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
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology,
and an Historical series.
Papers in the Bulletin are primarily the results of research carried out on the unique and
ever-growing collections of the Museum, both by the scientific staff of the Museum and by
specialists from elsewhere who make use of the Museum's resources. Many of the papers are
works of reference that will remain indispensable for years to come.
Parts are published at irregular intervals as they become ready, each is complete in itself,
available separately, and individually priced. Volumes contain about 300 pages and several
volumes may appear within a calendar year. Subscriptions may be placed for one or more of
the series on either an Annual or Per Volume basis. Prices vary according to the contents of
the individual parts. Orders and enquiries should be sent to:
Publications Sales,
British Museum (Natural History),
Cromwell Road,
London SW75BD,
England.
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
Keeper of Entomology: Laurence A. Mound
Assistant Editor: W. Gerald Tremewan
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
1
D
S a" a CJ '-J
s .S ^5
u
5 5 '5 3 3 |
'
"-> <0 c o S- -S 2
O
<L>
2 rf a I ri 2^8
| 1 II 1 IjSI
<u
~o
11
SB/SL,
Sc
^
<u
'C
a
O ^
u
a "^
d
td
TD
t O
c .S
^w C
i C
Ivinate
-i 1 ^ g, s |
Si ^ 2^ | -a
? o ,0 ^ o -a- s
"c
Q. 3
3
fc O & t., ^S> a ^ ^
rt
^ 00
Q.
<D
Xij ^"^^^^Sij O
"E,
J*. 1)
U
cJu i>hjiJ>^2 i)
C/3
K"
J^
"o
^ & .. *t ss eo
u-= u u uurs S
SB e m n c
DO .O(ijO<UO "o
-C
u
S ^o^'So ^
1
c
OU HJH>U H
a
o
"o
O)
.22
'G
J"
B
* 1,
D
t/5
B C
T3
C
1
a C ^
1 J -o 2
3
wo
22
'S
D
a,
C/5
<U
s
^s>
c
<+*
l c . 1 ^ "5 o- j 6
1 -g e I -:. * e
| 1 | B go ^Ipj^l R ! 1
|^Co- - 6 J lco^ : |S'-' 2|8 ^
(^apc^ ,s EC>(^-O o^c '-.5^^ rjo
iX^r: OJON o ^--g^^j^^^J3r<Sc U 2o ^^ <u ?58,-^'^ ^^ om
4-
(Al
<u
^S i *> ^^ ^=SJ'Sj^ur"2 ^g" 'S^'^^S^C^S ~-~
able 1 Checklist o
pecies and subspeci
| |i Jill It ii i fi ilHllfiiill
H
1/3
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
al
p
in
8 2
1-, O 03
III
35 w .2
43 C ex
6
co
11
. U
00
o
O O
C C
IM ^ <o
O T3
t^ 3 uo
.2 C
Cj 43
^- O cfl
6
ill!
1-a 2
>- CO CO >,
-
&.-S a
L. M. PITKIN
1-3
4-7,
14-16
13,42,44
--D
10-12
17-19
20-22
14-16
20-22
23-
25
45
O
44
r O-
26
42
44
J-D-
21,22,24,25
27-29
34
31-33
1,45
43
r O -
42-4^
_ -n
11.
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
IJ
1^
0)
T c
E -E* J>
J2
~8J
"x
u
.2
I
"x
u
0)
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.
References
Agenjo, R. 1962. Resultados cientificos de una Pension de Estudias en el 'Museum National d'Histoire
Naturelle' de Paris, con la description de un genero y otra especie nuevos de lepidopteros espanoles,
dedicados al Excmo. Sr. D. Jesus Rubio y Garcia-Mina, Ministro de Education National. Eos. Revista
Espahola de Entomologia 38: 147-189, figs 1-6, pis 2-6.
1968. Catalogo ordenador de los Lepidopteros en Espana. Vigesimo novena familia bis. Gelechiidae.
Graellsia23, Appendix: [l]-[8].
Amsel, H. G. 1935. Weitere Mitteilungen iiber palastinensische Lepidopteren. Veroffentlichungen aus dem
Deutschen Kolonial- und LJbersee-Museum in Bremen 1: 223-277.
1950. Kleinschmetterlinge aus dem badischen Schwarzwald. Beitrage zur naturkundlichen Forschung
in Sudwestdeutschland 9: 26-28, figs 1-2.
1959. Portugiesische Kleinschmetterlinge gesammelt von Teodoro Monteiro, O.S.B. Anais da
Faculdade de Ciencias do Porto 41: 1-20, pis 1-2.
Anonymous 1969. European clover leaf tier. Plant Protection Bulletin. F.A.O. 17: 69.
Bradford, E. S. [1979]. Gelechiidae. In Emmet, A. M. (Ed.), A field guide to the smaller British
Lepidoptera pp. 115-132. London.
