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Full text of "Bulletin of the British Museum (Natural History)"

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 









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GELECHIID MOTHS 5 

characters. The division of the two major species-groups is corroborated by host-plant differ- 
ences. Host-plants of the montivaga-group are unknown. The host-plants of the maculatella- 
group are all in the epulvinate series, in the tribes Loteae, Coronilleae, Trifolieae, Vicieae and 
Galegeae, while the interuptella-group has Genisteae host-plants (see Table 1). 

One species in the interuptella-group, cytisella, has Ononis as a host-plant, in addition to 
several species of Genisteae. Although Ononis is a member of the Trifolieae, it has some 
Genisteae-like morphological characters and it has been linked with the Genisteae by early 
authors (see the discussion of the taxonomic history of Ononis by Kupicha, 1977: 134). Ononis 
also has biochemical affinities with the Genisteae, to which it has similar isoflavenoids (see 
Gomes etalii, 1981: 472, 473). It is possible that these characteristics make the plant acceptable 
to the moth which otherwise selects Genisteae. In addition the host-plant must be acceptable to 
the larva. 

Although Mirificarma species within a species-group share host-plants of the same tribe or 
group of tribes, there is little evidence that the close relationship of species is reflected in the 
close relationship of their host-plants. In the maculatella-group, eburnella and flavella both 
occur mainly on Trifolieae, including Trifolium. However, eburnella also occurs on Coronilleae, 
the host-plant tribe of maculatella, a species to which eburnella is less closely related. M. 
maculatella and its sister-species denotata use plants of different tribes, Coronilleae and 
Galegeae respectively. 

In the interuptella-group , the closely related mulinella and ulicinella both occur on Ulex, 
although on different species, but mulinella shares Cytisus nigricans, Calicotome spinosa and 
Genista with cytisella; Genista germanica and G. tinctoria with lentiginosella, and Cytisus 
scoparius with interuptella. M. mulinella and ulicinella form a close group with cabezella which 
differs considerably from these two in its host-plant Adenocarpus , an outlier in the Genisteae. 

The distribution of a host-plant has not always restricted the distribution of the moth, which 
may have adapted to different plants in different parts of its range. For example, cytisella was 
described from specimens bred on Cytisus nigricans in Germany, and has subsequently been 
bred on this, for example, by Klimesch (1961: 651) in Austria. C. nigricans does not occur in 
France (Tutin etalii, 1968: 86) and cytisella has been recorded there on Calicotome and Ononis. 
The ability of some Mirificarma species to utilize a range of host-plant genera suggests that they 
have adapted to different hosts rather than that they have evolved in conjunction with their 
hosts. 

The systematic position of Mirificarma 

Many species currently in Mirificarma were originally described in Gelechia, and subsequently 
listed in this genus by Gaede (1937). However, Gelechia has been widely used to include many 
unrelated species and has been referred to as the microlepidopterist's 'waste-paper box' 
(Chambers, 1872: 147, and subsequent authors). Mirificarma is clearly separated from Gelechia 
(sensu Saltier, 1960) by the presence of a filament on the male genitalia, as well as other 
characters (see also remarks, p. 15). 

Following the original placing of flavella in Acompsia Hiibner, the related species eburnella 
(as formosella Hiibner) and pallidipulchra were also listed in this genus by Meyrick (1925: 142) 
and Gaede (1937), possibly because Acompsia and Mirificarma have similar wing venation. 
Acompsia and Mirificarma differ, however, in many characters including the male eighth tergite 
and sternite which are laterally fused in the former but separated in the latter. The three species 
have also been placed in Rhinosia Treitschke, either in the original descriptions or by 
subsequent authors including Meess (in Spuler, 1910: 344), apparently because of a strong 
superficial similarity to species then in Rhinosia but now transferred to Orophia Hiibner 
(Oecophoridae). Orophia ferrugella ([Denis & Schiffermiiller]), formerly in Rhinosia, has 
sometimes been misidentified as flavella (see p. 27). 

In the original description, Mirificarma was placed between Neofaculta Gozmany and Lita 
Treitschke, both currently in the subfamily Gelechiinae, without indication as to whether its 
position was deliberate or arbitrary. Sattler (1960: 41) also placed it among genera currently in 
the Gelechiinae although its placement between Lita and Aroga Busck was arbitrary. 



