial palpus two-segmented.
Thorax. In side view moderately deep with metapleurum distinct, clearly broadening ventrad and,
together with propodeum, broadly in contact with hind coxa and dorsally with both mesoscutum and
scutellum flat. In dorsal view posterior margin of pronotum strongly concave; visible part of mesoscutum
about one-half broader than long with notaular lines present anteriorly (the area around each notaular line
slightly concave), posterior margin of mesoscutum almost straight; axillae meeting medially; scutellum
nearly one and one-half times as long as broad and about one-third longer than mesoscutum, with its apex
blunt; propodeum medially a little less than one-third length of scutellum. Mesoscutum with moderately
270 J. S. NOYES & M. HAY AT
deep, raised, squamiform-reticulate or reticulate sculpture, that on scutellum similar but distinctly more
longitudinally elongate; propodeum medially almost smooth, but with some shallow reticulate sculpture;
dorsum of thorax with moderately dense short, recumbent, fairly inconspicuous, translucent setae.
Forewing hyaline, wing about three times as long as broad; linea calva not interrupted and nearly closed
near posterior margin of wing; filum spinosum absent; venation yellowish, submarginal vein with an
inconspicuous apical hyaline break and with parastigma clearly swollen, much broader than proximal
two-thirds of submarginal vein ; marginal vein about five or six times as long as broad and a little longer than
either postmarginal or stigmal veins which are subequal in length; costal cell about 10-11 times as long as
broad, with a single line of setae dorsally in distal half. Hindwing about three-quarters as long as forewing,
about six times as long as broad, with marginal fringe about one-third maximum wing width. Mid tibial spur
shorter than basal mid tarsal segment.
Caster. A little shorter than thorax, cereal plates at about midway along its length; hypopygium reaching
apex of gaster; paratergites not distinct in slide-mounted material available; last tergite a little longer than
half length of mid tibia; gonostyli fused to second valvifers and about one-sixth as long as ovipositor which
is a little more than half as long as mid tibia.
O". Differs from female as follows. Eye with posterior margin convex, with setae very sparse and
inconspicuous, clearly separated from occipital margin by nearly diameter of posterior ocellus; eye smaller
so that malar space is nearly half length of eye and frontovertex clearly more than half head width; ocelli
forming an obtuse angle, with posterior ocellus slightly closer to occipital margin than to eye margin, being
separated from the latter by a little more than its own diameter; antennal scrobes more broadly
semicircular, meeting dorsally and separated from anterior ocellus by not more than its diameter, antennal
torulus separated from mouth margin by slightly more than its own length and from other torulus by about
its own length, its lower margin only very slightly below ventral margins of eyes; antenna with scape about
as long as width of frontovertex and about four times as long as broad, broadest near base, pedicel conical
and subquadrate, much less than half length of any of funicle segments which are cylindrical and at least
about three times as long as broad, setae on flagellum about four times as long as diameter of segments,
clava entire, longer than any funicle segment, longitudinal sensillae on all flagellar segments; sculpture of
head clearly shallower than in female, the frontovertex distinctly more shiny. Thorax in profile with
mesoscutum slightly concave, in dorsal view scutellum about same length as mesoscutum; propodeum
medially about one-fifth length of scutellum; forewing with postmarginal vein a little longer than either
marginal or stigmal veins which are subequal in length; costal cell a little narrower than in female; hindwing
about two-thirds length of forewing, with marginal fringe about one-half maximum wing width; mid tibial
spur nearly as long as basal mid tarsal segment. Gaster about as long as thorax; genitalia with digiti long
with apical hooks, nearly half length of aedeagus, aedeagus about one-half length of mid tibia or twice
length of mid tibial spur.
COMMENTS. This genus belongs to the Charitopidini (Tetracneminae). The general body
structure suggests a close affinity with Charitopus from which it can be separated in the female by
the colour and structure of the antenna and incomplete notaular lines (complete in Charitopus);
the male can be separated by the unbranched antenna and incomplete notaular lines.
Eotopus beneficus (Shafee) comb. n.
(Figs 150, 325-327)
It is quite clear from Shafee's (1981) original description that his material had been in alcohol
and therefore parts of his description need augmenting or correcting as follows.
$. Length: 0.92-1. 59 mm.
Colour. Head dark metallic green with some coppery or purple reflections, particularly on frontovertex,
antenna from yellowish to testaceous yellow; pronotum and mesoscutum varying from almost entirely
orange with a darker longitudinal metallic green stripe to almost entirely metallic green with orange area at
extreme sides outside notaular lines, axillae from entirely orange through metallic green to deep metallic
purple, scutellum metallic green to metallic green mixed purple and blue; sides of thorax from almost
entirely orange to almost entirely dark brown (especially the mesopleurum); propodeum medially dark
brown, laterally dark brown mixed to a lesser or greater extent with orange; wings hyaline but very lightly
stained yellow; legs, including coxae, pale orange-yellow; gaster mostly orange with tergites laterally
strongly metallic purple.
INDO-PACIFIC ENCYRTIDAE 271
Head. As in original description except that posterior ocellus is separated from occipital margin by at
least slightly more than and usually nearly twice its own diameter. Relative measurements of head: length
60, width (facial view) 70, width (side view) 38, minimum frontovertex width 24, malar space 12, eye length
48, eye width 33, POL 12, OOL 4, scape length 40, scape width 7, antenna as in Fig. 325. There is a little
variation in the relative width of the frontovertex, but it is usually about one-third maximum head width;
Shafee states that the scape is 'slightly more than four times as long as wide', but in no specimens has it been
found to be as broad, usually about five and one-half times as long as broad; there is also some variation in
the relative proportions of the flagellar segments, the distal segments sometimes at least one-half longer
than broad whereas usually they are only about one-quarter longer than broad; there is also some variation
in the size of the ocelli so that POL and OOL may be a little different from that given above.
Thorax. Forewing very nearly three times as long as broad, not two and one-half times as given by
Shafee. There is some variation in the relative length of the postmarginal vein which is normally about as
long as the stigmal, but in some specimens it is clearly a little shorter, both types occurring occasionally in a
single specimen. Relative measurements: forewing length 165, width 56, hindwing length 119. Forewing
base as in Fig. 150.
Caster. Relative lengths: last tergite 45, ovipositor 52, gonostyli approx. 7, [mid tibia 86]. Genitalia as in
Fig. 326.
Cf . Differs from female as follows. Colour. Very much as female except that lower parts of face often
coloured orange, gaster more or less entirely orange but with apical one-half to one-third dorsally brown;
ocelli forming a distinctly obtuse angle; for antenna see Shafee (1981: fig. j); relative measurements: head
length 40, head width (facial view) 14, minimum width of frontovertex 28, malar space 12, eye length 23,
eye width 17, POL 12, OOL 6, scape length 19. Genitalia as in Fig. 327; relative lengths: aedeagus 34, mid
tibial spur 16. There is some variation in the relative width of the frontovertex so that in some specimens it
is a little broader than length of scape, also POL may be a little less than twice OOL depending on the
relative size of the ocelli and width of frontovertex.
DISTRIBUTION. India.
BIOLOGY. Reared from Icerya pilosa Green (Homoptera, Margarodidae) on Saccharum offici-
narum Linnaeus (Shafee, 1981).
MATERIAL EXAMINED
India: 1 9> determined as Ericydnus beneficus Shafee and probably a paratype but no data or
determination labels;! $, Uttar Pradesh, Aligarh, on grass, 23. ii. 1979 (M. Hayat&M. Verma);! $,Uttar
Pradesh, Aligarh, 20.viii.1979 (M. Verma); 1 $, Tamil Nadu, Shembaganur, x.1979 (J. S. Noyes); 2 $, 7
Cf, Kerala, Periyar Animal Sanctuary, 5-15. x.1979 (/. S. Noyes); 3 <j>, 7 cf , Tamil Nadu, 3 km E. of
Manjaler Dam, 5-18. x. 1979 (/. 5. Noyes), 1 cf, Tamil Nadu, Anamalai Animal Sanctuary, 21.x. 1979(7. S.
Noyes); 9 $, 6 cf, Tamil Nadu, Mudumalai Animal Sanctuary, 23-24.X.1979 (/. S. Noyes); 26 $, 8 cf,
Karnataka, Mudigere, 26.x-4.xi. 1979 (/. S. Noyes); 13 $, 6 cf, Karnataka, 25 km W. of Mudigere,
28.x-3.xi. 1979 (/. 5. Noyes); 1 cf , Karnataka, Bannerghatta N. P., 5. xi. 1979 (Z. Boucek&J. S. Noyes); 1
Cf, Kerala, Calicut University Area, xi.1979 (Z. Boucek); 1 cf, Hyderabad, Patancheru, ICRISAT,
vii-ix.1980, Malaise trap (Bernays & Woodhead) (BM, HC, USNM, UCR, ZI, PPRI).
EPANUSIA Girault
(Key couplet: 107)
Epanusia Girault, 1915cr: 154. Type-species: Epanusia bifasciatus Girault, by original designation.
DISTRIBUTION AND SPECIES. Australia only; two species: beenleighi Girault (1923e: 5) and
bifasciata Girault (19150: 154).
BIOLOGY. Unknown.
COMMENTS. The two included species are very close but are distinct. They can be separated on
the shape of the scape, but more easily by the extent of the infuscate areas of the forewing; the
forewing of beenleighi is infuscate to its base, whilst that of bifasciata is largely hyaline in the
basal cell.
The genus belongs to the same group as Cryptanusia (see comments under Cryptanusia) .
272 J. S. NOYES & M. HAY AT
EPIBLATTICIDA Girault
(Key couplets: 197, 367, 506)
Epiblatticida Girault, 1915o: 117. Type-species: Epiblatticida Iambi Girault, by original designation.
Neasteropaeus Girault, 19150: 109. Type-species: Neasteropaeus caudatus Girault, by original designation.
Syn. n.
Blatticidella Girault, x.!923c: 144. Type-species: Blatticidella aereitibiae Girault, by monotypy.
[Homonym of Blatticidella Gahan & Pagan, iv. 1923.] Syn. n.
Microencyrtus Girault, 1923c: 147. Type-species: Microencyrtus minutissimus Girault, by monotypy.
Syn. n.
Magellanana Girault, 19396: 324. [Replacement name for Blatticidella Girault.] Syn. n.
DISTRIBUTION AND SPECIES. Australia, New Caledonia and New Zealand only; five described
species: aereitibiae (Girault, 1923c: 144) (comb. n. from Blatticidella) (Australia), argentipes
(Girault, 19256: 99) (comb. n. from Epitetracnemus) (Australia) , caudatus (Girault, 19150: 109)
(comb. n. from Neasteropaeus) (Australia), Iambi Girault (19150: 117) (Australia) and minutis-
simus (Girault, 1923c: 147) (comb. n. from Microencyrtus), also undetermined material from
New Caledonia and New Zealand (BMNH, BPBM, DSIR).
BIOLOGY. Hyperparasites of Psyllidae (Homoptera) via other Encyrtidae.
COMMENTS. Closely related to Coccidoctonus (see comments, p. 254).
EPIDINOCARSIS Girault
(Key couplets: 174, 221. Fig. 94)
Epidinocarsis Girault, 19136: 83. Type-species: Epidinocarsis tricolor Girault, by original designation.
Apoanagyrus Compere, 1947a: 18. Type-species: Apoanagyrus calif ornicus Compere, by original desig-
nation. Syn. n.
DISTRIBUTION AND SPECIES. Fifteen species, New World, Palaearctic, Oriental, Australasian and
Pacific; seven species from review area: anamalaianus (Mani & Kaul in Mani etal., 1974: 63)
(comb. n. from Anagyrus) (India), auratiscutum Girault (19150: 144) (Australia), californicus
(Compere, 19470: 18) (comb. n. from Apoanagyrus) (Hawaiian Is.), cuneinota Girault (19150:
144) (Australia), marquesanus (Timberlake, 1941: 220) (comb. n. from Anagyrus) (Marquesas
Is.), rotundiceps (Girault, 19320: 3) (comb. n. from Dinocarsis) (Australia) and tricolor Girault
(19136: 83) (Australia).
REFERENCE. Partial revision: Kerrich (1982: 407-416).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Two of the above species may be misplaced here, but we have placed them in this
genus in an attempt to achieve a degree of consistency wrthin the key. One is marquesanus which
shows an affinity with punctifrons (Timberlake) which we place in Doliphoceras only because of
a difference in the sculpture of the scutellum (a character used to separate these two genera).
The other is rotundiceps which could also be placed in Anagyrus because the sculpture of the
head and mesoscutum is somewhat intermediate between the two genera (see comments under
Anagyrus).
We deliberated for some time before synonymising these two genera, but since we were
unable to find any real difference between tricolor and californicus (the respective type-species)
except colour we decided to do so here. Although Apoanagyrus is a fairly well-known name we
do not think that it is necessary to submit an application to the International Commission on
Zoological Nomanclature to ask for suppression of Epidinocarsis in favour of Apoanagyrus . Our
reasons for this are that we do not think the use of Epidinocarsis will lead to undue confusion in
the literature and also that a detailed study of this group of genera on a world-wide basis may
result in the synonymy of Epidinocarsis (and thus Apoanagyrus) and Doliphoceras with
Anagyrus (see also comments under Anagyrus).
INDO-PACIFIC ENCYRTIDAE 273
As a result of this new generic synonymy we also propose the following transfers of
extra-limital species from Apoanagyrus to Epidinocarsis: bermudensis Kerrich, diversicornis
Howard, elgeri Kerrich, gaudens Kerrich, lopezi De Santis, malenotus De Santis, montivagus
De Santis and trinidadensis Kerrich (all comb. n.).
EPISTENOTERYS Girault
(Key couplets: 364, 390)
Epistenoterys Girault, 1915a: 149. Type-species: Epistenoterys marmoratipes Girault, by monotypy.
Gounodia Girault, 1940: 149. Type-species: Gounodia mellea Girault, by monotypy. Syn. n.
DISTRIBUTION AND SPECIES. Two species, Australia only: marmoratipes Girault (19150: 149) and
mellea (Girault, 1940: 149) (comb. n. fom Gounodia}, also undetermined material from
Australia containing at least one further species (BMNH).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The above two species appear to differ only in coloration and minor morphological
characters which we regard as specific and not generic differences. Perhaps the most significant
difference between the two is the length of the malar space relative to the eye. In mellea it is only
a little shorter than the eye whereas in marmoratipes it is a little less than two-thirds the length of
an eye.
The genus is related to Aphycus, Cirrhencyrtus Timberlake and possibly Pseudaphycus (tribe
Aphycini, subtribe Aphycina).
EPITETRACNEMUS Girault
(Key couplets: 140, 488. Fig. 72)
Epitetracnemus Girault, 1915a: 164. Type-species: Epitetracnemus sexguttatipennis Girault, by original
designation.
Anabrolepis Timberlake, 1920: 431. Type-species: Anabrolepis extranea Timberlake, by original desig-
nation. Syn. n.
DISTRIBUTION AND SPECIES. Five species, cosmopolitan; three from review area: extraneus
(Timberlake, 1920: 434) (comb. n. from Anabrolepis) (Hawaiian Is.), sexguttatipennis (Girault,
19150: 164) (Australia) and zetterstedti (Westwood; Mercet, 1921: 678) (comb. n. from
Encyrtus) (New Zealand), also at least one further species amongst material from India, S.
China, New Caledonia and Australia (BMNH, BPBM).
REFERENCE. Review of most species: Tachikawa (1955).
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. The following extra-limital species are also transferred from Anabrolepis to
Epitetracnemus: japonicus Ishii and lindingaspidis Tachikawa (both comb. n.).
This genus belongs to the tribe Habrolepidini, subtribe Habrolepidina (Encyrtinae) and is
closely related to Adelencyrtus (see comments under Adelencyrtus) .
EPITETRALOPHIDEA Girault
(Key couplets: 194, 519. Fig. 115)
Epitetralophidea Girault, 1915a: 176. Type-species: Epitetralophidea bicinctipes Girault, by original
designation.
Ectromomyiella Girault, 1915a: 160. Type-species: Ectromomyiella articulus Girault, by original desig-
nation. Syn. n.
DISTRIBUTION AND SPECIES. Three species, all Australian: articulus (Girault, 19150: 160)
(comb. n. from Ectromomyiella}, bicinctipes Girault (19150: 176) (= Epitetralophidea bicinc-
274 J. S. NOYES & M. HAYAT
tipes emersoni Girault, 1923c: 142 syn. n.) and magnithorax (Girault, 123c: 146) (comb. n. from
Ooencyrtus) .
BIOLOGY. Unknown.
COMMENTS. Girault unfortunately described articulus from a single male. However, amongst
material on the same slide as the type of Casca nigra Girault, Ablerus speciosus Girault,
Perissoptems inexplicabilis Girault and Ooencyrtus magnithorax Girault, are some males which
appear to be identical to the holotype of articulus. It would seem reasonable to assume that all
this material may have been reared from the same host and put on one slide (under two separate
coverslips) . Therefore it is likely that the encyrtid males under one coverslip are the same species
as the encyrtid females, Ooencyrtus magnithorax, under the other. Since magnithorax is here
considered to be congeneric with bicinctipes we have no hesitation in regarding Ectromomyiella
as a synonym of Epitetralophidea.
Epitetralophidea appears to be very close to Coccidencyrtus (tribe Habrolepidini) from which
it can be separated by the uninterrupted linea calva and the two-segmented funicle in the male.
The latter suggests that it may also be closely related to Adelencyrtus from which it differs in
having the mandible with a single tooth and a broad truncation, that of Adelencyrtus having four
teeth or occasionally two teeth and a truncation.
ERENCYRTUS Mahdihassan
(Key couplets: 178, 394. Fig. 103)
Erencyrtus Mahdihassan, 1923: 71. Type-species: Erencyrtus dewitzi Mahdihassan, by monotypy.
DISTRIBUTION AND SPECIES. Four species, Afrotropical, Oriental, Australasian; two from review
area: dewitzi Mahdihassan; Ferriere (1935: 396) (India, Pakistan) and keatsi (Girault, 19390: 21)
(comb. n. from Mesastymachus) (Australia).
REFERENCES. Annecke & Mynhardt (1970a), Prinsloo & Mynhardt (1982: 38-41).
BIOLOGY. Parasites of lac insects (Homoptera, Keriidae).
COMMENTS. The genus is easily recogniseable in that the antenna of the male has a very short
two-segmented funicle and the clava is extremely long (more than twice as long as the scape and
pedicel together) and unsegmented.
Placed in the tribe Microteryini, subtribe Microteryina by Trjapitzin (19736).
ERICYDNUS Walker
(Key couplet: 211)
Ericydnus Walker, 1837: 363. Type-species: Ericydnus paludatus Walker, by designation of Westwood
(1840: 71).
Grandoriella Domenichini, 1951: 171. Type-species: Grandoriella lamasi Domenichini, by original
designation.
DISTRIBUTION AND SPECIES. Thirteen species, cosmopolitan; none known from review area, but
undescribed species examined from India, New Guinea and Australia (BMNH, BPBM).
REFERENCE. World revision: Kerrich (1967: 167-180).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Two other species previously placed in Ericydnus, i.e. chrysos (Walker, 1839: 34)
and megalarus (Walker, 18386: 477), do not belong in the Encyrtidae (see Kerrich, 1967: 179).
The species from the Australasian and Oriental regions differ from the known species of
Ericydnus in that the antennal toruli are placed relatively higher on the head with their ventral
margins being nearly level with the ventral margins of the eyes. They also have a sharper
occipital margin and the relative length of the gaster is less. We do not consider that these
INDO-PACIFIC ENCYRTIDAE 275
differences warrant separate generic status, but if future workers regard these species as
belonging to a separate genus they should perhaps consider using Grandoriella as a generic name
for this group.
The genus is placed in the tribe Ericydnini (Tetracneminae).
ETHORISgen.n.
(Key couplet: 315. Figs 156, 187, 328-331)
Type-species: Ethoris dahmsisp. n. Gender: feminine.
$ . Head. In frontal view about as long as broad, in profile less than twice as long as broad and more or less
gradually and evenly rounded anteriorly, but more strongly so at top of antennal scrobes. Eyes with
posterior margin almost straight but very slightly concave, about one-third longer than broad, clothed in
fairly dense translucent setae each a little longer than the diameter of a facet; eye reaching occipital margin
which is more or less sharp but not strongly so. Malar space about half as long as eye with sulcus present.
Frontovertex about half head width or a little less; ocelli nearly forming an equilateral triangle, the
posterior ones about equidistant from occipital and eye margins or a little nearer the latter, separated from
eyes by slightly less than to much less than their own diameters. Antennal scrobes shallow, separated
dorsally by interantennal prominence which is confluent with frontovertex, fairly sharp at this point and
extends almost all the way to mouth margin, scrobes very short, only reaching about one-quarter way from
toruli to anterior ocellus; antennal torulus separated from mouth margin by nearly twice its length and from
other torulus by about its own length, its lower margin only a little below the lower margins of eyes; clypeus
with margin straight or very slightly produced medially. Antennal scape subcylindrical, about five or six
times as long as broad and clearly longer than minimum width of frontovertex; pedicel conical, about
one-third length of scape and subequal in length to any of the funicle segments all of which are clearly
longer than broad and are cylindrical; clava about two-fifths as long as and not wider than funicle,
three-segmented, apically rounded and almost pointed with sutures parallel. Frontovertex with shallow,
raised, reticulate sculpture, becoming more irregular and elongate at top of scrobes and between scrobes
and eyes and on genae ; setae on frontovertex sparse , about as long as diameter of an ocellus . Mandible with
three acute teeth; maxillary palpus relatively long, four-segmented, apical segment nearly one and
one-half times as long as mandible and a little shorter than its apical seta; labial palpus three-segmented.
Thorax. In side view moderately deep, mesoscutum slightly convex, scutellum clearly more convex than
mesoscutum; metapleurum and propodeum very narrowly in contact with hind coxa. In dorsal view
pronotum with hind margin moderately concave; visible part of mesoscutum about twice as broad as long,
notaular lines absent, hind margin more or less straight, only slightly convex; axillae meeting; scutellum
about as long as mesoscutum, about as broad as long and apically rounded; propodeum medially about
one-fifth length of scutellum. Mesoscutum with shallow, raised, squamiform-reticulate sculpture, axillae
similar but a little finer and deeper; scutellum with conspicuously deeper, reticulate sculpture, regular
medially but more elongate towards sides, extreme apex nd sides smooth; mesopleurum almost smooth,
but with some irregular, very shallow sculpture; propodeum medially smooth; dorsum of thorax with fairly
numerous, moderately long, pale brown, inconspicuous setae. Forewing hyaline, about two and one-half
times as long as broad; linea calva not interrupted or closed; filum spinosum present; submarginal vein with
an apical hyaline break; costal cell about 10 or 11 times as long as broad; marginal vein about five or six
times as long as broad, slightly shorter than to one and one-half times as long as stigmal and clearly shorter
than postmarginal vein; postmarginal and stigmal veins forming an unusually acute angle. Hindwing a little
less than two-thirds as long as forewing, about four and one-half times as long as broad, with marginal
fringe about one-quarter as long as maximum width of wing. Mid tibial spur about as long as basal mid
tarsal segment.
Caster. About as long as thorax; cereal plates in basal half; hypopygium with apex about two-thirds along
gaster; last tergite about two-thirds as long as mid tibia; ovipositor very slightly exserted, a little shorter
than mid tibia; gonostyli free, about one-quarter as long as ovipositor.
O" . Unknown.
COMMENTS. We are unable to place the genus according to Trjapitzin's (19736) classification of
the Encyrtinae. It may be close to either Ageniaspis (Copidosomatini, Ageniaspidina) or less
probably to Rhytidothorax. The relatively high position of the antennal toruli and the very long
terminal segments of the maxillary palpus should separate it from either of these genera; the
276 J. S. NO YES & M. HAYAT
conformation of the antenna, the sculpture and coloration from Ageniaspis and the less
prominent hypopygium and short propodeum from Rhytidothorax .
The type-species is named in honour of Mr E. C. Dahms (QM).
Ethoris dahmsisp. n.
(Figs 156, 187, 328-331)
$. Length: 1-05-1 -14 mm (holotype, 1-05 mm).
Colour. Head dark metallic green with slight coppery sheen between anterior ocellus and antennal
scrobes; scape yellowish, pedicel, funicle and basal segment of clava dark brown, apical two segments of
clava white; face of pronotum medially, anterior margin of mesoscutum medially dark brown; scutellum
except sides dark metallic green, remainder of thorax, including legs, pale orange with metanotum and
dorsum of propodeum darker brownish orange; gaster dorsally brown, ventrally pale orange.
Head. Relative measurements (holotype): head length 55, head width (facial view) 58, head width (side
view) 33, minimum frontovertex width 24, malar space 19, eye length 37, eye width 28, POL 9, OOL 4,
diameter of anterior ocellus 5, scape length 33, scape width 6, proportions of antennal segments as in Fig.
328, head in facial view as in Fig. 331, mandible as in Fig. 329.
Thorax. Relative measurements (holotype): forewing length 155, width 61; hindwing length 102, width
22-5; base of forewing as in Fig. 156, venation as in Fig. 187. The angle between the stigmal and
postmarginal veins is a little variable and may be slightly greater than in Fig. 187.
Gaster. Relative lengths (paratype): last tergite 63, ovipositor 85, gonostyli 23, [mid tibia 94]. Genitalia
as in Fig. 330.
Cf . Unknown.
DISTRIBUTION. Sulawesi, India.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $ , Sulawesi: Tengah, nr Morowali, ii. 1980, Ranu River area, lowland rain forest, Malaise trap
(M. /. D. Brendell) (BMNH).
Paratypes. Sulawesi: 1 $, same data as holotype, iii.1980. India: 1 9, Hyderabad, Patancheru,
ICRISAT, vii-ix.1980, Malaise trap (Bernays & Woodhead) (BMNH).
COMMENTS. Also found in Zimbabwe and Cameroun (BMNH) and may be the same as dahmsi
from which it differs slightly in coloration, relative size and position of ocelli and relative
proportion of stigmal vein of forewing to marginal vein.
EUCOMOMORPHELLA Girault
(Key couplet: 459)
Eucomomorphella Girault, 1923c: 100. Type-species: Eucomomorphella emersoni Girault, by monotypy.
DISTRIBUTION AND SPECIES. Australia only, one species: emersoni Girault (1923c: 100).
BIOLOGY. Unknown.
COMMENTS. The genus is probably related to Prionomastix (tribe Prionomasticini, subtribe
Prionomasticina) and differs from this and related genera by having three teeth in the mandible
and the hypopyium not reaching half way along gaster (see also comments under Anagyrodes
andEncyrtus).
EUGAHANIA Mercet
(Key couplet: 143. Fig. 78)
Eugahania Mercet, 1926: 43. Type-species: Bothriothorax fumipennis Ratzeburg, by original designation.
DISTRIBUTION AND SPECIES. Four species, Palaearctic, Oriental and Australasian; two species
from review area: ishiharai Tachikawa (1956: 164) (India) and latiscapus (Ishii, 1925: 27)
INDO-PACIFIC ENCYRTIDAE 277
(India), also undetermined specimens from India, Vietnam, Taiwan, Indonesia, Irian Jaya and
Papua New Guinea (BMNH, BPBM, RMNH).
REFERENCE. Key to species: Hayat & Khanna (1977).
BIOLOGY. Parasites of nymphs of Cicadellidae (Homoptera).
COMMENTS. Placed in the tribe Eugahaniini (Encyrtinae) (see also comments under Anagyrodes
and Encyrtus) .
EUR YRHOPALUS Howard
(Key couplet: 501)
Euryrhopalus Howard, 18986: 237. Type-species: Euryrhopalus schwarzi Howard, by monotypy.
Synaspidia Timberlake, 1924: 397. Type-species: Synaspidia pretiosa Timberlake, by original designation.
DISTRIBUTION AND SPECIES. Nine species, New World; one species from review area: propinquus
Kerrich (1967: 240) (Hawaiian Is.).
REFERENCE. World revision: Kerrich (1967: 235-246).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The genus belongs to the tribe Aenasiini (see comments under Aenasius). A key
separating Euryrhopalus from related genera is provided by Kerrich (1967: 188-190).
EUSEMION Dahlbom
(Key couplet: 112. Fig. 51)
Eusemion Dahlbom, 1857: 293. Type-species: Encyrtus corniger Walker, by subsequent monotypy,
Thomson, 1876: 154.
DISTRIBUTION AND SPECIES. Two species, Palaearctic; one of these from New Zealand: cor-
nigerum (Walker; Annecke, 1967: 103).
BIOLOGY. Parasites of Coccidae (Homoptera).
COMMENTS. Placed in the tribe Cerapterocerini (Encyrtinae). A key to separate Eusemion from
related genera is provided by Annecke (1967: 100-101).
EXORISTOBIA Ashmead
(Key couplets: 201, 334, 449, 531. Fig. 238)
Exoristobia Ashmead, 1904a: 15. Type-species: Exoristobia philippinensis Ashmead, by monotypy.
Parasyrpophagus Girault, 1915a: 105. Type-species: Parasyrpophagus funeralis Girault, by original
designation. Syn. n.
Parageniaspis Masi, 1917ft: 154. Type-species: Parageniaspis macrocerus Masi by monotypy. Syn. n.
Mirsyrpophagus Girault, 1923a: 49. Type-species: Mirsyrpophagus columbi Girault, by monotypy. Syn. n.
DISTRIBUTION AND SPECIES. Six species, Afrotropical, Oriental and Australasian; three from
review area: columbi (Girault, 1923a: 49) (comb. n. from Mirsyrpophagus) (Australia),
funeralis (Girault, 19150: 105) (comb. n. from Parasyrpophagus) (Papua New Guinea,
Australia) and philippinensis Ashmead (1904a: 15) (Pakistan to Papua New Guinea), also
undetermined material from Thailand to New Hebrides (BMNH, BPBM).
REFERENCE. Subba Rao (1970).
BIOLOGY. Parasites of Syrphidae, Tachinidae and Phoridae (Diptera).
COMMENTS. The single extant female of Parageniapsis macrocerus Masi in the collection of the
BMNH is here designated LECTOTYPE. It belongs to the genus Exoristobia (comb. n.). The
278 J. S. NOYES & M. HAY AT
female syntype in the ZMCU belongs to Cerchysiella, whilst the single male syntype (BMNH) is
very probably the male of macrocerus.
One species (BMNH) which has been reared from Phoridae associated with pitcher plants
(see Beaver, 1979) is very distinct, having the dorsum of the thorax extremely hairy and the
prothoracic spiracle very enlarged and prominent, being clearly visible at low magnification in
dry-mounted material.
The two genera described by Girault differ only slightly from each other and from
philippinensis in the shape of the mandible, but since this can vary even in a single specimen it
is not considered to be of generic value. The genus may belong in the Microteryini but we are un-
able to place it with any degree of certainty according to Trjaptizin's (1973ft) classification
of the Encyrtinae. Trjapitzin & Gordh (1978ft) place it in the tribe Cheiloneurini, subtribe
Epiencyrtina which must be incorrect.
FULGORIDICIDA Perkins
(Key couplets: 339, 431, 437)
Fulgoridicida Perkins, 1906: 250. Type-species: Fulgoridicida dichroma Perkins, by monotypy.
DISTRIBUTION AND SPECIES. Five species, all Australian: cervantesi Girault (1923: 47), dichroma
Perkins (1906: 250), minuta Girault (1915: 148), nigricorpus Girault (19150: 148) and simplicis-
capus Girault (19150: 148), also at least one further species from Papua New Guinea and
Australia (BMNH, BPBM).
BIOLOGY. Parasites of eggs of Eurybrachidae (Homoptera).
COMMENTS. Girault (19150: 147) transferred Anagyrus saintpierrei Girault to Fulgoridicida, but
this is probably not correct. It is more likely that saintpierrei is an aberrant species of
Coelopencyrtus .
The genus is quite close to Ooencyrtus (Microteryini, subtribe Ooencyrtina) but differs mainly
in having fairly deep punctate-reticulate sculpture on the head and mesoscutum and a bidentate
mandible (that of Ooencyrtus has one or two teeth and a truncation or, rarely, three teeth).
GAHANIELLA Timberlake
(Key couplets: 190, 435. Figs 113, 227, 324)
Gahaniella Timberlake, 1926: 23. Type-species: Gahaniella californica Timberlake, by original desig-
nation.
DISTRIBUTION AND SPECIES. Three species, New World; one species from review area: saissetiae
Timberlake (1926: 27) (Hawaiian Is.).
REFERENCE. Kerrich (1953: 800-802).
BIOLOGY. Hyperparasites of Coccidae and Pseudococcidae (Homoptera) via other Encyrtidae,
possibly also primary parasites of Coccidae.
COMMENTS. Placed in the tribe Microteryini by Trjapitzin & Gordh (1978ft). The genus can be
recognised by the relatively high placement of the antennal toruli (Fig. 113) and the subequal
pedicel and funicle segments (Fig. 227).
GENTAKOLA gen. n.
(Key couplet: 155. Figs 84, 85, 332-339)
Type-species: Comperiella trifasciata Saraswat. Gender: feminine.
9 Head. Prognathous, occipital foramen situated in dorsal one-third of occiput; head in dorsal view a little
longer than broad and subrectangular with occipital margin distinctly concave, in side view also subrec-
INDO-PACIFIC ENCYRTIDAE 279
tangular, about one-half longer than broad and with genae dorso-anteriorly produced above and to side of
the antennal toruli (Fig . 332) . Eye with posterior margin clearly convex , about one-half longer than broad ,
with fairly sparse inconspicuous setae each very slightly longer than the diameter of a facet and about half
as long as those on frontovertex, eye not quite reaching occipital margin which is sharp. Malar space about
half length of eye and with sulcus absent. Frontovertex slightly more than one-third head width; ocelli
forming a very slightly acute angle, posterior ocellus separated from occipital and eye margins by slightly
less than its own diameter. Antennal scrobes not deep but bounded laterally by the dorso-anterior
projection of the genae and dorsally by the sharp angle resulting from the face being sharply inflexed at this
point, thus the scrobes more or less semi-circular and reaching about three-eighths way from antennal
toruli to anterior ocellus; antennal torulus separated from mouth margin by not more than half its length
and from other torulus by about twice its length, its dorsal margin well below ventral level of eyes; clypeal
margin very shallowly excised between toruli. Antennal scape much longer than minimum width of
frontovertex, clearly broadened and flattened, subrectangular about twice as long as broad, pedicel
conical, about one-third length of scape or funicle, the latter a little shorter than the scape, all funicle
segments transverse, at least about twice as broad as long and broadly oval in cross section, slightly
broadening distally so that sixth segment is clearly as broad as clava which is two-segmented, with apex
rounded and a little shorter than funicle; longitudinal sensillae on all flagellar segments except the first two.
Frontovertex entirely smooth and shiny, genal process with shallow irregular rugose sculpture, fron-
tovertex with a few inconspicuous setae, each a little longer than diameter of an ocellus. Mandible narrow
with three apical teeth, the middle one slightly the longest, maxillary palpus four-segmented, labial palpus
three-segmented (a little obscure in the only slide preparation available and may be two-segmented).
Thorax. In side view very slightly dorso-ventrally flattened with mesoscutum and scutellum quite flat,
metapleurum laterally obscure and, together with propodeum, quite broadly in contact with hind coxa. In
dorsal view pronotum quite long, slightly longer than half mesoscutum and with posterior margin slightly
concave; visible part of mesoscutum slightly less than twice as broad as long, without notaular lines and
with its posterior margin slightly produced backwards medially; axillae meeting; scutellum about as long as
broad, apically rounded with a very narrow apical flange which projects slightly over propodeum medially
and slightly longer than mesoscutum; propodeum medially about one-fifth length of scutellum. Pronotum
with shallow, raised, transverse rugose to squamiform-reticulate sculpture; mesoscutum with very shallow,
transverse, rugose sculpture, scutellum completely smooth and polished, propodeum medially with
shallow, irregular, raised reticulate sculpture; mesopleurum smooth; dorsum of thorax with sparse, short,
recumbent dark setae. Forewing slightly bent upwards at about middle as in Compendia, with three
longitudinal fuscous streaks, about three times as long as broad; linea calva not clearly defined and not
interrupted, there being at most only one or two setae on surface of wing proximad of it (Fig. 334); filum
spinosum present and directed towards junction of marginal and submarginal veins; submarginal vein with
an apical hyaline break and with parastigma a little swollen; marginal vein about two and one-half to three
times as long as broad, about one-half longer than postmarginal and subequal to stigmal vein, apex of
venation not reaching half way along wing; costal cell about 11 or 12 times as long as broad, with a single
line of setae dorsally in apical one-third. Hindwing also slightly infuscate, about two-thirds as long as
forewing, about four times as long as broad and with marginal fringe about one-quarter of wing width. Mid
tibial spur very slightly longer than basal mid tarsal segment.