46 L. M. PITKIN
Bradley, J. D. 1966. Type specimens of Microlepidoptera in the University Museum, Oxford, described by
Haworth. Entomologist's Gazette 17: 129-140.
Bruand d'Uzelle, C. T. 1859. Observations entomologiques faites en 1858, relativement aux Plogophora
scita, Sericornis astrana et Gelechia vicinella. Annales de la Societe Entomologique de France (1858) (3)
6: 477-483.
( apuse, I. 1964. Uber drei Arten palaarktischer Gelechiidae: Carpatolechia dumitrescui n. g., n. sp.
Aproaerema aureliana n. sp. und Mirificarma formosella (Hb.) n. comb. (Lepidoptera: Gelechiidae).
Entomologisk TidskriftSS: 12-19.
Caradja, A. 1920. Beitrag zur Kenntnis der geographischen Verbreitung der Mikrolepidopteren des
palaearktischen Faunengebietes nebst Beschreibung neuer Formen. Ill Teil. Deutsche Entomologische
Zeitschrift, Iris 34: 75-179.
Chambers, V. T. 1872. Micro-lepidoptera [continued]. Canadian Entomologist 4: 146-150.
Chretien, P. 1915. Contribution a la connaissance des Lepidopteres du nord de 1'Afrique. Annales de la
Societe Entomologique de France 84: 289-374, figs 1-11.
- 1925. La legende de Graellsia isabellae. Appendice. Amateur de Papillons 2: 241-247, 257-263.
Curtis, J. 1827. British Entomology; being illustrations and descriptions of the genera of insects found
in Great Britain and Ireland: containing coloured figures from nature of the most rare and beautiful
species, and in many instances of the plants upon which they are found. 4, nos 147-194, pis 147-194.
London.
- 1829 [-1831]. A guide to an arrangement of British insects; being a catalogue of all the named species
hitherto discovered in Great Britain and Ireland, vi + [128] pp. London.
[Denis, M. & Schiffermuller, I.] 1775. Ankiindung eines systematischen Werkes von den Schmetterlingen
der Wienergegend. 323 pp., 3 pis. Wien.
Disque, H. 1908. Versuch einer microlepidopterologischen Botanik. Deutsche Entomologische Zeitschrift,
Iris 21: 34-80, 81-147.
Dumont, C. 1931. Contribution a 1'etude des Lepidopteres du nord de 1'Afrique (suite). Description et
ethologie de trois especes nouvelles. Bulletin de la Societe Entomologique de France 1931: 148-153,
figs 1-2.
Duponchel, P.-A.-J. 1842-[1844]. Nocturnes, supplement aux tomes quatrieme et suivants. In Godart,
J.-B., Histoire naiurelle des Lepidopteres ou papillons de France. Supplement 4. 534 pp., pis 51-90.
Paris.
Eckstein, K. 1933. Die Schmetterlinge Deutschlands mil besonderer Beriicksichtigung ihrer Biologic und
wirtschaftlichen Bedeutung. 5. Die Kleinschmetterlinge Deutschlands. 223 pp., 32 pis. Stuttgart.
Fuchs, A. 1902. Neue Geometriden und Kleinf alter des europaischen Faunengebiets. Stettiner Entomolo-
gische Zeitung 63: 317-330.
Gaede, M. 1937. Gelechiidae. Lepidopterorum Catalogus 79, 630 pp.
Gomes, C. M. R., Gottlieb, O. R., Gottlieb, R. G. & Salatino, A. 1981. Phytochemistry in perspective:
chemosystematics of the Papilionoideae. In Polhill, R. M. & Raven, P. H. (Eds), Advances in legume
systematics. Part 2: pp. i-v + 427-1094. Kew.
Gozmany, L. A. 1955. Notes on some Hungarian Gelechioidea and Coleophoridae. Annales Historico-
Naturales Musei Nationalis Hungarici (S.N.) 6: 307-320, figs 1-24.
- 1958. Microlepidoptera IV. Fauna HungariaeW: 1-295, 1-8 [index], figs 1-145.
Guenee, A. 1849. In Lucas, H., Exploration scientifique de I'Algerie pendant les Annees 1840, 1841, 1842.
Zoologie. Histoire naturelle des animaux articules 3, 527 pp. Ibid. 4, Lepidopteres, pis 1-4. Paris.
Hartig, F. 1964. Microlepidotteri della Venezia Tridentina e delle regioni adiacenti. Parte III. (Fam.
Gelechiidae-Micropterygidae). Studi Trentini di Scienze Naturali 41 (34): 1-292.
Hartig, F. & Amsel, H. G. 1951. Lepidoptera Sardinica. Fragmenta Entomologica 1: 3-152.