O L. M. PITKIN 

The subfamilies of Gelechiidae are mostly poorly defined and the relationships of the genera 
still require much investigation. However, Mirificarma does appear to share some characters 
with certain other genera within the Gelechiidae. The eighth abdominal tergite and sternite of 
the male are separated into free flaps in Mirificarma. This is usual in the Gelechiinae (tribes 
Gelechiini, Teleiodini and Gnorimoschemini) whereas the tergite and sternite are normally 
fused laterally in the rest of the Gelechiidae. The long narrow scales of the coremata of the 
eighth tergite of Mirificarma are similarly placed to those occurring in many Gelechiinae, 
attached closely to the tergite. Coremata occur in a few genera outside the Gelechiinae, and are 
probably homologous in some cases, such as in Deltophora Janse in the Aristoteliinae, although 
in this genus the coremata are usually loosely attached to the tergite. The coremata of the male 
eighth sternite of Mirificarma, on the dorsal membrane, consist of enlarged scales with a 
rounded, often inflated structure. Some Gelechiinae have scales similarly situated, as seen in 
Gelechia scotinella Herrich-Schaffer, G. hippophaella (Schrank), G. nigra (Haworth) and 
Teleiodes myricariella (Frey), in which the scales are large and sometimes long although not 
inflated. The scales are deciduous, consequently fresh preparations need to be made to assess 
how widespread they might be in the Gelechiinae and whether they might occur outside the 
Gelechiinae. The scales are most similar to those of Mirificarma in Psoricoptera gibbosella 
(Zeller), currently in the Hypatiminae although with a number of similarities to Gelechia near 
which it might be better placed. 

In common with many Gelechiinae, the posterior apophyses of the female genitalia are often 
long in Mirificarma, although this character is not restricted to this subfamily. 

Within the Gelechiinae, Mirificarma does not appear to be closely related to the Gnorimos- 
chemini, in which the signum of the female genitalia is a strong hook-like structure. The signum 
of Mirificarma varies interspecifically but is never of this form. M. montivaga, which has the least 
apomorphies of the genus, has a signum similar to that frequently seen in Gelechia and the 
Teleiodini. This type of signum is also found in Neofriseria Sattler, currently placed in the 
Gelechiini although not in close relationship with Gelechia, and Psoricoptera gibbosella, 
currently in the Hypatiminae although, as mentioned above, probably more closely related to 
Gelechia. Mirificarma does not show evidence of a close relationship with the Teleiodini, in 
which the male genitalia have frequent modifications not shared by Mirificarma, such as a 
narrow, elongate uncus. The male eighth segment varies in shape in the Teleiodini and usually 
differs from that of Mirificarma. The gnathos is absent in some Teleiodini and if present, usually 
also differs from that of Mirificarma. 

The possession of a filament on the male genitalia distinguishes Mirificarma from all other 
Gelechiidae. However, the pair of sclerites which support the filament in Mirificarma, extending 
anteriorly from the base of the sacculus and valva and lying between the vinculum and tegumen, 
appear to be homologous with similar structures in species of Gelechia, particularly G. sabinella 
Zeller, G. nigra, G. rhombella ([Denis & Schiffermiiller]) and G. senticetella Staudinger. The 
sclerites in these species are fused anteriorly, at the region from which the filament would arise in 
Mirificarma. This region is usually membraneous in Gelechia, and never a well-sclerotized 
swelling as in montivaga which has the least-developed filament of Mirificarma. A membraneous 
sac is present around the filament and supporting sclerites of Mirificarma and this is sometimes 
also present around the sclerites of Gelechia. Slight sclerotized extensions from the base of the 
sacculus or valva may occur in other genera, such as Chionodes Hiibner, but it is the 
development of these into the long sclerites that appears to be a synapomorphy of Mirificarma 
and Gelechia. 

Thus I consider Mirificarma to belong in the Gelechiinae, as delimited above, on account of 
the structure of the male eighth abdominal segment. Within this subfamily I consider it to be 
close to Gelechia, with which it shares the presence of the sclerites described above. 