Caster. Very slightly shorter than thorax; cereal plates in anterior one-half; hypopygium more or less
reaching apex of gaster; paratergites absent; last tergite about as long as mid tibia; gonostyli free, about
one-quarter length of ovipositor which is nearly as long as mid tibia.
O". Differs from female as follows. Occipital foramen not quite in dorsal one-third of occiput, head about as
long as broad in facial view, with only very slight dorso-anterior projections of genae; eye about
one-quarter longer than broad, with setae not longer than diameter of a facet and clearly not reaching
occipital margin which is sharp; malar space about half as long as eye; frontovertex a little more than head
width; ocelli forming a right angle, antennal torulus separated from mouth margin by nearly its own length,
from other torulus by slightly more than twice its own length; antennal scape distinctly shorter than width
of frontovertex, subrectangular, stout, about twice as long as broad; pedicel a little less than half length of
scape, subquadrate, longer than any funicle segment, all funicle segments transverse, the sixth being the
longest and broadest and almost quadrate, clava entire, about half length of funicle; longest setae on
flagellum about twice as long as diameter of corresponding segment; frontovertex almost entirely smooth
but with some very shallow rugose-reticulate sculpture immediately above the scrobes. Mesoscutum with
very shallow squamiform-reticulate sculpture. Forewing not bent at middle, hyaline and about two and
one-half times as long as broad; apex of venation reaching about two-fifths along wing. Mid tibial spur
distinctly longer than basal mid tarsal segment. Gaster a little shorter than thorax; genitalia with digiti
280 J. S. NOYES & M. HAY AT
about one-fifth length of aedeagus which is slightly longer than half mid tibia or two and one-half times mid
tibial spur.
COMMENTS. The genus superficially resembles Comperiella, but the structure of the gaster, the
relatively long propodeum, the clearly tridentate mandible, the forewing venation and the filum
spinosum being directed towards the junction of the submarginal and marginal veins suggest an
affinity with Cerchysiella and Zaommoencyrtus which are placed in the tribe Bothriothoracini,
subtribe Coenocercina. It can be separated from other members of the subtribe by the shape of
the head, two-segmented clava and infuscate forewing, the latter being bent upwards at the
middle.
Gentakola trifasciata (Saraswat) comb. n.
(Figs 84, 85, 332-339)
9- The female can easily be recognised from Saraswat's (in Saraswat & Mukerjee, 1975: 51) original
description. There seems to be some variation in colour; in one dry-mounted specimen the head is largely
green or greenish blue with anterior genal protuberance more or less deep purple and the clypeus and
interantennal prominence quite strongly orange; the scutellum is more strongly blue than in Saraswat's
description. Also the mandible is quite clearly tridentate (not quadridentate) and the clava is two-
segmented (not three-segmented) (Fig. 85). These discrepancies may have arisen as a result of Saraswat
having based his description on uncleared slide-mounted material, the clava of the dry-mounted specimen
examined having the appearance of being three-segmented. Genitalia as in Fig. 333, hypopygium as in Fig.
335.
Cf . Length: 0-71 mm. Body generally dark purplish brown except scutellum which is slightly metallic green;
leg coloration more or less as for female. For other characters see Figs 336-339.
DISTRIBUTION. India.
BIOLOGY. Unknown.
MATERIAL EXAMINED
India: 1 $, 1 cf , Tamil Nadu, Coimbatore, 25.ix-l.x.l979 (/. S. Noyes); 1 $, Karnataka, Bannerghatta
N. P., 5.xi.l979 (Z. Boucek&J. S. Noyes); 1 cf , Delhi, IARI area, x.1979 (Z. Boucek) (BMNH).
GYRANUSOIDEA Compere
(Key couplets: 149, 172, 268. Fig. 93)
Gyranusoidea Compere, 1947a: 17. Type-species: Gyranusa citrina Compere, by original designation.
DISTRIBUTION AND SPECIES. Fifteen species, cosmopolitan except for Palaearctic; 7 species from
review area: advena Beardsley (1969: 303) (Hawaiian Is.), albiclavata (Ashmead, 1905ft: 404)
(comb. n. from Aphycus) (Philippines), ceroplastis (Agarwal, 1965: 73) (India), flava Shafee,
Alam & Agarwal (1975: 21) (India), indica Shafee, Alam & Agarwal (1975: 22) (India), mirzai
(Agarwal, 1965: 46) (comb. n. from Anagyrus) (India) and phenacocci (Beardsley, 1969: 299)
(Hawaiian Is.), also undetermined material from Taiwan, Philippines, Australia and New
Britain (BMNH, BPBM, UCR, HC).
REFERENCE. Key to some species: Shafee etal. (1975: 21).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The genus belongs to the tribe Anagyrini, subtribe Anagyrina (Tetracneminae) and
is extremely difficult to separate from Anagyrus on one hand and Leptomastidea on the other.
We have separated it from Anagyrus solely by the postmarginal vein of the forewing being at
least one-quarter longer than the stigmal whereas in Anagyrus it is not or hardly longer. This is
an extremely doubtful character for separating the two genera, but we have found it more
convenient and easier to define than the shape of the scape or sculpture of the head and thorax.
From Leptomastidea we have separated it basically on the head and thorax having fine punctate
sculpture of velvety or granulate appearance and the forewing being more or less hyaline,
IN DO-PACIFIC ENCYRTIDAE 281
whereas species placed in Leptomastidea have smoother, shallower sculpture and the forewing
often with two or more fuscous fasciae. Use of the above characters has resulted in the
undesirable transfer of some species from other genera to Gyranusoidea, but we have found this
necessary to achieve a degree of consistency within the key. We feel certain that an in-depth
study of this group of genera on a world basis will result in the eventual synonymy of most genera
of the Anagyrina. This is beyond the scope of the present work and therefore we have tried to
retain most of the genera as valid even though it has meant separating some of them on weak
characters such as those above (see also comments under Anagyrus).
HABROLEPIS Forster
(Key couplet: 97. Fig. 48)
Habrolepis Forster, 1856: 34. Type-species: Encyrtus nubilipennis Walker, by original designation.
Gymnoneura Risbec, 1951: 157. Type-species: Gymnoneura bambeyi Risbec, by monotypy.
DISTRIBUTION AND SPECIES. Twenty-one species, cosmopolitan; three species from review area:
dalmani (Westwood; Annecke & Mynhardt, 1970: 134) (New Zealand) , neocaledonensis Fabres
(1974: 56) (New Caledonia) and rouxi Compere (1936a: 495) (Hawaiian Is.), also undetermined
material, containing at least one further species, from Australia and Samoa (ANIC, CNC).
REFERENCE. Annecke & Mynhardt (19706: 128-146).
BIOLOGY. Parasites of Diaspididae and Asterolecaniidae (Homoptera).
COMMENTS. Placed in the tribe Habrolepidini, subtribe Habrolepidina (Encyrtinae).
HALIGRA gen. n.
(Key couplet: 532. Figs 253, 254, 340-345)
Type-species: Haligra concolorsp. n. Gender: feminine.
$. Head. In facial view about as long as broad and in profile about three-fifths as broad as long and
gradually curved dorsally, below top of antennal scrobes almost straight. Eye with posterior margin
straight, about one-third longer than broad, with numerous conspicuous, translucent setae, each about as
long as the diameter of a facet, eye almost touching occipital margin which is sharp. Malar space slightly
longer than half eye length, with sulcus present. Frontovertex about one-third head width; ocelli forming a
slightly acute angle, the posterior ocellus about equidistant from eye and occipital margins. Antennal
scrobes shallow, meeting dorsally and almost semicircular, reaching about half way from toruli to anterior
ocellus; antennal torulus separated from mouth margin by a little less than its own length and from other
torulus by slightly more than its own length, its upper margin about half its length below level of lowest eye
margin, clypeal margin very shallowly excised between toruli. Antennal scape clearly longer than
minimum width of frontovertex, fairly slender, about five times as long as broad; pedicel conical, about
one-third as long as scape and clearly much longer than any of the funicle segments which are all strongly
transverse, the first the smallest and sixth the largest; clava three-segmented with a short oblique
truncation apically, the outer suture converging slightly with the inner, clava about as long as and much
broader than the funicle; longitudinal sensillae on fifth and sixth funicle segments and clava; longest setae
on flagellum about as long as or a little longer than corresponding segment. Head almost totally smooth and
shiny but with some extremely shallow, raised irregular sculpture on the frontovertex, immediately above
scrobes and below this more or less completely smooth and polished except on genae posterior to malar
suture and on temples which have very shallow, raised, reticulate sculpture; setae on frontovertex dark and
conspicuous, each at least about twice as long as diameter of an ocellus, those on lower parts of face and
interantennal prominence about the same. Mandible with three acute teeth, the middle one clearly the
longest; maxillary palpus four-segmented, labial palpus two-segmented.
Thorax. In side view robust with mesoscutum and scutellum conspicuously convex, the metapleurum
and propodeum not quite meeting the hind coxa, although mesopleurum clearly separated from basal
segment of gaster. In dorsal view pronotum almost completely hidden, its posterior margin very concave;
visible part of mesoscutum about twice as broad as long, with notaular lines absent and its posterior margin
almost straight; axillae separated by about half the length of an axilla in dorsal view; scutellum a little
282 J. S. NOYES & M. HAY AT
broader than long, apically rounded; propodeum medially about one-sixth length of scutellum. Mesoscu-
tum with longitudinally elongate, raised, reticulate sculpture of moderate depth, the cells more or less
arranged in lines which slightly converge posteriorly; scutellum with finer, striate-reticulate sculpture of
about the same depth or a little deeper than mesoscutum; propodeum with irregular raised sculpture
medially and some sculpture along its anterior margin nearly reaching spiracles; mesopleurum almost
completely smooth but with some extremely shallow irregular sculpture. Forewing more or less completely
hyaline but very faintly suffused brownish, between two and one-half to three times as long as broad; linea
calva not interrupted but partially closed by two lines of setae; filum spinosum present; submarginal vein
with an apical hyaline break, parastigma not conspicuously swollen; costal cell over 20 times as long as
broad, with a single line of setae dorsally in its distal one-third; marginal vein about three times as long as
broad, nearly twice as long as stigmal which is about as long as postmarginal. Hindwing very slightly
suffused brownish as in fore wing, about two-thirds length of forewing, about five times as long as broad
with marginal fringe about half as long as maximum wing width. Mid tibial spur about as long as or a little
shorter than basal mid tarsal segment.
Caster. A little longer than thorax with apex of hypopygium reaching to about two-thirds along gaster;
ovipositor not or hardly exserted; cereal plates situated about halfway along gaster; paratergites absent;
last tergite about three-quarters as long as mid tibia; ovipositor about as long as mid tibia, gonostyli free,
about one-fifth as long as ovipositor.
Cf . Unknown.
COMMENTS. We are unable to place this genus according to Trjapitzin's (19736) classification of
the Encyrtinae. However, it does bear some resemblance to Forcipestricis Burks and the two
may be related, although the present genus lacks the pits on the scutellum which are so
characteristic of Forcipestricis.
Haligra concolorsp. n.
(Figs 253, 254, 340-345)
$. Length: approx. 0-78-0-87 mm (holotype, 0-87 mm).
Colour. Body generally very dark brown or black; head quite shiny, antenna dark brown; mesoscutum
slightly shiny, scutellum matt except the almost vertical apical surface which is completely smooth and
shiny, legs dark brown with apices of fore tibia, mid tibia, hind femur and tibia and fore and hind tarsi
testaceous, apical three-quarters of mid tibia and tarsus testaceous-yellow; gaster with some slight brassy
reflections.
Head. Relative measurements (holotype): head length 52, head width (facial view) 53, head width (side
view) 30, minimum width of frontovertex 17-5, malar space 20, eye length 34, eye width 26, POL 9, OOL 3,
scape length 26, scape width 5-5, proportions of antenna as in Fig. 253, mandible as in Fig. 341.
Thorax. Sculpture of mesoscutum as in Fig. 340, of scutellum as in Fig. 254. Relative measurements
(holotype): forewing length 130, width 51, other proportions of forewing as in Figs 342, 344; hindwing
length 92, width 18. The paratype has the forewing nearly three times as long as broad and the postmarginal
vein on right wing a little longer than stigmal, whilst on left wing it is a little shorter.
Gaster. Relative lengths (paratype): last tergite 45, ovipositor 64, [mid tibia 63]; genitalia as in Fig. 345,
hypopygium as in Fig. 343.
Cf . Unknown.
DISTRIBUTION. India.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $, India: Uttar Pradesh, Aligarh, 11. ii. 1979, on grass (M. Hayai) (BMNH).
Paratype. India: 1 $, Uttar Pradesh, Aligarh, ll.ii.1978, on grass (M. Hayai) (HC).
HAMBLETONIA Compere
(Key couplet: 122. Figs 62-64)
Hambletonia Compere, 1936a: 172. Type-species: Hambletonia pseudococdna Compere, by original
designation.
IN DO-PACIFIC ENCYRTIDAE 283
DISTRIBUTION AND SPECIES. One species, New World; also Hawaiian Is. and Taiwan: pseudo-
coccina Compere (1936a: 173).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the tribe Chrysoplatycerini, subtribe Chrysoplatycerina (Trjapitzin &
Gordh, 1978ft: 648), although it might possibly be more closely related to Taftia (subtribe
Taftiina). It can be easily recognised by the conspicuous long setae arising from the dorsal
surface of the pedicel.
HAMUSENCYRTUS Subba Rao & Hayat
(Key couplets: 84, 170. Figs 38, 39, 92, 346)
Hamusencyrtus Subba Rao & Hayat, 1979: 2. Type-species: Scelioencyrtus mymaricoides Compere, Subba
Rao & Kaur, by original designation.
Neoxanthoencyrtus Avasthi & Shafee, 1980: 535. Type-species: Scelioencyrtus mymaricoides Compere,
Subba Rao & Kaur, by original designation.
DISTRIBUTION AND SPECIES. Two species, possibly synonymous, India and Pakistan only: indicus
(Shafee, Alam & Agarwal, 1975: 33) (India) and mymaricoides (Compere, Subba Rao & Kaur,
1960: 46) (India, Pakistan), also further undetermined material, possibly including undescribed
species, from India (BMNH, HC).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The genus can be placed in the tribe Anagyrini, subtribe Rhopina (Tetracneminae)
and is possibly closest to Asitus. It can be distinguished from this genus by having a two-
segmented clava (solid in Asitus) , distinct axillae (fused with scutellum in Asitus) and forewing
venation not quite reaching anterior margin of wing (reaches anterior margin in Asitus).
HELEGONA TOPUS Perkins
(Key couplets: 283, 337)
Helegonatopus Perkins, 1906: 257. Type-species: H elegonatopus pseudophanes Perkins, by monotypy.
Chalcerinys Perkins, 1906: 258. Type-species: Chalcerinys eximia Perkins, by monotypy. Syn. n.
Schedioides Mercet, 1919a: 96. Type-species: Schedioides formosus Mercet, by monotypy.
Euchalcerinys Timberlake, 1922a: 161. Type-species: Euchalcerinys apicicornis Timberlake, by original
designation. Syn. n.
Hazmburkia Hoffer, 1954: 172. Type-species: Hazmburkia dimorpha Hoffer, by original designation.
Masencyrtus Hoffer, 1960: 98. Type-species: Masencyrtus concupiens Hoffer, by original designation.
Paludencyrtus Hoffer, 1965: 16. Type-species: Paludencyrtus nikolskajae Hoffer, by original designation.
DISTRIBUTION AND SPECIES. Fifteen species, cosmopolitan; five species from review area:
apicicornis (Timberlake, I922a: 165) (comb. n. from Euchalcerinys) (Hawaiian Is.), eximius
(Perkins, 1906: 259) (comb. n. from Chalcerinys) (Hawaiian Is.), ponomarenkoi Trjapitzin
(19640: 143) (India), pseudophanes Perkins (1906: 258) (Hawaiian Is.) and pulchricornis Hayat
& Verma (1978: 355) (India), also undetermined material from Java (BPBM).
REFERENCE. Szelenyi (1972ft: 348-352).
BIOLOGY. Hyperparasites of Auchenorrhyncha (Homoptera) via Dryinidae (Hymenoptera).
COMMENTS. We have examined one female and one male of what are probably syntypes of
Chalcerinys eximia Perkins (BMNH); they are close to, if not the same as pulchricornis and
therefore we propose the synonymy of Chalcerinys and Helegonatopus. We favour use of
Helegonatopus as the valid generic name since it is the better known of the two, although
Chalcerinys is the type-genus of the tribe Chalcerinyini.
We have also examined the holotype of Euchalcerinys apicicornis Timberlake (BPBM) and
are confident that it also belongs in the genus Helegonatopus.
The genus belongs in the tribe Chalcerinyini (Encyrtinae).
284 J. S. NOYES & M. HAY AT
HEMILEUCOCERUS Hoffer
(Key couplet: 294)
Hemileucocerus Hoffer, 1976: 101. Type-species: Hemileucocerus insignis Hoffer, by original designation.
DISTRIBUTION AND SPECIES. Only one described species known, Europe; at least two further
species from India, Laos and Borneo (BMNH, HC, BPBM).
BIOLOGY. Unknown.
COMMENTS. We have not seen any material authoritatively determined as Hemileucocerus
insignis Hoffer, but material examined from the Canary Is. (BMNH) agrees well with Hoffer's
generic description. This material is congeneric with that from India, Laos and Borneo.
Hemileucocerus belongs in the same group as Aseirba and Austroencyrtus and can be
separated from these genera by the characters given in the key (see also comments under
Aseirba).
HENGATA&n.n.
(Key couplet: 307. Figs 154, 347-354)
Type-species: Hengata spinosa sp. n. Gender: feminine.
$ . Head. In facial view about as long as broad, in profile about twice as long as broad, anteriorly more or
less evenly and gradually curved, the lower part of the interantennal prominence quite clearly visible. Eye
with posterior margin a little concave, about one-half longer than broad, covered with numerous, fairly
inconspicuous translucent setae, each nearly as long as the diameter of a facet, eye very slightly
overreaching occipital margin which is sharp. Malar space about half length of eye, with sulcus present.
Frontovertex about two-fifths head width; ocelli forming a right angle, the posterior ones separated from
eye or occipital margin by a little less than their own diameters. Antennal scrobes shallow and semicircular,
meeting dorsally and extending slightly more than halfway from toruli to anterior ocellus; antennal torulus
separated from mouth margin and other torulus by about one and a half times its own length, its dorsal
margin a little above the ventral margin of eyes, clypeal margin shallowly excised below toruli. Antennal
scape longer than mimimum width of frontovertex, almost cylindrical, about five times as long as broad;
pedicel conical about one-third length of scape and slightly longer than any funicle segment, all of which are
subequal and a little longer than broad; clava three-segmented, the sutures subparallel, its apex more or
less pointed; longitudinal sensillae on all flagellar segments; longest setae about as long as diameter of
corresponding segment. Frontovertex almost smooth, but with extremely shallow, raised, regular,
reticulate sculpture, much more elongate and irregular between antennal scrobes and eyes and on genae
and temples; frontovertex and interantennal prominence with a few moderately long, dark setae; genae
and clypeal margin with translucent setae. Mandible with two teeth and a truncation; maxillary palpus
four-segmented, labial palpus three-segmented.
Thorax. In side view robust with mesoscutum and scutellum fairly convex, the mesopleurum posteriorly
enlarged and touching basal segment of gaster, thus clearly separating the propodeum and metapleurum
from the hind coxa. In dorsal view pronotum with posterior margin broadly concave; visible part of
mesoscutum nearly twice as broad as long, broadly concave, notaular lines absent, a very shallow median
longitudinal ridge in posterior half, posterior margin clearly convex and projecting above axillae medially;
axillae separated by about two-thirds width of an axilla in dorsal view; scutellum convex, about as broad as
long and about as long as mesoscutum, its apex rounded; propodeum medially not more than about
one-tenth length of scutellum, laterally with only two or three setae to the outside of the spiracle.
Mesoscutum with very shallow, raised, squamiform-reticulate sculpture, becoming more shallow post-
eriorly, scutellum anteriorly, and axillae with extremely shallow, raised, reticulate sculpture, posterior half
or so of scutellum completely smooth ; propodeum almost completely smooth ; dorsum of thorax clothed in
sparse, moderately long, conspicuous, dark setae. Forewing hyaline, about two and one-half times as long
as broad; linea calva not interrupted and open; filum spinosum present; submarginal vein with an apical
hyaline break, with parastigma not conspicuously swollen; costal cell nearly 15 times as long as broad, with
a single line of setae dorsally in its apical one-third or so; marginal vein about twice as long as broad,
subequal to stigmal and postmarginal veins. Hindwing about two-thirds length of forewing, about four and
one-half times as long as broad, with marginal fringe about one-quarter as long as maximum wing width.
Mid tibia! spur a little shorter than basal mid tarsal segment.
INDO-PACIFIC ENCYRTIDAE 285
Caster. Slightly longer than thorax; cereal plates at about midway along its length; hypopygium with its
apex reaching to nearly two-thirds along gaster; paratergites absent; last tergite about half as long as mid
tibia; ovipositor hardly exserted, a little shorter than mid tibia, gonostyli free, about one-quarter length of
ovipositor.
Cf . Similar to female except body generally darker, antenna and genitalia. Differs as follows. Head clearly
broader than long; malar space nearly two-thirds as long as eye; frontovertex nearly half as wide as head;
posterior ocelli a little further from eye margin than from occipital margin; antennal scape about as long as
width of frontovertex with a very large thorn-like process arising from its ventral margin, from the apex of
which is a strong, apically hooked bristle, scape thus only about twice as long as greatest width, pedicel
conical, subquadrate, about as long as first funicle segment but clearly shorter than those following which
are all longer than broad and subequal, clava entire; longitudinal sensillae on all flagellar segments except
the first two, longest setae on funicle very nearly twice as long as diameter of segments. Forewing with
costal cell about 12 times as long as broad, with marginal, stigmal and postmarginal veins proportionately a
little shorter than in female. Genitalia with aedeagus about half as long as mid tibia, digiti each armed with
one apical hook, about one-sixth length of aedeagus.
COMMENTS. The expanded mesopleurum, structure of the mandible, and mesoscutum dorsally
separating the axillae indicate that this genus is related to Ooencyrtus and Fulgoridicida
(Microteryini, Ooencyrtina), also Amira (Amirini). It can be separated from these and all other
genera by the shallow but distinct median longitudinal ridge or carina on the mesoscutum. The
peculiar thorn-like process on the scape of the male is also another distinguished character.
Hengata spinosa sp. n.
(Figs 154, 347-354)
$. Length: 0-71-0-95 mm (holotype, 0-95 mm).
Colour. Body generally pale brownish yellow; frontovertex a little darker, with greenish reflections,
clypeus dark brown; scape yellow, pedicel and flagellum brown, clava white; face of pronotum, mesoscu-
tum anteriorly, scutellum in posterior half, mesopleurum posteriorly, propodeum and apical half of gaster,
dark brown, the posterior half of scutellum with a slight greenish or purplish sheen; legs yellow, mid coxa
brown; base of gaster and proximal half of venter yellowish. In some specimens the darker brown areas are
less intense or less extensive and occasionally the propodeum is yellowish; occasionally the base of the hind
coxa is dark brown.
Head. Relative measurements (holotype): head length 56, head width (facial view) 57, head width (side
view) 26, minimum width of frontovertex 21, malar space 20, eye length 37, eye width 25, POL 9, OOL 3,
scape length 33, scape width 6, other proportions of antenna as in Fig. 348; mandible as in Fig. 347.
Thorax. Relative measurements (holotype): forewing length 147, width 58, hindwing length 98, width
22. Base of forewing as in Fig. 154, venation as in Fig. 349.
Gaster. Relative lengths (paratype): last tergite 43, ovipositor 75, [mid tibia 87] ; genitalia as in Fig. 351,
hypopygium as in Fig. 350.
Cf . Length: 0-71-0-94 mm. Similar to female except following. Body completely dark brown, legs and
gaster as in female. Antenna as in Fig. 352; genitalia as in Figs 353, 354. Relative measurements (paratype
1): head width 74, minimum frontovertex width 33, scape length 31, forewing length 172, forewing width
73, hindwing length 114, hindwing width 28, mid tibia length 70, aedeagus length 32; relative measure-
ments (paratype 2); scape length 20, scape width (to apex of thorn-like process) 10, POL 9-5, OOL 3.
(Paratype 1 on a slide; paratype 2 dry-mounted on a card.)
DISTRIBUTION. Indonesia (Sulawesi).
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $, Indonesia: Sulawesi, Tengah, Ranu River area, nr Morowali, ii.1980, lowland rain forest,
Malaise trap (M. J. D. Brendell) (BMNH).
Paratypes. Indonesia: 4 $ , 6 Cf , same data as holotype; 4 $ , 4 O" , same data as holotype but iii.1980; 10
$, 11 Cf, same data as holotype but 27.i.-20.iv.l980 (M. J. D. Brendell) (BMNH, USNM, UCR, ZI,
PPRI,HC).
286 J. S. NO YES & M. HAY AT
HESPERENCYRTUS Annecke
(Key couplet: 136)
Hesperencyrtus Annecke, 1971a: 86. Type-species: Paraphaenodiscus lycoeniphila Risbec, by original
designation.
DISTRIBUTION AND SPECIES. One species known, Afrotropical; also reported from India:
lycoeniphila (Risbec, 1951: 147).
BIOLOGY. Parasites of the pupae of Lycaenidae (Lepidoptera).
COMMENTS. The material referred to by Hayat & Subba Rao (1981: 113) should be in the
collections of the BMNH. We have been unable to locate it and therefore it must be assumed
that this material was originally misidentified or has been lost.
The genus belongs in the tribe Microteryini and probably the subtribe Microteryina
(Encyrtinae).
HETEROCOCCIDOXENUS Ishii
(Key couplets: 280, 419. Fig. 172)
Heterococcidoxenus Ishii, 1940: 103. Type-species: Heterococcidoxenus javensis Ishii, by original des-
ignation.
Microsphenus Kerrich, 1963: 365. Type-species: Bothriothorax schlechtendali Mayr, by original des-
ignation.
DISTRIBUTION AND SPECIES. Two species, Palaearctic and Australasian; one from review area:
javensis Ishii (1940: 103) (Java).
BIOLOGY. Parasites of Scolytidae (Coleoptera).
COMMENTS. Placed in the tribe Bothriothoracini, subtribe Bothriothoracina (Encyrtinae).
HEXENCYRTUS Girault
(Key couplets: 222, 290)
Hexencyrtus Girault, 1915a: 105. Type-species: Hexencyrtus albidava Girault, by original designation.
Calliencyrtus De Santis, 1960: 61. Type-species: Calliencyrtus bucculentus De Santis, by original des-
ignation. Syn. n.
DISTRIBUTION AND SPECIES. Two species, Neotropical, Oriental and Australasian; one from
review area: albidava Girault (19150: 105) (= Hexencyrtus fumosipennis Girault, 1915a: 106
syn. n.), also further undetermined material from Vietnam and Papua New Guinea (BPBM).
BIOLOGY. Unknown.
COMMENTS. We have compared a specimen of Hexencyrtus albidava (compared with holotype)
with a specimen determined as Calliencyrtus bucculentus by De Santis. They are very close and
certainly belong in the same genus.
The genus is difficult to place according to Trjapitzin's (19736) classification of the Encyr-
tinae, but it almost certainly belongs to the same generic group as Parastenoterys and Rhytido-
thorax (see also comments under Parastenoterys).
HOLANUSOMYIA Girault
(Key couplet: 158. Figs 83, 355, 356)
Holanusomyia Girault, 1915c: 165. Type-species: Holanusomyia pulchripennis Girault, by original
designation.
DISTRIBUTION AND SPECIES. One species, Philippines only: pulchripennis Girault (1915c: 165).
INDO-PACIFIC ENCYRTIDAE 287
BIOLOGY. Unknown.
COMMENTS. The genus is related to Yasumatsuiola which is placed in the tribe Dinocarsiini
(Trjapitzin, 1977: 155). It is easily recognised by the abnormally long stigmal vein of the
forewing. We have seen other material from Taiwan (UCR, BPBM) which could possibly be
placed in this genus but refrain from doing so until a more detailed study of the group, to which
this genus belongs, can be undertaken.
HOLCOTHORAX Mayr
(Key couplet: 67)
Holcothorax Mayr, 1876: 691. Type-species: Encyrtus testaceipes Ratzeburg, by designation of Gahan &
Pagan (1923: 72).
DISTRIBUTION AND SPECIES. Two species, European; neither from review area, but one un-
described species from India (BMNH).
BIOLOGY. Polyembryonic parasites of the larvae of Gracillariidae and Nepticulidae (Lepidop-
tera) .
COMMENTS. The species from India is intermediate between Holcothorax and Paraleurocerus
Girault, having sculpture of the head and thorax similar to the latter, but with a five-segmented
funicle. It is probable that these two genera and Ageniaspis will eventually be considered
synonymous.
Holcothorax is placed in the tribe Copidosomatini, subtribe Ageniaspidina (see comments
under Ageniaspis) .
HOMALOPODA Howard
(Key couplet: 54)
Homalopoda Howard in Riley, Ashmead & Howard, 1894: 90. Type-species: Homalopoda cristata
Howard, by monotypy.
DISTRIBUTION AND SPECIES. One described species from the Neotropics, Sri Lanka and the
Hawaiian Is.: cristata Howard; Noyes (1979: 157), also undetermined material from Java
(BMNH).
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. We have not examined any of the material determined as cristata from either Sri
Lanka or the Hawaiian Is. and therefore cannot confirm its identity.
The genus belongs to the tribe Habrolepidini, subtribe Habrolepidina.
HOMALOTYLUS Mayr
(Key couplets: 34, 239, 354, 388. Figs 6, 146, 198)
Homalotylus Mayr, 1876: 752. Type-species: Encyrtus flaminius Dalman, by designation of Ashmead
(19006: 377).
Nobrimus Thomson, 1876: 116. Type-species: Encyrtus flaminius Dalman, by designation of Timberlake
(1919a: 134).
Mendozaniella Brethes, 1913: 97. Type-species: Mendozaniella mirabilis Brethes, by monotypy.
Hemaenasoidea Girault, 1916c: 307. Type-species: Hemaenasoidea oculata Girault, by monotypy.
Anisotylus Timberlake, 1919a: 170. Type-species: Homalotylus similis Ashmead, by original designation.
Syn. n.
Lepidaphycus Blanchard, 1936: 13. Type-species: Lepidaphycus bosqi Blanchard, by monotypy.
Neoaenasioidea Agarwal, 1966: 71 . Type-species: Neoaenasioidea indica Agarwal, by original designation.
DISTRIBUTION AND SPECIES. Twenty-eight species, cosmopolitan; 12 species from review area:
albidavatus (Agarwal, 1970: 27) (India), ferrierei Hayat, Alam & Agarwal (1975: 67) (India),
288 J. S. NO YES & M. HAYAT
flaminius (Dalman, 1820: 340) (India, S. China, Java, Australia), indicus (Agarwal, 1966: 73)
(India), mexicanus Timberlake (19190: 155) (India), microgaster Girault (1917g: 134) (Austra-
lia), mundus Gahan (1920: 343) (Philippines), nigritus (Agarwal, 1970: 27) (India), nipaecocci
(Subba Rao, 1967: 1) (India), oculatus (Girault, 19166: 308) (Philippines), orci Girault
(19170: 3) (Java), terminalis (Say; Timberlake, 19190: 148) (India), also much undetermined
material from throughout the region (BMNH, BPBM, GC, HC).
REFERENCES. Revision: Timberlake (19190: 133-170); review of Indian species: Hayat et al.
(1975: 64-69).
BIOLOGY. Parasites of coccinellid larvae (Coleoptera, Coccinellidae).
COMMENTS. We do not believe that the retention of Anisotylus as a valid genus is realistic; at
most, the characters used to separate it from Homalotylus (bidentate mandible and venation)
can be considered as valid only on a specific or perhaps species-group level but certainly not at
generic level. This proposal is further supported by the fact that, where known, all species of
Anisotylus are parasitic on coccinellid larvae.
Timberlake (19190: 141) synonymised both orci and microgaster with flaminius. Mr P. B.
Jensen (pers. comm.) has studied the type of flaminius and informed us that the species may
have been misinterpreted. For this reason we are maintaining these species as distinct until the
matter can be resolved.
The genus is placed in the tribe Homalotylini, subtribe Homalotylina (see also comments
under Aphycus).
HUNTERELLUS Howard
(Key couplets: 81, 152, 214)
Hunterellus Howard, 1908: 240. Type-species: Hunterellus hookeri Howard, by monotypy.
Australzaomma Girault, I925b: 97. Type-species: Australzaomma brunnea Girault, by monotypy. Syn. n.
DISTRIBUTION AND SPECIES. Five species, cosmopolitan; four from review area: brunneus
(Girault, 19256: 96) (comb. n. from Australzaomma) (Australia), hookeri Howard (1908: 241)
(India, Malaysia, Hawaiian Is.), mysorensis (Mani, 1941: 29) (comb. n. from Ixodiphagus)
(India) and sagarensis Geevarghese (1977: 49) (India).
BIOLOGY. Parasites of nymphs of Ixodidae (Acarina).
COMMENTS. We have not examined the type of Ixodiphagus mysorensis but it would seem
reasonable to assume from the description and distribution that it is a species of Hunterellus.
The genus belongs to the tribe Ixodiphagini (Encyrtinae).
HYPERGONA TOPUS Timberlake
(Key couplets: 90, 474, 485. Figs 42, 43, 226)
Hypergonatopus Timberlake, 1922a: 142. Type-species: Echthrogonatopus hawaiiensis Perkins, by orig-
inal designation.
Aulonops Timberlake, 1922a: 158. Type-species: Aulonops bifasciata Timberlake, by original designation.
Syn. n.
DISTRIBUTION AND SPECIES. Eight species, all from Hawaiian Is. : bifasciatus (Timberlake, 19220:
159) (comb. n. from Aulonops), brunneipes Timberlake (19220: 154), flavipes Timberlake
(19220: 155), hawaiiensis (Perkins, 1912: 17), hemipterus Timberlake (19220: 157), molokaien-
sis (Ashmead, 1901: 322), oahuensis Timberlake (19220: 153) and vulcanus Timberlake (19220:
152).
REFERENCE. Revision: Timberlake (19220: 142-161).
BIOLOGY. Hyperparasites of Auchenorrhyncha (Homoptera) via Dryinidae (Hymenoptera).
INDO-PACIFIC ENCYRTIDAE 289
COMMENTS. We have examined the holotype ofAulonops bifasciata (BPBM) and conclude that
it must be regarded as belonging to the genus Hypergonatopus .
The genus is placed in the tribe Chalcerinyini by Trjapitzin (19736) but we think it would
probably be better placed in the Cheiloneurini. It is possible that future study will show that the
Chalcerinyini could be considered as a subtribe within the Cheiloneurini.
INDAPHYCUS Hay at
(Key couplet: 62. Figs 24, 357)
Indaphycus Hayat, 1981ft: 20. Type-species: Indaphycus plan us Hayat, by original designation.
DISTRIBUTION AND SPECIES. One species, India only: planus Hayat (19816: 21).
BIOLOGY. Unknown.
COMMENTS. It is possible that Pseudectroma bryanti Girault will run to Indaphycus in the key
since this species may have a solid clava. It will probably be possible to determine its correct
generic placement only when fresh material is available for study.
The genus is apparently close to Acerophagus and Pseudectroma (tribe Aphycini, subtribe
Aphycina) from which it can be separated using the characters given in the key, notably by the
elongate pronotum.
ISODROMOIDES Girault
(Key couplets: 453, 472)
Isodromoides Girault, 1914a: 30. Type-species: Isodromoides triangularis Girault, by original designation.
Neocopidosomyia Girault, 1915a: 95. Type-species: Neocopidosomyia viridiscutellum Girault, by original
designation. Syn. n.
DISTRIBUTION AND SPECIES. One species, Australia only: triangularis Girault (19140: 30)
(= Neocopidosomyia viridiscutellum Girault, 1915a: 95 syn. n.), also at least one further
species from Australia (BMNH).
BIOLOGY. Hyperparasites of Epipyropidae (Lepidoptera) parasitic on Fulgoridae (Homoptera).
COMMENTS. The genus appears to be related to Ooencyrtus (tribe Microteryini, subtribe
Oencyrtina) from which it can be separated by having a solid clava (that of Ooencyrtus is
three-segmented).