Hauder, F. 1918. Einige Kleinschmetterlings-Aberrationen. Entomologische Zeitschrift 31: 97-98, 102-
103.
Hofmann, E. 1875. Die Kleinschmetterlingsraupen - Microlepidoptera - in systematischer Reihenfolge nach
dem Cataloge von Dr. Staudinger & Dr. Wocke 1871. Zugleich als Erganzung von S. v. Praun's
Microlepidoptera. iv + 221 pp., 10 pis. Niirnberg.
Hruby, K. 1964. Prodromus Lepidopter Slovenska. 962 pp., 14 maps. Bratislava.
Hiibner, J. 1793. Sammlung auserlesener Vdgel und Schmetterlinge, mil ihren Namen. 16 pp., 100 pis.
Augsburg.
- 1796-[1836]. Sammlung europaischer Schmetterlinge 8, 78 pp. (1796), 71 pis (1796-[1836]).
Augsburg.
1816-[1825]. Verzeichniss bekannter Schmettlinge. 431 pp. Augsburg.
1822. Systematisch-alphabetisches Verzeichniss aller bisher bey den Furbildungen zur Sammlung
GELECHIID MOTHS 47
europaischer Schmetterlinge angegebenen Gattungsbenennungen; mit Vormerkung auch augsburgischer
Gattungen. vi + 81 pp. Augsburg.
Humphreys, H. N. & Westwood, J. O. [1843-J1845. British moths and their transformations. 2. xix +
268 pp., pis 57-124. London.
Karsholt, O. & Nielsen, E. S. 1976. Systematisk fortegnelse over Danmarks sommerfugle. 128 pp.
Klampenborg, Denmark.
Klimesch, J. 1961. Ordnung Lepidoptera. I. Teil: Pyralidina, Tortricina, Tineina, Eriocraniina und
Micropterygina. In Franz, H. , Die Nordost-Alpen im Spiegel ihrer Landtierwelt. 2: 481-789.
Klots, A. B. 1956. Lepidoptera. In Tuxen, S. L. (Ed.), Taxonomist's glossary of genitalia in insects.
Pp. 97-111, figs. 121-132. Copenhagen.
Koc.ak, A. 0. 1982. Additions and corrections to the names published in 'Systematic and Synonymic List of
the Lepidoptera of France, Belgium and Corsica' by Leraut, 1980. Priamus 2: 97-133.
Krogerus, H., Opheim, M., Schantz, M. von, Svensson, I & Wolff, N. L. 1971. Catalogus Lepidopterorum
Fenniae et Scandinaviae. Microlepidoptera. 40pp. Helsinki.
Kupicha, F. K. 1977. The delimitation of the tribe Vicieae (Leguminosae) and the relationships of Cicer L.
Botanical Journal of the Linnean Society of London 74: 131-162.
Kuznetzov, V. I. & Stekol'nikov, A. A. 1978. Systematic position and phylogenetic relationships of the
superfamily Coleophoroidea (Lepidoptera: Oecophoridae, Coleophoridae, Ethmiidae) treated on the
base of functional morphology of the male genitalia. Entomologiche Obozrenie 57: 131-149, figs 1-11.
[In Russian.]
1979. The systematic position and phylogenetic relationships of the superfamily Coleophoroidea
(Lepidoptera: Oecophoridae, Coleophoridae, Ethmiidae) as revealed by the functional morphology of
the male genitalia. Entomological Review (1978) 57: 91-103, figs 1-11. [English translation of Kuznetzov
& Stekol'nikov, 1978.]
Lempke, B. J. 1976. Naamlijst van de Nederlandse Lepidoptera. 100 pp. [Amsterdam.]
Lhomme, L. [1946-1948]. Catalogue des Lepidopteres de France et de Belgique. 2 (4): 489-648. Douelle
(Lot).
[1948-1949]. Catalogue des Lepidopteres de France et de Belgique. 2 (5): 649-808.
Mann, J. 1866. Aufzahlung der im Jahre 1865 in der Dobrudscha gesammelten Schmetterlinge. Verhand-
lungen der Zoologisch-Botanischen Gesellschaft in Wien 16: 321-360.
Mariani, M. 1941, 1943. Fauna Lepidopterorum Italiae. Parte 1 - Catalogo ragionato dei Lepidotteri
d'ltalia. Giornale di Scienze Naturali ed Economiche di Palermo 42 (3): 1-79 (1941), 81-237 (1943), 1 pi.
Meess, A. 1910. See Spuler, A. 1903-1910.
Meyrick, E. 1895. A handbook of British Lepidoptera. [vi] + 843 pp. London.
1925. Lepidoptera Heterocera. Fam. Gelechiadae. Genera Insectorum 184: 1-290, pis 1-5.
Milliere, P. 1859-1864. Iconographie et description de chenilles et Lepidopteres inedits 1, 424 pp., 50 pis.