Classification of Mirificarma species 

The cladogram (Fig. 2) is based on the list of characters (Table 2) of Mirificarma and its 
presumed close relative Gelechia. Further material of Mirificarma and related genera would be 



GELECHIID MOTHS 



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monticolella 

interuptella 

flavonigrella 

burdonella 

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mulinella 



Fig. 2 Cladogram derived from the character matrix for the species of Mirificarma. Numbers refer to 
characters listed in Table 2. Black squares (with numbers below) denote presumed apomorphies, open 
squares (with numbers above) denote presumed convergences, black circles (with numbers below) 
denote presumed character-reversals. 

required to resolve fully the relationships within Mirificarma. Autapomorphies of individual 
species are not included except where they are presumed to involve character-reversal or 
convergence. The females of four species are unknown. Intermediate states are scored as 
apomorphies, with the exception of those discussed in the additional comments on characters 39 
and 45. Apparent convergences occur in characters 9, 13, 14, 21, 22, 26, 34, 35, 40, 42-45. 



GELECHIID MOTHS 



Polarities of characters have been estimated by comparison with the out-group (Gelechia and 
other Gelechiinae), but this has not always been possible and therefore the polarities of some 
character states are dubious. 

From the analysis of the characters examined (Figs 1 , 2) montivaga has the least apomorphies 
of the genus, since it is plesiomorphic in characters 1-3, whereas these are apomorphic in all 
other species of Mirificarma. These other species can be divided into two groups, the macu- 
latella-group and the interuptella-group. The interuptella-group is defined by the apomorphic 
states of characters 4-7 and 14-16, although the last three characters are reversed in some 
species (see Figs 1,2). The maculatella-group is defined by the apomorphy of character 37 and 
the reversal of 8. The division of the two species-groups is corroborated by their host-plant 
relationships (see p. 5). 

Within the maculatella-group, the relationships oirhodoptera are not fully resolved, hence the 
trichotomy in Fig. 2. In order to resolve the relationships within the interuptella-group, reversal 
of characters 14-16, 20-22, 24 and 25 has been presumed. 

The 'Remarks' on each species that follow (p. 19 onwards) reflect the primary concern of this 
paper, namely species diagnosis and therefore stress similarities and differences between species 
but do not necessarily indicate their relationships. 

Table 2 Characters used in the construction of a cladogram for Mirificarma. 



Character 



State 



Plesiomorphic 



Apomorphic 



1 Male filament 

2 Ductusbursae 

3 Aedeagus 

4 Vinculum 



5 Uncus 



6 Tegumen lateral margins 



7 Membrane between papillae 
anales and eighth 
abdominal segment of 
female 

8 Hind edge of vinculum 



9 Hind wing veins Rs and MI 
10 Sacculus 



very weakly developed 

extremely long 

bulbous in basal half 

without sclerites extending 
from saccus, although 
sometimes with short faint 
sclerites extending from 
anterior margin of 
vinculum (sclerites of 
montivaga are more similar 
to this than to the 
apomorphic state) 

large, slightly narrower 
than tegumen 



with pair of small rounded 
processes (often weak in 
montivaga) 

with two invaginations 



broadly projecting but 
without distinctly 
demarcated median 
projection 

separate 

broad, at least basally 
[this character has four 
apomorphic states - 
see also 20, 26, 33] 



well developed 

comparatively short 

uniformly slender 

with pair of sclerites or 
single sclerite extending 
from saccus towards hind 
edge of vinculum 



small, usually considerably 
narrower than tegumen 
(small but narrowing 
evenly from tegumen 
in monticolella) 