ISODROMUS Howard
(Key couplet: 389. Figs 200, 201)
Isodromus Howard, 1887: 488. Type-species: Isodromus iceryae Howard, by monotypy.
Parataneostigma Girault, 1915d: 275. Type-species: Parataneostigma nigriaxillae Girault, by monotypy.
DISTRIBUTION AND SPECIES. Fourteen species, cosmopolitan; one species from review area:
axillaris Timberlake (19196: 185) (China, Hawaiian Is.), also undetermined material from
Pakistan, Philippines and Australia (BMNH, BPBM).
REFERENCE. Revision: Timberlake (19196: 176-190).
BIOLOGY. Parasites of larvae of Chrysopidae and Hemerobiidae (Neuroptera).
COMMENTS. Placed in the tribe Homalotylini, subtribe Homalotylina (see also comments under
Aphycus).
KAKAOBURRA Girault
(Key couplet: 3 19)
Kakaoburra Girault, 1922a: 44. Type-species: Kakaoburra fera Girault, by monotypy.
290 J. S. NOYES & M. HAY AT
DISTRIBUTION AND SPECIES. Australia only, two described species: angeliconini (Girault,
1924a: 6) (comb. n. from Echthrobaccha) andfera Girault (19220: 44), also possibly two further
species from Australia and New Zealand (BMNH, DSIR).
BIOLOGY. Unknown.
COMMENTS. The single extant syntype (probably the holotype) otKakaoburrafera is in very poor
condition (QM) ; fragments of the head and one forewing on a slide and legs on a card are all that
remain. However, the wing venation, arrangement of the setae in the basal cell and relatively
short scape are very characteristic and we believe that we have interpreted the genus correctly by
assigning angeliconini to it.
Kakaoburra, as we understand it, is probably very close to Mayridia (subtribe Mayridiina)
which Trjapitzin (19736) has incorrectly placed in the tribe Miraini (see comments under
Ectroma, Mir a and Mayridia).
KATAKAgen.n.
(Key couplets: 223, 319. Figs 132, 358-367)
Type-species: Kataka mudigerensis sp. n. Gender: feminine.
$?. Head. In facial view (Fig. 358) about one-fifth broader than long, in profile slightly more than one and
one-half times as long as broad and anteriorly more or less gradually and evenly curved although slightly
flatter from top of scrobes to lowest level of toruli. Eye with posterior margin straight, about one-half
longer than broad, much shorter than minimum width of frontovertex; naked and separated from occipital
margin by at least about the diameter of an ocellus; occipital margin rounded. Malar space about
three-quarters as long as eye, with sulcus present although not well marked. Frontovertex slightly less than
two-thirds head width; ocelli forming an obtuse angle of about 120-130 the posterior ones about
equidistant from eye and occipital margin. Antennal scrobes very shallow, semicircular, meeting dorsally,
reaching slightly less than half way from antennal torulus to anterior ocellus; antennal torulus separated
from mouth margin by nearly twice its own length and from other torulus by about one-quarter more than
its own length, its lowest margin a little below lowest eye margin; clypeal margin shallowly but broadly
excised below toruli. Antennal scape about two-thirds as long as minimum width of frontovertex and about
four times as long as broad; pedicel conical, about half length of scape and a little longer than any funicle
segment, all of which are longer than broad, the first the longest and sixth shortest; clava three-segmented,
apically rounded with sutures parallel and about one-third length of funicle; longitudinal sensillae on all
flagellar segments, longest setae slightly shorter than diameter of corresponding segment. Sculpture on
frontovertex shallow, raised reticulate, fairly regular and almost hexagonal, on cheeks and between toruli
and eyes more elongate and tending towards squamiform-reticulate; setae on frontovertex and genae fairly
numerous, dark and short, each not longer than about the diameter of an ocellus. Mandible with two teeth
and a truncation or obscurely tridentate with upper tooth broadly rounded; maxillary palpus four-
segmented, labial palpus three-segmented.
Thorax. In side view moderately robust with mesoscutum and scutellum flat, the hind margin of the
mesopleurum clearly separating the hind coxa from the metapleurum and propodeum and more or less in
contact with basal gastral segment (Fig. 361) . In dorsal view with posterior margin of pronotum moderately
concave; visible part of mesoscutum about one-half broader than long, with notaular lines absent, its
posterior margin slightly concave medially; axillae meeting, scutellum a little broader than long and a little
longer than mesoscutum with apex acute; propodeum medially about one-seventh as long as scutellum and
medially with some shallow, irregular, reticulate sculpture. Dorsum of thorax with fairly regular, shallow,
raised, almost hexagonal sculpture, covered in numerous, conspicuous dark brown setae. Forewing more
or less hyaline but almost imperceptibly infused pale brown, about three times as long as broad; linea calva
not interrupted but closed near posterior margin of wing by two lines of setae; filum spinosum present;
submarginal vein without an apical hyaline break, parastigma not thickened; costal cell about 10 times as
long as broad, with one or two lines of setae dorsally along its entire length; marginal vein about four times
as long as broad, a little longer than stigmal and at least twice as long as postmarginal. Hindwing about
two-thirds as long as forewing and about five times as long as broad, marginal fringe about one-third as long
as wing width. Mid tibial spur a little longer than basal mid tarsal segment.
Caster. A little shorter than thorax; cereal plates in basal half; hypopygium with apex reaching to about
four-fifths along gaster, clothed in very long setae apically; paratergites absent; last tergite a little shorter
INDO-PACIFIC ENCYRTIDAE 291
than two-thirds mid tibia; ovipositor slightly shorter than mid tibia, gonostyli free, about one-sixth as long
as ovipositor.
Cf Very similar to female but differs as follows. Antennal scrobes very shallow, almost non-existant;
antennal toruli a little higher on head, their lowest margins about level with lowest eye margins; antennal
scape less than two-thirds as long as minimum width of frontovertex, about three and one-half times as long
as broad, pedicel about as long as each funicle segment all of which are clearly longer than broad; clava
entire, about twice as long as a funicle segment; longest setae on flagellum about as long as diameter of
segments. Forewing with linea calva open or closed by a single line of setae on dorsal surface. Genitalia
with aedeagus about half as long as mid tibia, digiti each with apical hook and about one-seventh as long as
aedeagus.
COMMENTS. The relatively high placement of the antennal toruli, rounded occipital margin, wing
venation and flattened thoracic dorsum indicate that this genus may be related to Mayridia (see
comments under Mayridia). It can be separated from this genus in having the setae on the dorsal
surface of the forewing extending to the base, the mandible having the third (upper) tooth more
or less truncate, relatively smaller eyes and posteriorly enlarged mesopleurum (in profile, the
hind coxa is in contact with the propodeum in Mayridia).
Kataka mudigerensis sp. n.
(Figs 132, 358-367)
$. Length: 0-98-1-22 mm (holotype, 1-22 mm).
Colour. Head, dorsum of thorax and gaster black with some slight green, brassy and purple reflections
on head, purple reflections on mesoscutum, axillae and basal two-thirds or so of scutellum; apical one-third
or so of scutellum green, gaster basally with a metallic green sheen, strongly coppery-purple over
remainder; antennal scape, mesopleurum and legs yellowish orange, antennal pedicel and flagellum
brown.
Head. Facial view as in Fig. 358. Relative measurements (holotype): head length 58, head width (facial
view) 69, head width (side view) 35, minimum frontovertex width 40, malar space 25, eye length 33, eye
width 22, POL 22, OOL 8, scape length 30, scape width 8, other proportions of antenna as in Fig. 359,
mandible as in Fig. 360.
Thorax. Relative measurements (holotype): forewing length 170, width 61, proportions of veins as in
Figs 132, 362; hindwing length 119, width 24. Basal cell of forewing with setae to base and nearly twice as
dense as in distal half of wing; propodeum with 10-12 very long translucent setae immediately outside each
spiracle, each seta about twice as long as diameter of spiracle.
Gaster. Relative lengths (paratype): last tergite 23, ovipositor 31-5, gonostyli 4-5, [mid tibia 38];
genitalia as in Fig. 364, hypopygium as in Fig. 363.
Cf . Length: 0-89-0-92 mm. Generally similar to female except for antenna (Fig. 367) and genitalia (Figs
365, 366). Relative measurements (paratype): scape length 33, scape width 10, minimum width of
frontovertex 48, length of mid tibia 89, length of aedeagus 40.
DISTRIBUTION. India.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $, India: Karnataka, Mudigere, 26.x-4.xi. 1979 (J. S. Noyes) (BMNH).
Paratypes. India: 4 $, 3 cT, same data as holotype (BMNH).
LAKSHAPHAGUS Mahdihassan
(Key couplet: 97. Figs 46, 77, 368)
Lakshaphagus Mahdihassan, 1931: 170. Type-species: Microterys hautefeuilli Mahdihassan, by original
designation.
Cheilonicetus Shafee, Alam & Agarwal, 1975: 55. Type-species: Cheilonicetus daulai Shafee, Alam &
Agarwal, by original designation.
292 J. S. NO YES & M. HAY AT
DISTRIBUTION AND SPECIES. Three species, all from India: daulai (Shafee, Alam & Agarwal,
1975: 58),fusiscapus (Agarwal, 1965: 63) and hautefeuilli (Mahdihassan; Hayat, 1981: 22).
REFERENCE. Review: Hayat (19816: 21-23).
BIOLOGY. Parasites of Asterolecaniidae and Keriidae (Homoptera).
COMMENTS. The genus is superficially very similar to Atropates Howard and Anisophleps
Fidalgo. It can be separated from Atropates using the characters provided by Hayat (19816), and
from Anisophleps by the following: forewing lacking subapical fuscous band, mesoscutum
lacking posterior depression, whereas in Anisophleps the forewing has a subapical fuscous band
and the mesoscutum has a transverse posterior depression as in Diversinervus .
Lakshaphagus is probably close to Cerapterocerus (tribe Cerapterocerini) from which it
differs in lacking a flattened flagellum and weaker, less well-defined infuscate pattern of the
forewing.
LAMENNAISIA Girault
(Key couplets: 322, 532. Fig. 188)
Lamennaisia Girault, 1922a: 40. Type-species: Lamennaisia quadridentata Girault, by monotypy.
Mercetencyrtus Trjapitzin, 1963: 886. Type-species: Encyrtus ambiguus Nees, by original designation.
Syn. n.
Sabirella Agarwal, Agarwal & Khan, 1980: 30. Type-species: Sabirella indica Agarwal, Agarwal & Khan,
by original designation. Syn. n.
DISTRIBUTION AND SPECIES. Four species, cosmopolitan; three from review area: ambigua (Nees;
Mercet, 1921: 283) (comb. n. from Encyrtus) (India), indica (Agarwal, Agarwal & Khan, 1980:
30)(comb. n. from Sabirella) (India) and quadridentata Girault (19220: 40) (Australia), also
undetermined material from India and S. China to Australia, New Zealand and Hawaiian Is.
(BMNH, BPBM, DSIR, HC).
BIOLOGY. Unknown, but possibly some of the New Zealand material was reared from larvae of
Lathridiidae (Coleoptera).
COMMENTS. We are unable to place the genus according to Trjapitzin's (19736) classification of
the Encyrtinae.
LEEFMANSIA Waterston
(Key couplet: 383)
Leefmansia Waterston, 19286: 527. Type-species: Leefmansia bicolor Waterston, by original designation.
DISTRIBUTION AND SPECIES. One species only, Australasia: bicolor Waterston (19286: 528)
(Moluccas, Bismarck Archipelago, New Hebrides).
BIOLOGY. Parasites of eggs of Tettigoniidae (Orthoptera).
COMMENTS. The genus is superficially extremely similar to Microterys, but we believe that it
should be treated as distinct. It can be separated from Microterys by the characters given in the
key, notably the axillae being separated by the posterior margin of the mesoscutum. This
character, together with the fact that the only included species is parasitic in the eggs of other
insects, leads us to believe that the genus is more closely related to Ooencyrtus (Microteryini,
Ooencyrtina) than to Microterys.
LEPTOMASTIDEA Mercet
(Key couplets: 149, 245, 276. Fig. 163)
Leptomastidea Mercet, 19166: 112. Type-species: Leptomastidea aurantiaca Mercet, by monotypy.
INDO-PACIFIC ENCYRTIDAE 293
Tanaomastix Timberlake, 1918: 362. Type-species: Paraleptomastix abnormis Girault, by original desig-
nation.
Leptanusia De Santis, 1964: 80. Type-species: Leptomastidea pseudococci Brethes, by original desig-
nation.
DISTRIBUTION AND SPECIES. Seventeen species, cosmopolitan; two from review area: abnormis
(Girault, 19156: 184) (Hawaiian Is.) and shafeei Hayat & Subba Rao (1981: 114) ( (= indica
Shafee, Alam & Agarwal, 1975: 24) (India), also at least four further species amongst
undetermined material from India, Philippines and Australia (BMNH).
REFERENCE. Mercet (1924: 252-258).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Leptomastidea is in the tribe Anagyrini, subtribe Anagyrina (= Leptomastideina
syn. n.). The genus is very close to Gyranusoidea on one hand and Leptomastix on the other. It
can be separated from Gyranusoidea by the sculpture of the head and thorax and the degree of
infuscation of the forewing (see comments under Gyranusoidea), and from Leptomastix by its
generally smaller size and relatively shorter funicle segments in relation to the pedicel (see also
Kerrich, 1982:402).
LEPTOMASTIX Forster
(Key couplets: 230, 276. Figs 164, 168)
Leptomastix Forster, 1856: 34. Type-species: Leptomastix histrio Mayr, by subsequent reference of Mayr
(1876: 730).
Sterrhocoma Forster, 1856: 36. Type-species: Sterrhocoma histrio Forster, by original designation.
Stenoterys Thomson, 1876: 115. Type-species: Stenoterys orbitalis Thomson, by monotypy.
DISTRIBUTION AND SPECIES. Thirty-three species, cosmopolitan; 13 from review area: aligarhensis
Khan & Shafee (1975: 194) (India), auraticorpus Girault (19150: 152) (Australia), bre-
vipedicelus Khan & Shafee (1975: 194) (India), brevis Hayat, Alam & Agarwal (1975: 14)
(India), dactylopii Howard; Dozier (1927: 267) (Pakistan, India, Hawaiian Is.), gunturiensis
Shafee ((1971: 49) (India), longicornis Khan & Shafee (1975: 195) (India), longiscapus Khan &
Agarwal (1976: 378) (India), nigritegulae Girault (1915a: 153) (Australia), nigrocoxalis Com-
pere (1928: 219) (India), salemensis Hayat, Alam & Agarwal (1975: 17) (India), singularis
Shafee (1971: 50) (India) and trilongifasciata Girault (1916c: 479) (Java), also undetermined
material, undoubtedly including several undescribed species, from India, Bangladesh, Laos,
China, Java, Philippines and Papua New Guinea (BMNH, BPBM, RMNH, GC, HC).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Leptomastix belongs to the tribe Anagyrini, subtribe Anagyrina and is most closely
related to Leptomastidea and Apoleptomastix . It can be separated from these genera by the
characters given in the key (see also Kerrich, 1982: 402).
LEUROCERUS Crawford
(Key couplet: 112. Fig. 53)
Leurocerus Crawford, 1911: 276. Type-species: Leurocerus ovivorus Crawford, by original designation.
DISTRIBUTION AND SPECIES. Two species known, both from review area: hongkongensis Subba
Rao (1971: 220) (S. China, Hong Kong) and ovivorus Crawford (1911: 277) (Philippines,
Sumatra).
BIOLOGY. Parasites of eggs of Amathusiidae and Satyridae (Lepidoptera).
COMMENTS. The genus is placed in the tribe Microteryini, subtribe Ooencyrtina which may be
correct, although it does appear to have some affinities with Pentelicus (see comments under
Pentelicus), Proleurocerus (placed in the tribe Proleurocerini by Trjapitzin, 19736), Zozoros,
294 J. S. NO YES & M. HAY AT
Paksimmondsius and Proleuroceroides . It may be that these genera form a single group (or
tribe), but it would require a further, more detailed morphological study to determine their true
relationship. All these genera have similar forewing venation and a characteristic naked streak
from the apex of the postmarginal vein to the apex of the stigmal vein (this character is also found
in some genera of the Cerapterocerini and Cheiloneurini).
LUTHERISCA Ghesquiere
(Key couplets: 123, 483, 490. Figs 239, 240)
Lutheria Girault, 1919a: 166. Type-species: Lutheria ajanea Girault, by original designation. [Homonym
of Lutheria Hofsten, 1907.]
Lutherisca Ghesquiere, 1946: 369. [Replacement name for Lutheria Girault.]
DISTRIBUTION AND SPECIES. One species, Singapore and possibly Borneo: ajanea (Girault, 1919a:
167).
BIOLOGY. Unknown.
COMMENTS. The single, extant, syntype of Lutheria ajanea (BMNH) is here designated
LECTOTYPE. It is lacking the head which presumably was put on a microscope slide by
Girault and has since been lost. The specimen from Borneo (BPBM) is almost certainly con-
specific, but differs slightly from the lectotype in colour and sculpture of the dorsum of the
thorax.
The genus is closely related to Taftia (tribe Chrysoplatycerini, subtribe Taftiina) and can be
separated from this genus by the characters given in the key, notably the relatively larger clava
(Fig. 239).
MAHENCYRTUS Masi
(Key couplets: 253, 304, 350, 417. Figs 147, 210)
Mahencyrtus Masi, 19176: 157. Type-species: Mahencyrtus occultans Masi, by monotypy.
Tyndarichoides Mercet, 1921: 649. Type-species: Tyndarichoides metallicus Mercet, by original desig-
nation. [Homonym of Tyndarichoides Girault, 1920.] Syn. n.
Protyndarichus Mercet, 19236: 479. [Replacement name for Tyndarichoides Mercet.] Syn. n.
DISTRIBUTION AND SPECIES. Five species, cosmopolitan; three from review area, all Australian:
aereifemur (Girault, I922e: 150) (comb. n. from Echthrogonatopus) , gracilis (Girault, 1915a:
100) (comb. n. from Zarhopaloides) and longifasciatipennis (Girault, 1915a: 100) (comb. n. from
Zarhopaloides), also further material, containing at least one undescribed species, from India,
S. China, Hong Kong, Thailand, Malaysia, Java, Philippines and Australia (BMNH, BPBM).
BIOLOGY. Unknown.
COMMENTS. The holotype of Mahencyrtus occultans has been examined (BMNH). It is a typical
male of the genus previously known as Protyndarichus. The genus has been synonymised with
Parechthrodryinus by Trjapitzin & Gordh (19780) but after studying material belonging to these
two genera we believe that Protyndarichus and thus Mahencyrtus is a valid genus. It differs from
Parechthrodryinus by the characters given in the key, but most notably by general body shape
and also by the structure of the propodeum and scutellum. We propose the following transfers to
Mahencyrtus for extra-limital species: comara Walker (from Encyrtus) and nitidus Howard
(from Encyrtus) (both comb. n.).
The genus is placed in the tribe Cheiloneurini (Encyrtinae).
MANICNEMUS Hayat
(Key couplet: 242. Figs 145, 369, 370)
Manicnemus Hayat, 19816: 23. Type-species: Tetralophidea indica Mani & Saraswat, by original des-
ignation.
INDO-PACIFIC ENCYRTIDAE 295
DISTRIBUTION AND SPECIES. Afrotropical, Oriental, Australasian; one described species: indicus
(Mani & Saraswat; Hayat, 1981&: 24) (India), also undetermined material from S. China, Hong
Kong, Indonesia, Irian Jaya and Bismarck Archipelago (BPBM).
BIOLOGY. Unknown.
COMMENTS. The genus belongs in the tribe Charitopidini (Tetracneminae) and can be separated
from related genera by the strongly infuscate forewings, relatively short marginal vein of the
fore wing and very transverse head in dorsal view.
MARXELLA Girault
(Key couplet: 48)
Marxella Girault, 1932a: 6. Type-species: Marxella richteri Girault, by monotypy.
DISTRIBUTION AND SPECIES. One species, Australia only: richteri Girault (1932a: 6).
BIOLOGY. Unknown.
COMMENTS. The genus is very close to Acerophagoides and Coccidoxenoides (Tetracneminae,
tribe Pauridiini) but differs in having a three-segmented funicle in the female, whereas the other
genera have a five- and six-segmented funicle respectively.
MASHHOODIA Shafee
(Key couplets: 266, 407. Fig. 159)
Mashhoodia Shafee, 1972: 159. Type-species: Mashhoodia indica Shafee, by original designation.
DISTRIBUTION AND SPECIES. One species, India only: indica Shafee (1972: 160).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The genus belongs to the tribe Anagyrini, subtribe Anagyrina (Tetracneminae) and
differs from other included genera in the characteristic venation of the forewing (more or less
punctiform marginal vein and relatively long postmarginal and stigmal veins) and the patterns of
light and dark setae on the forewing (Fig. 159).
MASHHOODIELLA Hayat
(Key couplets: 257, 378. Fig. 194)
Mashhoodiella Hayat, 1972: 209. Type-species: Mashhoodiella echthromorpha Hayat, by original des-
ignation.
DISTRIBUTION AND SPECIES. One species, India only: echthromorpha Hayat (1972: 210).
BIOLOGY. Parasites of Coccidae (Homoptera).
COMMENTS. Placement of this genus is difficult but it should belong in the tribe Aphycini
(Encyrtinae) and appears to have some characters in common with Parablastothrix (see
comments under Parablastothrix).
MA Ffl/D/A Mercet
(Key couplets: 191, 333, 435, 473)
Mayridia Mercet, 1921: 426. Type-species: Mayridia pulchra Mercet, by original designation.
Superprionomitus Mercet, 1921: 376. Type-species: Superprionomitus procerus Mercet, by original
designation.
Indoencyrtus Hayat & Verma, 1978: 361. Type-species: Indoencyrtus caeruleus Hayat & Verma, by
original designation.
DISTRIBUTION AND SPECIES. Twenty-four species, cosmopolitan except perhaps the Neotropics;
296 J. S. NOYES & M. HAY AT
two species from review area: caerulea (Hayat & Verma, 1978: 362) (India) andpulchra Mercet
(1921: 431) (India), also undetermined material, containing at least one undescribed species,
from India, Sri Lanka, Hong Kong, Malaysia and Australia (BMNH, BPBM, HC).
BIOLOGY. Parasites of Aclerdidae and Pseudococcidae (Homoptera) and Cecidomyiidae
(Diptera).
COMMENTS. Trjapitzin (1973ft) places the genus in the subtribe Mayridiina which he incorrectly
places in the tribe Miraini. Mir a belongs to the subfamily Tetracneminae whilst Mayridia
belongs in the Encyrtinae. In our view, the most realistic solution to the problem this creates
would be the transfer of the subtribe Echthroplexiellina (also incorrectly included in the
Miraini) to the Aphycini and the Mayridiina to the Cheiloneurini.
MENISCOCEPHALUS Perkins
(Key couplets: 425, 441, 476)
Meniscocephalus Perkins, 1906: 249. Type-species: Meniscocephalus eximius Perkins, by monotypy.
Helmecephala Noyes, 1980: 200. Type-species: Helmecephala albisetosa Noyes, by original designation.
Syn. n.
DISTRIBUTION AND SPECIES. Four described species, Neotropical, Oriental, Australasian; two
species from review area: exflores (Trjapitzin, 19826: 1603-1604) (comb. n. from Helmecephala)
(Indonesia) and eximius Perkins (1906: 249) (Malaysia, Australia), also further undetermined
material, including several undescribed species, from India to Australia (BMNH, BPBM, QM).
BIOLOGY. Parasites of nymphs of Cicadellidae (Homoptera).
COMMENTS. We are unable to place the genus according to Trjapitzin's (1973ft) classification, but
it may be related to genera included in the tribe Prionomasticini (see Noyes, 1980: 202) or, as
Trjapitzin (1982ft) suggests, it may belong in the tribe Cheiloneurini, subtribe Tyndarichina.
MESANUSIA Girault
(Key couplet: 203)
Blatticida Girault, 1915a: 94. Type-species: Blatticida ashmeadi Girault, by original designation.
[Homonym of Blatticida Ashmead, 1904.] Syn. n.
Mesanusia Girault, I922d: 208. Type-species: Mesanusia latiscapus Girault, by monotypy.
Blatticidella Gahan & Pagan, 1923: 22. [Replacement name for Blatticida Girault.] Syn. n.
DISTRIBUTION AND SPECIES. Two described species, Australia only: ashmeadi (Girault, 1915a: 94)
(comb. n. from Blatticida) and latiscapus Girault (1922d: 208), also one further undescribed
species from Papua New Guinea (BPBM).
BIOLOGY. Parasites of eggs of cockroaches (Orthoptera, Blattodea) and Tettigoniidae (Orthop-
tera).
COMMENTS. Mesanusia speciosa (see p. 353) does not belong in this genus and we are unable to
place it with any degree of certainty.
Mesanusia is related to Ooencyrtus (Microteryini, subtribe Ooencyrtina) from which it can be
separated by the very large, obliquely truncate clava and relatively longer marginal vein of the
forewing.
MESASTYMACHUS Girault
Mesastymachus Girault, 1923c: 142. Type-species: Mesastymachus silvae Girault, by monotypy.
DISTRIBUTION AND SPECIES. One species, Australia only: silvae Girault (1923c: 142).
BIOLOGY. Unknown.
INDO-PACIFIC ENCYRTIDAE 297
COMMENTS. Unfortunately the type(s) oisilvae appear to be lost and therefore the generic and
specific names will have to be regarded as nomina dubia. However, when the Australian
encyrtid fauna is better known, it should be possible to recognise the genus and possibly even the
species from Girault's meagre description.
MESOCALOCERINUS Girault
(Key couplet: 255)
Mesocalocerinus Girault, 1922d: 206. Type-species: Mesocalocerinus gemmus Girault, by monotypy.
DISTRIBUTION AND SPECIES. One species, Australia only: gemmus Girault (I922d: 206).
BIOLOGY. Unknown.
COMMENTS. Very close to Cheiloneurus and quite possibly should be considered synonymous.
However, we are retaining it as a distinct genus until the genera belonging to this difficult group
can be studied in more detail. For the present it can be separated from Cheiloneurus on the
combination of the basal cell of the forewing being setose to the base, the scutellum lacking a
subapical tuft of setae and the basal segment of the gaster being orange and contrasting with the
dark remainder (in Cheiloneurus the basal cell is usually bare in its proximal half or so, the
scutellum usually has a subapical tuft of setae and the gaster is usually unicolorous; the
combination of these characters is never the same as in Mesocalocerinus).
MESORHOPELLA Girault
(Key couplet: 71)
Mesorhopella Girault, 1923c: 145. Type-species: Mesorhopella emersoni Girault, by monotypy.
DISTRIBUTION AND SPECIES. One species, Australia only: emersoni Girault (1923c: 145).
BIOLOGY. Unknown.
COMMENTS. The genus is extremely close to Pararhopella from which it differs by the characters
given in the key. Both genera probably belong in the tribe Trechnitini and can be distinguished
from related genera by the five-segmented funicle, very elongate clava (longer than funicle),
absence of notaular lines on the mesoscutum and relatively small size, not being much larger
than 0-75 mm (excluding ovipositor).
METAPHAENODISCUS Mercet
(Key couplet: 480. Fig. 237)
Metaphaeno discus Mercet, 1921: 626. Type-species: Metaphaeno discus nemoralis Mercet, by original
designation.
Keatsia Girault, 19280: 1. Type-species: Keatsia umbilicata Girault, by monotypy. Syn. n.
Ramalia Ferriere, 1953: 27. Type-species: Tetralophidea maxima Mercet, by original designation.
DISTRIBUTION AND SPECIES. Ten species, Palaearctic, Afrotropical, and Australasian; four from
review area: aligarhensis Hayat (19816: 25) (India), proximus (Hayat, 19816: 26) (India),
umbilicatus (Girault, 19280: 1) (comb. n. from Keatsia) (Australia) and yasumatsui Myartseva &
Trjaptzin (1979: 1238), also further undetermined material from S. China, Java and Australia
(BMNH, BPBM).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Belongs to the tribe Aenasiini (see comments under Aenasius) and distinguished
from related genera by the very broad, menisciform head and the relatively long marginal and
short postmarginal and stigmal veins of the fore wings.
298 J. S. NOYES & M. HAY AT
METAPHYCUS Mercet
(Key couplets: 180, 377. Fig. 104)
Metaphycus Mercet, 1917a: 138. Type-species: Aphycus zebratus Mercet, by monotypy.
Mercetiella Dozier, 1926: 98. Type-species: Mercetiella reticulata Dozier, by original designation.
Oaphycus Girault, 1932a: 5. Type-species: Aphycus sanguinithorax Girault, by original designation.
Syn. n.
Melanaphycus Compere, 1947a: 5. Type-species: Pseudococcobius fumipennis Timberlake, by original
designation.
Anaphycus Sugonjaev, 1960: 372. Type-species; Aphycus nitens Kurdjumov, by original designation.
Notoencyrtus De Santis, 1964: 211. Type-species: Notoencyrtus guttofasciatus De Santis, by original
designation.
DISTRIBUTION AND SPECIES. About 200 described species, cosmopolitan; 38 from review area:
agarwali Hayat & Subba Rao (1981: 115) (= latiscapus Shafee, Alam & Agarwal, 1975: 84)
(India), alberti (Howard; Annecke & Mynhardt, 1981: 38) (Hawaiian Is.), angustifrons
Compere (1957: 227) (Taiwan), argenteus (Girault, 1936: 1) (comb. n. from Aphycus) (Aus-
tralia), atriphragma (Girault, 1936: 1) (comb. n. from Aphycus) (Australia), bicinctitibiae
(Girault, 1932c: 1) (comb. n. from Aphycopsis) (Australia), bowenesis (Girault, 19320: 4)
(comb. n. from Aphycus) (Australia), buderimi (Girault, 1936: 1) (comb. n. from Aphycus)
(Australia), cerococci (Shafee, Alam & Agarwal, 1975: 83) (India), citricola Annecke &
Mynhardt (1971: 333) (Pakistan), claviger (Timberlake, 1916: 620) (New Zealand), crotolariae
(Shafee, Alam & Agarwal, 1975: 85) (India), flavus (Howard; = hesperidum Mercet, 1916: 784)
(India, Hawaiian Is.), fuscidorsum (Gahan, 1919: 521) (comb. n. from Aphycus) (India),
gontsharenkoi Trjapitzin & Khlopunov (1976: 98) (Vietnam), helvolus (Compere; Annecke &
Mynhardt, 1981: 42) (Pakistan, Bangladesh); indicus Shafee, Alam & Agarwal (1975: 79)
(India), ichneumon (Girault, 1936: 1) (comb. n. from Aphycus) (Australia), keatsi (Girault,
19320: 4) (comb. n. from Aphycus) (Australia), latiscapus Alam (1972: 134) (India), lichtensiae
(Howard; Compere & Annecke, 1961: 35) (Pakistan, India, Sri Lanka), longiclavatus (Shafee,
Alam & Agarwal, 1975: 88) (India) , lounsburyi (Howard, 18986: 244) (Hawaiian Is.), maculatus
Agarwal (1965: 89) (India) , malabarensis (Mukerjee in Saraswat & Mukerjee, 1975: 46) (India),
memnonicus Compere (1940: 46) (Australia), mexicanus (Howard, 18986: 247) (Hawaiian Is.),
nigrivarius (Girault, 19296: 313) (comb. n. from Aphycus) (Australia) , parkeri (Girault, 19320:
1) (comb. n. fromAenasomyiella) ( Australia), portoricensis (Dozier, 1926: 100) (Hawaiian Is.),
sanguinithorax (Girault, 19150: 112) (comb. n. from Aphycus) (Australia), semialbus (Girault,
1932a: 5) (comb. n. from Aphycus) (Australia) , stanleyi (Compere; Compere & Annecke, 1961:
33) (Hawaiian Is.), timberlakei (Ishii, 1923: 108) (New Zealand), tricinctus (Girault, 19310: 4)
(comb. n. from Aphycus) (Australia), turneri (Girault, 19320: 5) (comb. n. from Aphycus)
(Australia), varius (Girault, 19150: 178) (comb. n. from Aenasioidea) (Australia), verdini
(Girault, 1936: 1) (comb. n. from Aphycus) (Australia), zebratus (Mercet, 19170: 138) (India),
also much undetermined material from throughout the region (BMNH, BPBM, DSIR, QM,
ANIC,GC,HC).
REFERENCES. Timberlake (1916: 587-639); Afrotropical species, with descriptions of some
relevant species: Annecke & Mynhardt (1971; 1972; 1981); review of some Indian species:
Shafee etal. (1975:78-88).
BIOLOGY. Parasites of Coccidae, Diaspididae, Keriidae, Asterolecaniidae and Eriococcidae
(Homoptera).
COMMENTS. Mesanusia speciosa Girault (see p. 353) may also belong in this genus, but this will
only be confirmed when fresh material is collected.
Metaphycus argenteus and iohneumon may be synonymous, also crotolariae may be synony-
mous with fuscidorsum.
The genus is placed in the tribe Aphycini, subtribe Paraphycina (Encyrtinae) (see comments
under Paraphycus) .
INDO-PACIFIC ENCYRTIDAE 299
MICROTEK YS Thomson
(Key couplets: 94, 102, 139, 237, 261, 383)
Sceptrophorus Forster, 1856: 34. Type-species: Encyrtus sceptriger Forster, by designation of Ashmead
(19006: 381) [Suppressed in favour of Microterys Thomson: Opinion 1110, 1978, Opin. Decl. int.
Commn zool. Norn. 35: 99-100.]
Microterys Thomson, 1876: 155. Type-species: Encyrtus sylvius Dalman, by designation of Ashmead
(1900: 390).
Apentelicus Fullaway, 1913: 26. Type-species: Apentelicus kotinskyiFu\\away, by original designation.
DISTRIBUTION AND SPECIES. Very nearly 150 described species, cosmopolitan; 20 from review
area: anomalococci Shafee, Alam & Agarwal (1975: 69) (India), aristotelea (Girault; Prinsloo,
19766: 414) (Australia), australicus Prinsloo (19766: 420) (Australia), dauseni Compere (1926:
35) (Pakistan), ditaeniatus Huang (1980: 432, 434) (S. China), flavus (Howard; = frontalis
Mercet, 1921: 413) (Pakistan, India, Sri Lanka, Australia, New Zealand, Hawaiian Is.),
garibaldia (Girault; Prinsloo, 19766: 417) (Australia), gilberti (Girault; Prinsloo, 19766: 411)
(Australia), hesperidum (Trjapitzin & Khlopunov, 1976: 101) (Vietnam), indicus Subba Rao
(1977: 13) (India), kerrichi Shafee, Alam & Agarwal (1975: 69) (India), lichtensiae (Howard in
Howard & Ashmead, 1896: 636) (Sri Lanka), longifuniculus (Girault, 19326: 1) (Australia),
mirzai Shafee, Alam & Agarwal (1975: 67) (India), newcombi (Girault; Prinsloo, 19766: 413)
(Malaysia, Australia), nietneri (Motschulsky, 1859: 170) (Sri Lanka) , purpureiventris (Girault;
Prinsloo, 19766: 417) (Australia), sinicus Jiang (1982: 180, 186) (S. China), spinozai (Girault;
Prinsloo, 19766: 415) (Australia) and triguttatus (Girault; Prinsloo, 19766: 414) (Australia), also
much undetermined material, including several undescribed species, from throughout the
region (BMNH, BPBM, QM, ANIC, USNM, HC).
REFERENCES. World catalogue: Rosen (1976); review of some species: Shafee et al. (1975:
65-71); review of Australian species: Prinsloo (19766).
BIOLOGY. Parasites of Coccidae, Kermococcidae and Lecaniodiaspididae (Homoptera).
COMMENTS. One undescribed species from Brunei (BMNH) has an apical tuft of setae on the
scutellum similar to that found in Cheiloneurus .
The types of Microterys coeruleus Bingham (HDOU) have been examined by Z. Boucek who
informs us that the female belongs to Tetrastichus (Eulophidae) and the male to Anastatus
(Eupelmidae).
Placed in the tribe Microteryini, subtribe Microteryina (Encyrtinae).
MIRA Schellenberg
(Key couplet: 77)
Mir a Schellenberg, 1803: 68. Type-species: Mir a mucora Schellenberg, by monotypy.
Dicelloceras Menzel, 1855: 270. Type-species: Dicelloceras vibrans Menzel, by monotypy.
Euryscapus Forster, 1856: 32. Type-species: Encyrtus platycerus Dalman, by original designation.