Paris.
Miiller-Rutz, J. 1913-1914. xxxiv. Gelechiidae. Pp. 453-500. In Vorbrodt, K. & Muller-Rutz, J., Die
Schmetterlinge der Schweiz 2: 727 pp. , 2 tables. Bern.
Nickerl, 0. 1908. Beitrage zur Insekten- Fauna Bohmens. 6. Die Motten Bohmens (Tineen). [viii] + 161 pp.
Prag.
Opheim, M. 1978. The Lepidoptera of Norway, check-list. Part III Gelechioidea (first part). 30 pp.
Trondheim.
Piskunov, V. 1. 1981. 50. Gelechiidae. In Zagulaev, A. K. etalii, [Key to insects in the European part of the
U.S.S.R. 4: Lepidoptera, pt 2]. Pp. 649-748, figs 608-675. Leningrad. [In Russian.]
Polhill, R. M. & Raven, P. H. (Eds) 1981. Advances in legume systematics. Part 1. xvi + 425 + xxi pp. Kew.
Prose, H. 1957. II. Spezieller Teil: Systematische Liste. B. Microlepidoptera. In Daniel, F. & Wolfsberger,
J., Die Fohrenheidegebiete des Alpenraumes als Refugien warmeliebender Insekten. II. Der Sonnen-
berghang bei Naturns im Vintschgau (Siidtirol). Mitteilungen der Miinchener Entomologischen Gesell-
schaft 47: 21-121, 2 maps.
Rebel, H. 1901. Famil. Pyralidae - Micropterygidae. In Staudinger, O. & Rebel, H., Catalog der
Lepidopteren des palaearctischen Faunengebietes 2, 368 pp. Berlin.
- 1903. Studien iiber die Lepidopterenfauna der Balkenlander. I. Teil. Bulgarien und Ostrumelien.
Annalen des (K.K.) Naturhistorischen (Hof) Museums 18: 123-347, pi. 3.
1914. Zweiter Beitrag zur Lepidopterenfauna Unter-Aegyptens. Deutsche Entomologische Zeit-
schrift, Iris 28: 258-270, pi. 4; figs 1-8.
1930. Zwei neue Gelechiidae aus Korsika. Zeitschrift des Osterreichischen Entomologen-Vereines,
Wien 15: 25-26.
1931. In Rebel, H. & Zerny, H., Die Lepidopterenfauna Albaniens (mit Berucksichtigung der
48 L. M. PITKIN
Nachbargebiete). Denkschriften der Kaiserlichen Akademie der Wissenschaften 103: 37-161, figs 1-10,
pi. 1, map. [Dated 1931 on wrapper of reprint.]
Rothschild, N. C. 1913. Adatok Magyarorszag lepkefaunajahoz (Beitrage zur Lepidopterenfauna
Ungarns). Rovartani Lapok 20: 66-91, pi. 2.
Sattler, K. 1960. Generische Gruppierung der europaischen Arten der Sammelgattung Gelechia (Lepidop-
tera, Gelechiidae). Deutsche Entomologische Zeitschrift (N.F.) 7: 10-118, figs 1-138.
1961 . Uber Mirificarma cabezella (Chret. , 1925). Zeitschrift der Wiener Entomologischen Gesellschaft
46: 86-88, figs 1-4.
- 1976. A taxonomic revision of the genus Ornativalva Gozmany, 1955 (Lepidoptera: Gelechiidae).
Bulletin of the British Museum (Natural History) (Entomology) 34: 87-152, 27 pis, 27 figs.
1979. A taxonomic revision of the genus Deltophora Janse, 1950 (Lepidoptera: Gelechiidae). Bulletin
of the British Museum (Natural History) (Entomology) 38: 263-322, 112 figs.
Schille, F. 1931. Fauna motyli Polski. 2. Prace Monograficzne Komisji Fizjograficznej 7: 1-358.
Schiitze, K. T. 1931. Die Biologie der Kleinschmetterlinge unter besonderer Berucksichtigung ihrer
Ndhrpflanzen und Erscheinungszeiten. 235 pp. Frankfurt.
Sorhagen, L. 1886. Die Kleinschmetterlinge der Mark Brandenburg und einiger angrenzenden Landschaf-
ten mil besonderer Berucksichtigung der Berliner Arten. \ + 368 pp. Berlin.
Spuler, A. 1903-1910. Die Schmetterlinge Europas. 2: [vi] + 523 pp, figs 1-151, 27-239, 21 pis. Stuttgart.
Stainton, H. T. 1865. The natural history of the Tineina 9, [vii] + 276 pp., pis 1-8. London, Paris, Berlin.