comparatively even 



with single invagination 



with distinctly demarcated 

median projection, usually 

comparatively narrow, or 

hind edge a narrow 

projection 
on common stalk 
moderately slender, straight 

or slightly curved, with 

one or more small 

irregular projections at apex 



10 



L. M. PITKIN 



Table 2 - Continued 



Character 



State 



Plesiomorphic 



Apomorphic 



11 Median projection of 
hind edge of vinculum 

12 Projection near apex 
of aedeagus 



13 Pair of membraneous 
sacs between apophyses 
anteriores and antrum 

14 Antrum 

15 Saccus 

16 Aedeagus/tegumen 
length ratio 

17 Apophyses anteriores 

18 Female eighth 
abdominal segment 

19 Filament extent 



20 Sacculus 



21 Uncus 



22 Filament 



23 Patch of sclerotization 
at posterior margin of 
female seventh 
abdominal segment 

24 Gnathos 

25 Uncus 

26 Sacculus 



27 Aedeagus apex 

28 Apophyses anteriores 

29 Female eighth sternite 



30 Filament 



not sharply rectangular 



relatively broad [this 
character has three 
apomorphic states - 
see also 32, 37] 

absent 



comparatively short 
short or moderately long 
low (at most 1-3) 

long (at least 0-6 mm) 
slightly sclerotized 



far beyond hind edge 
of vinculum 

broad, at least basally 
[this character has four 
apomorphic states - 
see also 10, 26, 33] 

not or scarcely 
constricted at base 

gently curved, slightly 

helical or straight 

[this character has 

two apomorphic states - 

see also 30] 
faint, diffuse, merging 

with median longitudinal 

band or absent 

a large hook, or complex 
broad or not narrowing 

from base towards apex 
not stouter apically [this 

character has four 

apomorphic states - 

see also 10, 20, 33] 
not cylindrical 
rod-like 
without strongly contrasting 

areas of sclerotization 

narrowing evenly towards 
apex [this character has 
two apomorphic states - 
see also 22] 



sharply rectangular, 
scarcely emarginate, 
membraneous 

slight, very narrow, 
distinctly sclerotized 



present 



extremely long 
extremely long 
high (1-7-1 -8) 

short (less than 0-6 mm) 
well sclerotized (either 

uniformly, or with patches 

of very strong sclerotization) 
at most a comparatively 

short distance beyond hind 

edge of vinculum 
slender, moderately curved, 

with even apex 



constricted at base (more so 
in constricta than in 
lentiginosella) 

with apical half kinked, 
basal half straight 



well defined 



small simple hook-shape 
small and narrowing from 

base towards apex 
stouter towards apex (club- 
shaped or slender and 
spatulate) 

almost cylindrical 

lobe-like 

strongly sclerotized 
laterally, contrasting with 
weakly sclerotized median area 

with basal half 
dorsoventrally expanded 
and laterally compressed 



GELECHIID MOTHS 



11 



31 Posterior margin of 
female seventh 
abdominal segment 

32 Projection near apex 
of aedeagus 



33 Sacculus 



34 Signum 

35 Signum 



36 Filament-supporting 
sclerites 

37 Projection near apex 
of aedeagus 



38 Fore wing pattern 



39 Median longitudinal 
band of sclerotization 
of female seventh 
abdominal segment 

40 Antrum 

41 Fore wing pattern 



42 Fore wing pattern 



43 Gnathos 



44 Saccus 

45 Female frenulum 



with, at most, diffuse 
band of scales 

small, sclerotized, evenly 
tapering [this character 
has three apomorphic 
states - see also 12, 37] 

broad, at least basally 
[this character has four 
apomorphic states - 
see also 10, 20, 26] 



present 

with very many small 
spines or without spines 

not twisted 

small, evenly tapering 
[this character has 
three apomorphic states 
-see also 12, 32] 

small spots or uniform 
[this character has 
three apomorphic states - 
see also 41, 42] 

if present, not or slightly 
constricted 



curved or constricted 

towards apex 
small spots or uniform 

[this character has 

three apomorphic states 

see also 38, 42] 
small spots or uniform 

[this character has three 

apomorphic states - 

see also 38, 41] 
curved 



broad, at least basally 
three setae 



with distinct, dense band 
of scales 

weakly sclerotized, rounded 



evenly slender, straight and 
smooth (the particularly 
slender sacculus of 
mulinella and ulicinella 
represents a further 
development - 33x) 

absent 

with large spines (small 
spines sometimes also 
present) 

twisted (weakly twisted in 
flavella) 

large, angular, hooked, 
narrowing to a point 



zig-zag yellowish markings 



strongly constricted 



straight and not constricted 

towards apex 
large spots 



median longitudinal stripe 
(broken or unbroken) 



almost straight (short in 
ulicinella, long in 
fasciata) 

relatively slender 

usually two setae 



Additional comments on the characters listed in Table 2 

5 M. ocellinella and fasciata are intermediate, more similar to the plesiomorphic than to the 
apomorphic state. 

8 M. montivaga is classed as intermediate since the hind edge of the vinculum sometimes 

approaches the apomorphic state. 