Lonchocerus Dahlbom, 1857: 292. Type-species: Encyrtus platycerus Dalman, by subsequent reference of
Thomson (1876: 130).
Euzkadia Mercet, 1921: 552. Type-species: Euzkadia integrate Mercet, by original designation.
DISTRIBUTION AND SPECIES. Four described species, Palaearctic; one undescribed species from
Australia (BMNH, UCR).
REFERENCE. Revision of European species: Boucek (19776: 141-146).
BIOLOGY. Unknown.
COMMENTS. The genus has been incorrectly placed in the subfamily Encyrtinae by Trjapitzin
(19736). It is actually very close to the genera included in the tribe Charitopidini (Tetracnemi-
nae).
300 J. S. NO YES & M. HAYAT
MONSTRANUSIA Trjapitzin
(Key couplet: 118)
Monstranusia Trjapitzin, 19646: 243. Type-species: Monstranusia mirabilissima Trjapitzin, by original
designation.
DISTRIBUTION AND SPECIES. Two species, Palaearctic, Afrotropical, Oriental; both known from
Oriental region: antennata (Narayanan, 1960: 122) (India) and mirabilissima Trjapitzin (19646:
245) (Pakistan).
BIOLOGY. Unknown.
COMMENTS. Placed in the tribe Anagyrini, subtribe Anusiina (Tetracneminae) by Trjapitzin
(19730), but possibly this may be incorrect since it also shows many characters in common with
the Tetracnemini, e.g. forewing venation and infuscation.
MOZARTELLA Girault
(Key couplet: 65)
Mozartella Girault, 1926a: 1. Type-species: Mozartella beethoveni Girault, by monotypy.
DISTRIBUTION AND SPECIES. One species, Australia only: beethoveni Girault (19260: 1).
BIOLOGY. Apparently reared from plant galls.
COMMENTS. The genus superficially appears to be very close to Pseudectroma and Acerophagus
(tribe Aphycini, subtribe Aphycina) but differs in the mandible having two teeth and a broad
truncation instead of three teeth. This character may indicate a relationship with Aphyco-
morpha which has been placed in the tribe Aphycini by Trjapitzin (19736) but which we now
believe belongs in the Microteryini (see comments under Aphycomorpha).
MULUENCYRTUSgen. n.
(Key couplet: 458. Figs 215, 216, 371-373)
Type-species: Muluencyrtus nudipennis sp. n. Gender: masculine.
9- Head. In frontal view slightly broader than long and in side view about twice as long as broad and
anteriorly more or less gradually and evenly rounded. Eye relatively small with sparse, short, inconspi-
cuous setae each not longer than the diameter of a facet, posterior margin of eye straight, eye a little less
than one-third longer than broad, not quite reaching occipital margin which is sharply carinate behind
ocelli. Malar space slightly longer than eye, sulcus absent, but more or less indicated by a change of
sculpture; mouth opening relatively small, about two-fifths head width. Frontovertex about two-fifths head
width; ocelli in a right angle, the posterior ones a little closer to eye margin than to occipital margin,
separated from the latter by about their own major diameters. Antennal scrobes only a little longer than
toruli, gently curved inwards but not meeting dorsally, reaching about half way from antennal toruli to
anterior ocellus; antennal torulus separated from mouth margin by slightly more than its own length and
from other torulus by about one and one-half times its own length, its dorsal margin a little below ventral
eye margin; clypeus very shallowly excised, but broadly concave. Antennal scape distinctly longer than
width of frontovertex, subcylindrical, slightly more than four and one-half times as long as broad, with a
pair of short flanges on its lower surface either side of pedicel and clothed in conspicuous long setae ; pedicel
cylindrical, about one-third length of scape, shorter than any of the first three funicle segments but longer
than any of the last three ; clava entire , transversely truncate about one-third length of funicle ; longitudinal
sensillae on all flagellar segments except perhaps the first two; setae on flagellum moderately long, the
longest being slightly longer than diameter of first funicle segment. Frontovertex above scrobes with
shallow, raised, squamiform-reticulate sculpture; between scrobes and eyes and interantennal prominence
dorsally with rough, raised, irregular sculpture, this continuing onto lower parts of face but becoming
shallower and more longitudinally elongate; setae on frontovertex and, to a less extent, on lower parts of
INDO-PACIFIC ENCYRTIDAE 301
face but borne in shallow, but distinct punctures; setae moderately sparse, short and inconspicuous on
frontovertex above toruli, but longer and more conspicuous on genae and lower parts of interantennal
prominence. Mandible broad with two teeth and a broad truncation; maxillary palpus four-segmented,
labial palpus three-segmented.
Thorax. Moderately robust in side view, with mesoscutum more or less flat but slightly convex anteriorly
and scutellum very slightly convex; the metapleurum and propodeum together narrowly in contact with
hind coxa. In dorsal view pronotum moderately concave posteriorly, with spiracles clearly visible laterally;
visible part of mesoscutum slightly less than twice as broad as long, with notaular lines absent, its hind
margin slightly produced distad; axillae slightly separated; scutellum slightly shorter than mesoscutum and
slightly broader than long, its apex more or less acute; propodeum medially a little less than one-quarter
length of scutellum. Dorsum of thorax with very shallow, raised, squamiform-reticulate sculpture;
mesopleurum more or less smooth and shiny; propodeum more or less smooth; dorsum of thorax clothed
with very short, inconspicuous dark setae. Fore wing infuscate, about two and one-half times as long as
broad; linea calva not interrupted or closed; dorsal surface of forewing with extremely sparse discal setae,
apart from a few setae in basal cell and filum spinosum almost naked dorsally, those on dorsal surface
clearly much less dense than those ventrally; submarginal vein without an apical hyaline break, parastigma
not swollen; costal cell about nine or ten times as long as broad, with only one seta dorsally near apex;
marginal vein about twice as long as broad, about as long as postmarginal and a little longer than half
stigmal vein. Hindwing about two-thirds length of forewing, very slightly infumate, about three and
one-half times as long as broad ; marginal fringe about one-ninth as long as maximum wing width . Mid tibial
spur about one-half as long as basal mid tarsal segment; all legs very smooth; hind and mid tarsi very
characteristic, being very smooth and shiny and gradually tapering towards apex; hind tibia with apex
strongly oblique (Fig. 216).
Gaster. About as long as thorax; cereal plates in apical one-third; hypopygium with apex reaching to
about three-quarters along gaster ; ovipositor not or only slightly exserted ; paratergites presumably absent.
CT. Unknown.
COMMENTS. The affinities of this interesting new genus are not at all clear, but the head shape and
structure, forewing venation and positioning of the cereal plates suggest that it should be placed
near Encyrtus, Olypusa or perhaps Eucomomorphella. It can be easily separated from these
genera by the nearly naked dorsal surface of the forewing and from Encyrtus and Olypusa by the
smaller eyes and presence of mandibular teeth. The presence of mandibular teeth suggest that it
is closest to Eucomomorphella, but this genus has a very long postmarginal vein and three-
segmented clava (see comments under Anagyrodes).
Muluencyrtus nudipennis sp. n.
(Figs 215, 216, 371-373)
$. Length, 2-06 mm.
Colour. Body, including legs and antenna, reddish brown to dark reddish brown; apex of clava whitish
yellow; between each eye and occiput more or less orange-brown; mesoscutum with a slight bluish sheen;
scutellum slightly purplish brassy; scutellum laterally either side of apex, metanotum and propodeum
medially brown; forewing infuscate as in Fig. 373.
Head. Frontal view as in Fig. 372. Relative measurements: head length 85, head width (frontal view)
100, head width (side view) 47, minimum width of frontovertex 39, POL 16, OOL 5, malar space 46, length
of eye 43, width of eye 37, scape length 55, other proportions of antenna as in Fig. 371.
Thorax. Relative measurements: forewing length 220, hindwing length 153, hindwing width 45; forewing
as in Fig. 373, venation as in Fig. 215, hind tibia and tarsus as in Fig. 216.
Gaster. Relative lengths: last tergite 46, [mid tibia 132].
Cf . Unknown.
DISTRIBUTION. Sarawak.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $, Sarawak: Gunong Mulu National Park, 26. v. 1978 (N. M. Collins} (BMNH).
302 J. S. NO YES & M. HAYAT
NASSAUIA Girault
(Key couplet: 50)
Nassauia Girault, 1932a: 5. Type-species: Nassauia atoma Girault, by monotypy.
DISTRIBUTION AND SPECIES. Two species, Australia only: atoma Girault (1932a: 5) and secunda
Girault (1932c: 3).
BIOLOGY. Reared from 'coccids' (Girault, 1932c: 3).
COMMENTS. The genus is near Metaphycus (tribe Aphycini, subtribe Paraphycina) and differs
from all related genera in having a four-segmented funicle.
NATHISMUSIAgen. n.
(Key couplet: 37. Figs 214, 374-376)
Type-species: Nathismusiasouthwoodisp. n. Gender: feminine.
$ . Head. In facial view about one-quarter wider than long, with mouth opening very wide, about half head
width, in side view head about two-thirds longer than wide and anteriorly more or less gradually curved.
Eyes very small with posterior margin straight, almost oval, a little longer than broad, naked; eye separated
from occipital margin, which is rounded, by at least about twice the diameter of an ocellus. Malar space
about as long as an eye with sulcus absent but marked by a slight change of sculpture. Frontovertex about
three-quarters head width, ocelli forming an obtuse angle of about 110, the posterior ocelli separated from
occipital margin by a little more than their diameter but from eye margin by about five times their diameter.
Antennal scrobes extremely shallow, broadly semicircular, more or less meeting dorsally and reaching
about halfway from toruli to anterior ocellus; antennal torulus separated from mouth by a little less than its
own length and from other torulus by about one and one-half times its own length, its upper margin a little
above the lower eye margin; clypeus broadly but shallowly excised below and between toruli. Antennal
scape a little longer than half minimum width of frontovertex, stout, a little more than three times as long as
broad; pedicel conical, nearly half length of scape and clearly longer than any funicle segment, all
subquadrate, the first the longest, the sixth the shortest; clava three-segmented, its apex rounded, sutures
between segments subparallel; longitudinal sensillae on all flagellar segments; longest setae not much more
than one-third as long as diameter of segments. Frontovertex with extremely shallow, raised reticulate
sculpture, this becoming more elongate on lower parts efface and genae; setae on head moderately dense,
very short, inconspicuous. Mouth parts not clearly visible in single specimen available.
Thorax. Moderately deep in side view, but conspicuously dorso-ventrally flattened, the mesoscutum and
scutellum almost completely flat, the anterior face of the pronotum almost perpendicular, the meta-
pleurum and propodeum fairly broadly in contact with hind coxa. In dorsal view hind margin of pronotum
shallowly concave ; visible part of mesoscutum slightly more than twice as broad as long, with notaular lines
absent and its hind margin almost straight, only very slightly produced backwards; axillae nearly touching
in middle, not clearly separated from scutellum which is nearly one-third longer than mesoscutum and
apically rounded; propodeum medially less than one-seventh as long as scutellum. Dorsum of thorax with
similar sculpture to frontovertex but a little deeper on posterior part of mesoscutum than on scutellum;
mesopleurum with shallow, raised, slightly elongate, reticulate sculpture; propodeum almost completely
smooth, but with some irregular sculpture near spiracle and laterally; setae on dorsum of thorax sparse,
dark and short, very inconspicuous. Forewing lightly infused greyish brown, with a small dark brown spot
immediately below marginal vein, wing nearly three times as long as broad with marginal fringe very short,
nearly absent; linea calva completely obliterated by setae on both surfaces of wing; filum spinosum absent;
setae in disc of forewing very dense , short and evenly distributed except near extreme base where they are
virtually absent; submarginal vein with an indistinct hyaline break, parastigma hardly swollen; costal cell
about 11 times as long as broad and with one line of setae dorsally in its proximal one-third and two or three
lines of setae in its distal two-thirds or so; marginal vein thick, about twice as long as wide, longer than
stigmal vein which in turn is longer than postmarginal. Hindwing lightly infused pale greyish brown, about
two-thirds as long as forewing; marginal setae about one-seventh as long as maximum wing width. Mid
tibial spur a little shorter than mid basal tarsal segment.
Caster. Slightly longer than thorax with cereal plates in posterior half; hypopygium reaching apex of
gaster; last tergite a little less than one-third as long as mid tibia; paratergites not visible, probably absent;
ovipositor hardly exserted.
INDO-PACIFIC ENCYRTIDAE 303
Cf . Unknown.
COMMENTS. Placement of this genus is difficult, although it must certainly belong to the
Encyrtinae. It may be related to Coelopencyrtus (tribe Copidosomatini, subtribe Ceolopencyr-
tina) since there are some similarities in head shape, fore wing venation and structure of the
gaster. However, it can easily be separated on the relatively small eyes, high placement of
antennal toruli, lack of linea calva and very short last abdominal tergite.
The type-species of the genus is named in honour of Professor Sir T. R. E. Southwood.
Nathismusia southwoodi sp. n.
(Figs 214, 374-376)
$. Length (holotype): 1-16 mm.
Colour. Generally dark brown; antenna yellowish brown; wings suffused very pale greyish brown, dark
brown immediately below marginal vein of forewing; all coxae dark brown, femora dark brown, fore and
mid femora apically yellow, tibiae and tarsi yellow, pretarsi dark brown.
Head. Relative measurements: head length 45, head width (facial view) 56, head width (side view) 27,
minimum width of frontovertex 42, malar space 18, eye length 17-5, eye width 14, POL 13-5, OOL 16-5,
scape length 23, scape width 7, proportions of antenna as in Fig. 374, head in facial view as in Fig. 375.
Thorax. Relative measurements: forewing length 165, forewing width 62, venation as in Figs 214, 376;
hindwing length 115, hindwing width 27. Base of forewing as in Fig. 214.
Gaster. Relative lengths: last tergite 21, [mid tibia 62].
Cf . Unknown.
DISTRIBUTION. India.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $, India: Hyderabad, Patancheru, ICRISAT, vii-ix.1980, Malaise trap (Bernays &
Woodhead) (BMNH).
NEABROLEPOIDEUS Girault
(Key couplet: 255)
Neabrolepoideus Girault, 1917g: 140. Type-species: Neabrolepoideus bioculatus Girault, by original
designation.
DISTRIBUTION AND SPECIES. One described species, Australia only: bioculatus Girault (1917g:
141), also one further undescribed species from Australia (BMNH).
BIOLOGY. Unknown.
COMMENTS. The genus belongs to the tribe Cheiloneurini (Encyrtinae) and is very close to
Cheiloneurus from which it more or less differs only by the setae of the forewing reaching nearer
the base, the pattern of infuscation of the forewing, the scutellum being slightly more convex and
lacking the apical tuft of setae, and the body (especially the antenna) being more slender (see
comments under Cheiloneurus).
NEANAGYRUS Girault
(Key couplets: 495, 523)
Neanagyrus Girault, 1915a: 174. Type-species: Neanagyrus capitatus Girault, by original designation.
Anisodromus Riek, 1962c: 283. Type-species: Anisodromus tarsius Riek, by original designation. Syn. n.
DISTRIBUTION AND SPECIES. Three species, all Australian: capitatus Girault (19150: 174), niger
(Riek, 1962c: 285) (comb. n. from Anisodromus) and tarsius (Riek, 1962c: 283) (comb. n. from
Anisodromus).
304 J. S. NO YES & M. HAY AT
BIOLOGY. Parasites of lerp-forming Psyllidae (Homoptera).
COMMENTS. The genus is very close to Psyllaephagus (Trechnitini, subtribe Metaprionomitina)
and perhaps should be considered as a species-group within that genus since it differs only in the
structure of the antenna.
NEASTYMACHUS Girault
(Key couplets: 181, 312, 414. Fig. 101)
Neastymachus Girault, 1915a: 86. Type-species: Neastymachus auraticorpus Girault, by monotypy.
Nikolskiella Trjapitzin, 19626: 560. Type species: Microterys luteus Nikol'skaya, by original designation.
Syn. n.
Pseudmicroterys Shafee, Alam & Agarwal, 1975: 71. Type-species: Pseudmicroterys burski Shafee, Alam
& Agarwal, by original designation. Syn. n.
DISTRIBUTION AND SPECIES. Provisionally eight species, Palaearctic, Afrotropical, Oriental,
Australasian; five from review area: angustifrons (Shafee, Alam & Agarwal, 1975: 76) (comb. n.
from Pseudmicroterys) (India), auraticorpus Girault (1915a: 86) (Australia), burski (Shafee,
Alam & Agarwal, 1975: 73) (comb. n. from Pseudmicroterys) (India), cerococci (Shafee, Alam
& Agarwal, 1975: 76) (comb. n. from Pseudmicroterys) (India) and delhiensis (Subba Rao, 1957:
387) (comb. n. from Microterys) (India), also undetermined material from Sri Lanka, Hong
Kong and Malaysia (BMNH, BPBM).
REFERENCE. Review of some Indian species: Shafee etal. (1975: 71-78).
BIOLOGY. Parasites of Aclerdidae and Asterolecaniidae (Homoptera).
COMMENTS. Neastymachus auraticorpus Girault is very close to luteus Nikol'skaya (comb, n.),
differing only slightly in the relative proportions of the antennal segments, forewings and width
of fronto vertex.
We have examined paratypes of Nikolskiella japonica Tachikawa (1970: 100). The species
does not appear to belong in Neastymachus nor to any genus known to us. We have not seen any
material of Nikolskiella secunda Trjapitzin (1962ft: 565) and therefore defer formally transfer-
ring it to Neastymachus.
The genus is placed in the tribe Microteryini, subtribe Microteryina (Encyrtinae).
NEBLATTICIDA Girault
(Key couplet: 130)
Neblatticida Girault, 1915a: 96. Type-species: Neblatticida fasciatipes Girault, by original designation.
DISTRIBUTION AND SPECIES. Three species, all Australian: fasciatipes Girault (1915a: 96), lotae
(Girault, 1922ft: 106) (comb. n. from Baeoanusia) and perfuscipennis (Girault, 19150: 164)
(comb. n. from Baeoanusia).
BIOLOGY. Unknown.
COMMENTS. The generic placement of perfuscipennis is difficult. Girault originally included it in
Baeoanusia, no doubt because of the similarity in the structure of the mandible. It is very
probable that perfuscipennis belongs to neither Neblatticida nor Baeoanusia if these genera are
defined as narrowly as they are at present. However, perfuscipennis will run to Neblatticida in
the key to genera as it stands and therefore this necessitates the transfer of this species to
Neblatticida.
Both Neblatticida and Baeoanusia belong to the Cheiloneurini (Encyrtinae) and a detailed
study is required in order to achieve some idea of the natural grouping of the species, particularly
those in Australia, before the generic limits can be determined with any degree of certainty. We
feel that when such a study can be undertaken, very many of the genera recognised at present
will fall in synonymy (see also comments under Cheiloneurus) .
INDO-PACIFIC ENCYRTIDAE 305
NEGENIASPIDIUS Trjapitzin
(Key couplet: 322)
Negeniaspidius Trjapitzin, 1982a: 39. Type-species: Encyrtus nobilis Nees, by original designation.
DISTRIBUTION AND SPECIES. Only one described species, Palaearctic, Afrotropical and Oriental:
nobilis (Nees; = Coccidencyrtus pretiosus Mercet, 1921: 281) (India).
BIOLOGY. Unknown.
COMMENTS. It is possible that the species occurring in southern Africa (Zambia, Zimbabwe) is
distinct from nobilis (BMNH).
We are unable to place Negeniaspidius according to Trjapitzin's (19736) classification of the
Encyrtinae although quite possibly it is related to Lamennaisia.
NEOCHARITOPUSHayat, Alam & Agarwal
(Key couplet: 502. Figs 243, 244, 380)
Neocharitopus Hayat, Alam & Agarwal, 1975: 24. Type-species: Charitopus orientalis Agarwal, by
original designation.
Insleyia Prinsloo & Annecke, 1979: 377. Type-species: Insleyia crassa Prinsloo & Annecke, by original
designation. Syn. n.
DISTRIBUTION AND SPECIES. Two species, Afrotropical, Oriental; one from review area: orientalis
(Agarwal, 1965: 91) (India).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. We have compared some paratypes of Insleyia crassa with material of Neocharitopus
orientalis collected in India and find that they must be considered congeneric. Further study,
using fresh material of orientalis for comparison, may reveal that the two species are also
synonymous. The material of orientalis at hand is not in sufficiently good condition for reliable
comparison at this level.
The genus belongs in the tribe Charitopidini (Tetracneminae) and is superficially very similar
to Manicnemus from which it can be separated by the hyaline forewings and relatively short
propodeum (not more than about one-fifth length of scutellum). The forewings of Manicnemus
are strongly infuscate and the propodeum is about half as long as the scutellum.
NEOCLADELLA Girault
(Key couplets: 110, 186. Figs 110, 111)
Noecladella Girault, 1915a: 99. Type-species: Neocladella compressipes Girault, by original designation.
Pteromalencyrtus Girault, 1915a: 116. Type-species: Pteromalencyrtus quadridentatus Girault, by original
designation. Syn. n.
DISTRIBUTION AND SPECIES. One species, Australia only: compressipes Girault (19150: 99)
(= Pteromalencyrtus quadridentatus Girault, 1915a: 116 syn. n.).
BIOLOGY. Unknown.
COMMENTS. We favour the use of Neocladella as the valid generic name since the holotype of
compressipes is in much better condition than that of quadridentatus.
Placement of the genus is difficult , but the wing venation suggests that it very probably belongs
to the tribe Habrolepidini, subtribe Comperiellina. It is easily separated from genera included in
this subtribe by the quadridentate mandible, relatively high placement of the antennal toruli on
the head, relatively small scape and hyaline wings.
306 J. S. NOYES & M. HAY AT
NEOCLADIA Perkins
(Key couplet: 36. Fig. 213)
Neocladia Perkins, 1906: 251. Type-species: Neodadia howardi Perkins, by monotypy.
Phyllotibia Risbec, 1954: 1071. Type-species: Phyllotibia senegalensis Risbec, by monotypy.
DISTRIBUTION AND SPECIES. Seven species, Afrotropical, Oriental, Australasian; four from
review area: howardi Perkins (1906: 251) (Australia), indica (Agarwal, 1970: 25) (India),
shadsakus (Mani & Kaul in Mani etal. , 1973: 74) (India) and violacea Masi (1926: 272) (stat. n.
from subspecies of howardi) (Taiwan), also further undetermined material from India to Hong
Kong and Australia (BMNH, BPBM).
\J\ & I ^ f i/[ r
BIOLOGY. Parasites of nymphs of Cicadellidae (Homoptera).
COMMENTS. We have examined the holotype of Neocladia howardi violacea Masi (IPK). It
represents a valid species differing from howardi in the forewing venation (shorter marginal
vein and less curved stigmal).
Placed in the tribe Neocladiini (Encyrtinae) which is possibly too narrowly defined by
Trjapitzin (19736) (see comments under Anagyrodes). It can be separated from other closely
related genera by the combination of the enormously expanded and flattened hind tibia,
sickle-shaped mandible, forewing with relatively long marginal vein, three-segmented clava and
hypopygium more or less reaching the apex of the gaster.
NEODISCODES Compere
(Key couplets: 129, 210)
Neodiscodes Compere, 1931: 272. Type-species: Neodiscodes martinii Compere, by original designation.
DISTRIBUTION AND SPECIES. Seven species, Afrotropical, Oriental; four species from review area:
indicus Narayanan & Subba Rao (1960: 75) (India, Pakistan), lepelleyi Kerrich (1953: 794)
(India, Sri Lanka), parvus Kerrich (1967: 228) (S. China) and subbaraoi Kerrich (1967: 232)
(Hong Kong, Java).
REFERENCE. Revision: Kerrich (1967: 228-235).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Here placed in the tribe Aenasiini (Tetracneminae) (see comments under Aena-
sius). It is very closely related to Aenasius, but can be separated by the relatively narrower
frontovertex (see key, also Kerrich, 1967: 188-190).
NEODUSMETIA Kerrich
(Key couplet: 80. Figs 34, 377, 378)
Neodusmetia Kerrich, 1964a: 76. Type-species: Dusmetia sangwani Subba Rao, by original designation.
DISTRIBUTION AND SPECIES. One species known, New World, Afrotropical, Oriental and
Australasian: sangwani (Subba Rao, 1957: 385) (Pakistan, India, Bangladesh, Philippines,
Australia, Hawaiian Is.).
BIOLOGY. A parasite otAntonina graminis (Maskell) and has been successfully introduced into
various parts of the world to control this mealybug (Homoptera, Pseudococcidae).
COMMENTS. We are unable to place the genus according to Trjapitzin's (1973a) classification of
the Tetracneminae. It may be related to the genera of the Dinocarsini.
NEOPLATYCER US Subba Rao
(Key couplet: 116. Fig. 56)
Neoplatycerus Subba Rao, 1965b: 150. Type-species: Neoplatycerus tachikawai Subba Rao, by original
designation.
INDO-PACIFIC ENCYRTIDAE 307
DISTRIBUTION AND SPECIES. One species, India only: tachikawai Subba Rao (I965b: 151), also at
least three further undescribed species from India and Malaysia (BMNH).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The genus belongs to the tribe Chrysoplatycerini, subtribe Chrysoplatycerina
(Tetracneminae). A key to related genera is provided by Kerrich (1978: 113-114).
NEORHOPUS Girault
(Key couplet: 56)
Neorhopus Girault, 1917g: 139. Type-species: Neorhopus australicus Girault, by original designation.
DISTRIBUTION AND SPECIES. One species, Australia only: australicus Girault (1917g: 140)
(= Neorhopus australicus aureus Girault, 1917g: 140 syn. n.).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Very close to Rhopus (tribe Anagyrini, subtribe Rhopina), but can be distinguished
by having a five-segmented funicle, whereas Rhopus has a six-segmented funicle.
NEZARHOPALUS Girault
(Key couplet: 198)
Nezarhopalus Girault, 1915a: 109. Type-species: Nezarhopalus caudatus Girault, by original designation.
DISTRIBUTION AND SPECIES. One species, Australia only: caudatus Girault (1915a: 109).
BIOLOGY. Unknown.
COMMENTS. The holotype of caudatus is in very poor condition and mounted on a slide (QM). A
critical assessment of the validity of the genus will not be possible until freshly collected material
can be carefully compared with it. The wings of the holotype cannot be located and the relative
position of the apex of the hypopygium cannot be determined. These are considered to be
important diagnostic characters and therefore we prefer to retain Nezarhopalus as valid for the
present. However, the genus appears to be close to Coccidoctonus (tribe Microteryini, subtribe
Syrphophagina) and caudatus may be very close to C. lowelli. From the parts available for
comparison these two species differ only slightly in the relative width of the scape.
OLYPUSAgen. n.
(Key couplet: 460. Figs 217-219, 379)
Type-species: Olypusahirsutasp. n. Gender: feminine.
$. Head. In frontal view about one-quarter wider than long, in profile about twice as long as broad and
more or less gradually and evenly rounded anteriorly. Eye with posterior margin straight with extremely
short, sparse, inconscipuous setae, eye about one-third longer than broad and more or less reaching
occipital margin which is very sharp. Malar space about one-half as long as eye, with sulcus absent.
Frontovertex about two-fifths head width; ocelli forming an obtuse angle, the posterior ones about
equidistant from eye and occipital margins. Antennal scrobes broadly semicircular, just meeting dorsally
and delimited dorsally by a more or less complete carina and reaching a little less than half way from
antennal toruli to anterior ocellus; antennal torulus separated from mouth margin and from other torulus
by a little more than its own length, its lower margin a little above the ventral margins of the eyes; mouth
margin broadly concave. Antennal scape subcylindrical, about four or five times as long as broad and about
as long as minimum width of frontovertex; pedicel conical, about one-third as long as scape and a little
shorter than first funicle segment which is clearly longer than broad; funicle segments cylindrical,
shortening and broadening slightly so that sixth funicle segment is a little transverse; clava entire, apically
rounded and slightly less than one-third as long as funicle; longitudinal sensillae on all flagellar segments,
longest setae not more than about one-half as long as diameter of segments. Frontovertex with shallow,
raised, irregular, transverse, reticulate sculpture, this becoming finer and more irregular on lower parts of
308 J. S. NO YES & M. HAY AT
face and on interantennal prominence; frontovertex with numerous short translucent setae, each a little
shorter than the diameter of the facet, lower parts efface and interantennal promimence with fairly dense,
downwardly directed pale setae. Mandible apically rounded, edentate; maxillary palpus four-segmented,
the apical segment longer and larger than the other three together; labial palpus two-segmented.
Thorax. In side view robust, both mesoscutum and scutellum only very slightly convex, almost flat;
metapleurum and propodeum together broadly in contact with hind coxa. In dorsal view posterior margin
of pronotum broadly concave; visible part of mesoscutum slightly less than twice as broad as long with
notaular lines absent, its posterior margin clearly convex; axillae meeting medially; scutellum a little longer
than broad, apically rounded with an indistinct apical flange and very indistinctly carinate longitudinally in
middle (this only visible if viewed in correct position and correct light); propodeum medially nearly
one-third length of scutellum. Pronotum, mesoscutum and axillae with shallow, raised, squamiform-
reticulate sculpture; scutellum with more longitudinally elongate and slightly deeper reticulate sculpture;
propodeum with shallow, raised, reticulate sculpture; mesopleurum almost smooth with very shallow,
raised, irregular, longitudinally elongate sculpture; pronotum, mesoscutum and axillae with fairly dense,
short, inconspicuous dark setae, scutellum clothed in dense conspicuous white setae which become
progressively longer towards apex of scutellum, propodeum on outside of spiracles with dense, short, pale
setae. Forewing almost hyaline, but clearly suffused dark brown, slightly paler nearer base and with a few
paler streaks or areas, wing only a little more than twice as long as broad; linea calva very narrow, closed
towards hind margin by several lines of setae on dorsal surface; filum spinosum present; submarginal vein
without an apical hyaline break and with parastigma a little but distinctly broader than proximal part of
vein; costal cell about eight times as long as broad, with setae dorsally about as dense as in disc of wing;
marginal vein about four times as long as broad; stigmal vein long and curved, about twice as long as
marginal vein and subequal to postmarginal vein, a narrow naked streak extending from apex of
postmarginal to apex of stigmal vein. Hindwing hyaline, about two-thirds lengths of forewing and about
two and one-half times as long as broad, costal cell about seven times as long as broad, marginal setae less
than one-twentieth maximum wing width. Mid tibial spur slightly shorter than mid basal tarsal segment.
Caster. About two-thirds as long as thorax; cereal plates in basal one-third; hypopygium reaching to
about four- fifths along gaster, with several long setae medially at its apex; ovipositor not exserted.
O" . Apart from genitalia and antenna very similar to female. Differs as follows. Antenna yellow with sixth
funicle segment a little longer than broad. Genitalia with aedeagus spatulate, i.e. a little broadened
subapically, digiti very stout, only about three times as long as broad.
COMMENTS. Belongs to the same group of genera as Encyrtus, Prionomastix and Aethognathus
(see comments under Anagyrodes); separated from related genera by the combination of the
solid clava in the female, relatively widely separated antennal toruli, densely hairy forewings,
moderately long marginal vein, subequal postmarginal and stigmal veins, structure of scutellum,
lack of apical scutellar tuft and the relatively well-advanced cereal plates (in the other genera
they are generally about half way along gaster or in posterior half).
Olypusa hirsuta sp. n.
(Figs 217-219, 379)
$. Length: 1-82-2-24 mm (holotype, 2-14 mm).
Colour. Head, thorax and gaster black or very dark brown except a very narrow area between eye and
occipital margin dorsally, on occiput immediately below this and a small quadrate area on each side of
pronotum which are white; legs dark brown except knees, fore and mid tarsi, apex and base of mid femur
and tibia, apex of mid tibia, mid tibial spur and basal tarsal segment which are yellowish white to
yellow-orange, mid tibial spur sometimes dark brown; forewing generally suffused dark brown except
proximal half of basal cell, costal cell, a small area at apex of venation and an indistinct longitudinal streak
from linea calva which is more or less hyaline (Fig. 217).
Head. Relative measurements (holotype): head length 44, head width (facial view) 54, head width (side
view) 24, minimum frontovertex width 22-5, malar space 15, eye length 30, eye width 23, POL 12-5, OOL
3, scape length 23, scape width 5, proportions of antennal segments as in Fig. 219, head in facial view as in
Fig. 218.
Thorax. Relative measurements (holotype): forewing length 126, width 58, venation and base of
forewing as in Fig. 217; hindwing length 91, hindwing width 33.
INDO-PACIFIC ENCYRTIDAE 309
C?. Similar to female except antenna which is yellowish, foretibia which may be completely yellowish, and
genitalia (see generic description). Antenna as in Fig. 379.
DISTRIBUTION. Papua New Guinea.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $, Papua New Guinea: Wau, 1100-1200 m, vi.1968 (N. L. H. Krauss) (BPBM).
Paratypes. Papua New Guinea, 1 $, Torricelli Mts, Mobitei, 750 m, 1-15. iv. 1959 (J. W. Brandt)
[Specimen lacking head]; 1 cT, Kokoda-Pitoki, 400 m, 23.iii.1956 (/. L. Gressitt); 1 $, New Ireland, SW,
Gilingil Plantation, 2 m, 5.vii.l956 (/. L. Gressiti) (specimen lacking head); 1 cf , New Britain, Warongoi
Valley, Gazelle Peninsula, 100 m, 25.V.1956 (/. L. Gressitt) (specimen lacking gaster) (BPBM).
OOENCYRTUS Ashmead
(Key couplets: 91, 260, 283, 313, 339, 381, 393, 431, 438, 494, 514.
Figs 152, 177, 181, 182, 246, 381)
Ooencyrtus Ashmead, 19006: 381. Type-species: Encyrtus clisiocampae Ashmead, by original designation.
Echthrodryinus Perkins, 1906: 252. Type-species: Echthrodryinus destructor Perkins, by monotypy. Syn.
n.
Schedius Howard, 1910: 2. Type-species: Schedius kuvanae Howard, by original designation.
Tetracnemella Girault, 1915a: 170. Type-species: Tetracnemella australiensis Girault, by original desig-
nation. Syn. n.
Xesmatia Timberlake, 1920: 424. Type-species: Xesmatia flavipes Timberlake, by original designation.
Syn. n.
Pseudolitomastix Risbec, 1954: 1068. Type-species: Litomastix creona Risbec, by original designation.
DISTRIBUTION AND SPECIES. About one-hundred species, cosmopolitan; 37 from review area:
alboantennatus (Subba Rao, 1971: 222) (comb. n. from Pentalitomastix) (Java), australiensis
(Girault, 19150: 170) (comb. n. from Tetracnemella) (Australia), batocerae Ferriere (1936: 333)
(Malaysia), bicolor Girault (19150: 78) (Australia), cochereaui Prinsloo & Annecke (19780: 41)
(New Caledonia), corbetti Ferriere (1931: 284) (Malaysia), crassulus Prinsloo & Annecke
(19780: 42) (Samoa), destructor (Perkins, 1906: 253) (comb. n. from Echthrodryinus) (Austra-
lia), erionotae Ferriere (1931: 284) (India, Malaysia, Java, Mariana Is., Hawaiian Is.), euxoae
(Girault, 19270: 2) (comb. n. from Schedius) (Australia), ferrierei Shafee, Alam & Agarwal
(1975: 97) (India), flavipes (Timberlake, 1920: 425) (comb. n. from Xesmatia) (Hawaiian Is.),
guamensis Fullaway (1946: 205) (Mariana Is., Hawaiian Is.), hyalinipennis (Dodd, 1917: 354)
(comb. n. from Tetracnemella) (Australia), inconspicuus (Girault, 19150: 141) (comb. n. from
Coccidoxenus) (Australia) , javanicus Mercet (19220: 152) (Ja\a)Johnsoni (Howard, 18980: 18)
(Hawaiian Is.), lacteiclavus Girault (19326: 1) (Australia), larvarum (Girault, 19196: 58)
(comb. n. from Paracopidosomopsis) (Java), leucocerus Mercet (19220: 150) (Java), major
Ferriere (1931: 285) (Java), malayensis Ferriere (1931: 282) (India, Malaysia, Indonesia,
Papua New Guinea), metallicus Girault (19140: 37) (Australia), ovidivorus (Girault, 19250: 2)
(comb. n. from Echthrodryinus) (Australia) , pacificus Waterston (1915: 307) (Fi]i) , pallidipes
(Ashmead, 19040: 15) (comb. n. from Aphidencyrtus) (Philippines), papilionidis (Girault,
1932c: 3) (comb. n. from Stenoteropsis) ( Australia), papilionis Ashmead (19050: 4) (Pakistan,
India, Philippines), p/iog/ Trjapitzin, Myartseva & Kostjukov (1977: 671) ( Vietnam), podon-
tiae Gahan (1922: 51) (Java), segestes Trjapitzin (1965: 320) (Indonesia), shakespearei (Girault,
19230: 48) (comb..n. from Coccidoxenus) (Australia), sphingidarum Timberlake (1941: 223)
(Marquesas Is.), submetallicus (Howard; Noyes, 1979: 160) (Hawaiian Is.), swezeyi Fullaway
(1946: 206) (Mariana Is.), tricolor (Girault, 19150: 140) (comb. n. from Coccidoxenus)
(Australia), and xanthogaster (Girault, 19150: 150) (comb. n. from Echthrodryinus) (Australia),
also much undetermined material from throughout the region (BMNH, BPBM, CNC, GC).