1867. The natural history of the Tineina 10, xi + 304 pp., pis 10-16. London, Paris, Berlin.
Staudinger, 0. 1859. Diagnosen nebst kurzen Beschreibungen neuer andalusischer Lepidopteren. Stettiner
Entomologische Zeitung 20: 211-259.
1871. Beitrag zur Lepidopterenfauna Griechenlands. Horae Societatis Entomologicae Rossicae 7:
3-304, pis 1-3.
Stephens, J. F. 1829. A systematic catalogue of British insects, etc. 2. Insecta Haustellata. 388 pp. London.
1834-1835. Illustrations of British entomology, etc. Haustellata 4. 436 pp., pis 33-41. London.
Treitschke, F. 1832. Die Schmetterlinge von Europa 9 (1), viii + 262 pp. Leipzig.
- 1833. Die Schmetterlinge von Europa 9 (2), 294 pp. Leipzig.
Turati, E. 1924. Spedizione Lepidotterologica in Cirenaica 1921-1922. Atti della Societa Italiana di Scienze
Naturali e del Museo Civico di Storia Naturale 63: 21-191, figs 1-7, pis 1-6.
Tutin, T. G., Heywood, V. H., Surges, N. A., Moore, D. M., Valentine, D. H., Walters, S. M. & Webb,
D. A. (Eds), 1964-1980. Flora Europaea. 5 vols. Cambridge.
Villers, C. de 1789. Caroli Linnaei Entomologia Faunae Suecicae descriptionibus aucta 2: xvi + 656 pp.,
pis 4-6. Lugduni.
Walsingham, Lord 1904. Algerian Microlepidoptera. Entomologist's Monthly Magazine 40: 214-223,
265-273.
Wocke, M. 1861. II. Microlepidoptera. In Staudinger, O. & Wocke, M., Catalog der Lepidopteren
Europa 's und der angrenzenden Lander, xvi + 192 pp. Dresden.
- 1871. II. Microlepidoptera. In Staudinger, O. & Wocke, M., Catalog der Lepidopteren des
europaeischen Faunengebiets. xxxviii + 426 pp. Dresden.
Wolff, N. L. 1971. Lepidoptera. Zoology of ^Iceland 3 (45): 193 pp., figs 1-33, pis 1-15.
Zeller, P. C. 1839. Versuch einer naturgemassen Eintheilung der Schaben. Isis von Oken 1839: 167-220.
1847. Bemerkungen iiber die auf einer Reise nach Italien und Sicilien beobachteten Schmetterlingsar-
ten. IX. Isis von Oken 1847: 801-859.
Zerkowitz, A. 1946. The Lepidoptera of Portugal. Journal of the New York Entomological Society 54:
51-87,115-165,211-261.
Zerny, H. 1936. Die Lepidopterenfauna des Grossen Atlas in Marokko und seiner Randgebiete. Memoires
de la Societe des Sciences Naturelles (et Physiques) du Maroc 42: 1-163, figs 1-4, pis 1, 2.
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,
and an Historical series.
Papers in the Bulletin are primarily the results of research carried out on the unique and
ever-growing collections of the Museum, both by the scientific staff of the Museum and by
specialists from elsewhere who make use of the Museum's resources. Many of the papers are
works of reference that will remain indispensable for years to come.
Parts are published at irregular intervals as they become ready, each is complete in itself,
available separately, and individually priced. Volumes contain about 300 pages and several
volumes may appear within a calendar year. Subscriptions may be placed for one or more of
the series on either an Annual or Per Volume basis. Prices vary according to the contents of
the individual parts. Orders and enquiries should be sent to:
Publications Sales,
British Museum (Natural History),
Cromwell Road,
London SW75BD,
England.
^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
00
o
-a
ed
.n
c
"O
u
c
Q
cd
6
P. C. BARNARD
1 1 1 1 l + t 1 1 1 I++I + I+ +
1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 + 1
2 I I
4 I I
00 I I
I I I I I I I I I I I + I I I I
I I I I I I I I I ++ I I I I I
I I I I I I I I I I + I I I I I
I I I I
I I I + I I I I + I I I I
I I I I ++ I I I I I I I I
"-> I I I I I I I I I I + I I I I I I I I
2 + + + + + + + +I i i i 1 + 1 i i i i
' I I I I I I + I I I I I I I I I I I
^ I I I I I I + I I I I I I I I I I I I
S++++++IIIIIIIIIIIII
^++++++1111111111111
I I I I I + I I I I I I I I I I I I I
m + + + + +l I I I I I++I I I I 1 +
s I I I ++ I I I I I I I I I i I I I I
s I I I I + I I I I I I I I I I I I I I
^ + + +11 + 111 c-- I I I I I I I I I
J i i + i i i i i i i i i i i i i i i i
D.
00
c-as
i J f !
C-Q v> -i-
1 J i
"
a
& > a " .