9 This character is not very reliable since both states are found in some species, although one 

state is always greatly predominant. 

16 Aedeagus/tegumen length ratio is intermediate in lentiginosella (1-4-1-5). 
18 This character is difficult to observe (since the sclerotization is more obvious in well-stained 

specimens) but trends of difference between species can be observed. M. montivaga has the 



12 L. M. PITKIN 

least sclerotization. M. lentiginosella and cytisella have larger areas of sclerotization than in 
species with the plesiomorphic state but smaller areas or weaker sclerotization than in the 
other species with the apomorphic state. 

19 The species with the longest filaments of the apomorphic state, in relation to tegumen length, 
are marked as intermediate. 

21 M. interuptella is intermediate between the two states, with the base of the uncus constricted, 

usually slightly, in a minority of specimens. 

22 M. interuptella is marked as intermediate since the filament is kinked throughout its length. 
26 The saccus offlavella is occasionally stouter apically but is usually more uniformly stout than 

in the apomorphic state. 
35 It is impossible to assess this character in denotata and the other species in which the signum is 

absent, or in eburnella which has a barely discernible signum. 
37 The aedeagus projection of montivaga (plesiomorphic state) is always small in relation to 

aedeagus width. M. rhodoptera and eburnella are intermediate with an aedeagus projection 

of the derived shape but small. M. scissella is intermediate with a more rounded projection 

than the derived state. 

39 The derived state is not to be confused with the slight constriction of the band of 

sclerotization in the several species marked as intermediate and not scored on the 
cladogram. 

40 M. denotata is classed as intermediate since its antrum narrows towards the apex more than 

that oiflavella and eburnella. It is not appropriate to score this character in species with an 
extremely short antrum. 

42 In the species marked as intermediate both states are present in different specimens. 

45 Too few specimens have been examined to be certain of the character state of every species, 
since this character is sometimes very variable intraspecifically. As this character is 
unstable, only the species which usually have two setae are scored as apomorphic. Species 
with an equal occurrence of two and three setae or with a tendency towards three setae are 
classed as intermediate and are not scored on the cladogram. This character seems to show a 
trend towards a relationship between mulinella, ulicinella and cabezella, which have the 
apomorphic state, although this is also shared by cytisella. 

The structure of the filament of the male genitalia 

In his original description of Mirificarma, Gozmany referred to the filament as 'a very long 
filamental prong' accompanying the aedeagus. It is orientated longitudinally and is near to the 
aedeagus in the undissected genitalia, but it has no physical connection with the aedeagus. The 
aedeagus lies between the ventral surface and the dorsal membrane of the vinculum, whereas 
the filament is more dorsal, lying between the dorsal membrane of the vinculum and the 
tegumen. Saltier (1960: 41) compared the filament of Mirificarma with processes of the transtilla 
in Bryotropha Heinemann and Filatima Busck, but these structures are not homologous with the 
filament. 

The filament is very weakly developed in montivaga, in which it is merely a slight sclerotized 
swelling where the supporting sclerites meet. In all other Mirificarma species it is a long, 
well-sclerotized tube, usually cylindrical, although narrowing gradually, posteriorly, and usually 
very narrow at the posterior end; usually gently curved or, if very long, sometimes slightly 
helical. Ocasionally the filament is straight, as in ocellinella, or modified in shape, for example 
with a strong kink near the apex in lentiginosella and with a projecting lobe in constricta and 
ulicinella. The filament varies in length between species, being shortest in relation to tegumen 
length in flavonigrella and ulicinella. In the latter, the filament does not usually reach the 
anterior of the tegumen and extends posteriorly only as far as the hind edge of the vinculum. The 
filament is longest in relation to tegumen length in pallidipulchra, in which it extends very far 
beyond the anterior of the tegumen and extends posteriorly beyond the uncus. 