REFERENCE. Key to Indo-Malayan species: Trjapitzin et al. (1977: 672-674).
BIOLOGY. Parasites of eggs of various insects, notably Lepidoptera and Heteroptera, and of
310 J. S. NOYES & M. HAYAT
spiders (Araneida). Several species are also hyperparasites of other Hymenoptera (Dryinidae
and Braconidae) parasitising Lepidoptera and Auchenorrhyncha (Homoptera). One group
(guamensis-group) of species are parasites of Aphididae (Homoptera) and puparia of Syrphidae
(Diptera).
COMMENTS. The species previously included in Echthrodryinus (see Gordh & Trjapitzin, 1978)
almost certainly represent a polyphyletic group since they appear to be more closely related,
morphologically, to widely separated species of Ooencyrtus than they do to each other, e.g.
Ooencyrtus bucculatricis (Howard) (comb, n.) is more closely related to Ooencyrtus johnsoni
(Howard) than it is to destructor (the type-species of Echthrodryinus}. Thus in our view the
change from egg parasitism to larval parasitism or hyperparasitism mush have occurred more
than once and probably via different routes. Therefore we can find no reason for retaining
Echthrodryinus as a distinct genus whose type-species or other included species cannot be
reliably separated from species of Ooencyrtus other than by an apparent difference in biology.
The guamensis-group is an apparently monophyletic group of species found in South America
(see Noyes, 1980: 194), Africa, India and Mariana Is. which parasitise syrphid puparia (and also
possibly aphids) . The species of this group can be distinguished from other species of Ooencyrtus
by being slightly larger and the colour of the head and thorax always being black or dark brown
with a very slight blue or green sheen, the mesoscutum clothed in very conspicuous white or
translucent setae and the marginal vein of the forewing always being punctiform and the stigmal
vein relatively long. For the present, we prefer to leave these species in Ooencyrtus than propose
a new genus to accommodate them.
One group consists of species (bicolor, lacteiclavus and metallicus) which are parasites of
spiders' eggs. For the present we are leaving these in Ooencyrtus although it may be considered
that they belong to a genus apart. They can be distinguished from other species of Ooencyrtus by
the infuscate forewings and largely pale brown or orange-brown thorax instead of the usual
metallic or dark brown colour typical of Ooencyrtus. They differ also in general body shape,
especially that of the head.
It is possible that Scotteus Masi (1917b) is a synonym of Ooencyrtus. The holotype of
ochroleucus has been examined (BMNH) but unfortunately the body is missing. However, the
remaining parts (forewing, antenna and legs) and Masi's description indicate that it may be a
species of Ooencyrtus with an obliquely truncate clava (species with an obliquely truncate clava
are known to occur in the Neotropics).
The genus is placed in the tribe Microteryini, subtribe Ooencyrtina (Encyrtinae) and can be
separated from other closely related genera (including Trichomasthus and Fulgoridicidd) by the
characters given in the key.
OVALOENCYRTUSgen. n.
(Key couplets: 133, 470. Figs 67, 68, 225, 382-384)
Type-species: Ovaloencyrtus fijiensis sp. n. Gender: masculine.
$ . Head. In frontal view slightly broader than long, in profile slightly less than twice as long as broad and
anteriorly evenly rounded to top of antennal scrobes, below this almost straight to mouth margin. Eye with
moderately conspicuous hairs, each clearly longer than the diameter of a facet, posterior margin of eye
imperceptibly concave, eye about one-quarter longer than broad and reaching occipital margin which is
sharp. Malar space nearly half as long as eye, with sulcus present. Frontovertex about one-quarter head
width; ocelli forming a slightly acute angle, the posterior ones nearly touching eye and separated from
occipital margin by about their own major diameters. Antennal scrobes fairly long and narrow, more or
less meeting dorsally and reaching about half way from antennal toruli to anterior ocellus, interantennal
prominence dorsally acute; antennal torulus separated from mouth margin by a little less than its own
length and from other torulus by about its own length, its dorsal margin about level with ventral margins of
eyes; clypeus fairly broadly and deeply excised medially. Antennal scape clearly longer than width of
frontovertex and broadened and flattened, about three times as long as broad; pedicel conical, about
one-third length of scape and clearly longer than any of the funicle segments which are cylindrical and
INDO-PACIFIC ENCYRTIDAE 311
slightly broadening distally, the first four of which are a little longer than broad or quadrate, the fifth and
sixth transverse; clava three-segmented, a little longer than funicle, with a strongly oblique apical
truncation with the sutures strongly convergent towards base ventrally; longitudinal sensillae on all
flagellar segments except first two, longest setae about as long as diameter of first segment. Frontovertex
above antennal scrobes almost completely smooth except for the shallow piliferous punctures, top of
scrobes and between scrobes and eyes with shallow, raised, rugose-reticulate to squamiform-reticulate
sculpture, interantennal prominence and genae with squamiform-reticulate sculpture, piliferous punctures
on lower parts of face deeper than on frontovertex; setae on frontovertex sparse but conspicuous, each
longer than diameter of an ocellus. Mandible with two very small teeth and a very broad truncation;
maxillary palpus four-segmented, labial palpus three-segmented.
Thorax. In side view moderately robust with mesoscutum and scutellum only slightly convex, almost flat;
propodeum and metapleurum together quite widely separated from hind coxa by the posterior margin of
the mesopleurum which is quite broadly in contact with basal segment of gaster. In dorsal view pronotum
broadly concave but not strongly so; visible part of mesoscutum about four-fifths broader than long with
notaular lines absent and posterior margin convex and slightly projecting above axillae and thus these
appear broadly separated; axillae more or less meeting; scutellum about as long as mesoscutum and a little
broader than long, with apex rounded; propodeum medially about one-fifth length of scutellum. Mesoscu-
tum and axillae with shallow, raised, squamiform-reticulate sculpture, scutellum more or less entirely
smooth and polished but in anterior one-third or so with extremely shallow reticulate sculpture;
mesopleurum smooth and shiny but bordered posteriorly, ventrally and anteriorly by some shallow,
irregular, reticulate to rugose sculpture; propodeum with some fairly deep, raised irregular sculpture
medially and laterally. Forewing lightly infuscate, pale brown in middle beneath venation and near base,
nearly three times as long as broad; linea calva not interrupted or closed; filum spinosum present;
submarginal vein with an indistinct apical hyaline break, parastigma not swollen; costal cell about 10 times
as long as broad, with a single line of setae dorsally in apical one-third; marginal vein about four times as
long as broad, about twice as long as postmarginal and a little shorter than stigmal. Hindwing about
three-quarters as long as forewing, about five times as long as broad, with marginal setae about one-third as
long as maximum wing width. Mid tibial spur shorter than basal mid tarsal segment.
Gaster. Shorter than thorax; cereal plates in anterior half; hypopygium reaching to about four-fifths
along gaster; last tergite about two-thirds as long as mid tibia, paratergites absent; ovipositor a little shorter
than mid tibia, gonostyli free, about one-quarter as long as ovipositor.
C?. Unknown.
COMMENTS. Closely related to Paratetralophidea from which it can be separated by the
characters given in the key. Also probably related to Xenoencyrtus and Ooencyrtus (tribe
Microteryini, Ooencyrtina) and separated from both by the very long, strongly obliquely
truncate clava, relatively long marginal and stigmal veins of the forewing and sharp occipital
margin.
Ovaloencyrtusfijiensissp. n.
(Figs 67, 68, 225, 382-384)
$. Length: 0-95-1-13 mm (holotype, 1-13 mm).
Colour. Head strongly shining metallic green, purple or blue, mesoscutum shining purple, scutellum
strongly shining deep blue or green with strong purple reflections; propodeum and mesopleurum dark
brown; antenna very dark brown, almost black; all coxae, fore femur, fore tibia and tarsus to a less extent,
dark brown, remainder of legs orange-brown; gaster dark brown, basally orange-brown; forewing
generally suffused pale brownish, darker in middle across wing from apical one-third of venation, proximal
part of basal cell, except extreme base, more or less hyaline (Fig. 68).
Head. Relative measurements (holotype): head length 64, head width (facial view) 75, head width (side
view) 35, minimum width of frontovertex 20, POL 8, OOL 1-5, malar space 21, length of eye 46, width of
eye 38 , scape length 45 , maximum scape width 14 , other proportions of antenna as in Fig. 67 , mandible as in
Fig. 225. There is a little variation in the relative width of the frontovertex; in smaller specimens it is slightly
wider and thus correspondingly the eyes are a little smaller.
Thorax. Relative measurements (holotype): forewing length 174, width 65; hindwing length 123, width
26; base of forewing as in Fig. 68, forewing venation as in Fig. 382.
Gaster. Relative lengths (paratype): ovipositor 80, last tergite 60, [mid tibia 88] ; genitalia as in Fig. 383,
hypopygium as in Fig. 384.
312 J. S. NOYES & M. HAYAT
Cf . Unknown.
DISTRIBUTION. Fiji.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $, Fiji: Viti Levu, Nausori Highlands, 500-700 m, xi.1976 (N. L. H. Krauss) (BPBM).
Paratypes. Fiji: 1 <j>, same data as holotype, 26.iii.1970; 1 $, Ovalau, Levuka, 0-200 m, xii.1969
(N. L. H. Krauss); 1 $, Lami, 20-200 m, iii.1976 (N. L. H. Krauss); 1 $, Lau, Lakomba, 3.ix.l924
(E. H. Bryan Jr) (BPBM, BMNH).
OVIDOENCYRTUS Girault
(Key couplet: 492)
Ovidoencyrtus Girault, 1924a: 7. Type-species: Ovidoencyrtus pallidipes Girault, by monotypy.
DISTRIBUTION AND SPECIES. One species, Australia only: pallidipes Girault (1924a: 7).
BIOLOGY. Parasites of eggs of Reduviidae (Heteroptera).
COMMENTS. We are unable to place the genus according to Trjapitzin's (19736) classification of
the Encyrtinae.
PAKSIMMONDSIUS Ahmad & Ghani
(Key couplet: 134. Fig. 73)
Paksimmondsius Ahmad & Ghani, 1974: 391. Type-species: Paksimmondsius pakistanensis Ahmad &
Ghani, by original designation.
DISTRIBUTION AND SPECIES. One species, Pakistan only: pakistanensis Ahmad & Ghani
(1974: 392).
BIOLOGY. Parasites of Kermesidae (Homoptera).
COMMENTS. Paksimmondsius appears to be related to Leurocerus and Zozoros and can be
separated from these genera by the characters given in the key (see also comments under
Leurocerus and Zozoros) .
PAPUNA gen. n.
(Key couplet: 346. Figs 190, 385-389)
Type-species: Papuna nemis sp. n. Gender: feminine.
$. Head. In facial view about one-quarter broader than long, in profile about one-half longer than broad
and gradually and evenly curved dorsally, most strongly curved about level with top of scrobes, below this
point almost straight. Eye with posterior margin more or less straight, only a little longer than broad,
conspicuously hairy and overreaching occipital margin which is more or less rounded, at least not sharp.
Malar space about half length of eye, with sulcus present. Frontovertex a little less than one-third head
width; ocelli fairly large, forming an acute angle which is almost an equilateral triangle, the posterior
ocellus separated from occipital margin by slightly less than its own diameter and considerably closer to the
eye margin. Antennal scrobes moderately deep and elongate, meeting dorsally and reaching about
two-thirds from antennal toruli to anterior ocellus; antennal torulus separated from mouth margin by
about three-quarters its own length and from other torulus by about one and one-half times its own length,
its dorsal margin about level with lowest margin of eye, clypeal margin shallowly excised between toruli.
Antennal scape clearly longer than minimum width of frontovertex, slightly flattened and broadened,
slightly more than three times as long as broad; pedicel conical, slightly more than one-third length of scape
and clearly much longer than any of the funicle segments which are all transverse and nearly equal in length
but become much broader apically; clava three-segmented with a strong oblique, apical truncation and
much broader than and a little longer than funicle, the sutures strongly converging; longitudinal sensillae
on all segments of flagellum; longest setae about as long as diameter of first funicle segment. Frontovertex
INDO-PACIFIC ENCYRTIDAE 313
with very shallow, raised reticulate sculpture of small mesh, more squamiform-reticulate between eyes and
antenna! toruli and lower parts of face where it also becomes more longitudinally elongate; frontovertex
and lower parts efface clothed in fairly dense, conspicuous setae, each about as long as the diameter of an
ocellus. Mandible with two teeth and a truncation or obscurely tridentate; maxillary palpus four-
segmented, labial palpus three-segmented.
Thorax. In side view with mesoscutum and scutellum quite flat, the metapleurum together with the
propodeum quite broadly in contact with hind coxa. In dorsal view posterior margin of pronotum quite
strongly concave and slightly angled medially; visible part of mesoscutum about one-half broader than
long, notaular lines absent, its posterior margin angled outwards medially; axillae meeting; scutellum a
little shorter than mesoscutum and a little broader than long, its apex rounded; propodeum medially about
one-eighth as long as scutellum, laterally with fairly dense setae almost completely surrounding the
spiracle, except posteriorly. Dorsum of thorax with sculpture similar to frontovertex, but perhaps of
slightly larger mesh, mesopleurum almost smooth but with some very shallow, irregular sculpture;
propodeum medially with numerous, shallow, incomplete carinae, laterally with irregular, raised, rugose
sculpture; dorsum of thorax with setae of similar length, colour and density to those on frontovertex.
Forewing more or less hyaline but very faintly suffused brownish, nearly three times as long as broad; linea
calva not interrupted or closed; filum spinosum present; submarginal vein with an apical hyaline break,
parastigma very slightly and conspicuously swollen; costal cell about 15 times as long as broad, with a single
line of setae dorsally in its apical half; marginal vein about six times as long as broad, subequal to stigmal,
both a little shorter than postmarginal. Hindwing about two-thirds as long as forewing, about four times as
long as broad, with marginal setae about one-fifth as long as maximum wing width. Mid tibial spur slightly
shorter than basal mid tarsal segment.
Caster. Without exserted part of ovipositor about as long as thorax, exserted part of ovipositor about
one-third as long as gaster; cereal plates in anterior half of gaster; hypopygium with apex a little more than
halfway along gaster; last tergite produced apically, very pointed, slightly longer than mid tibia; ovipositor
about twice as long as mid tibia or gonostyli; gonostyli free.
Cf. Unknown.
COMMENTS. Placement of this genus according to Trjapitzin's classification of the Encyrtinae is
difficult. The structure of the antenna is very similar to that found in the Tyndarichina
(Cheiloneurini, see Trjapitzin & Gordh, 1980), but differs from these in the structure of the
mandible, wing venation and the mesopleurum not being posteriorly enlarged. It is most
probable that this genus is related to Pseudencyrtus (Microteryini, Pseudencyrtina), having a
similar structure of the gaster (e.g. elongate last tergite), thorax, mandible and head. It differs
from Pseudencyrtus in forewing venation (e.g. postmarginal vein longer than stigmal, smaller
angle between stigmal and postmarginal veins), narrower hindwing (in Pseudencyrtus it is not
much more than three times as long as broad), narrower frontovertex (Pseudencyrtus has the
frontovertex more than one-third head width) and antenna.
I 'up unn ut'iiii.v sp. n.
(Figs 190, 385-389)
$. Length (excluding ovipositor): 1-98-2-03 mm (holotype, 2-03 mm).
Colour. Frontovertex deep metallic blue-green with purple reflections around eyes, below top of toruli
more green with blue or purple reflections, antennal scrobes above toruli coppery; antenna with scape
yellow, pedicel apically testaceous, basal part of pedicel and flagellum dark brown; mesoscutum dull
shining dark blue with purple reflections, axillae brownish purple with brassy reflections; scutellum dull
shining blue-green with apex distinctly more shiny green with brassy reflections; mesopleurum dark
chestnut-brown; legs with all coxae and femora dark brown, fore femur apically yellowish, mid femur with
a pale sub-basal ring, all tibiae and tarsi yellow; propodeum and gaster (including ovipositor sheaths) dark
brown, the gaster with strong brassy purple reflections; apex of ovipositor sheaths yellowish.
Head. Head in facial view as in Fig. 387, mandible as in Fig. 388. Relative measurements (holotype):
head length 85, head width (facial view) 103, head width (side view) 52, minimum frontovertex width 30,
malar space 34, eye length 61, eye width 54, POL 13, OOL 2, scape length 41, scape width 13, proportions
of antenna as in Fig. 385.
Thorax. Relative measurements (holotype): forewing length 278, width 100, base of forewing as in Fig.
386, venation as in Fig. 190; hindwing length 175, width 45.
314 J. S. NOYES & M. HAYAT
Caster. Relative lengths (paratype): last tergite 60, ovipositor 1 18, [mid tibia 53]; genitalia as in Fig. 389.
Cf . Unknown.
DISTRIBUTION. Papua New Guinea.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $, Papua New Guinea: Morobe Province, Lasanga I., x. 1979 (/. H. Martin) (BMNH).
Paratype. Papua New Guinea: 1 9 > Morobe Province, Buso Forest, x.1979 (/. H. Martin) (BMNH).
PARABLASTOTHRIX Mercet
(Key couplet: 15)
Parablastothrix Mercet, I9l7d: 538. Type-species: Parablastothrix vespertinus Mercet, by monotypy.
DISTRIBUTION AND SPECIES. Fourteen species, cosmopolitan; three species from review area:
magnioculus (Girault, 1923#: 47) (comb. n. from Schedius) (= Paracaenocercus albifemur
Girault, 19226: 103 syn. n.) (Australia), nepticulae Hedqvist (1976: 50) (Sri Lanka) and
unicinctipes (Girault, 1915a: 111) (comb. n. from Schedius) (Australia), also further undeter-
mined material from Bismark Archipelago, Solomon Is. and Australia (BMNH, BPBM, QM,
ANIC).
REFERENCE. Key to Holarctic species: Loginovskaya (1981: 160-162).
BIOLOGY. Parasites of larvae of Lyonetiidae and Nepticulidae (Lepidoptera).
COMMENTS. Trjapitzin & Gordh (19786) erected the new subtribe Parablastothrichina within the
tribe Copidosomatini (Encyrtinae) to accommodate Parablastothrix and Calometopia. How-
ever, we think that the genera of this subtribe (including also Mashhoodiella) are morphologi-
cally closer to the Aphycina (tribe Aphycini) than they are to the Copidosomatini.
PARABLATTICIDA Girault
(Key couplets: 42, 175, 195, 212. Figs 390-395)
Parablatticida Girault, 1915a: 117. Type-species: Parablatticida pachyscapha Girault, by original designa-
tion.
Holanusia Girault, 1915a: 162. Type-species: Holanusia convexus Girault, by original designation. Syn. n.
Symphycus Masi, 19176: 149. Type-species: Symphycus aphycoides Masi, by monotypy. Syn. n.
Geniaspidius Masi, 19176: 155. Type-species: Geniaspidius viduus Masi, by monotypy. Syn. n.
Amaurilyma Graham, 1958: 153. Type-species: Encyrtus brevicornis Dalman, by original designation.
Syn. n.
Desobius Noyes, 1980: 192. Type-species: Desobius convexus Noyes, by original designation. Syn. n.
DISTRIBUTION AND SPECIES. Seven species, Neotropical, Palaearctic, Afrotropical, Oriental and
Australasian; one from review area: pachyscapha Girault (1915a: 117) (= Holanusia convexus
Girault, 19150: 162 syn. n.) (Australia), also much undetermined material, including many
undescribed species, from India and Taiwan to Samoa and Australia (BMNH, BPBM, ANIC,
HC).
BIOLOGY. Unknown.
COMMENTS. Examination of material from throughout the region has shown that the species
previously placed in Desobius fall within our concept of the range of variation within Parablatti-
cida. We have found that differences in sculpture of the mesoscutum and relative position of the
hypopygium are not consistent and therefore regard Desobius as a synonym of Parablatticida.
The holotype male of Geniaspidius viduus Masi (BMNH) belongs to Parablatticida (comb, n.)
and to the same species-group as trinidadensis (nom. n. for convexus Noyes, 1980 nee Girault,
1915), characterised by the striate-reticulate sculpture of the mesoscutum and generally smaller
size.
INDO-PACIFIC ENCYRTIDAE 315
We have examined the four syntypes of Symphycus aphycoides Masi (BMNH, ZMUC,
MCSN). The female from ZMUC is here designated LECTOTYPE and has been so labelled. It
is in several fragments glued on a card rectangle but is more or less complete. It falls within our
concepts of the limits of Parablatticida and therefore we believe that the two genera are
synonymous. The single male syntype is not congeneric.
Parablatticida brevicornis (Dalman) (comb, n.) is very close to pachyscapha Girault and
differs only very slightly in the relative width of the scape and frontovertex. More detailed study
may show that these differences fall within the range of variation of brevicornis.
We are unable to place the genus, although it may be related to Exoristobia or Phaulo-
encyrtus.
PARACERAPTROCERUS Girault
(Key couplet: 113)
Paraceraptrocerus Girault, 1920c: 184. Type-species: Paraceraptrocerus africanus Girault, by original
designation.
DISTRIBUTION AND SPECIES. Fourteen species, Neotropical, West Palaearctic, Afrotropical,
Oriental; two species from review area: brevicaudatus (Subba Rao, 19650: 74) (India) and
italicus (Masi, 1917a: 80) (India).
REFERENCE. Revision: Annecke (1967: 130-156).
BIOLOGY. Parasites of Coccidae (Homoptera).
COMMENTS. The genus is very close to Anicetus (tribe Cerapterocerini) and possibly should be
considered synonymous. Annecke (1967: 100-101) provides a key to distinguish this genus from
its relatives.
PARACHALCERINYS Girault
(Key couplet: 497)
Parachalcerinys Girault, 19256: 97. Type-species: Parachalcerinys nonaericornis Girault, by monotypy.
DISTRIBUTION AND SPECIES. Two species, both Australian: coccidoxenoides Girault (1926c: 130)
and nonaericornis Girault (19256: 97).
BIOLOGY. Unknown.
COMMENTS. Girault (1926c: 130) stated that the type-species of Parachalcerinys was cocci-
doxenoides but since the description of Parachalcerinys nonaericornis was published the
previous year, nonaericornis must be taken as the type-species.
The two species included here may belong to different genera; coccidoxenoides possibly to
Australia (see comments under Australia) and nonaericornis to Psyllaephagus . However, until
fresh material can be carefully compared with the types of the two included species we prefer to
retain them in the present combination and thus treat Parachalcerinys as a valid genus.
PARACLADELLA Girault
(Key couplets: 422, 429)
Paradadella Girault, 1920J: 142. Type-species: Paracladella globosa Girault, by monotypy.
DISTRIBUTION AND SPECIES. Two species, Australia only: giorgionei Girault (1932a: 4) and
globosa Girault (1920d: 142).
BIOLOGY. Unknown.
COMMENTS. The holotype male of globosa (QM) is in extremely poor condition, but may be the
male of giorgionei.
316 J. S. NOYES & M. HAYAT
Paradadella belongs to the same group of genera as Anagyrodes (see comments under
Anagyrodes) and is probably most closely related to Neodadia (tribe Neocladiini). It can be
separated from related genera by having a mandible with a single long tooth (which may have a
short subapical second tooth), a solid clava and fore wing with a punctiform marginal vein.
PARACLAUSENIA Hayat
(Key couplets: 265, 502. Figs 151, 396, 397)
Paraclausenia Hayat, 1980: 637. Type-species: Paraclausenia herbicola Hayat, by original designation.
DISTRIBUTION AND SPECIES. One described species, India only: herbicola Hayat (1980: 639), also
further undetermined material, which may include an undescribed species, from India and S.
China (BMNH,BPBM).
BIOLOGY. Unknown.
COMMENTS. This genus belongs to the tribe Charitopidini (Tetracneminae).
PARAENASOMYIA Girault
(Key couplets: 302, 342, 433, 456. Fig. 178)
Paraenasomyia Girault, 1915a: 110. Type-species: Paraenasomyia orro Girault, by monotypy.
DISTRIBUTION AND SPECIES. Three species, Australia only: australiensis (Girault, 19146: 59)
(comb. n. from Copidosomd) (= Cerchysius bellulus Girault, 1915a: 84), johnsoni Girault
(1922/: 1) and orro Girault (19150: 110).
BIOLOGY. Parasites of galls of Cecidomyiidae (Diptera).
COMMENTS. Girault (\9lle: 95) synonymised bellulus and australiensis. Comparison of the
descriptions of both species indicates that he must have inadvertently described the same
specimen twice under two different names.
Its biology, wing venation and general morphology indicate that the genus is probably related
to Pseudencyrtus (Microteryini, Pseudencyrtina).
PARALEPTOMASTIX Girault
(Key couplet: 459)
Paraleptomastix Girault, 1915o: 168. Type-species: Paraleptomastix ihoreauini Girault, by original
designation.
DISTRIBUTION AND SPECIES. One species Australia only: thoreauini Girault (19150: 168).
BIOLOGY. Unknown.
COMMENTS. Belongs to the same group of genera as Anagyrodes (see comments under
Anagyrodes) and is probably most closely related to Neodadia (tribe Neocladiini). It can be
separated from related genera by having a mandible with a long single tooth (possibly with a
smaller subapical tooth), a three-segmented clava, forewing with a relatively long marginal vein
and hypopygium reaching the apex of the gaster.
PARALITOMASTIX Mercet
(Key couplet: 498)
Paralitomastix Mercet, 1921: 438. Type-species: Encyrtus varicornis Nees, by original designation.
DISTRIBUTION AND SPECIES. Sixteen species, cosmopolitan; three from review area: bicolori-
cornis (Girault, 1915a: 104) (comb. n. from Cocddencyrtus) (Australia), ipswichia (Girault,
INDO-PACIFIC ENCYRTIDAE 317
1923d: 2) (comb. n. from Coccidencyrtus) (Australia) and varicornis (Nees; Mercet, 1921:
439) (Pakistan, India), also some undetermined material from Australia (BMNH).
BIOLOGY. Polyembryonic parasites of larvae of Pyralidae and Gelechiidae (Lepidoptera).
COMMENTS. Placed in the tribe Copidosomatini, subtribe Copidosomatina (Encyrtinae) and
should very probably be considered synonymous with Copidosoma. The genus can be dis-
tinguished from Copidosoma solely on the bicolorous antennal flagellum since several species
near Copidosoma koehleri Blanchard have sculpture on the scutellum similar to those species
placed in Paralitomastix (a character sometimes used to separate the two genera).
PARANA THRIX Myartseva
(Key couplets: 145, 227, 270)
Paranathrix Myartseva, 1980: 722. Type-species: Anathrix acanthococci Myartseva, by original desig-
nation.
DISTRIBUTION AND SPECIES. Two species, Palaearctic, Oriental and Australasian; one from
review area: thailandicus (Myartseva, 1979: 1746) (Thailand), also much undetermined mate-
rial, including several undescribed species, from Bangladesh to the Solomon Is. and Australia
(BMNH, BPBM, CNC).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The species listed as Dinocarsis sp. by Tandon & Srivastava (1980) probably belongs
to this genus (Subba Rao, pers. comm.).
Paranathrix belongs in the tribe Anagyrini, subtribe Anagyrina (Tetracneminae) and can be
separated from related genera by the characters given in the key, in particular the pattern of
silvery setae on the scutellum.
PARAPHAENODISCUS Girault
(Key couplets: 92, 136, 380)
Paraphaenodiscus Girault, I9l5a: 93. Type-species: Paraphaenodiscus verus Girault, by original desig-
nation.
DISTRIBUTION AND SPECIES. Nine species, Afrotropical, Oriental and Australasian; three from
review area, all Australian: parus (Girault, 19150: 93) (comb. n. from Encyrtus), verus Girault
(19150: 93) and wundti (Girault, 19150: 92), also two further, undescribed, species from India
and Malaysia (BMNH).
REFERENCE. Revision of southern African species: Prinsloo (19760); Prinsloo & Myndardt
(1982).
BIOLOGY. Parasites of Coccidae (Homoptera).
COMMENTS. The genus is near Microterys (Microteryini, subtribe Microteryina) and can be
separated by the characters given in the key, in particular the presence of an apical flange on the
scutellum which is absent in Microterys (see Prinsloo, 19760: 161).
PARAPHYCUS Girault
(Key couplets: 166, 285)
Paraphycus Girault, 19150: 97. Type-species: Paraphycus abnormiscapus Girault, by original designation.
DISTRIBUTION AND SPECIES. One species, Australia only: abnormiscapus Girault (19150: 97).
BIOLOGY. Unknown.
COMMENTS. The genus is the type-genus of the subtribe Paraphycina Hoffer, 1955 (tribe
318 J. S. NOYES & M. HAYAT
Aphycini). However, this is based on a misidentification of the genus by Mercet (1921: 232) who
incorrectly included the species flavovarius in Paraphycus. This species has since been trans-
ferred to Xenaphycus Trjapitzin.
The systematic position of Paraphycus cannot be accurately ascertained since the holotype
female of abnormiscapus is lacking its head. The parts that remain suggest that it quite possibly
may belong in the tribe Aphycini, but doubtfully to the subtribe Paraphycina as understood at
present. The genus should be easily recognisable from the parts that remain and Girault's
description.
PARARHOPELLA Girault
(Key couplet: 71)
Pararhopella Girault, 1923c: 144. Type-species: Metalonella longfellowi Girault, by original designation.
DISTRIBUTION AND SPECIES. Two species, both Australian: longfellowi (Girault, 19150: 77) and
maculatipes Girault (1923c: 144); possibly one further species from Australia (BMNH).
BIOLOGY. Unknown.
COMMENTS. The genus is close to Mesorhopella Girault (see comments under Mesorhopella).
PARASCHEDIUS Mercet
(Key couplets: 259, 311. Figs 180, 398, 399)
Paraschedius Mercet, 1925ft: 328. Type-species: Paraschedius ductor Mercet, by original designation.
DISTRIBUTION AND SPECIES. Four species, Palaearctic; two undescribed species from India and
Java (BMNH, BPBM).
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. Paraschedius can probably best be placed in the tribe Habrolepidini, subtribe
Comperiellina (Encyrtinae).
PARASTENOTER YS Girault
(Key couplet: 466)
Parastenoterys Girault, 1915a: 165. Type-species: Parastenoterys punctatus Girault, by original desig-
nation.
DISTRIBUTION AND SPECIES. One species, Australia only: punctatus Girault (19150: 165).
BIOLOGY. Unknown.
COMMENTS. The genus belongs to the same group as Rhytidothorax , Tachinaephagus and
possibly Hexencyrtus which could probably be accommodated within the subtribe Coenocercina
of the Bothriothoracini (Encyrtinae). This group can be characterised by the normally relatively
long propodeum and fore wing with a fairly long marginal vein, stigmal vein and postmarginal
vein, the stigmal vein usually fairly straight and forming an angle of less than 45 with the
postmarginal. The hypopygium often reaches the apex of the gaster. Parastenoterys can easily be
recognised because of the characteristic sculpture of the mesopleurum, the extremely elongate,
relatively heavily sculptured propodeum and infuscate forewings.
PARATETRACNEMOIDEA Girault
(Key couplet: 280. Figs 170, 171)
Paratetracnemoidea Girault, 1915: 166. Type-species: Paratetracnemoidea breviventris Girault, by orig-
inal designation.
Rhinoencyrtus Mercet, 1918: 234. Type-species: Rhinoencyrtus malenotti Mercet, by monotypy. Syn. n.
INDO-PACIFIC ENCYRTIDAE 319
INCLUDED SPECIES. Two species, Palaearctic, Afrotropical, Oriental, Australasian; one species
from review area: breviventris Girault (19150: 166) (Australia), also undetermined material
from India and Vietnam (BMNH, BPBM).
BIOLOGY. Unknown.
COMMENTS. Trjapitzin (19730) incorrectly places the genus in the Tetracneminae as type-genus
of the tribe Rhinoencyrtini. Examination of material by phase-contrast shows the absence of
paratergites and also that the ovipositor structure is similar to that found in genera of the
Copidosomatini (Encyrtinae). Furthermore, the venation and arrangement of the sensillae at
the apex of the stigmal vein suggest an affinity with the Copidosomatini. General body shape and
morphology is also not unsimilar to Cowperia (Bothriothoracini) and it may be that the present
genus shows an unsuspected link between these two tribes.
PARATETRALOPHIDEA Girault
(Key couplets: 133, 463. Figs 69-71)
Paratetralophidea Girault, 1915a: 168. Type-species: Paratetralophidea ornatipennis Girault, by original
designation.
DISTRIBUTION AND SPECIES. One species, Australia only: ornatipennis Girault (1915a: 169), also
at least two further species from Australia and Indonesia (BMNH, BPBM).
BIOLOGY. The species from Indonesia (Seram I.) has been reared from heteropteran eggs
possibly belonging to the family Coreidae (Heteroptera).
COMMENTS. The genus is closely related to Ovaloencyrtus and probably also Xenoencyrtus and
Ooencyrtus (Microteryini, subtribe Ooencyrtina) from which it can be separated by the
characters given in the key.
PARECHTHRODRYINUS Girault
(Key couplet: 417. Figs 206, 207)
Parechthrodryinus Girault, 1916c: 480. Type-species: Parechthrodryinus convexus Girault, by original
designation.
DISTRIBUTION AND SPECIES. Eight species, Palaearctic, Afrotropical, Oriental and Australasian;
five species from review area: albidavatus (Shafee, Alam & Agarwal, 1975: 63) (India),
davicornis (Cameron, 1913: 101) (India, Sri Lanka), convexus Girault (1916c: 480) (Java),
hemiaspidoproctis (Subba Rao, 1967: 5) (India) and nigridavatus (Shafee, Alam & Agarwal,
1975: 60) (India).
REFERENCE. Review of Indian species: Shafee etal. (1975: 59-63).
BIOLOGY. Parasites of Coccidae and Keriidae (Homoptera).
COMMENTS. Placed in the tribe Cheiloneurini (Encyrtinae), it is very close to Tyndarichus from
which it can be very difficult to separate if the biology is not known (see characters given in key)
(see also comments under Tyndarichus).
PARECTROMOIDELLA Girault
(Key couplets: 83, 119, 147, 157, 245, 386. Figs 79, 400)
Parectromoidella Girault, 1915a: 175. Type-species: Parectromoidella thackerayi Girault, by monotypy.
Eucheiloneuropsis Girault, 1922ft: 104. Type-species: Eucheiloneuropsis lotae Girault, by original desig-
nation.
DISTRIBUTION AND SPECIES. Nine species, all Australian: abnormis (Girault, 1917g: 136) (comb,
n. from Dinocarsis), acaciae Girault (1931: 1), holbeini (Girault, I923e: 6) (comb. n. from
320 J. S. NOYES & M. HAY AT
Dinocarsis), laticincta (Girault, 19320: 3) (comb. n. from Epanusid) , lotae (Girault, 19226: 105),
lowelli (Girault, 19226: 105) (comb. n. from Eucheiloneuropsis) , pacorus (Walker, 1839: 39)
(comb. n. from Encyrtus), regalis (Girault, 19226: 106) (comb. n. from Eucheiloneuropsis) and
thackerayi Girault (1915a: 175), also many other species from New Caledonia, Australia and
New Zealand (BMNH, BPBM, UCR, DSIR).
REFERENCE. Noyes (1978: 551-552).
BIOLOGY. Unknown.
COMMENTS. The single extant male of Encyrtus pacorus Walker is here designated LECTO-
TYPE (BMNH) and has been so labelled; it belongs to Parectromoidella.
The holotype female of Dinocarsis abnormis cannot be located but from the description it
must belong to Parectromoidella.
The genus belongs to the same group as Epanusia, Cryptanusia and Cyrtocoryphes (see
comments under Cryptanusia).
PARECTROMOIDES Girault
(Key couplets: 326, 447)
Parectromoides Girault, I9l5a: 171. Type-species: Parectromoides magniscutellum Girault, by original
designation.