-c C ? oo^>^s 3 S -S
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
^
uei i ici i
a.
U
o
l_
O)
1
a
c
a
L ^
**
12
11 ^
TUSCQIU +
26 32 2 ,
- 34
W '
i-?- 3 . 3
1 - ulmen +
19 24
-n 3 . 1
pei IUCIQQ *
^O
on
i_
a>
e
JU
^
27
11
21 23 25
1 parva
1
H 5
mnnrvri
ttononnl one. ie
279
22
J8_
exsiliens
- apicalis
10
- gratiosa
petiolata
17
15 16
13
distincta
bifasciata
proluta
!2 Q.
"a ^
5 2
Q. O)
O
a a.
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).
References
Banks, N. 1913. Synopses and descriptions of exotic Neuroptera. Transactions of the American Entomo-
logical Society 39: 201-242.
- 1920. New neuropteroid insects. Bulletin of the Museum of Comparative Zoology, Harvard 64:
299-362.
1931. Some neuropteroid insects from the Malay peninsula. Journal of the Federated Malay States
Museums 16: 377-409.
1939. Notes and descriptions of Oriental Oestropsychinae (Trichoptera). Psyche, Cambridge 46:
52-61.
1940. Report on certain groups of neuropteroid insects from Szechwan, China. Proceedings of the
United States National Museum 88: 173-220.
Barnard, P. C. 1980. A revision of the Old World Polymorphanisini (Trichoptera: Hydropsychidae).
Bulletin of the British Museum (Natural History) (Entomology) 41: 59-106.
Beatty, J. 1982. Classes and cladists. Systematic Zoology 31: 25-34.
Betten, C. 1909. Notes on the Trichoptera in the collection of the Indian Museum. Records of the Indian
Museum 3: 233-242, 5 pis.
Boon, P. J. 1979. Adaptive strategies of Amphipsyche larvae (Trichoptera: Hydropsychidae) downstream
of a tropical impoundment. In Ward, J. V. & Stanford, J. A. (Eds), The ecology of regulated streams, pp.
237-255. New York.
Brauer, F. 1875. Beschreibung neuer und ungenugend bekannter Phryganiden und Oestriden. Verhand-
lungen der Zoologisch-Botanischen Gesellschaft in Wien 25: 69-78.
Chutter, F. M. 1963. Hydrobiological studies on the Vaal River in the Vereeniging area. Part I.
128 P. C. BARNARD
Introduction, water chemistry and biological studies on the fauna of habitats other than muddy bottom
sediments. Hydrobiologia21: 1-89.
1968. On the ecology of the fauna of stones in the current in a South African river supporting a very
large Simulium (Diptera) population. Journal of Applied Ecology 5: 531-561.
Corbet, P. S. 19580. Some effects of DDT on the fauna of the Victoria Nile. Revue de Zoologie et de
Botanique Africaines 51 ': 73-95.
19586. Larvae of certain East African Trichoptera. Revue de Zoologie et de Botanique Africaines 58:
203-213.
1966. Parthenogenesis in caddisflies (Trichoptera). Canadian Journal of Zoology 44: 981-982.
Corbet, P. S. & I jonnelaiid, A. 1955. The flight activity of East African Trichoptera. Universitet i Bergen
Arbok 1955: 1-49.
Flint, O. S. & Bueno Soria, J. 1982. Studies of neotropical caddisflies, XXXII: the immature stages of
Macronema variipenne Flint & Bueno, with the division of Macronema by the resurrection of Macroste-
mum (Trichoptera: Hydropsychidae). Proceedings of the Biological Society of Washington 95:
358-370.
Flint, O. S. & Wallace, J. B. 1980. Studies of neotropical caddisflies XXV: the immature stages of
Blepharopus diaphanus and Leptonema columbianum (Trichoptera: Hydropsychidae). Proceedings of
the Biological Society of Washington 93: 178-193.
Gibbs, D. G. 1973. The Trichoptera of Ghana. Deutsche Entomologische Zeitschrift (N.F.) 20: 363-424.
Gourou, P. 1970. L'Afrique. 88 pp. Paris.
Hafiz, H. A. 1937. Notes on the larva and pupa of Amphipsyche indica Martynov (Insecta: Trichoptera).
Records of the Indian Museum 39: 113-116.
Hennig, W. 1966. Phylogeneticsystematics. 263 pp. Urbana.
Hickin, N. E. 1955. Larvae of some East African Trichoptera. Proceedings of the Royal Entomological
Society of London (A) 30: 155-163.
Hwang Chi-ling 1957. Descriptions of Chinese caddis flies (Trichoptera). [In Chinese, with brief English
summary.] Acta Entomologica Sinica 7: 373^04.