The filament is open anteriorly and has a small opening near the posterior end. This opening is 
almost terminal in flavella and ocellinella whereas it is situated further from the posterior in 



GELECHIID MOTHS 13 

maculatella and cabezella. In scissella a second tiny posterior opening is present. In ocellinella 
and denotata a tiny projection occurs from the posterior opening of the filament. In maculatella a 
similar projection is situated more posteriorly. The filament surface of all species appears 
smooth except for occasional minute spines. 

The filament arises from a pair of lateral sclerotized strips, referred to here as filament- 
supporting sclerites. These extend from the base of the sacculus and, to a less extent, the valva, 
and meet anteriorly, at which point they fuse to form the filament. The filament is attached only 
at its anterior, and the length of the sclerites thus varies with the anterior extent of the filament. 
In many species the filament-supporting sclerites appear to be a smooth continuation of the 
sacculus. The sclerites are usually parallel-sided over much of their length, at least in the species 
in which they are long, but they are widened, often considerably and irregularly, towards the 
sacculus and valva. This expanded area of the sclerite is often faintly rugose and may represent 
an area of muscle attachment. This feature is not confined to Mirificarma since similarly placed, 
although even fainter, rugose patches are present in Gelechia nigra. The filament-supporting 
sclerites of Mirificarma are widened anteriorly, where they meet. 

The filament and supporting sclerites are covered ventrally by a membraneous sac which is 
attached to the sclerites and the anterior end of the filament. The sclerites are spiralled round 
each other in some species and in these the membrane is similarly twisted. A membraneous sac is 
usually also present dorsally, from the tegumen. These membranes are flimsy and the dorsal 
membrane in particular tends to disintegrate during the preparation of genitalia slides. The 
dorsal membrane is relatively well developed in cabezella and mulinella, for example, but it is 
most obvious in ulicinella. 

The function of the filament is unknown. There is some correlation between its length and that 
of the female apophyses anteriores. There appears to be no correlation between the length of the 
filament and that of the aedeagus such as the correlation discussed between the saccus and the 
aedeagus (see p. 15). 

MIRIFICARMA Gozmany 

Helina Guenee, 1849: 411. Type-species: Carcina flammella Hiibner, [1825], by monotypy. [Junior 

homonym of Helina Robineau-Desvoidy, 1830 (Diptera).] 
Mirificarma Gozmany, 1955: 308, 309 [keys], 313. Type-species: Tinea maculatella Hiibner, 1796, by 

original designation. 

O", $ (4-5) 5-0-11-0 mm. Head without frontal modifications. Ocellus present. Proboscis well developed; 
squamose, particularly at base. Maxillary palpus with four segments. Labial palpus recurved, first segment 
much shorter than second; third segment approximately same length as second or, rarely, slightly shorter; 
second segment without brush below or, rarely, with slight to moderate brush. Antenna without pecten on 
scape. Metascutum with paired group of narrow hair-like scales. Fore wing sometimes with zig-zag pattern 
of yellowish markings; sometimes with large or small dark spot across fold and another at end of cell; 
sometimes with dark, median, longitudinal, broken or unbroken stripe; a few species without any of these 
patterns. Fore wing with veins R 4 and R 5 on long common stalk (Fig. 3); distance R\-R2 slightly greater 
than, to four times R^-R^', M^ and Cui separate. Hind wing with veins Rs and MI on common stalk or 
separate (Figs 3,4). Frenulum of female with three or sometimes two long setae. 