DISTRIBUTION AND SPECIES. Two species, Australia only: magniscutellum Girault (19150: 171)
and varipes (Girault, 19150: 166) (comb. n. from Parastenoterys), also further undetermined
material from Australia and New Zealand (BMNH, DSIR, QM, ANIC).
BIOLOGY. Unknown.
COMMENTS. Close to Clausenia (here placed in the tribe Charitopidini, Tetracneminae) and
superficially very similar. It differs mainly in having very much deeper, more irregular sculpture
on the head and dorsum of thorax, the forewing with a filum spinosum present (very unusual in
the Tetracneminae) and the gaster relatively shorter and more apically rounded (acute in
Clausenia).
PARENCYRTOMYIA Girault
(Key couplets: 232, 372, 463. Fig. 137)
Parencyrtomyia Girault, 1915a: 111. Type-species: Parencyrtomyia niveidava Girault, by original designa-
tion.
DISTRIBUTION AND SPECIES. One species, Australia only: niveidava Girault (19150: 111), also
undetermined material, including at least one undescribed species, from Vietnam, Solomon Is.
and Papua New Guinea (BPBM).
BIOLOGY. Unknown.
COMMENTS. We are unable to place the genus according to Trjapitzin's (19736) classification of
the Encyrtinae, but possibly it belongs to the same, group of genera as Tachinaephagus,
Rhytidothorax and Parastenoterys (see comments under Parastenoterys). It differs from these
genera in having a relatively shorter propodeum and the hypopygium not extending more than
halfway along the gaster. It may also be related toAseirba, Hemileucocerus and Austroencyrtus
(see comments under Aseirba).
PASULINIAgen.n.
(Key couplet: 440. Figs 228-230, 401-406)
Type-species: Pasulinia gentha sp. n. Gender: feminine.
INDO-PACIFIC ENCYRTIDAE 321
$. Head. In facial view clearly broader than long, in profile about one-half longer than broad, almost
straight from mouth margin to about half way up antennal scrobes and then gradually curved inwards in a
near semicircle to occipital margin. Eye with posterior margin very slightly concave, almost straight, only
slightly longer than broad, with sparse inconspicuous setae each not longer than the diameter of a facet and
not clearly separated from occiput by a more or less rounded occipital margin. Malar space about half eye
length, with sulcus present. Frontovertex between one-fifth and one-quarter head width; ocelli forming an
angle of about 45, the posterior ones nearly four times their own diameter from occipital margin and a little
less than their diameters from eye margin. Antennal scrobes shallow, meeting dorsally and reaching
slightly more than halfway from antennal toruli to anterior ocellus; antennal torulus separated from mouth
margin by about its own length and from other torulus by about one and one-half times its own length, its
dorsal margin about level with ventral eye margins; clypeal margin almost straight but extremely shallowly
excised medially. Antennal scape much longer than minimum width of frontovertex, subcylindrical, nearly
five times as long as broad, pedicel conical and slightly more than one third length of scape and about twice
as long as any of the funicle segments, the first three of which are a little longer than broad, subquadrate,
the last three of which are clearly transverse, the funicle distinctly widening distally; clava three-
segmented, more or less obliquely truncate , the outer suture clearly converging with inner one, clava about
two-thirds length of funicle; longitudinal sensillae on all flagellar segments except the first. Frontovertex
with shallow, raised, reticulate sculpture, at top of scrobes more or less squamiform-reticulate and between
eyes and genae becoming more longitudinally elongate; setae on frontovertex dark, sparse and not
conspicuous. Mandible with one very small lower tooth and a very broad truncation, almost edentate;
maxillary palpus three-segmented, labial palpus two-segmented.
Thorax. In side view slightly dorso-ventrally flattened with mesopleurum and propodeum narrowly in
contact with hind coxa and dorsally with mesoscutum and scutellum very flat. In dorsal view with posterior
margin of pronotum very concave, strongly angled centrally; visible part of mesoscutum about one-half
broader than long, with notaular lines absent, with posterior margin almost straight but produced a little
posteriorly; axillae more or less meeting; scutellum very slightly broader than long, a little shorter than
mesoscutum, with apex broadly rounded; propodeum medially about one-quarter length of scutellum.
Mesoscutum with moderately deep, raised reticulate sculpture; scutellum with distinctly deeper, raised,
reticulate sculpture of slightly smaller mesh; propodeum medially with shallow, irregular, raised reticulate
sculpture; mesopleurum with moderately deep, raised, reticulate sculpture. Forewing hyaline, but faintly
suffused yellow in middle one-third or so, wing about three times as long as broad, linea calva not
interrupted or closed, filum spinosum present; venation yellowish brown; submarginal vein without a
conspicuous hyaline break and not swollen apically; marginal vein about four or five times as long as broad,
clearly longer than stigmal; postmarginal vein almost absent; costal cell nearly 25 times as long as broad,
with a single line of setae in its apical one-sixth. Hindwing about four-fifths length of forewing, about five
and one-half times as long as broad, with marginal fringe about one-half maximum wing width. Mid tibial
spur a little shorter than basal mid tarsal segment.
Gaster. About as long as thorax with cereal plates in anterior half, ovipositor hardly exserted,
hypopygium reaching to about two-thirds to three-quarters along gaster; paratergites absent, last tergite
about two-thirds length of mid tibia; gonostyli free, about one-quarter as long as ovipositor which is nearly
as long as mid tibia.
O". Unknown.
COMMENTS. The shape of the mandible and head, and the forewing venation suggest that this
genus is related to Coccidencyrtus (tentatively placed in the Habrolepidini by Trjapitzin, 1973),
but also possibly related to Zaomma and Mahencyrtus (placed in the Cheiloneurini). Pasulinia
can be separated from Coccidencyrtus by the relatively narrow frontovertex and structure of the
clava (long and apically rounded with sutures parallel in Coccidencyrtus), and from the genera of
the Cheiloneurini by the structure of the mandible.
Pasulinia gentha sp. n.
(Figs 228-230, 401-406)
?. Length: 0-86-1-03 mm (holotype, 1-03 mm).
Colour. Head black with purple reflections, slightly brassy or greenish on lower parts of face; antenna
with scape testaceous yellow, pedicel and flagellum pale brown to dark brown; pronotum black with some
slight brassy reflections, mesoscutum shining blue-green edged purplish, anteriorly more greenish, axillae
322 J. S. NOYES & M. HAY AT
black with green, brassy or purple reflections, scutellum shining green slightly mixed coppery anteriorly
and apically; forewing hyaline, slightly suffused pale yellow in middle one-third; legs excluding fore coxa
completely yellow to slightly dusky orange, fore coxa brown; gaster dark purplish brown with some slight
brassy reflections, basal tergite and venter mostly orange; exserted part of gonostyli orange-brown.
Head. Relative measurements (holotype): head length 49, head width (facial view) 58, head width (side
view) 33, minimum frontovertex width 14, malar space 19, eye length 36, eye width 32, POL 4-5, OOL 2,
scape length 27, scape width 6, other proportions of antenna as in Fig. 230; mandible as in Fig. 229; head in
side view as in Fig. 228.
Thorax. Relative measurements (holotype): forewing length 132, width 45, other proportions of
forewing as in Fig. 401; hindwing length 104, width 18; forewing as in Fig. 401, sculpture of mesoscutum
and scutellum as in Figs 402, 403, mid tibia and tarsus as in Fig. 405.
Gaster. Relative lengths (paratype): last tergite 34, ovipositor 45, gonostyli 11, [mid tibia 50] ; genitalia as
in Fig. 406, hypopygium as in Fig. 404.
Cf . Unknown.
DISTRIBUTION. India, Sulawesi, Papua New Guinea.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype Q, Sulawesi: Tengah, nr Morowali, Ranu River Area, ii.1980, lowland rain forest, Malaise
trap (M. J. D. Brendell) (BMNH).
Paratypes. Sulawesi: 1 9, Tengah, same data as holotype, iii.1980 (M. J. D. Brendell). Papua New
Guinea: 1 $, East Highlands, Kundiawa, 6. i. 1965, Malaise trap (/. Sedlacek). India: 1 $, Kerala, Periyar
Animal sanctuary, 5-15.X.1979 (/. S. Noyes) (BMNH, BPBM).
PENTACLADOCERUS Erdos
(Key couplet: 355)
Pentacladocerus Erdos, 1963: 287. Type-species: Pentadadocerus matranus Erdos, by original desig-
nation.
DISTRIBUTION AND SPECIES. Three species, Palaearctic, plus one unidentified specimen from
India (BMNH).
REFERENCE. Re view of species: Trjapitzin (1968: 111-114).
BIOLOGY. Unknown.
COMMENTS. Placed in the Bothriothoracini, subtribe Coenocercina and separated from the
nearest related genera (Cerchysiella and Zaommoencyrtus) by the character given in the key.
PENTELICUS Howard
(Key couplets: 121, 207. Fig. 126)
Pentelicus Howard, 1895: 611. Type-species: Pentelicus aldrichi Howard, by monotypy.
Hemaenasius Ashmead, 19006: 374. Type-species: Hemaenasius confusus Ashmead, by original desig-
nation. Syn. n.
Epaenasomyia Girault, \9\ld: 3. Type-species: Epaenasomyia varicornis Girault, by original designation.
Syn. n.
Cowperella Girault, 1935: 4. Type-species: Cowperella aeneifrons Girault, by monotypy. Syn. n.
DISTRIBUTION AND SPECIES. Five species, Holarctic and Australian; one species from review area:
aeneifrons (Girault, 1935: 4) (comb. n. from Cowperella) (Australia), also undetermined
material from India and Taiwan to Australia (BMNH, BPBM).
REFERENCES. Notes on species: Trjapitzin & Gordh (1979), Khlopunov (1979).
BIOLOGY. Unknown.
COMMENTS. We can see no reason for retaining Hemaenasius as a distinct genus from Pentelicus.
INDO-PACIFIC ENCYRTIDAE 323
The only major difference between the two genera is the presence of deep piliferous punctures
on the head and dorsum of thorax in Pentelicus and their absence in the described species of
Hemaenasius . The undetermined material mentioned above shows virtually a complete range
from almost totally smooth sculpture to deeply punctured sculpture.
The classification of Trjaptzin & Gordh (1978ft) is curious. They place Hemaenasius in
the subtribe Hemaenasiina (Discodini) and Pentelicus in the subtribe Bothriothoracina
(Bothriothoracini), probably because of the difference in sculpture of the described species.
Pentelicus (as understood here) is probably related to Leurocerus, Proleurocerus , etc. (see
comments under Leurocerus) and can be distinguished from these and related genera by the
presence of a very shallow median longitudinal ridge or carina along the scutellum.
PHA ULOENCYRTUS Girault
(Key couplet: 201. Fig. 120)
Phauloencyrtus Girault, 1940: 150. Type-species: Phauloencyrtus mirisimilis Girault, by monotypy.
DISTRIBUTION AND SPECIES. One species only, Australasia: mirisimilis Girault (1940: 50)
(Sarawak, Australia).
BIOLOGY. Unknown.
COMMENTS. We are unable to place the genus, but the very hairy eyes and structure of the
antenna suggest that it may be related to Exoristobia or Parablatticida.
PHILOSINDIA gen. n.
(Key couplet: 160. Figs 89, 90, 407-410)
Type-species: Philosindia longicornis sp. n. Gender: feminine.
$. Head. In facial view a little broader than long, in profile about twice as long as broad, gently curved to
top of antennal toruli then strongly -curved at top of toruli and almost straight below this, thus the straight
part being nearly twice as long as the curved part. Eye with posterior margin straight or very slightly
concave, about one-quarter longer than broad, almost naked but with a few sparse, short setae, each much
shorter than the diameter of a facet, eye reaching or slightly overreaching occipital margin which is sharp.
Malar space a little shorter to distinctly longer than half length of eye, with sulcus absent or present.
Frontovertex from one-quarter to about one-half head width; ocelli large, more or less forming a right
angle; posterior ocellus much less than to about its own diameter from occipital or eye margin, or much
closer to eye margin. Antennal scrobes almost non-existent, not meeting dorsally, separated by a fairly
sharp interantennal prominence and reaching slightly less than half way from toruli to anterior ocellus;
antennal torulus separated from mouth margin by at least one and one-half times its own length and from
other torulus by about two-thirds its own length, its ventral margin clearly above or rarely well below
ventral margins of eyes; clypeal margin broadly but shallowly excised in middle. Antennal flagellum long,
about two to three times as long as head width; scape shorter or slightly longer than minimum width of
frontovertex, about three times as long as broad; pedicel conical and subquadrate, clearly much shorter
than any of the funicle segments which are subequal in size; clava three-segmented or with segments
separated and similar in appearance to funicle so that flagellum has an undifferentiated, nine-segmented
appearance; longitudinal sensillae very distinct, present on all flagellar segments; longest setae on
flagellum about as long as diameter of segments. Frontovertex with raised, reticulate sculpture of moderate
mesh, almost hexagonal in front of anterior ocellus, becoming more longitudinally elongate between
scrobes and eyes rnd on lower parts of face, interantennal prominence with similar sculpture to
frontovertex but d ; itinctly shallower; head with fairly long translucent or dark setae, those on frontovertex
about as long as or longer than the diameter of the ocelli. Mandible with two teeth and a truncation or
obscurely tridentate; maxillary palpus long, four-segmented, labial palpus three-segmented.
Thorax. In side view robust with mesoscutum and scutellum moderately convex, the metapleurum and
propodeum together narrowly in contact with hind coxa or slightly separated from it by posterior margin of
mesopleurum. In dorsal view with posterior margin of pronotum moderately concave; visible part of
mesoscutum nearly twice as broad as long, notaular lines absent, with its posterior margin almost straight
but slightly angled outwards in centre; axillae more or less meeting; scutellum about as long as
324 J. S. NO YES & M. HAY AT
mesoscutum, clearly convex, about as long as broad, its apex rounded; propodeum medially about
one-eighth to one-tenth as long as scutellum. Dorsum of thorax with conspicuous, long, dark setae and
shallow irregular, raised, reticulate sculpture, a little shallower than on head except occasionally on
scutellum which may be conspicuously deeper; propodeum medially quite smooth or with very few carinae,
around spiracles with shallow, raised, irregular sculpture, mesopleurum with very shallow, fine, elongate,
reticulate sculpture. Forewing hyaline or almost imperceptibly infuscate in basal two-thirds or so, about
two and one-half times as long as broad; linea calva not closed nor interrupted; filum spinosum present;
submarginal vein with an indistinct, apical, hyaline break, parastigma not or hardly swollen; costal cell
about 13 to 14 times as long as broad, with a single line of setae dorsally in distal half; marginal vein about
three to four times as long as broad, about as long as stigmal which in turn is as long as or a little shorter than
postmarginal vein. Hindwing about two-thirds as long as forewing, about three and one-half to five times as
long as broad, marginal fringe about one-sixth as long as wing width. Mid tibial spur about as long as basal
segment of mid tarsus.
Caster. Shorter than thorax; cereal plates in anterior half; hypopygium reaching to about one-half to
two- thirds along gaster; paratergites absent, last tergite about one-half to two-thirds as long as mid tibia;
ovipositor not exserted to slightly exserted with exserted part about one-quarter length of gaster,
ovipositor at least about as long as mid tibia, gonostyli fused to second valvifers and about one-fifth length
of ovipositor or longer.
Cf . Unknown.
COMMENTS. This genus belongs to the tribe Microteryini, subtribe Microteryina (Encyrtinae)
and can be distinguished from all other included genera by the extraordinarily long antenna,
relatively high placement of antennal toruli and the long postmarginal vein of the forewing.
Philosindia longicornis sp. n.
(Figs 89, 90, 407-410)
$. Length: 1-14-1 -27 mm (holotype, 1-24 mm).
Colour. Body generally dusky yellowish orange, gaster a little darker; dorsal margin of scape slightly
brownish apically, dorsal surface of flagellar segments and whole of two apical flagellar segments brownish;
head with translucent setae, scape, mesoscutum and scutellum with conspicuous dark setae; forewing
venation yellow.
Head. Malar sulcus present but indistinct, absent towards mouth margin; antennal flagellum a little more
than twice as long as maximum head width (2-08-2-23); posterior ocelli a little closer to eye margin than to
occipital margin ; antennal toruli with lower margins clearly above lower margins of eyes (Fig. 89) . Relative
measurements (holotype): head width (facial view) 80, head width (side view) 36, head length 70,
minimum frontovertex width 33, maximum diameter of posterior ocellus 8, malar space 20, eye length 46,
eye width 38, POL 14, OOL 3, scape length 29, scape width 10-5, overall length of flagellum 178,
proportions of antenna as in Fig. 407.
Thorax. Scutellum with sculpture similar to that of mesoscutum (Figs 409, 410) and head, not distinctly
deeper; propodeum medially smooth, about one-tenth as long as scutellum. Relative measurements
(holotype): forewing length 240, width 99, venation as in Fig. 90; hindwing length 156, width 44. The
relative width of the hindwing can vary; in one paratype it is almost exactly four times as long as broad.
Gaster. Ovipositor not exserted. Relative lengths (paratype): last tergite 22-5, ovipositor 39, gonostylus
7-5, [mid tibia 42]. Genitalia as in Fig. 408.
C? . Unknown.
DISTRIBUTION. Hong Kong.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $, Hong Kong: N.T., Taipokau, 27.viii,1965, light trap (Lee Kit Ming & Hui Wai Ming)
(BPBM).
Paratypes. Hong Kong: 3 $, same data as holotype; 1 $, same locality and collectors, 3-4.vii.1964
(BMNH, BPBM).
COMMENTS. A further eight species from India, Philippines, Papua New Guinea and Solomon Is.
(BMNH, BPBM, AMNH, USNM). They can be distinguished on several characters but mainly
INDO-PACIFIC ENCYRTIDAE 325
by the relative lengths of antenna to head width, position of antennal toruli in relation to eyes,
relative size of eye, coloration of head, length of last tergite of gaster in relation to mid tibia, and
relative length of exserted part of ovipositor.
PLAGIOMERUS Crawford
(Key couplet: 51)
Plagiomerus Crawford, 1910: 89. Type-species: Plagiomerus diaspidis Crawford, by original designation.
Parahomalopoda Girault, 1915c: 170. Type-species: Parahomalopoda peruviensis Girault, by original
designation.
DISTRIBUTION AND SPECIES. Six species, New World, Oriental, Pacific; four from review area:
bangaloriensis Shafee, Alam & Agarwal (1975: 102) (India), diaspidis Crawford (1910: 90)
(Hawaiian Is.), dorceto Trjapitzin (I969b: 1252) (S. China) and hospes Timberlake (1920: 428)
(Hawaiian Is.), also undetermined material from Taiwan and Java (BPBM).
REFERENCES. Review of some species: Shafee etal. (1975: 101-103); Beardsley (1976: 223).
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. The genus is placed in the Habrolepidini, subtribe Habrolepidina (Encyrtinae). It
can be separated from related genera by having hyaline wings and a four-segmented funicle with
the first joint being shorter than the fourth (see comments under Coccidencyrtus) .
PL A TYRHOPUS Erdos
(Key couplets: 152, 206)
Platyrhopus Erdos, 1955: 40. Type-species: Platyrhopus delitescens Erdos, by original designation.
DISTRIBUTION AND SPECIES. Two species, Palaearctic; neither from review area, but one
undescribed species from India (BMNH).
REFERENCE. Herthevtzian & Trjapitzin (1974).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the Anagyrini, subtribe Rhopina (Tetracneminae) and separated from
related genera by the characters given in the key.
PRALEUROCERUS Agarwal
(Key couplet: 117. Figs 57, 58, 411)
Paraleurocerus Agarwal, 1966: 68. Type-species: Paraleurocerus viridis Agarwal, by original designation.
[Homonym of Paraleurocerus Girault, 1915.]
Praleurocerus Agarwal, 1974: 394. [Replacement name for Paraleurocerus Agarwal.]
DISTRIBUTION AND SPECIES. One species, India and Sri Lanka only: viridis (Agarwal, 1966: 70).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The genus is related to Dinocarsis Forster (Tetracneminae, Dinocarsini) and can
easily be recognised by the flattened antennal flagellum and the thin flange at the apex of the
scutellum (Figs 57, 58).
PRIONOMASTIX Mayr
(Key couplets: 224, 278, 373, 429)
Prionomastix Mayr, 1876: 725. Type-species: Encyrtus morio Dalman, by monotypy.
Liocarus Thomson, 1876: 115, 121. Type-species: Encyrtus morio Dalman, by monotypy.
326 J. S. NO YES & M. HAYAT
Chestomorpha Ashmead, 1900ft: 370. Type-species: Chestomorphia biformis Ashmead, by original
designation.
Aprionomastix Girault, 19130: 68. Type-species: Aprionomastix fasciatipennis Girault, by original
designation.
DISTRIBUTION AND SPECIES. Seven species, cosmopolitan; none recorded from review area, but
several undetermined species from India, Thailand, Vietnam, Malaysia, Sarawak and Philip-
pines (BMNH, BPBM).
REFERENCE. Review of world species: Annecke (1962).
BIOLOGY. Parasites of nymphs of Membracidae (Homoptera).
COMMENTS. Placed in the Prionomasticini, subtribe Prionomasticina (Encyrtinae) (see also
comments under Anagyrodes).
PRIONOMITOIDES Girault
(Key couplet: 287)
Prionomitoides Girault, 1915a: 118. Type-species: Prionomitoides viridiscutellum Girault, by original
designation.
DISTRIBUTION AND SPECIES. One species, Australia only: viridiscutellum Girault (19150: 118).
BIOLOGY. Unknown.
COMMENTS. We are unable to place the genus confidently according to Trjapitzin's classification
of the Encyrtinae. In general appearance it superficially resembles species of Ooencyrtus or
Psyllaephagus, but differs by having a tridentate mandible and the hypopygium clearly reaching
the apex of the gaster. It may belong to the tribe Aphycini.
PROCHEILONEURUS Girault
(Key couplets: 94, 137, 384)
Procheiloneurus Girault, 19200: 39. Type-species: Procheiloneurus triguttatipennis Girault, by original
designation.
Raphaelana Girault, 1926ft: 66. [Unnecessary replacement name for Procheilonerus Girault.] Syn. n.
DISTRIBUTION AND SPECIES. Four species, all Australian: divinus (Girault, 1926ft: 69) (comb. n.
from Eusemionelld) , flaviscutellum Girault (19240: 5), perbellus Girault (19220: 43) and
triguttatipennis Girault (19200: 39), also several other species, near perbellus, from Australia
(BMNH, UCR, QM, ANIC).
BIOLOGY. Unknown.
COMMENTS. Girault (19266) unnecessarily proposed the replacement name Raphaelana for
Procheiloneurus Girault which he thought was a junior homonym of Prochiloneurus Silvestri,
1915 . According to Article 56a of the International Code of Zoological Nomenclature a one letter
difference is sufficient to prevent homonymy. Therefore the original name must stand.
The species placed within this genus almost certainly represent a polyphyletic group within the
Cheiloneurini. Further study will probably indicate that flaviscutellum and possibly also
triguttatipennis should belong in Cheiloneurus (and thus Procheiloneurus will become a junior
synonym of Cheiloneurus). However, in this case it will probably become necessary to describe
at least one new genus to accommodate perbellus and divina. For the present we are separating
Procheiloneurus from Cheiloneurus by the characters given in the key, in particular the presence
of two areas of dark setae in the basal cell of the forewing (Cheiloneurus has only one) and/or the
presence of a white, rectangular spot on each side of the pronotum (absent in Cheiloneurus}.
INDO-PACIFIC ENCYRTIDAE 327
PROCHILONEURUS Silvestri
(Key couplets: 103, 125, 254, 257, 357, 362)
Prochiloneurus Silvestri, 19156: 350. Type-species: Prochiloneurus pulchellus Silvestri, by original
designation.
Achrysopophagus Girault, 19150: 89. Type-species: Achrysopophagus oviductus Girault, by original
designation.
Parachrysopophagus Agarwal, 1965: 65. Type-species: Achrysopophagus insolitus Alam, by original
designation. [As subgenus of Achrysopophagus .]
Neoprochiloneurus Viggiani, 1966: 95. Type-species: Prochiloneurus bolivari Mercet, by original des-
ignation.
ProchiloneuroidesHayat, Alam & Agarwal, 1975: 61. Type-species: Prochiloneurus comperei Viggiani, by
original designation.
DISTRIBUTION AND SPECIES. Thirty-two species, cosmopolitan; 20 from review area: aegyptiacus
Mercet (1929: 360) (India), agarwali Hayat (19810: 23) (India), albifuniculus (Hayat, Alam &
Agarwal, 1975: 62) (India), albioviductus (Girault, 1925ft: 92) (comb. n. from Cheiloneurus)
(Australia), annulatus (Ferriere, 1951: 190) (comb. n. from Achrysopophagus) (Indonesia),
aureipleurum (Girault, 19320: 4) (comb. n. from Achrysopophagus) (Australia), clavatus
(Girault, 19150: 89) (Australia), comperei Viggiani (1970: 68) (India), hayati Shafee, Alam &
Agarwal (1975: 53) (India), indicus Shafee, Alam & Agarwal (1975: 49) (India), insolitus
(Alam, 1961: 235) (India), io (Girault, 1920c: 187) (Java, Philippines), javanicus (Ferriere,
1951: 188) (comb. n. from Achrysopophagus) (Indonesia), nigricornis (Girault, 1920c: 187)
(comb. n. from Achrysopophagus) (Hong Kong, Philippines), nigriflagellum (Girault, 19320: 6)
(comb. n. from Achrysopophagus) (Australia), oviductus (Girault, 19150: 89) (Australia), rex
(Girault, 1920c: 188) (Java, Philippines, Hawaiian Is.), taurus (Girault, 19230: 49) (comb. n.
from Achrysopophagus) (Australia), testaceus (Agarwal, 1965: 68) (India) and valparianus
Mani & Kaul in Mani et al. (1974: 66) (India), also undetermined material, containing several
undescribed species, from India, Hong Kong, Papua New Guinea, Australia, New Hebrides and
Hawaiian Is. (BMNH, BPBM, CNC, QM, ANIC, HC).
REFERENCE. Review of Indian species: Hayat (19810: 22-26).
BIOLOGY. Hyperparasites, via other encyrtids, of various families of Coccoidea (Homoptera),
mainly Pseudococcidae and Coccidae, and also Coccinellidae (Coleoptera).
COMMENTS. Placed in the tribe Cheiloneurini. It is most closely related to Cheiloneurus and
Tineophoctonus and can be separated from these by having the hypopygium reaching the apex of
the gaster, the ovipositor well exserted and the gaster apically rounded (not gradually tapered as
in species of the other genera with an exserted ovipositor).
PROLEUROCEROIDES Shafee, Alam & Agarwal
(Key couplet: 401. Figs 205, 416)
Proleuroceroides Shafee, Alam & Agarwal, 1975: 42. Type-species: Proleuroceroides pyrillae Shafee,
Alam & Agarwal, by original designation.
DISTRIBUTION AND SPECIES. Two species, both known only from India and possibly synonymous:
pyrillae (Crawford, 1916: 102) (comb. n. from Ooencyrtus) and pyrillae Shafee, Alam &
Agarwal (1975: 42), also undetermined material from Sulawesi (BMNH).
BIOLOGY. Parasites of eggs of Lophopidae (Homoptera).
COMMENTS. The holotype female of Ooencyrtus pyrillae Crawford has been examined (USNM).
It is very close to, if not the same as, pyrillae Shafee, Alam & Agarwal and for this reason we are
not proposing a replacement name for the latter.
Proleuroceroides is closely related to Proleurocerus (Encyrtinae, tribe Proleurocerini) and
can be separated by the characters given in the key, notably by the dorsum of the thorax being
non-metallic (metallic in Proleurocerus) (see also comments under Leurocerus).
328 J. S. NOYES & M. HAY AT
PROLEUROCERUS Ferriere
(Key couplet: 124. Figs 59, 60)
Proleurocerus Ferriere, 1935: 402. Type-species: Proleurocerus fulgoridis Ferriere, by original des-
ignation.
Arachnosinis Compere & Zinna, 1955: 112. Type-species: Arachnosinis zululandiae Compere & Zinna, by
original designation.
DISTRIBUTION AND SPECIES. Two species, Afrotropical and Oriental; only one known from review
area: fulgoridis Ferriere (1935: 403) (India), and one, undescribed species from India (BMNH).
BIOLOGY. Parasites of eggs of spiders (Araneida) and Eurybrachidae (Homoptera).
COMMENTS. Placed in the tribe Proleurocerini (Encyrtinae) but probably also related to
Leurocerus , Pentelicus, etc. (see comments under Leurocerus).
PROTYNDARICHOIDES Noyes
(Key couplets: 303, 318, 320, 397, 443. Figs 157, 231-234)
Protyndarichoides Noyes, 1980: 224. Type-species: Protyndarichoides nigriceps Noyes, by original
designation.
DISTRIBUTION AND SPECIES. Two described species, Neotropical, European, Afrotropical,
Oriental and Australasian; one species from review area: cinctiventris (Girault, 19346: 1)
(comb. n. from Echthrogonatopus) (Australia), also many undescribed species from India,
Bangladesh, S. China, Malaysia, Papua New Guinea, New Caledonia and New Zealand
(BMNH, BPBM, GC, DSIR).
BIOLOGY. Unknown, but has been found in association with scolytid beetles (Coleoptera,
Scolytidae) on Pinus sp. in France (BMNH).
COMMENTS. The material from New Zealand and France is very close to cinctiventris and may be
this species.
We are unable to place the genus satisfactorily but it may belong in the Cheiloneurini, as
suggested previously (Noyes, 1980: 225).
PSEUDAPHYCUS Clausen
(Key couplet: 66. Figs 25-27)
Pseudaphycus Clausen, 1915: 41. Type-species: Aphycus angelicus Howard, by original designation.
Psilomirinus Brethes, 1916: 424. Type-species: Psilomirinus flavidulus Brethes, by original designation.
DISTRIBUTION AND SPECIES. Twenty-five species, cosmopolitan; two from review area: orientalis
Ferriere (1937: 317) (Philippines) and utilis Timberlake (1923: 323) (Hawaiian Is.), also further
undetermined material from S. China and Cook Is. (BMNH, BPBM, DSIR).
REFERENCE. World revision: Gahan (1946).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Pseudaphycus utilis Timberlake is possibly out of place in this genus, having the
dorsum of the thorax convex and clothed in conspicuous dark setae. However, at present we do
not regard these differences as sufficient to warrant separation into another genus.
Placed in the Aphycini, subtribe Aphycina (Encyrtinae). It is very close to Acerophagus and
Pseudectroma and can be separated from these genera by the characters given in the key.
INDO-PACIFIC ENCYRTIDAE 329
PSEUDECTROMA Girault
(Key couplet: 63. Fig. 28)
Pseudectroma Girault, 1915a: 161. Type-species: Pseudectroma auricorpus Girault, by original des-
ignation.
Timberlakia Mercet, I925a: 9. Type-species: Acerophagus europaeus Mercet, by original designation.
Syn. n.
DISTRIBUTION AND SPECIES. Seven described species, Neotropical, European, Afrotropical,
Australasian; three species from review area, all Australian: auricorpus Girault (1915a: 161),
bryanti Girault (1922e: 150) and obscura Girault (1923c: 143), also further undetermined
material from Malaysia and Cook Is. (BMNH, DSIR).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The clava of bryanti may be solid and thus the species could run to either
Acerophagus or Indaphycus in the key. However, we are retaining it in its original combi-
nation pending closer examination of the single extant syntype (QM) or more freshly collected
material.
The genus belongs to the Aphycini, subtribe Aphycina (Encyrtinae) and is probably closest to
Acerophagus. Generally it can be distinguished from Acerophagus by having a two-segmented
clava, whereas Acerophagus usually has three segments. The relative width of the frontovertex
is probably a more reliable character: in Acerophagus, at its narrowest point, it is not wider than
the scape length, whereas in Pseudectroma it is at least about one-quarter wider than the scape
length. Prinsloo (1982) has suggested that these and some related genera may eventually by
synonymised and we echo these sentiments. However, he also described two new species from
South Africa (under Timberlakia}, each with a relatively narrow frontovertex. Although both of
these species have a two-segmented clava, they may actually belong in Acerophagus as defined
here.
PSEUDOCOCCOBIUS Timberlake
(Key couplets: 153, 167, 360, 390. Fig. 197)
Pseudococcobius Timberlake, 1916: 563. Type-species: Aphycus terryi Fullaway, by original designation.
Australrhopoideus Girault, 1926ft: 58. Type-species: Australrhopoideus melleicorpus Girault, by mono-
typy. Syn. n.
Pezaphycus Nowicki, 1926: 105. Type-species: Pezaphycus obenbergeri Nowicki, by original designation.
Syn. n.
DISTRIBUTION AND SPECIES. Four species, Europe, Australia, Pacific; three species from review
area: melleicorpus (Girault, 19266: 58) (comb. n. from Australrhopoideus) (Australia), quin-
queguttatus (Girault, 19256: 93) (comb. n. from Aphycus) (Australia) and terryi (Fullaway,
1913: 281) (Hawaiian Is.), also undetermined material from Hong Kong (BPBM).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Pseudococcobius has, in the past, been treated as a synonym of Aphycus, although
Pezaphycus has generally been regarded as a distinct genus (even though their respective
type-species are extremely close). We have reinstated it as a good genus since it does appear to
differ significantly from species that we regard as belonging to Aphycus, notably in the very
different head shape and relatively smaller eye with a convex posterior margin (that in Aphycus
is slightly concave, thus giving the eye a kidney-shaped appearance). There are also differences
in structure of the thorax (Pseudococcobius has notaular lines always reaching about one-third
to one-half way across the mesoscutum and the sculpture of the thoracic dorsum shallow and
smooth). In general the antenna is also shorter, with the clava about as long as or longer than the
funicle, whereas in Aphycus the clava is shorter than the funicle.
Pseudococcobius belongs to the Aphycini, subtribe Aphycina (Encyrtinae).
330 J. S. NO YES & M. HAY AT
PSYLLAEPHAGUS Ashmead
(Key couplets: 142, 181, 204, 218, 252, 289, 301, 345, 433, 448, 456, 462, 495, 525.
Figs 74, 75, 102, 121,122)
Psyllaephagus Ashmead, 1900ft: 382. Type-species: Encyrtus pachypsyllae Howard, by original designa-
tion.
Mirocerus Ashmead, 1904c: 309. Type-species: Mirocerus peyelae Ashmead, by original designation.
Calocerineloides Girault, 1913e: 111. Type-species: Calocerineloides ramosa Girault, by original desig-
nation. Syn. n.
Epanagyrus Girault, 1915a: 160. Type-species: Epanagyrus punctatiscutum Girault, by original desig-
nation. Syn. n.
Anagyropsis Girault, 1917g: 136. Type-species: Anagyrus purpureus Girault, by original designation.
Syn. n.
Metaprionomitus Mercet, 1921: 260. Type-species: Metaprionomitus intermedius Mercet, by original
designation.
Shakespearia Girault, 1928a: 3. Type-species: Shakespearia flabellata Girault, by monotypy.
Psyllencyrtus Tachikawa, 1955: 63. Type-species: Psyllencyrtus syntomozae Tachikawa, by original
designation.
Calluniphilus Erdos, 1961: 413. Type-species: Calluniphilus vendicus Erdos, by monotypy.
Ooencyrtoides Hoffer, 1963: 568. Type-species: Ooencyrtus albopilosus Hoffer, by original designation.
Propsyllaephagus Blanchard in De Santis, 1964: 235. Type-species: Propsyllaephagus trellesi Blanchard,
by original designation.
Mercetia Bakkendorf, 1965: 139. Type-species: Copidosoma lusitanicum Mercet, by original designation.
Kaszabicyrtus Szelenyi, 1971: 389. Type-species: Kaszabicyrtus acutigastris Szelenyi, by original des-
ignation.