Jacquemart, S. 1957. Trichoptera des Lacs Kivu et Edouard. Exploration Hydrobiologique des Lacs Kivu,
Edouard et Albert (1952-1954), Resultats Scientifiques 3 (2): 67-129, 2 pis.
1963. Trichoptera. South African Animal Life 11: 337-415.
Kimmins, D. E. 1955. Results of the Oxford University expedition to Sarawak, 1932. Order Trichoptera.
Sarawak Museum Journal 6: 374-442.
1957a. New and little-known species of African Trichoptera. Bulletin of the British Museum (Natural
History) (Entomology) 6: 1-37.
1957ft. Lectotypes of Trichoptera from the McLachlan collection now in the British Museum (Natural
History). Bulletin of the British Museum (Natural History) (Entomology) 6: 91-126.
1962. New African caddis-flies (Order Trichoptera). Bulletin of the British Museum (Natural History)
(Entomology) 12: 81-121.
1963. On the Trichoptera of Ethiopia. Bulletin of the British Museum (Natural History) (Entomology)
13: 117-170.
Lepneva, S. G. 1947. The larvae of caddisflies of the subfamily Macronematinae (Trichoptera, Hydro-
psychidae) in the U.S.S.R. [In Russian.] Entomologicheskoe Obozrenie 29: 139-151.
- 1970. Trichoptera. II (1). Larvae and pupae of Annulipalpia. Fauna SSSR 88: iv+638 pp.
Marlier, G. 1962. Genera des Trichopteres de 1'Afrique. Annales du Musee Royal de I'Afrique Centrale
(Serie 8, Sciences Zoologiques) 109: 1-263.
1978. Sur une collection de Trichopteres de I'Afrique occidentale. Revue de Zoologie Africaine 92:
283-302.
Martynov, A. V. 1935. On a collection of Trichoptera from the Indian Museum. Part I. Annulipalpia.
Records of the Indian Museum 37: 93-209.
- 1936. On a collection of Trichoptera from the Indian Museum. Part II. Integripalpia. Records of the
Indian Museum 38: 239-306.
McLachlan, R. 1872. Non-odonates (pp. 47-71). In Selys-Longchamps, M. E. & McLachlan, R.,
Materiaux pour une faune nevropterologique de 1'Asie septentrionale. Seconde partie. Annales de la
Societe Entomologique de Belgique 15: 25-77.
1878. A monographic revision and synopsis of the Trichoptera of the European fauna. Part VII, pp.
349-428. London.
Mosely, M. E. 1942. Chinese Trichoptera: a collection made by Mr. M. S. Yang in Foochow. Transactions
of the Royal Entomological Society of London 92: 343-362.
MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 129
Navas, L. 1914. Les insectes de 1'Orient recueillis par M. le baron J. de Guerne. Entomologicheskoe
Obozrenie 14: 319-322.
1922. Insectos exoticos. Broteria (Serie Zoologia) 20: 49-63.
1923. Insecta nova. Memorie dell' Accademia Pontifica del Nuovi Lincei (2) 6: 19-27.
- 1927. Nevropteres d'Egypte et de Palestine. Bulletin. Societe Entomologique d'Egypte 19 (1926):
192-216.
- 1931a. Nevropteres et insectes voisins - Chine et pays environnants. Deuxieme serie (1). Notes
d'Entomologie Chinoise 1 (7): 1-10.
1931ft. Insectos de la India. Revista de la Academia de Ciencias Exactas, Fisico-Quimicas y Naturales
de Zaragoza 14 (1930): 74-92.
1934. Insecta orientalia XIII. Memorie dell' Academia Pontifica del Nuovi Lincei (3) 1: 217-228.
1935. Decades de insectos nuevos. 27. Broteria (Serie trimestral) 4 (31): 97-107.
Nelson, G. & Platnick, N. I. 1981. Systematics and biogeography: cladistics and vicariance. xiii+567 pp.
New York.
Nielsen, A. 1957. A comparative study of the genital segments and their appendages in male Trichoptera.
Biologiske Skrifter, Danske Videnskab ernes Selskab 8 (5): 1-159.
Platnick, N. 1. 1979. Philosophy and the transformation of cladistics. Systematic Zoology 28: 537-546.
Resh, V. H. & Unzicker, J. D. 1975. Water quality monitoring and aquatic organisms: the importance of
species identification. Journal of the Water Pollution Control Federation 47: 9-16.
Savigny, M. J. C. L. de 1813. Description del'Egypte. Histoire Naturelle, vol. 2 (Nevropteres, pi. 3,fig. 18).
Paris.
Schmid, F. 1958. Trichopteres de Ceylan. Archivfiir Hydrobiologie 54: 1-173.
1980. Genera des Trichopteres du Canada et des Etats adjacents. Les Insectes et Arachnides du
Canada 7: 1-296.