GENITALIA cf Eighth tergite and sternite separated into free flaps. Eightn tergite with pair of coremata 
consisting of dense brushes of long, thin, hair-like scales, inserted laterally at anterior of tergite and 
attached to ventral membrane of tergite. Dorsal membrane of sternite with coremata consisting of two 
groups of large, rounded, often inflated, grape-like scales covering most of sternite. Genitalia withdrawn 
inside eighth segment. Uncus rounded, often large. Gnathos well-sclerotized, usually simple hook. 
Gnathos base with small membraneous area densely covered with microtrichia. Tegumen with or without 
lateral pair of rounded processes. Anterior margin of tegumen with deep V- or occasionally U-shaped 
emargination. Valva long, usually reaching uncus, slender, simple, slightly swollen at rounded apex. 
Sacculus clearly separated from valva. Long pair of sclerotized strips present, extending from base of valva 
and sacculus and meeting anteriorly. Long sclerotized tube (filament) present, arising from where pair of 
sclerotized strips join; extending posteriorly. In montivaga, filament extremely small; merely slight 
swelling of sclerotization. Hind edge of vinculum usually with medially emarginate median projection. 
Vinculum extends far posteriorly, hind edge reaching anterior of gnathos in a few species. Vinculum with 



14 



L. M. PITKIN 





Figs 3, 4 Wing venation of Mirificarma species, d* . 3, M. maculatella (Hiibner). 4, hind wing of M. 
interuptella (Hubner). 

or without pair of narrow sclerites or, exceptionally, single narrow sclerite, extending from saccus towards 
hind edge of vinculum. Saccus ranging from very slight projection to extremely long. Aedeagus usually 
cylindrical, slender; occasionally inflated basally; without cornuti. Projection near apex of aedeagus 
ranging from scarcely discernible to large, hook-like. Anterior part of ductus ejaculatorius often with 
sclerotized lamina. 

GENITALIA. $. Dorsal and ventral abdominal surfaces sometimes form broad, median, longitudinal 
sclerotized band. Posterior margin of seventh segment sometimes with more strongly sclerotized patches. 
Anterior margin of seventh sternite usually unmodified but in one species (pallidipulchrd) with pair of 
sclerites and membraneous sac. Papilla analis usually longer than broad. Invagination of membrane 
between eighth tergite and papillae anales usually present, immediately opposite invagination, sometimes 
present, of membrane between sternite and papillae anales. Eighth segment with predominantly lateral 
areas of sclerotization; occasionally almost completely sclerotized. Apophysis anterior usually rod-like, 
occasionally short , broad lobe ; length 0-2-1 -3mm; apophysis posterior two to six times length of apophysis 
anterior in each species. Sclerotized antrum present, usually long, nearly as long as apophysis anterior to 
much longer; occasionally extremely short. Ductus bursae often much shorter than corpus bursae, 
sometimes longer, exceptionally six times length of corpus bursae. Ductus seminalis arises from posterior 
part of ductus bursae, usually very near antrum. Corpus bursae with minute spines on inner surface 



; GELECHIID MOTHS 15 

sometimes sparse, sometimes extending into ductus bursae. Corpus bursae with or without signum. 
Signum, if present, usually oval or round, usually spinose; spines, if present, directed inside corpus bursae; 
signum sometimes curved inwards, never hook-like. 

REMARKS. The fore wing venation of Mirificarma has been checked in nine specimens and is 
similar to that of many Gelechiidae but appears to differ from that of many Gelechia species. In 
Mirificarma the fore wing veins A/ 3 and Cu\ are separate whereas these are frequently on a 
common stalk in Gelechia. 

The coremata of the male eighth abdominal segment of Mirificarma are readily lost during the 
preparation of genitalia slides although the scale bases can still be seen. The membraneous area 
of the base of the gnathos is much smaller in Mirificarma than in Gelechia and the Gnorimosche- 
mini in both of which it is an extensive sac. This structure is seen in some genera outside the 
Gelechiinae, for example Dichomeris Hiibner and Acompsia. The strongly sclerotized hook- 
shaped gnathos usual in Mirificarma occurs in various Gelechiidae but not in Gelechia. 
Mirificarma is distinguished by the presence of the filament of the male genitalia (see also the 
discussions on pp. 5, 6). 

The narrow sclerites of the vinculum which extend from the saccus towards the hind edge of 
the vinculum in the interuptella-group of Mirificarma are not usually present in the Gelechiidae, 
although they occur in Psoricoptera gibbosella which may be related to Gelechia and Mirificarma 
(see p. 6). In the maculate '//a-group the sclerotized anterior margin of the vinculum is 
occasionally continued towards the median across the posterior of the saccus as a pair of very 
faint short sclerites. Similar, though longer and very distinct, sclerites occur in the Gnorimosche- 
mini. These sclerites, unlike those of the interuptella-group, do not appear to arise from the 
saccus itself. The vinculum in Mirificarma is longest in interuptella, sometimes almost reaching 
the gnathos base of this species. 