DISTRIBUTION AND SPECIES. About 150 species, cosmopolitan; 107 species from review area, all
from Australia except aligarhensis which is from India: *abyssus Riek (1962d: 710), aeneoculex
(Girault, 19296: 312) (comb. n. from Coccidoxenus) , albiclava (Girault, 19150: 135) (comb. n.
from Anagyrus) , alienus Riek (1962 d: 707), aligarhensis Shafee, Alam & Agarwal (1975: 100),
anna (Girault, 1938: 83) (comb. n. from Anagyropsis), aquilus Riek (I962d: 699), *arctatus Riek
(I962d: 713), arduus Riek (1962d: 714), *argutus Riek (1962d: 715), arsanes (Walker, 1839: 38)
(comb. n. from Encyrtus), ascitus Riek (1962d: 704), *asser Riek (1962d: 712), *atavus Riek
(1962d: 701), *atratus Riek (I962d: 699), attenuatus Riek (1962d: 711), auricorpus (Girault,
19150: 133) (comb. n. from Anagyrus), australiensis Girault (19140: 29) (comb. n. from
Anagyrus), avus Riek (1962d: 706), *basileus Riek (I962d: 721), blandus Riek (I962d: 721),
*bliteus Riek (1962d: 722), boletus Riek (1962d: 718), "bolus Riek (I962d: 719), *brachiatus
Riek (1962d: 726), brevicornis (Girault, 1926c: 129) (comb. n. from Coccidoxenus), broccus
Riek (I962d: 716), bruchus Riek (1962d: 720), burnsi (Girault, 19210: 2) (comb. n. from
Anagyropsis), carinatus Riek (1962d: 735), cellinini (Girault, 19150: 134) (comb. n. from
Anagyrus), channingi (Girault, 1913e: 111) (comb. n. from Anagyrus) , cicada (Girault, 19150:
137) (comb. n. from Anagyrus) (= Paraenasomyia dubia Girault, 19230: 48 syn. n.), cinctorum
(Girault, 19230: 47) (comb. n. from Paraenasomyia), *clarus Riek (19620 1 : 745), compactus
(Girault, 19230: 50) (comb. n. from Coccidoxenus), concisus Riek (1962d: 747), cornuatus Riek
(I962d: 731), *cornuphagus Riek (1962d: 754), *dignus Riek (1962d: 724), *discretus Riek
(1962d: 723), dius (Girault, 19150: 137) (comb. n. horn Anagyrus) , dyari (Girault, 19150: 137)
(comb. n. from Anagyrus), * emarginatus Riek (1962d: 731), emersoni (Girault, 1913e: 113)
(comb. n. from Anagyrus), *excisus Riek (1962d: 737), *exiguus Riek (19620 1 : 748), *facetus
Riek (1962d: 745), *facilis Riek (1962d: 741), *faustus Riek (19620 1 : 74Q),flabellatus (Girault,
19280: 3), *fundus Riek (1962d: 742), *funiculus Riek (I962d: 738), gemitus Riek (I962d: 755),
gram (Girault, 19150: 135) (comb. n. from A nagyrus) , guttatipes (Girault, 19150: 134) (comb. n.
from Anagyrus), hardyi (Girault, 1922f: 1) (comb. n. from Blastothrix) , hegeli (Girault, 19150:
136) (comb. n. from Anagyrus), hirtus Riek (1962d: 744), howardi (Girault, 19150: 134)
(comb. n. from Anagyrus), irvingi (Girault, 19220: 44) (comb. n. from Anagyropsis), *longis-
simus Riek (1962d: 733), longistylus (Girault, 19296: 312) (comb. n. from Anagyropsis),
mazzinini (Girault, 19150: 133) (comb. n. from Anagyrus), mercurius (Girault, 19220: 41)
INDO-PACIFIC ENCYRTIDAE 331
(comb. n. fromAnagyropsis), minutellus (Girault, 19150: 171) (comb. n. from Tetracnemella) ,
*neoxenus Riek (I962d: 752), novipurpureus (Girault, 19150: 136) (comb. n. from Anagyrus) ,
pallidipes (Girault, 19150: 81) (comb. n. from Aenasiella), *paradoxus Riek (I962d: 705),
parvus Riek (I962d: 749), pegasus (Girault, 19230: 48) (comb. n. from Paraenosomyia), penni
(Girault, 1913e: 112) (comb. n. from Anagyrus}, perplexus Riek (1962d: 750), positus Riek
(I962d: 729), probus Riek (\962d: 736), prolatus Riek (19620 1 : 734), punctatiscutum (Girault,
19150: 160) (comb. n. from Epanagyrus), purpureus (Girault, 19150: 133) (comb. n. from
Anagyrus}, * quadriannellus Riek (I962d: 751), quadricyclus Riek (19620 1 : 751), ramosus
(Girault, 1913e: 111) (comb. n. from Calocerineloides) , resolutus Riek (\962d: 730), richteri
(Girault, 1923c: 142) (comb. n. fromAnagyropsis), mbensi (Girault, 19320: 1) (comb. n. from
Coccidoxenus}, semicitripes (Girault, 19266: 66) (comb. n. from Coccidoxenus) , similis Riek
(I962d: 738), smaragdus (Girault, 19390: 19) (comb. n. from Anagyropsis), spondyliaspidis
(Girault, 19390: 19) (comb. n. from Anagyropsis), spongitus (Girault, 19150: 136) (comb. n.
from Anagyrus), subgiganteus (Girault, 19150: 138), (comb. n. from Anagyrus} (= Psyllaepha-
gus usticius Riek, 19620": 695 syn. n.), suburbis (Girault, 19266: 67) (comb. n. from Blastothrix)
(= Psyllaephagus fuscus Riek, I962d: 753 syn. n.), terraefilius (Girault, 1938: 83) (comb. n. from
Anagyropsis), turbulentus (Girault, 19200: 48) (comb. n. from Anagyropsis) , turneri (Girault,
19256: 100) (comb. n. from Blastothrix), *undnatus Riek (19620 1 : 691), unus Riek (19620 1 : 697),
*utilis Riek (I962d: 693), viridiscutellum (Girault, 19150: 171) (comb. n. from Tetracnemella),
westralis Riek (I962d: 714, worcesteri (Girault, 19150: 139) (comb. n. from Coccidoxenus),
wundti (Girault, 19150: 140) (comb. n. from Coccidoxenus), xenus Riek (1962O 1 : 714), *xi Riek
(19620 1 : 728), xuthus (Walker, 1839: 38) (comb. n. from Encyrtus), *ypsilon Riek (1962d: 732),
and zameis (Walker, 1839: 39) (comb. n. from Encyrtus), also much undetermined material
from throughout the region (BMNH, BPBM, DSIR, CNC, QM, ANIC, HC).
REFERENCES. Revision of some Australian species: Riek (19620 7 ); key to Palaearctic species:
Trjapitzin (1981).
BIOLOGY. Parasites or hyperparasites of nymphs of Psyllidae (Homoptera).
COMMENTS. The single extant syntype female of Encyrtus arsanes Walker (BMNH) is here
designated LECTOTYPE. The single extant syntype male of Encyrtus zameis Walker (BMNH)
is here designated LECTOTYPE; it is lacking both pairs of wings. There are two syntypes of
Encyrtus xuthus Walker in the BMNH; one is in very poor condition and the other does not quite
fit Walker's description. They both belong to Psyllaephagus, but we are not selecting a lectotype
for this species at present.
Riek (1962d) described a number of species from Australia. Unfortunately he failed to label
the holotypes (or paratypes) of any of his species even though he cited these in his descriptions.
During a visit to ANIC, Canberra, one of us (JSN) selected a primary type from those specimens
of the type-series of each species where the data of more than one specimen agreed with the data
of the holotype of that species as published by Riek . The names of species for which this has been
done are preceded in the above list by an asterisk (*). These specimens are here designated
LECTOTYPE and have been labelled as such.
Several species here placed in other genera may actually belong in Psyllaephagus, e.g.
Aenasiella sidneyi (Girault) and other species placed in that genus and also Parachalcerinys
nonaericornis Girault. Psyllaephagus is so enormously complex in Australia that it is exceed-
ingly difficult to define its limits and there are possibly as many as 1 ,000 species to be found there.
Psyllaephagus belongs to the tribe Trechnitini, subtribe Metaprionomitina and is largely
characterised by its brightly metallic green or blue-green colour, punctiform marginal vein of the
forewing, the mandible having one or two teeth and a broad truncation, and the hypopygium not
extending more than two-thirds along the gaster. However, there are exceptions to each of these
characters.
332 J. S. NO YES & M. HAY AT
PSYLLAPHYCUS Hayat
(Key couplet: 411)
Psyllaphycus Hayat, 1972: 207. Type-species: Psyllaphycus diaphorinae Hayat, by original designation.
DISTRIBUTION AND SPECIES. One species, India only: diaphorinae Hayat (1972: 208).
BIOLOGY. Parasites of nymphs of Psyllidae (Homoptera).
COMMENTS. Placed in the Microteryini, subtribe Syrphophagina (Encyrtinae). It can easily be
distinguished from related genera by the bright yellow colour of the body and the mandible
having a single tooth and a broad truncation.
RAFFAELLIA Girault
(Key couplet: 69)
Raffaellia Girault, I922d: 205. Type-species: Raff ae Ilia Sidney i Girault, by monotypy.
Raffaellisca Ghesquiere, 1946: 369. [Unnecessary replacement name for Raffaellia Girault.] Syn. n.
DISTRIBUTION AND SPECIES. One species, Australia only: sidneyi (Girault, 1922d: 205).
BIOLOGY. Unknown.
COMMENTS. Very close to Copidosomopsis (tribe Copidosomatini, subtribe Copidosomatina)
from which it can be separated using the characters given in the key.
RHOPALENCYRTOIDEA Girault
(Key couplets: 219, 300, 369. Fig. 123)
Rhopalencyrtoidea Girault, 1915a: 101. Type-species: Rhopalencyrtoidea purpureicorpus Girault, by
original designation.
DISTRIBUTION AND SPECIES. Three species, all Australian: austrina Girault (19296: 313) , perplexa
(Girault, 1925a: 3) (comb. n. from Nezarhopalus) and purpureicorpus Girault (1915a: 101).
BIOLOGY. Unknown.
COMMENTS. Related to Coccidoctonus and Teleterebmtus (see comments under Coccidoctonus} .
It can be separated from other related genera by having the apex of the hypopygium more or less
reaching the apex of the gaster and not beyond, and the fore wing having the postmarginal vein
longer than the stigmal.
KHOPl/SForster
(Key couplets: 84, 170, 273, 395, 404. Figs 40, 91, 412-414)
Rhopus Forster, 1856: 34. Type-species: Encyrtuspiso Walker, by original designation.
Xanthoencyrtus Ashmead, 1902: 302. Type-species: Xanthoencyrtus nigrodavatus Ashmead, by
monotypy.
Scelioencyrtus Girault, 1915a: 161. Type-species: Scelioencyrtus nigridavus Girault, by original des-
ignation.
Mirastymachus Girault, 1915a: 166. Type-species: Mirastymachus europaeus Girault, by original des-
ignation.
Pholidoceras Mercet, 1918: 237. Type-species: Pholidoceras brachyptera Mercet, by monotypy.
Pholidocerodes Ferriere, 1956: 358. Type-species: Pholidoceras parvula Mercet, by original designation.
DISTRIBUTION AND SPECIES. Thirty-six species, cosmopolitan; 17 species from review area:
apterus (Timberlake, 19196: 201) (Hawaiian Is.), bridwelli (Timberlake, 1920: 420) (Hawaiian
Is.), desantisiellus Ghesquiere (1957: 18) (India), extradavus (Girault, 1922e: 149) (comb. n.
from Xanthoencyrtus} ( Australia), fullawayi (Timberlake, 19196: 204) (India, Hawaiian Is.),
garibaldia (Girault, 1933: 4) (comb. n. from Xanthoencyrtus) (Australia), gramineus Hayat
INDO-PACIFIC ENCYRTIDAE 333
(1970a: 110) (India), keatsi (Girault, 19150: 162) (comb. n. from Scelioencyrtus) (Australia),
laysanensis (Timberlake, I9l9b: 203) (Hawaiian Is.), longidavatus (Shafee, Alam & Agarwal,
1975: 31) (India), nigridavus (Girault, 19150: 161) (Australia), qadrii (Shafee, Alam &
Agarwal, 1975: 30) (India), sacchari (Alam, 1961: 239) (India), sanguineus (Timberlake, 1920:
416) (Hawaiian Is.), semiflavus (Timberlake, I9l9b: 204) (Hawaiian Is.), semiluteus (Timber-
lake, 1920: 419) (Hawaiian Is.) and tricolor (Girault, 19150: 162) (comb. n. from Scelioencyrtus)
(Australia), also much undetermined material from throughout the region (BMNH, BPBM,
DSIR,QM,ANIC,HC).
REFERENCES. Review of Hawaiian species: Timberlake (1920: 413-421); review of Indian
species: Shafee etal. (1975: 30-36).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. The species of this genus are exceedingly difficult to separate since coloration is not
at all reliable. Apart from the relative lengths of the funicle segments, distribution of setae
around the linea calva and in the basal cell and relative dimensions of the forewing, we have
found that the position and number of the dark erect seta(e) between the posterior ocellus and
eye to be of considerable use in separating the species.
Placed in the Anagyrini, subtribe Rhopina (Tetracneminae).
RHYTIDOTHORAX Ashmead
(Key couplets: 176, 216, 236, 306, 316, 331, 399, 450, 467, 530. Figs 99, 139, 415)
Rhytidothorax Ashmead, 1900ft: 377. Type-species: Rhytidothorax marlatti Ashmead, by original designa-
tion.
Anusomyia Girault, 1915a: 164. Type-species: Anusomyia auratiscutum Girault, by original designation.
Syn. n.
Ectromoides Girault, 1915a: 167. Type-species: Ectromoides purpureiscutellum Girault, by monotypy.
Syn. n.
Mesanusomyia Girault, I922a: 48. Type-species: Mesanusomyiafera Girault, by monotypy. Syn. n.
Swazencyrtus Prinsloo & Annecke, 1979: 379. Type-species: Swazencyrtus latiscapus Prinsloo & Annecke,
by original designation. Syn. n.
DISTRIBUTION AND SPECIES. Eleven species, New World, Afrotropical, Oriental, Australasian;
four species from review area, all Australian: aereiscutellum (Girault, 19150: 164) (comb. n.
from Anusomyia), auratiscutum (Girault, 19150: 164) (comb. n. from Anusomyia), ferus
(Girault, 19220: 48) (comb. n. from Mesanusomyia) and purpureiscutellum (Girault, 19150: 168)
(comb. n. from Ectromoides), also many undescribed species from India, Hong Kong and the
Philippines to Australia and New Caledonia (BMNH, BPBM, USNM, CNC, QM, ANIC).
BIOLOGY. Unknown.
COMMENTS. The synonymies proposed above may be difficult to accept, particularly if only the
type-species of each of the five genera are examined since they appear to be very morphologi-
cally diverse (except perhaps ferus and purpureiscutellum). However, we have examined
probably scores of species from all areas and find that most characters which may be used to
separate genera are totally unreliable, e.g. number of teeth on the mandible (the mandibles vary
from unidentate to tridentate), shape of the head, sculpture of head and dorsum of thorax,
relative length of postmarginal vein of forewing and relative position of the apex of the
hypopygium. All the species have these three important characters in common: a similar basic
type of wing venation, relatively long propodeum and, in particular, the structure of the
ovipositor. The latter is very unusual in the Encyrtinae in that the third valvulae (gonostyli) are
completely fused to the second valvifers (Fig. 415) and also no part of the female genitalia is
visible externally unless the ovipositor is partially or totally exserted in the egg-laying position.
We believe that to keep all of these genera separate at this stage could eventually lead to the total
confusion that now seems to exist in the Anagyrini (Tetracneminae) where new genera have
334 J- S. NO YES & M. HAYAT
been described for species which do not quite fit the narrowly defined and unnatural limits of
already existing genera. Furthermore we do not think that the morphological diversity of
Rhytidothorax, as defined here, is any greater than in Copidosoma.
The genus is close to Tachinaephagus and can be separated by the structure of the ovipositor.
In Tachinaephagus the third valvulae are free and visible externally. The two genera are
probably related to Parastenoterys (see comments under Parastenoterys).
RUANDEROMA gen. n.
(Key couplet: 228, Figs 134-136, 417-421)
Type-species: Ruanderomasankaranisp. n. Gender: feminine.
9- Head. In facial view slightly broader than long, in profile about twice as long as broad and anteriorly
more or less gradually and evenly curved. Eye with posterior margin slightly concave, slightly more than
one and one-half times as long as broad, more or less naked, with extremely few short hairs and very nearly
reaching occipital margin which is sharp. Malar space about one-third length of an eye, malar sulcus
present. Frontovertex slightly less than half head width; ocelli in an acute angle, nearly forming a right
angle, posterior ocellus about its own diameter from eye margin and about twice this from occipital margin.
Antennal scrobes meeting dorsally, very short, reaching about one-fifth way from antennal toruli to
anterior ocellus, sharply delimited dorsally by a transverse ridge which nearly runs from eye to eye;
antennal torulus separated from mouth margin by slightly less than its own length and from other torulus by
about its own length, its mid line about level with ventral margins of eyes; clypeal margin broadly and
shallowly excised. Antennal scape much longer than minimum width of front overtex, cylindrical, about
seven or eight times as long as broad, pedicel conical, a little less than one-fifth length of scape, less than
half as long as first funicle segment and not quite as long as sixth; funicle segments cylindrical, all longer
than broad, the first the longest and gradually shortening distally; clava three-segmented, about one-
quarter as long as funicle, with apex rounded but outer suture distinctly converging with inner; longitudinal
sensillae on all flagellar segments. Frontovertex with numerous, deep piliferous punctures each separated
by a little less than their own diameters and thus giving it a thimble-like appearance, the area between the
punctures with shallow, irregular, raised reticulate sculpture, below ridge at top of antennal scrobes more
regular and piliferous punctures distinct only on genae but here rather small; setae on frontovertex short,
hardly longer than diameter of an ocellus. Mandible with three teeth (Fig. 136), the inner and outer ones
rather short, the middle one quite long; maxillary palpus four-segmented, labial palpus three-segmented.
Thorax. In side view robust with mesoscutum and scutellum moderately convex, the metapleurum and
propodeum together quite broadly in contact with the hind coxa. In dorsal view posterior margin of
pronotum moderately concave, visible part of mesoscutum about two-thirds broader than long with
notaular lines in anterior half, its posterior margin slightly convex; axillae meeting; scutellum a little
shorter than mesoscutum with a distinct subapical carina and with its apex broadly rounded; propodeum
medially about half as long as scutellum with a pair of submedian carinae between which is some very
shallow irregular rugose sculpture. Mesoscutum, axillae and scutellum with irregular, very shallow, raised
reticulate sculpture; dorsum of thorax with numerous inconspicuous dark brown setae. Forewing infuscate
with base hyaline and several wedge-shaped hyaline marks, a little over three times as long as broad; linea
calva broadly closed by several lines of setae near posterior wing margin; filum spinosum absent;
submarginal vein without an apical hyaline break, slightly swollen apically; costal cell more than 30 times as
long as broad, with a single line of setae dorsally in its distal one-third or so; marginal vein about 12 times as
long as broad, subequal in length to postmarginal which is nearly twice as long as stigmal; submarginal vein
with eight or nine very long conspicuous setae on its ventral surface at about two-thirds along its length,
each seta at least three times as long as maximum diameter of submarginal vein at this point. Hindwing
lightly infuscate, almost hyaline, about two-thirds as long as forewing and about six times as long as broad,
its marginal fringe about one-quarter to one-third wing width. Mid tibial spur about as long as basal mid
tarsal segment.
Caster. Much shorter than thorax, cereal plates about midway along its length; hypopygium reaching
apex of gaster, paratergites not distinct in availabe slide-mounted material; last tergite slightly more than
one-third as long as mid tibia, ovipositor a little less than half as long as mid tibia, gonostyli fused to second
valvifers, about one-quarter as long as ovipositor.
Cf . Only available male is slide-mounted, but apparently differs from female as follows. Eye clearly smaller
than in female so that frontovertex distinctly more than half head width and malar space about half as long
INDO-PACIFIC ENCYRTIDAE 335
as an eye; transverse ridge above antennal scrobes not present, or if so then not distinct; antennal torulus
separated from mouth margin by about one and one-half times its own length , from other torulus by its own
length, its lowest margin only slightly below lower eye margins; antennal scape slightly broader than
minimum width of frontovertex, cylindrical, about five times as long as broad (may be a little less because
material is slide-mounted), pedicel about one-quarter length of scape, conical, a little longer than broad
but not more than half length of any funicle segment, first funicle segment longest, about six times as long as
broad, sixth shortest, about three times as long as broad, clava entire, about as long as first funicle segment,
setae on flagellum about as long as diameter of segments, longitudinal sensillae present on all flagellar
segments but first; forewing infuscate, but less strongly so than in female; aedeagus a little shorter than half
length of mid tibia or about one and one-half times as long as mid tibial spur, digiti a little less than one-fifth
length of aedeagus, with apical teeth present.
COMMENTS. At first glance it is not easy to place this genus in either of the recognised subfamilies
of the Encyrtidae since it superficially resembles both Callipteroma (Tetracneminae, Anagyrini)
and Ruandella (Encyrtinae, Microteryini). The structure of the ovipositor, gaster and wing
venation clearly point to it belonging to the Tetracneminae, but it cannot be placed in the tribe
Anagyrini because of the presence of notaular lines and clearly tridentate mandibles. The
presence of notaular lines, ovipositor structure and apparent absence of paratergites suggest
that the genus can be best placed in the Charitopidini although it is somewhat out of place here,
having strongly infuscate forewings and different venation.
The type-species of the genus is named in honour of Dr T. Sankaran (Commonwealth Institute
of Biological Control, Bangalore, India).
Ruanderoma sankarani sp. n.
(Figs 134-136, 417-421)
?. Length: 1-31-1-57 mm (holotype, 1-57 mm).
Colour. Holotype with head dark metallic green, between punctures with weak purple reflections; scape
testaceous yellow, pedicel and flagellum dark brown; pronotum, sides and venter of thorax dark
orange-brown, mesoscutum, scutellum and axillae dark brown with green, blue and brassy reflections;
forewing infuscate with pattern as in Fig. 134; legs orange with fore and mid coxae brown, mid tibia a little
paler than mid femur, hind femur and tibia which are a little brownish; gaster dark brown with purple and
brassy reflections, basal segment orange. There is some variation in colour: the purple colour between the
piliferous punctures of the frontovertex can be quite strong and one paratype has the thorax mostly orange
(including coxae) with only the mid line of the mesoscutum slightly metallic, the axillae and scutellum with
weak purple reflections, the head of this specimen is less strongly metallic green with a hint of orange, the
lower parts of the face being distinctly orange.
Head. Relative measurements (holotype): head length 36, head width (facial view) 41, head width (side
view) 18, minimum frontovertex width 18, malar space 9, eye length 28-5, eye width 17, POL 12, OOL 2,
scape length 25, scape width 3, other proportions of antenna as in Fig. 420. Mandible as in Fig. 136.
Thorax (Fig. 135). Relative measurements (holotype): forewing length 112, other proportions of
forewing as in Fig. 134; hindwing length 82, hindwing width 13.
Gaster. Relative lengths (paratype): last tergite 60, ovipositor 76, gonostyli 16, [mid tibia 166]; genitalia
as in Fig. 417.
Cf . Length: approx. 1-33 mm. Generally differs from female in structure of antenna (Fig. 421), size of eye
and genitalia (Figs 418, 419). Relative measurements (paratype): head width 78, minimum frontovertex
width 45, scape length 40, aedeagus length 42, mid tibia 103, mid tibial spur 27.
DISTRIBUTION. India.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $, India: Karnataka, 25 km W. of Mudigere, 28.x-3.xi. 1979 (/. S. Noyes) (BMNH).
Paratypes. India: 2 $, same data as holotype; 1 $, Bangalore, iii.1979 (T. Sankaran); 1 cf, Himachal
Pradesh, Manali, Bilaspur, 13.x. 1979 (Z. Boucek) (BMNH).
336 J. S. NOYES & M. HAY AT
RUSKINIANA Girault
(Key couplet: 97)
Ruskiniana Girault, 1923e: 5. Type-species: Ruskiniana sexguttatipennis Girault, by monotypy.
DISTRIBUTION AND SPECIES. One species, Australia only: sexguttatipennis (1923e: 5), also one
further species from Australia (BMNH).
BIOLOGY. Unknown.
COMMENTS. Girault described this genus and species from at least two specimens, both of which
appear to have been lost. However, three specimens (BMNH) agree totally with Girault's brief
description and our interpretation of the genus is based on these.
Ruskiniana belongs to the Habrolepidini, subtribe Habrolepidina (Encyrtinae). It is ex-
tremely close to Habrolepis and virtually can only be separated by the number of scale-like setae
at the apex of the scutellum (see key). Very probably they should be synonymised.
SAKENCYRTUS Hayat
(Key couplets: 76, 385. Figs 33, 195, 196, 422-426)
Sakencyrtus Hayat, 1981ft: 27. Type-species: Sakencyrtus mirus Hayat, by original designation.
DISTRIBUTION AND SPECIES. One species, India only: mirus Hayat (19816: 28), also at least two
undescribed species from India, Fiji (?) and Australia (BMNH, BPBM).
BIOLOGY. Unknown.
COMMENTS. The genus is close to Mira (see comments under Mira).
SAPRENCYRTUSgen. n.
(Key couplets: 234, 462. Figs 141, 221, 222, 427)
Type-species: Parasyrpophagus casuarinae Girault. Gender: masculine.
$ . Head. In frontal view a little wider than long, in side view about twice as long as broad and more or less
gradually and evenly anteriorly rounded. Eye more or less naked but with a few, sparse, inconspicuous
setae each no longer than the diameter of a facet; posterior margin of eye more or less straight, eye about
one-half longer than broad and more or less reaching occipital margin which is moderately acute. Malar
space about two-thirds length of an eye with sulcus absent but marked by a slight change of sculpture.
Frontovertex slightly more than one-third head width; ocelli more or less forming a right angle, the
posterior ones nearly touching eye margin but separated from occipital margin by clearly more than their
own major diameters. Antennal scrobes shallow, semicircular, meeting dorsally and reaching about
one-third way from antennal toruli to anterior ocellus; antennal torulus separated from mouth margin by
about is own length and from other torulus by about one-third more than its own length, its dorsal margin a
little below ventral level of eyes; clypeus shallowly emarginate. Antennal scape clearly longer than width of
frontovertex, slightly flattened, about five times as long as broad; pedicel conical, a little less than one-third
as long as the scape and clearly shorter than the first funicle segment; all funicle segments cylindrical,
longer than broad, clearly becoming broader and shorter distally so that the sixth is very nearly quadrate;
clava three-segmented, less than one-third length of funicle, with apex more or less rounded and sutures
more or less parallel; longitudinal sensillae on all flagellar segments except perhaps the first; longest setae a
little shorter than diameter of first funicle segment. Frontovertex with fine, shallow, raised, reticulate
sculpture, more longitudinally elongate between eyes and antennal scrobes, interantennal prominence
almost smooth; setae on frontovertex sparse, each not longer than diameter of anterior ocellus. Mandible
with three teeth; maxillary palpus four-segmented, labial palpus three-segmented.
Thorax. In side view moderately deep with both mesoscutum and scutellum only a little convex,
metapleurum and propodeum together broadly in contact with hind coxa. Pronotum in dorsal view with
posterior margin broadly concave; visible part of mesoscutum about one-third broader than long, notaular
lines absent, its posterior margin slightly convex but not projecting above axillae which meet medially;
scutellum fairly convex, nearly one-third longer than broad, apically quite pointed; propodeum medially
only a little less than one-fifth length of scutellum. Mesoscutum with shallow, raised, reticulate sculpture,
INDO-PACIFIC ENCYRTIDAE 337
scutellum similar but conspicuously shallower and smoother, propodeum medially with similar sculpture to
posterior part of scutellum but otherwise fairly smooth; mesopleurum with irregular, shallow, raised,
reticulate sculpture in anterior one-third, medially and posteriorly becoming smoother and alutaceous;
mesocutum and scutellum with a few sparse short setae; a few translucent setae on propodeum in front of
and to the outside of the spiracle. Forewing infuscate, about two and one-half to nearly three times as long
as broad; linea calva not interrupted or closed; basal cell moderately hairy but naked proximally; some
setae on proximal side of linea clava opposite filum spinosum flattened and scale-like as are some below
marginal vein; submarginal vein with parastigma hardly swollen and with a subapical hyaline break; costal
cell about 15 times as long as broad, with a few setae dorsally apically; marginal vein about five times as long
as broad, about two-thirds longer than stigmal and twice as long as postmarginal. Hindwing hyaline. Mid
tibial spur slightly shorter than mid basal tarsal segment.
Caster. About one-quarter longer than thorax (including propodeum), acute apically; hypopygium with
apex at about one-third along gaster; last tergite about three-quarters as long as gaster or a little longer than
mid tibia; ovipositor slightly exserted, the exserted part about one-tenth length of gaster and sheaths
slightly flattened from side to side.
Cf . Not available for description.
COMMENTS. This genus may possibly belong near Pseudencyrtus Ashmead (Microteryini,
subtribe Pseudencyrtina). This is suggested by the association with galls, the extremely long
gaster and the elongate last gastral tergite. However, the venation is more similar to that of
Syrphophagus. The genus can be distinguished from these and related genera principally by the
strongly infuscate forewing and presence of scale-like setae on the proximal side of the linea
calva opposite the filum spinosum.
Saprencyrtus casuarinae (Girault) comb. n.
(Figs 141, 221, 222, 427)
Parosyrpophagus casuarinae Girault, 19346: 3. LECTOTYPE $, AUSTRALIA (QM), here designated
[examined].
$. Length: 2-85-3-24 mm (lectotype, 2-85 mm).
Colour. Head dark metallic greenish blue, mesoscutum metallic green, scutellum metallic blue with
some green reflections, mesopleurum, propodeum and gaster purplish, base of gaster more shiny; antenna
with scape dark brownish and with a slight metallic green sheen , funicle segments dark brown , apex of fifth ,
whole of sixth and clava yellowish ; legs , including coxae , dark brown , femora and tibiae very slightly brassy
or metallic green; forewing infuscate as in Fig. 141.
Head. Mandible as in Fig. 221. Relative measurements (paralectotype): head length 114, head width
(frontal view) 123, head width (side view) 57, minimum frontovertex width 35, POL, 21, OOL 1-5, malar
space 49, eye length 71, eye width 51, scape length 72, other proportions of antenna as in Fig. 427. Girault,
in his description of the species, states that funicle segments five and six are one-half longer than wide, but
this conflicts with the intact specimen described here and the antenna figured. This may result from Girault
describing the antenna from a poorly mounted specimen on a slide.
Thorax. Relative measurements (paralectotype): forewing length 343, forewing width 129. Base of
forewing as in Fig. 222.
Gaster. Relative lengths (paralectotype): last tergite 176, [mid tibia 150].
O". Not available for description.
DISTRIBUTION. Australia.
BIOLOGY. Parasites or inquilines in galls of Cylindrococcus amplior Maskell (Homoptera,
Eriococcidae) on Casuarina stricta.
MATERIAL EXAMINED
Lectotype $, Australia: South Australia, Adelaide, from gall of Cylindrococcus amplior on Casuarina
strica,5.\ii. 1932(7. B. Cleland).
Australia: 2 9 (paralectotypes), South Australia, Adelaide, from gall of Cylindrococcus amplior on
Casuarina stricta, 5.vii.l932 (/. B. Cleland) (one lacking head). (The collector's name conflicts with that
given by Girault (19346: 3), i.e. A. L. Tonnoir.}
338 J. S. NO YES & M. HAY AT
COMMENTS. Although the genus and species is described here from two female syntypes, all the
syntypes were examined during a visit to the Queensland Museum, Brisbane. In his unpublished
manuscript (QM), Girault states that the species was described from one male and four females.
A specimen on loan from ANIC, Canberra in QM, Brisbane is here designated lectotype and
labelled as such by one of us (JSN).
SCHILLERIELLA Ghesquiere
(Key couplet: 76)
Schilleria Girault, 1932a: 1. Type-species: Schilleria pulchra Girault, by monotypy. [Homonym of
Schilleria Dahl, 1907.]
Schilleriella Ghesquiere, 1946: 369. [Replacement name for Schilleria Girault.]
DISTRIBUTION AND SPECIES. One species, Australia only: pulchra (Girault, 1932a: 1).
BIOLOGY. Unknown.
COMMENTS. Schilleriella appears to be related to Anusia Forster (tribe Anagyrini, subtribe
Anusiina).
SPANIOPTERUS Gahan
(Key couplet: 53. Figs 19, 20)
Spaniopterus Gahan, 19276: 149. Type-species: Spaniopterus crucifer Gahan, by original designation.
DISTRIBUTION AND SPECIES. One species, Java and Malaysia only: crucifer Gahan (1921 b: 150).
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. Placed in the tribe Habrolepidini, subtribe Comperiellina (Encyrtinae). It can be
distinguished from Comperiella by having a four-segmented funicle (Comperiella has a six-
segmented funicle).
STENOTEROPSIS Girault
(Key couplet: 530)
Stenoteropsis Girault, 1915a: 176. Type-species: Stenoteropsis abjectus Girault, by original designation.
DISTRIBUTION AND SPECIES. One species, Australia only: abjectus Girault (1915a: 176).
BIOLOGY. Unknown.
COMMENTS. Probably related to Helegonatopus (Chalcerinyini), from which it can be separated
by having the ovipositor slightly exserted and the sheaths a little swollen apically. The two
genera may be synonymous, but we are retaining Stenoteropsis as valid until fresh material can
be compared with the holotype of abjectus, which is in poor condition.
SYRPHOPHAGUS Ashmead
(Key couplets: 289, 301, 341, 343, 349, 398, 455, 526. Fig. 186)
Syrphophagus Ashmead, 19006: 397. Type-species: Encyrtus mesograptae Ashmead, by original desig-
nation.
Aphidencyrtus Ashmead, 19006: 398. Type-species: Encyrtus aphidiphagus Ashmead, by original desig-
nation. Syn. n.
Echthrobaccha Perkins, 1906: 253. Type-species: Echthrobaccha injuriosa Perkins, by monotypy.
Nesyrpophagus Girault, 1915a: 113. Type-species: Nesyrpophagus flavithorax Girault, by original desig-
nation. Syn. n.
Hexanusia Girault, 19220: 39. Type-species: Hexanusia nigricornis Girault, by monotypy. Syn. n.
Syrphidencyrtus Blanchard, 1940: 107. Type-species: Syrphidencyrtus bacchae Blanchard, by monotypy.
INDOPACIFIC ENCYRTIDAE 339
DISTRIBUTION AND SPECIES. About 60 species, cosmopolitan; 22 from review area: aeruginosus
(Dalman, 1820: 170) (India, Hawaiian Is.), aphidivorus (Mayr, 1876: 712, 713, 714) (comb. n.
from Encyrtus) (India, Hawaiian Is.), aquacyaneus (Girault, 19230: 50) (comb. n. from Cocci-
doxenus) (Australia), cinctipes (Girault, 19150: 110) (comb. n. from Neasteropaeus) (Austra-
lia), feralis (Girault, 19296: 315) (comb. n. from Paraenasomyia) (Australia), flavithorax
(Girault, 19150: 113) (comb. n. from Nesyrpophagus) (= Nesyrpophagus unguttatus Girault,
19150: 113 syn. n., = Hexanusia sanguinithorax Girault, 19276: 310 syn. n.) (Australia), hakki
Agarwal (19620: 248) (India), hofferi (Hayat, 1973: 35) (comb. n. from Aphidencyrtus) (India),
indicus Agarwal (19620: 246) (India), injuriosus (Perkins, 1906: 254) (Australia), kumaoensis
(Bhatnagar, 1952: 163) (comb. n. from Coccidencyrtus) (India), luciani (Girault, 19220: 42)
(comb. n. from Echthrobacchd) (Australia), merceti (Masi, 1926: 268) (comb. n. from Micro-
terys) (Taiwan), metallicus (Girault, 19140: 33) (comb. n. iiomAratus) (Australia), nigricornis
(Girault, 19220: 39) (comb. n. from Hexanusia) (Australia), obscurus (Girault, 1923c: 143)
(comb. n. from Neasteropaeus) (Australia), occidentalis (Girault, 1917e: 95) (comb. n. from
Cerchysius) ( Australia), parvus (Girault, 19230: 47) (comb. n. from Cerchysiopsis) (Australia),
perdubius (Girault, 1926c: 132) (comb. n. from Coccidoxenus) , puparia (Girault, 19296: 313)
(comb. n. from Epiblatticidd) (Australia), raffaellini (Girault, 1922d: 208) (comb. n. from
Habrolepoidea) (Australia) and varicornis (Girault, 1923c: 143) (comb. n. from Neasteropaeus)
(Australia), also much undetermined material from throughout the region (BMNH, BPBM,
QM, ANIC, USNM, HC, GC).
BIOLOGY. Parasites of Aphididae (primary or secondary), Psyllidae (Homoptera) and larvae of
Diptera, mostly of Syrphidae predatory on aphids.