Scott, K. M. F. 1970. Some notes on the Trichoptera of standing waters in Africa, mainly south of the
Zambezi. Hydrobiologia35: 177-195.
1975. The value of larval stages in systematic studies of the Trichoptera, with particular reference to
the Hydropsychidae from Africa south of the Sahara. Proceedings of the 1st Congress of the Entomo-
logical Society of South Africa 1975: 41-52.
(in press). On the Hydropsychidae (Trichoptera) of southern Africa with keys to African genera of
images, larvae and pupae and species lists. Annals of the Cape Provincial Museums (Natural History) 14
(8).
Seshadri, A. R. 1955. An extraordinary outbreak of 'caddice flies' (Trichoptera) in the Mettur-Dam
township area in Salem District, South India. Indian Journal of Entomology 17: 337-340.
Smith, A. G., Hurley, A. M. & Briden, J. C. 1981. Phanerozoic palaeocontinental world maps, vi+102 pp.
Cambridge.
Stanford, J. A. & Ward, J. V. 1981. Preliminary interpretations of the distribution of Hydropsychidae in a
regulated river. Proceedings of the 3rd International Symposium on Trichoptera, 1980, pp. 323-328.
Statzner, B. 1981. A progress report on Hydropsychidae from the Ivory Coast: characters for the specific
identification of larvae and population dynamics of four abundant species. Proceedings of the 3rd
International Symposium on Trichoptera, 1980, pp. 329-335.
I 'liner, G. 1907. Trichopteren. Monographic der Macronematinae. Collections Zoologiques du Baron
Edm. de Selys Longchamps 6 (2): 1121. Brussels.
1909. Einige neue exotische Trichopteren. Notes from the Leyden Museum 31: 125-142.
1910. Uber einige von Herrn E. Jacobson auf Java gesammelte Trichopteren. Notes from the Leyden
Museum 32: 47-66.
1913. Uber einige von Edw. Jacobson auf Java gesammelte Trichopteren. Notes from the Leyden
Museum 35: 78-101.
- 1923. Trichopteren aus dem agyptischen Sudan und aus Kamerun. Mitteilungen der Munchener
Entomologischen Gesellschaft 13: 9-20.
1924. Wissenschaftliche Ergebnisse der mit Unterstiitzung der Akademie der Wissenschaften in Wien
aus der Erbschaft Treitl von F. Werner unternomennen zoologischen Expedition nach dem Anglo-
Agyptischen Sudan (Kordofan) 1914. XII. Trichopteren und Ephemeropteren. Denkschriften der
Akademie der Wissenschaften, Wien 99: 1-9.
1927. Einige neue Trichopteren aus Asien. Entomologische Mitteilungen 16: 172-182.
1930. Trichopteren von den Philippinen und von den Sunda-Inseln. Treubia 11: 373-498.
1951. Kocherfliegen (Trichopteren) von den Sunda-Inseln (Teil I). Archiv fur Hydrobiologie,
Supplement 19: 1-528.
130
P. C. BARNARD
- 1957. Kocherfliegen (Trichopteren) von den Sunda-Inseln (Teil III). Archiv fur Hydroblologie,
Supplement 23: 109-470.
1963. Trichopteren (Kocherfliegen) aus Agypten. Archiv fiir Hydrobiologie 59: 257-271.
Weidner, H. 1964. Die entomologischen Sammlungen des Zoologischen Staatsinstituts und Zoologischen
Museums Hamburg. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 62:
55-100.
Wiley, E. O. 1981. Phylogenetics: the theory and practice of phylogenetic systematics. xvi+439 pp. New
York.
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
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four
scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology,
and an Historical series.
Papers in the Bulletin are primarily the results of research carried out on the unique and
ever-growing collections of the Museum, both by the scientific staff of the Museum and by
specialists from elsewhere who make use of the Museum's resources. Many of the papers are
works of reference that will remain indispensable for years to come.
Parts are published at irregular intervals as they become ready, each is complete in itself,
available separately, and individually priced. Volumes contain about 300 pages and several
volumes may appear within a calendar year. Subscriptions may be placed for one or more of
the series on either an Annual or Per Volume basis. Prices vary according to the contents of
the individual parts. Orders and enquiries should be sent to:
Publications Sales,
British Museum (Natural History),
Cromwell Road,
London SW75BD,
England.
World List abbreviation: Bull. Br. Mas. nat. Hist. (Ent.)
Trustees of the British Museum (Natural History), 1984
The Entomology series is produced under the general editorship of the
Keeper of Entomology: Laurence A. Mound
Assistant Editor: W. Gerald Tremewan
ISBN 565 06002 3
ISSN 0524-6431 Entomology series
Vol48No3ppl31-395
British Museum (Natural History)
Cromwell Road
London SW7 5BD Issued 28 June 1984
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 w 