COMMENTS. The apparent difference in biologies of Syrphophagus and Aphidencyrtus have been
virtually the only reason for regarding both genera as valid, the two being difficult to separate
reliably on morphology alone (see Trjapitzin, 1972). One Australian species, nigricornis
(Girault), would be considered a typical species of Syrphophagus by most encyrtid taxonomists
since morphologically it is very close to aeruginosus (Dalman). However, this species is regularly
reared from aphids! With this in mind, the ecological closeness of their hosts and the difficulty in
assigning many species to either genus without knowledge of their biologies, we here regard the
two as synonymous. Consequently, in addition to the above, we also propose the following new
combinations for extra-limital species known to us: africanus Gahan (from Aphidencyrtus),
cassatus Annecke (from Aphidencyrtus), inquisitor Howard (from Encyrtus), mamitus Walker
(from Encyrtus), quercicola Hoffer (from Aphidencyrtus), similis Prinsloo (from Aphidencyr-
tus), tachikawai Hoffer (from Aphidencyrtus) and taeniatus Forster (from Encyrtus) (all
Syrphophagus, comb. n.).
The holotype of Microterys merceti Masi has been examined (IPK). It belongs to Syrpho-
phagus.
We have not seen the holotype of Coccidencyrtus kumaoensis Bhatnagar, but from the
description it must be a Syrphophagus.
The genus is placed in the Microteryini, subtribe Syrphophagina (Encyrtinae) (see comments
under Coccidoctonus) .
SZELENYIOLA Trjapitzin
(Key couplet: 182)
Szelenyiola Trjapitzin, 1977: 160. Type-species: Szelenyiola nearctica Trjapitzin, by original designation.
DISTRIBUTION AND SPECIES. Two described species, New World and Australia: prospheris
(Ferriere, 1947: 629) (comb. n. from Ooencyrtus) (Australia).
BIOLOGY. Parasites of eggs of Buprestidae and Scolytidae (Coleoptera).
COMMENTS. Ferriere (1947) described the clava of prospheris as being three-segmented, but
examination of slide-mounted material shows that it is entire.
340 J. S. NOYES & M. HAY AT
Placed in the tribe Microteryini, subtribe Oobiina (Encyrtinae). Trjapitzin (1977) provides a
key to separate the genera of this subtribe.
TACHARDIAEPHAGUS Ashmead
(Key couplets: 308, 392, 409. Figs 184, 185)
Tachardiaephagus Ashmead, 1904c: 503. Type-species: Tachardiaephagus thoracicus Ashmead, by orig-
inal designation.
Lissencyrtus Cameron, 1913: 99. Type-species: Lissencyrtus troupi Cameron, by monotypy.
DISTRIBUTION AND SPECIES. Four species, Afrotropical, Oriental, Australasian; one from review
area: tachardiae (Howard in Howard & Ashmead, 1896: 637) (India, Sri Lanka, Malaysia,
Brunei), also undetermined material from Taiwan and the Philippines (BPBM).
REFERENCE. Prinsloo (1977: 57-69).
BIOLOGY. Parasites of Keriidae (Homoptera).
COMMENTS. Placed in the tribe Microteryini, subtribe Microteryina (Encyrtinae), it can be easily
distinguished from other genera found in review area by the structure and shape of the antennal
scrobes (Figs 184, 185).
TACHINAEPHAGUS Ashmead
(Key couplets: 236, 306, 316, 365, 399, 450. Figs 143, 144, 236, 428, 430)
Tachinaephagus Ashmead, 1904c: 304. Type-species: Tachinaephagus zealandicus Ashmead, by original
designation.
Phaenodiscoides Girault, 1915a: 82. Type-species: Phaenodiscoides australiensis Girault, by original
designation. Syn. n..
Tachinaephagus Girault, 1917g: 142. Type-species: Tachinaephagus australiensis Girault, by original
designation.
Australencyrtus Johnson & Tiegs, 1921: 118. Types-species Australencyrtus giraulti Johnson & Tiegs, by
original designation.
Australomalotylus Risbec, 1956: 170. Type-species: Australomalotylus rageaui Risbec, by monotypy.
DISTRIBUTION AND SPECIES. Ten species, Afrotropical, east Palaearctic, Oriental and Austral-
asian; seven from review area: australiensis (Girault, 19146: 59) (comb. n. from Phaenodiscus)
(= Phaenodiscoides australiensis Girault, 19150: 82 syn. n.) (Australia), ceylonicus (Subba Rao,
1972: 191) (Sri Lanka), javpnsis Subba Rao (1978: 71) (Indonesia), lutheri (Girault, 1924a: 6)
(comb. n. from Phaenod$coides) (Australia), lyperosae (Ferriere, 1933: 638) (comb. n. from
Cerchysius) (Java), maldyensis Subba Rao (1978: 72) (Malaysia) and zealandicus Ashmead
(1904c: 304) (Australia, New Caledonia, New Zealand), also many undescribed species
amongst material from India and S. China to Australia and Fiji (BMNH, BPBM, USNM,
CNC).
REFERENCE. Revision: Subba Rao (1978).
BIOLOGY. Parasites of larvae of Calliphoridae, Muscidae, Sarcophagidae and Tephritidae
(Diptera).
COMMENTS. Girault inadvertently described the same specimen twice as australiensis, once under
Phaenodiscus and once under Phaenodiscoides. This is evident from a comparison of the
original descriptions. We do not consider Phaenodiscoides as a valid genus since australiensis is
fairly typical of Tachinaephagus except that the antennae are a little longer than in most species
included in this genus.
We have examined a paratype of Cerchysius lyperosae Ferriere (BMNH); it is a species of
Tachinaephagus with a well-exserted ovipositor.
The genus is related to Rhytidothorax , Parastenoterys (see comments under these genera) and
INDO-PACIFIC ENCYRTIDAE 341
Nerissa Trjapitzin (1977: 165). The last is very close and may eventually be considered
synonymous with Tachinaephagus , differing only very slightly in the venation of the forewing.
TAFTIA Ashmead
(Key couplets: 402, 427. Figs 211, 212)
Taftia Ashmead, I904d: 137. Type-species: Taftia prodeniae Ashmead, by original designation.
DISTRIBUTION AND SPECIES. Two species, Philippines and Java only: prodeniae Ashmead (1904d:
137) (Philippines) and saissetiae Gahan (1920: 344) (Philippines, Java), also one further species
from Malaysia (BPBM).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the Chrysoplatycerini, subtribe Taftiina (Tetracneminae) which also
includes Lutherisca from which it can be separated using the characters given in the key. In all
probability Taftiina should be considered synonymous with Chrysoplatycerina.
TASSONIA Girault
(Key couplet: 515. Fig. 252)
Tassonia Girault, 1921a: 2. Type-species: Tassonia gloriae Girault, by monotypy.
DISTRIBUTION AND SPECIES. Two species, Oriental and Australasian only: gloriae Girault
(19210): 2) (= Neblatticida tassoniaeformis Girault, 19210: syn. n.) (Australia) and magni-
clava (Hayat & Subba Rao, 1981: 108) (comb. n. from Aphidencyrtus) (India), also further
material containing several undescribed species from India, Hong Kong, Malaysia and Java
(BMNH,BPBM,UCR).
BIOLOGY. Parasites of Aphididae (Homoptera).
COMMENTS. The genus is related to Syrphophagus (Microteryini, subtribe Syrphophagina). It
differs in several characters, notably in its generally smaller size, more convex thoracic dorsum,
shorter clavate antenna, thicker and subequal marginal, postmarginal and stigmal veins of the
forewing and the presence of a naked streak joining the apex of the postmarginal vein to the
stigmal (Fig. 252).
TELETEREBRATUS Compere & Zinna
(Key couplets: 199, 369. Figs 431, 432)
Teleterebratus Compere & Zinna, 1955: 108. Type-species: Teleterebratus perversus Compere & Zinna, by
original designation.
DISTRIBUTION AND SPECIES. Three species, Oriental and Australasian only: amplis (Girault,
1915a: 81) (comb. n. from Aenasielld) (Australia), daripennis (Girault, 19150: 101) (comb. n.
from Rhopalencyrtoidea) (Australia) and perversus Compere & Zinna (1955: 110) (China).
BIOLOGY. Parasites of Diaspididae and gall-forming Eriococcidae (Homoptera).
COMMENTS. We have not seen the types ofAgeniaspis indicus Narayanan, but from the very poor
description the species possibly belongs in Teleterebratus.
The genus appears to be related to Coccidoctonus (see comments under Coccidoctonus) .
TETRACNEMOIDEA Howard
(Key couplet: 53. Figs 21, 22)
Tetracnemoidea Howard, 18986: 232. Type-species: Tetracnemoidea australiensis Howard, by monotypy.
Tetracnemopsis Ashmead, 1900a: 358. Type-species: Tetracnemus westwoodii Cockerell, by original
designation.
342 J. S. NO YES & M. HAY AT
Ectromella Girault, 1915a: 142. Type-species: Ectromella bicolor Girault, by original designation. Syn. n.
Arhopoideus Girault, 1915a: 174. Type-species: Arhopoideus brevicornis Girault, by original designation.
Hungariella Erdos, 1946ft: 144. Type-species: Hungariella piceae Erdos, by original designation.
Antipodencyrtus Kerrich, 1964ft: 505. Type-species: Antipodencyrtus procellosus Kerrich, by monotypy.
Syn. n.
DISTRIBUTION AND SPECIES. Sixteen species, cosmopolitan; nine from review area: australiensis
Howard (18986: 232) (Australia), bicolor (Girault, 19150: 142) (comb. n. from Ectromella) (=
Arhopoideus tertius Girault, 1923c: 144 syn. n.) (Australia), brevicornis (Girault; Tachikawa,
1974: 23) (Australia), brouni (Timberlake, 1929: 6) (New Zealand), indica (Ayyar, 1932: 287)
(India), ipswichia (Girault, 1922/: 1) (Australia), procellosa (Kerrich, 19646: 505) (comb. n.
from Antipodencyrtus) (New Zealand) and secunda (Girault, 19150: 175) (Australia), also
undetermined material, containing several undescribed species from Papua New Guinea,
Tonga, Australia and New Zealand (BMNH, BPBM, DSIR, QM, ANIC).
REFERENCE. World review: Trjapitzin & Gordh (19800).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Recent collecting in New Zealand has revealed a wealth of material belonging to this
genus, including winged forms of procellosus or a very similar species. We do not think that the
slightly flattened body, or the difference in the number of branches in the antenna of the male,
are sufficient reasons for regarding Antipodencyrtus as distinct from Tetracnemoidea.
Placed in the tribe Tetracneminae, subtribe Arhopoideina (Tetracneminae).
TETRACNEMUS Westwood
(Key couplet: 361)
Tetracnemus Westwood, 1837: 258. Type-species: Tetracnemus diversicornis Westwood, by monotypy.
Tetracladia Howard, 1892: 367. Type-species: Tetracladia texana Howard, by designation of Ashmead
(1900: 358).
Tetralophidea Ashmead, 1900ft: 348. Type-species: Tetralophidea bakeri Ashmead, by original desig-
nation.
Tetralophiellus Ashmead, 1900ft: 357. Type-species: Tetralophiellus brevicollis Ashmead, by original
designation.
Paracalocerinus Girault, 1915a: 142. Type-species: Paracalocerinus australiensis Girault, by original
designation.
Masia Mercet, 1919ft: 470. Type-species: Masia bifasciatella Mercet, by original designation.
Anusiella Mercet, 1923a: 287. Type-species: Anusia heydeni Mayr, by original designation.
Placocerus Erdos, 19460: 1. Type-species: Placocerus calocense Erdos, by monotypy.
Comperencyrtus De Santis, 1964: 106. Type-species: Comperencyrtus maculipennis De Santis, by original
designation.
DISTRIBUTION and species. Twenty-one species, cosmopolitan; five from review area: australien-
sis (Girault, 19150: 142) (Australia), deccanensis (Mani & Kaul in Mani etal. , 1974: 65) (India),
diversicornis Westwood (= Masia pulchripennis Mercet, 19230: 289) (India), heterocornis Mani
& Saraswat in Mani et 0/., 1974: 75) (India) and peninsularis (Mani & Saraswat in Mani et al.,
1974: 73), also several undetermined species from India and Australia (BMNH, ANIC, QM,
HC).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. Placed in the Tetracnemini, subtribe Tetracnemina (Tetracneminae). The genus
can be easily recognised by the well-exserted ovipositor, darkened forewings with a relatively
long marginal and short postmarginal and stigmal veins, and the very flattened antennal
flagellum.
INDO-PACIFIC ENCYRTIDAE 343
THOMSONISCA Ghesquiere
(Key couplet: 191. Figs 114, 429)
Thomsoniella Mercet, 1921: 89. Type-species: Thomsoniella typica Mercet, by original designation.
[Homonym of Thomsoniella Signoret, 1880.]
Thomsonisca Ghesquiere, 1946: 369. [Replacement name for Thomsoniella Mercet.]
Heterencyrtus Hoffer, 1953: 86. Type-species: Heterencyrtus sumavicus Hoffer, by original designation.
Athesmus Erdos & Nowicky, 1955: 198. Type-species: Athesmus luctuosus Erdos & Nowicky, by original
designation.
Kosztarabia Erdos, 1957ft: 367. Type-species: Kosztarabia chionaspidis Erdos, by original designation.
Euussuria Chumakova, 1957: 539. Type-species: Euussuria pallipes Chumakova, by original designation.
Pakencyrtus Ahmad, 1970: 237. Type-species: Pakencyrtus pakistanensis Ahmad, by original designation.
DISTRIBUTION AND SPECIES. Six species, Palaearctic, Oriental; four from review area: amathus
(Walker; = Thomsoniella typica Mercet, 1921: 90), indica Hayat (1970b: 55) (India) , pakis-
tanensis (Ahmad, 1970: 238) (India, Pakistan) andsankarani Subba Rao (1979: 142) (India).
REFERENCE. Review: Subba Rao (1979: 139-144).
BIOLOGY. Parasites of Diaspididae (Homoptera).
COMMENTS. Placed in the tribe Thomsoniscini (Encyrtinae).
TINEOPHOCTONUS Ashmead
(Key couplet: 357)
Tineophoctonus Ashmead, 1900ft: 351. Type-species: Phaenodiscus armatus Ashmead, by original des-
ignation.
DISTRIBUTION AND SPECIES. Three species, New World, Europe, and one undescribed species
from Papua New Guinea (BPBM).
BIOLOGY. Parasites of gall-inhabiting Tineidae (Lepidoptera) , Cynipidae (Hymenoptera) and
larvae of Anobiidae and Cerambycidae (Coleoptera).
COMMENTS. The species from Papua New Guinea may be incorrectly placed in Tineophoctonus
since the antennal clava is obliquely truncate and the funicle segments are relatively shorter than
in the described species. However, in other characters it seems to comply with those of
Tineophoctonus .
Placed in the tribe Cheiloneurini (Encyrtinae) and closest to Cheiloneurus and Prochilo-
neurus. It can be separated from Cheiloneurus by the hypopygium reaching the apex of the gaster
and from Prochiloneurus by the gaster being apically acute.
TONG YUS gen. n.
(Key couplet: 150. Figs 81, 433-439)
Type-species: Tongyus nesus sp. n. Gender: masculine.
$. Head. In facial view slightly broader than long, in profile slightly less than twice as long as broad and
anteriorly gradually and more or less evenly curved except along length of antennal scrobes where it is
almost straight. Eye with posterior margin a little concave, almost straight, about one-third longer than
broad, covered with fairly dense long hairs, each hair about one and one-half times to twice as long as
diameter of a facet, eye reaching occipital margin which is sharp. Malar space about one-third eye-length,
sulcus present. Frontovertex about one-third head width; ocelli nearly forming an equilateral triangle, the
posterior ones a little nearer eye margin than occipital margin and separated from the latter by about their
own major diameters. Antennal scrobes moderately deep and meeting dorsally or separated by interanten-
nal prominence which is confluent with frontovertex, reaching about half way from antennal toruli to
anterior ocellus; antennal torulus separated from mouth margin by not more than two-thirds its own length
and from other torulus by slightly more than about one-half its own length, its dorsal margin about level
344 J. S. NO YES & M. HAY AT
with or a little above the lowest eye margin; clypeal margin shallowly excised. Antennal scape broadened
and flattened, a little more than twice as long as broad and clearly longer than minimum width of
frontovertex; pedicel conical, slightly longer than any funicle segment except perhaps the first; all funicle
segments longer than broad, the sixth only slightly so; clava three-segmented, apically rounded, but with
outer suture slightly oblique and converging with inner; flagellar segments slightly flattened, subyclindric-
al; longitudinal sensillae on all flagellar segments, longest setae clearly shorter than diameter of segments.
Frontovertex with very fine, raised, moderately deep, squamiform-reticulate sculpture (Fig. 436), with
scattered inconspicuous, translucent setae; eye margins with fairly conspicuous dark setae. Mandible
narrow with two acute apical teeth; maxillary palpus four-segmented, labial palpus three-segmented.
Thorax. In side view moderately deep, with mesoscutum and scutellum very slightly convex, the
metapleurum and propodeum together narrowly in contact with hind coxa. In dorsal view pronotum with
hind margin slightly concave; visible part of mesoscutum about twice as broad as long, notaular lines absent
posterior margin very clearly convex and produced backwards above axillae; axillae meeting medially;
scutellum about as long as mesoscutum, about as broad as long, with apex more or less pointed, sides
straight; propodeum medially short, not more than about one-ninth as long as scutellum. Mesoscutum with
sculpture similar to but clearly shallower than that on frontovertex; scutellum with same sculpture as
frontovertex; propodeum medially with very shallow, irregular, rugose sculpture, outside spiracles much
deeper and irregular, mesopleurum with shallow, very fine, raised, regular, reticulate sculpture; setae on
dorsum of thorax fairly dense, translucent or brown, quite conspicuous, particularly on scutellum.
Forewing at least partially infuscate, about two and one-half times as long as broad; linea calva closed in
posterior one-third; filum spinosum absent; submarginal vein with an apical hyaline break, parastigma not
swollen; costal cell about 14 times as long as broad, with a single line of setae dorsally in its apical one-third;
marginal vein about three or four times as long as broad, clearly shorter than stigmal which is as long as or a
little longer than postmarginal. Hindwing about two-thirds length of forewing, about four times as long as
broad, marginal fringe about one-ninth as long as maximum wing width. Mid tibial spur a little shorter than
basal mid tarsal segment.
Caster. About as long as thorax; cereal plates in basal one-half; hypopygium reaching apex of gaster;
paratergites present; last tergite slightly shorter than mid tibia; ovipositor hardly exserted, about
three-quarters as long as mid tibia; gonostyli fused to second valvifers, about one-eighth as long as
ovipositor.
Cf. Similar to female except body generally darker, antenna and genitalia. Differs as follows. Head
proportionately a little broader in frontal view; malar space about one-half length of eye; frontovertex
nearly half head width; ocelli nearly forming a right angle, the posterior ones almost equidistant from
occipital margin and eye, although a little closer to the former; antennal toruli separated from mouth
margin by much more than their own lengths, their lowest margins a little below lowest eye margins;
antennal scape shorter than minimum width of frontovertex, stout and slightly broadened and flattened, a
little less than three times as long as broad; pedicel conical, subquadrate, not more than half as long as any
funicle segment all of which are cylindrical and beset with long setae, the longest at least about four times as
long as diameter of segments; clava entire and gradually tapering to a point; longitudinal sensillae on all
flagellar segments; scale like sensillae on clava only. Forewing a little broader than in female; linea calva
interrupted and closed. Genitalia with aedeagus slightly less than half as long as mid tibia, digiti (excluding
apical spines) about one-fifth as long as aedeagus, each with a pair of long apical spines.
COMMENTS. Tongyus belongs in the Anagyrini, subtribe Anagyrina (Tetracneminae) and
appears to be most closely related to Anagyrus. In the female it can be separated from this and
other genera of this group by the combination of the slightly flattened flagellar segments,
converging sutures of the clava, sculpture of the head and dorsum of thorax, infuscation of
forewings and wing venation.
Tongyus nesus sp. n.
(Figs 81, 433-439)
$. Length: 1-11-1-75 mm (holotype, 1-60 mm).
Colour. Head and thorax brownish yellow, with scutellum, metanotum and propodeum largely dark
brown, gaster dark brown; antenna with scape more or less white but margined dark brown ventrally and
dorsally (Fig. 433), pedicel and flagellum dark brown; legs brownish yellow but mixed with dark brown,
especially apex of mid tibia and all of hind femur and tibia, hind coxa dark brown; forewing infuscate from
INDO-PACIFIC ENCYRTIDAE 345
base to about one-quarter along wing, an indistinct and incomplete fuscous fascia across wing from
marginal and stigmal veins (Fig. 81), remainder of forewing and hindwing hyaline.
Head. Antennal scrobes meeting dorsally, interantennal prominence at its upper level clothed in
numerous, fairly dense, white setae which continue down either side of prominence to mouth margin.
Relative measurements (holotype): head length 35, head width (facial view) 40, head width (side view) 18,
minimum frontovertex width 14, malar space 8, eye length 25, eye width 19, POL 5-5, OOL 2-5, scape
length 22, scape width 10, proportions of antenna as in Fig. 433.
Thorax. Mesoscutum anteriorly with dark setae, posteriorly, and axillae, with translucent or pale setae,
usually one or two dark setae scattered amongst pale setae, scutellum with dark setae. Relative
measurements (holotype): forewing length 105, width 40, proportions of veins as in Fig. 81; hindwing
length 69, width 18. Basal cell of forewing with setae evenly distributed and as dense as in disc distal to
venation.
Caster. Relative lengths (paratype): last tergite 47, ovipositor 40, [mid tibia 55]; genitalia as in Fig. 434,
hypopygium as in Fig. 435.
Cf . Length: 0-96-1-30 mm. Similar to female except following. Body completely dark brown except for
interantennal prominence and lower part of face below and to outside of antennal toruli, which are
brownish yellow; outer part of scape at base brownish yellow, remainder of antenna dark brown; prepectus
whitish; legs dark brown except fore femur and tibia, base of mid tibia and all tarsi which are testaceous
yellow, four apical tarsal segments of mid leg mixed dark brown, occasionally mid leg pale as in foreleg but
apex of mid tibia always dark brown. Antenna as in Fig. 437, forewing with linea calva interrupted by two
lines of setae and closed by a single line on dorsal surface, basal cell with proximal one-third or so naked;
genitalia as in Figs 438, 439. Relative measurements (paratype 1): head width 45, minimum frontovertex
width 22, scape length 18, forewing length 109, forewing width 48, hindwing length 73, hindwing width 21 ,
aedeagus length 21, mid tibia length 52. Relative measurements (paratype 2): scape length 35-5, maximum
scape width 14, POL 17, OOL 8. (Paratype 1 on slide; paratype 2 dry-mounted on card.)
DISTRIBUTION. Cook Is.
BIOLOGY. Unknown.
MATERIAL EXAMINED
Holotype $, Cook Is.: Raratonga, Totokuitu, xi.1978 (E. W. Valentine) (DSIR).
Paratypes. Cook Is.: 9 $, 13 cf , same data as holotype (DSIR, BMNH, USNM, PPRI, ZI).
COMMENTS. A second species from the Society Is. (BPBM) differs from nesus in the arrangement
of the scrobes and setae on the interantennal prominence, relative proportions of antennal
segments, pattern of infuscation of forewing and venation.
TRECHNITES Thomson
(Key couplet: 72. Fig. 31)
Trechnites Thomson, 1876: 118. Type-species: Trechnites fuscitarsis Thomson, by monotypy.
Psylledontus Crawford, 1910: 88. Type-species: Psylledontus insidiosus Crawford, by original designation.
Metallonella Girault, 1915a: 77. Type-species: Metallonella australiensis Girault, by original designation.
DISTRIBUTION AND SPECIES. Sixteen species, Holarctic, Afrotropical, Oriental, Australasian; five
species from review area: aligarhensis Hayat, Alam & Agarwal (1975: 90) (India), australiensis
(Girault, 1915a: 77) (Australia), manaliensis Hayat, Alam & Agarwal (1975: 88) (India),
secundus (Girault, 1915e: 281) (Sri Lanka) and viridiscutellum (Girault, 19150: 132) (comb. n.
from Encyrtomyia), also material from Nepal, Vietnam, Hong Kong, Borneo, New Caledonia
and Solomon Is. (BMNH, BPBM).
REFERENCES. Hayat etal. (1975: 87-92); Prinsloo (1981: 236).
BIOLOGY. Parasites of nymphs of Psyllidae (Homoptera).
COMMENTS. Placed in the tribe Trechnitini, subtribe Trechnitina (Encyrtinae). It is very close to
Coccidaphycus from which it can be separated by the characters given in the key.
346 J. S. NOYES & M. HAY AT
TRICHOMASTHUS Thomson
(Key couplets: 237, 341, 438, 469. Figs 138, 153, 220)
Trichomasthus Thomson, 1876: 142. Type-species: Encyrtus cyaneus Dalman, by subsequent designation
of Gahan & Pagan (1923: 148).
Coccidoxenus Crawford, 1913: 248. Type-species: Coccidoxenus portoricensis Crawford, by original
designation.
DISTRIBUTION AND SPECIES. About 50 species, cosmopolitan; only one species from review area:
mexicanus (Girault, 1917c: 21) (Hawaiian Is.), also several undetermined species from India,
S. China, Hong Kong, Borneo and Australia (BMNH, BPBM, ANIC).
BIOLOGY. Parasites of Coccidae, Diaspididae, Eriococcidae and Pseudococcidae (Homoptera).
COMMENTS. Both Tetracnemella and Stenoteropsis have been incorrectly synonymised with
Trichomasthus. Tetracnemella is here treated as a synonym of Ooencyrtus and Stenoteropsis as a
valid genus near Helegonatopus .
The genus is placed in the Microteryini, subtribe Microteryina (Encyrtinae). However, it
must be very much closer to Ooencyrtus (subtribe Ooencyrtina) than this infers since the two
genera occasionally can be difficult to separate.
TRJAPITZINELLUS Viggiani
(Key couplets: 326, 508)
Trjapitzinellus Viggiani, 1967: 166. Type-species: Trjapitzinellus semidaliphagus Viggiani, by original
designation.
DISTRIBUTION AND SPECIES. Six species, Holarctic, Oriental; possibly two undetermined species
from India (BMNH, HC).
REFERENCE. Key to Palaearctic species: Myartseva (1980).
BIOLOGY. Parasites of immature stages of Coniopterygidae (Neuroptera).
COMMENTS. Placed in the Bothriothoracini, subtribe Coenocercina (Encyrtinae).
TROPIDOPHR YNE Compere
(Key couplet: 154)
Tropidophryne Compere, 1931: 269. Type-species: Tropidophryne africana Compere, by monotypy.
DISTRIBUTION AND SPECIES. Five species, Afrotropical; one undescribed species from New
Britain (BPBM).
REFERENCES. Review: Prinsloo & Annecke (19786: 312-315); Kerrich (1978: 145-150).
BIOLOGY. Parasites of Pseudococcidae (Homoptera).
COMMENTS. We have not seen the female specimen recorded by Baker (1978: 56, Fig. 3), under
the name Zaplatycerus sp., but from his figure it is almost certainly a species of Tropidophryne.
The host given by Baker is also probably incorrect (Doleschalla sp.; Diptera, Tachinidae).
The genus belongs in the tribe Chrysoplatycerini, subtribe Chrysoplatycerina (Tetracnemi-
nae). A key to related genera is given by Kerrich (1978: 113-114).
TYNDARICHUS Howard
(Key couplets: 184, 415. Fig. 107)
Tyndarichus Howard, 1910: 5. Type-species: Tyndarichus navae Howard, by original designation.
DISTRIBUTION AND SPECIES. Seven species, Nearctic, Palaearctic, Afrotropical, Oriental,
IN DO-PACIFIC ENCYRTIDAE 347
Australasian; two species from review area: melanicis (Dalman, 1820: 345) (India) and parti-
cornis (Girault, 19240: 8) (comb. n. from Epiblattidda) (Australia), also undetermined material
from India, Sri Lanka, Nepal, Hong Kong, Java, Sulawesi, New Britain and Australia (BMNH,
BPBM).
BIOLOGY. Hyperparasites of larvae of Lepidoptera through other Encyrtidae (Hymenoptera).
COMMENTS. Placed in the tribe Cheiloneurini, subtribe Epiencyrtina (Encyrtinae) by Trjapitzin
& Gordh (1978&). It is very close to Parechthrodryinus , from which it can be very difficult to
separate if the biology is not known (see characters given in key). This subtribe may be out of
place in the Cheiloneurini and it is possible that its included genera are more closely related to
those placed in the subtribe Syrphophagina (tribe Microteryini) since there is some similarity in
forewing venation and general morphology, particularly the structure of the thorax.
TYNDARICOPSIS Gordh & Trjapitzin
(Key couplets: 184, 416. Figs 108, 109)
Tyndaricopsis Gordh & Trjapitzin, 1981: 48. Type-species: Tyndarichus davatus Eady, by original
designation.
DISTRIBUTION AND SPECIES. One species, New Guinea only: davatus (Eady, 1960: 669).
BIOLOGY. Hyperparasites of larvae of Pyralidae (Lepidoptera) via other Encyrtidae (Hymen-
optera).
COMMENTS. Closely related to Tyndarichus which has been placed in the Cheiloneurini, subtribe
Epiencyrtina (Encyrtinae) by Trjapitzin & Gordh (1978b) (see comments under Tyndarichus).
It can be separated from Tyndarichus and Parechthrodryinus by the characters given in the key.
WHITTIERIA Girault
Whittieria Girault, 1938: 82. Type-species: Whittieria pilosigena Girault, by original designation.
DISTRIBUTION AND SPECIES. One species, Australia only: pilosigena Girault (1938: 82).
BIOLOGY. Unknown.
COMMENTS. The genus must be related to Tachardiaephagus, Bennettisca Noyes, Aloencyrtus
Prinsloo and Allencyrtus Annecke & Mynhardt (Microteryini, subtribe Microteryina). Girault
mentions that the scrobes are 'deep gouges' (a typical character of this group). Also the dense
setation of the forewing, venation and structure of the mandible indicate a relationship with the
genera of this group. The true systematic position of the genus will not be known until fresh
material, including females, is studied.
XENANUSIA Girault
(Key couplets: 75, 107)
Xenanusia Girault, 1917g: 137. Type-species: Xenanusia pulchripennis Girault, by original designation.
DISTRIBUTION AND SPECIES. Two species, Australia only: flava (Girault, 1915a: 153) (comb. n.
from Anusia) and pulchripennis Girault (1917g: 138).
BIOLOGY. Unknown.
COMMENTS. Xenanusia flava may be out of place here and may require a new genus to
accommodate it. However, we feel that it is correctly placed within the group of genera to which
Xenanusia belongs.
The type-species of Xenanusia is remarkable in that superficially it closely resembles species
of Cerapterocerus or Cerapteroceroides . However, it belongs to the same group of genera as
Cryptanusia, Cyrtocoryphes and Parectromoidella (see comments under Cryptanusia).
348 J. S. NO YES & M. HAY AT
XENOENCYRTUS Riek
(Key couplets: 91, 249. Figs 44, 45)
Xenoencyrtus Riek, 1962a: 151. Type-species: Xenoencyrtus niger Riek, by original designation.
DISTRIBUTION AND SPECIES. Four species, Australia only: hemipterus (Girault, 1915a: 172),
hemipterus pentlandensis (Girault, 1915a: 173), hemipterus stigmatiferus (Girault, 1923c: 147),
megymeni (Dodd, 1917: 354), megymeni brachypterus (Dodd, 1917: 355), niger Riek (19620:
152) and rubricates Riek (1962a: 154), also much undetermined material from Australia
(BMNH,ANIC,QM).
REFERENCE. Revision: Riek (1962a).
BIOLOGY. Parasites of eggs of Pentatomidae (Heteroptera).
COMMENTS. The types of hemipterus pentlandensis and hemipterus stigmatiferus cannot be
located. (Girault actually described Ericydnus stigmatifera hemiptera (1923c: 147), but we feel
that somehow the specific and subspecific names must have become juxtaposed either by a
lapsus on Girault's part or by a type-setting error.)
It is probable that all the species included by Riek in this genus are all forms of the same
species.
The genus is close to Ooencyrtus, Ovaloencyrtus and Paratetralophidea (Microteryini,
subtribe Ooencyrtina) and can be separated from these genera by the characters given in the
key.
XENOSTRYXIS Girault
(Key couplets: 296, 348. Fig. 440)
Xenostryxis Girault, 19200: 41. Type-species: Xenostryxis margiscutellum Girault, by monotypy.
DISTRIBUTION AND SPECIES. One species, known only from Australia: margiscutellum Girault
(1920a: 41), possibly the same species distributed to India and southern Africa (BMNH, PPRI).
BIOLOGY. Unknown.
COMMENTS. Xenostryxis is probably related to either Neococcidencyrtus (tribe ?Habrolepidini)
or Thomsonisca (tribe Thomsoniscini) (Encyrtinae).
YASUMATSUIOLA Trjapitzin
(Key couplet: 158. Figs 82, 441)
Yasumatsuiola Trjapitzin, 1977: 153. Type-species: Yasumatsuiola orientalis Trjapitzin, by original
designation.
DISTRIBUTION AND SPECIES. One described species, Thailand: orientalis Trjapitzin (1977: 155),
also further undetermined material, which may include at least one undescribed species, from
India, Philippines and Australia (BMNH, BPBM).
BIOLOGY. Unknown.
COMMENTS. Placed in the tribe Dinocarsini (Tetracneminae).
ZAMENHOFELLA Girault
(Key couplet: 332)
Zamenhofella Girault, 1941:132. Type-species: Zamenhofella voltai Girault, by monotypy.
DISTRIBUTION AND SPECIES. One species, Australia only: voltai Girault (1941: 133).
BIOLOGY. Unknown.
INDO-PACIFIC ENCYRTIDAE 349
COMMENTS. We are unable to place this genus according to Trjapitzin's (19736) classification of
the Encyrtinae. It bears some resemblance to Austroencyrtus (which may be related to the
Bothriothoracini, see comments under Austroencyrtus).
ZAOMMA Ashmead
(Key couplet: 101. Fig. 49)
Zaomma Ashmead, 19006: 401. Type-species: Encyrtus argentipes Howard, by original designation.
Apterencyrtus Ashmead, 1905a: 5. Type-species: Apterencyrtus pulchricornis Ashmead, by original
designation.
Metallonoidea Girault, 1915c: 169. Type-species: Metallonoldea brittanica Girault, by monotypy.
Chiloneurinus Mercet, 1921: 646. Type-species: Chiloneurus microphagus Mayr, by original designation.
Richardsius Alam, 1957: 439. Type-species: Apterencyrtus thomsoniscae Alam, by original designation.
[As subgenus of Apterencyrtus.]
Metapterencyrtus Tachikawa, 1963: 213. Type-species: Metapterencyrtus mococc/Tachikawa, by original
designation.
DISTRIBUTION AND SPECIES. Thirteen species, cosmopolitan; one species from review area:
lambinus (Walker, 18380: 422) (India, Java, Philippines, New Zealand, Hawaiian Is.), also
undetermined material from Taiwan (UCR).
REFERENCES. Key to species: Gordh & Trjapitzin (19790); revision of Afrotropical species:
Prinsloo (1979).
BIOLOGY. Hyperparasites of Diaspididae (Homoptera) through other Encyrtidae (Hymenop-
tera).
COMMENTS. Ashmead (19050: 5) described Apterencyrtus pulchricornis from the Philippines.
This species has since been synonymised with microphagus Mayr (= lambinus) by Gahan (1951:
171), a synonymy which has been followed here. However, in the light of recent work by Gordh
& Trjapitzin (19790) and Prinsloo (1979), a more detailed study may show that the two species
are distinct.
ZAOMMOENCYRTUS Girault
(Key couplet: 214, 509. Figs 250, 251, 442-444)
Zaommoencyrtus Girault, 1916a: 46. Type-species: Zaommoencyrtus submicans Girault, by monotypy.
Bethylomimus Trjapitzin, 1962a: 430. Type-species: Bethylomimus liaoi Trjapitzin, by original desig-
nation.
DISTRIBUTION AND SPECIES. Five species, Holarctic, and also several undescribed species from
Papua New Guinea, Caroline Is., Solomon Is. and Fiji (BPBM, USNM).
REFERENCE. Review of Palaearctic species: Khlopunov (1981).
BIOLOGY. Parasites of larvae of Tenebrionidae and eggs and larvae of Cerambycidae (Cole-
optera).
COMMENTS. We have compared s 