The aedeagus is usually more slender than in Gelechia. There appears to be some correlation 
between the length of the saccus and the aedeagus in Mirificarma. In relation to the tegumen 
length, these structures are both exceptionally long in ocellinella and fas data; in lentiginosella 
they are slightly shorter, although still longer than in the remaining Mirificarma species. The 
saccus is shortest in rhodoptera and in this species the aedeagus is also the shortest. In the species 
with the longest saccus the aedeagus exceeds 1-5 times the length of the tegumen and the 
sclerotized saccus may serve to support the aedeagus where it extends beyond the tegumen 
anteriorly. The aedeagus is attached to the anterior of the saccus by retractor muscles in the 
representative of the Gelechiidae examined by Kuznetsov & Stekol'nikov (1978: 131) and there 
is a similarly attached muscle in Mirificarma fasciata. 

Sattler (1976: 93) mentions an apparent correlation between the length of the ductus 
ejaculatorius of the male and the ductus bursae of the female in the species of Ornativalva 
Gozmany, currently in the Gelechiini, in which these structures are exceptionally long. 
Likewise, in Mirificarma, the ductus ejaculatorius and the ductus bursae are both considerably 
longer in montivaga than in the rest of the genus. There is no obvious correlation in the other 
Mirificarma species, in which both structures are shorter. 

Mirificarma species with slight sclerotization of the female eighth sternite often have 
membraneous areas between the apophyses anteriores and the antrum, although the apophyses 
anteriores and the lateral margins of the antrum are usually continued into the sternite as narrow 
sclerites, reaching the sclerotized areas of the sternite. In montivaga there is little sclerotization 
of the sternite other than the continuation of the apophyses anteriores. The longitudinal band of 
the female abdomen in some species of Mirificarma is faint and is best observed in well-stained, 
freshly prepared specimens. I have included this feature in the species diagnoses only when it is 
characteristic. The band is also present in the male but is less useful as a character for species 
diagnosis. 

The sclerotized antrum of Mirificarma is usually long, whereas it is usually very short if present 
in Gelechia. Most Mirificarma species have a rounded, spinose signum. Only montivaga has a 
signum of the type common in Gelechia, diamond-shaped with a pair of serrated-edged ridges. 



16 L. M. PITKIN 

BIOLOGY. The known host-plants of Mirificarma are all Leguminosae and are discussed under 
each species and in the section on host-plant relationships. Although most of the species of 
Mirificarma are not known to have any economic significance, eburnella has been reported as an 
agricultural pest in the U.S.A., where it damages clover crops (see p. 28). 

The egg stage and site of egg deposition on the host-plant are unknown. The larva makes a 
spinning amongst the leaves of the host-plant or feeds in the shoots. In plants with small leaves, 
such as Ulex, the larva usually feeds in the flowers. Where details are known, pupation usually 
takes place in a slight cocoon amongst fallen leaves on the ground, although eburnella is 
recorded as pupating between the leaves on the host-plant. There are few records of the number 
of generations per year; two species are reputed to have only one generation but two are 
recorded for eburnella. Species may breed continuously where climatic conditions permit as is 
suggested by the wide range of dates on which ocellinella has been collected. 

DISTRIBUTION. Mirificarma species are distributed in the Palaearctic region between 55 N and 
30 S, extending east to c.45. One species, eburnella, also occurs in the U.S.A., where it has 
been introduced. 

Key to the species of Mirificarma 

Males 

1 Fore wing with large dark spots (Figs 9,10) 2 

- Fore wing with small dark spots or different markings 3 

2 Fore wing with large dark spot across fold narrowing gradually from middle towards costa; 

spot at end of cell comparatively distinct from patch at costa (Fig. 10). Portion of aedeagus 

apical to hook large (Fig. 65) maculatella (p. 23) 

Fore wing with large dark spot across fold narrowing sharply from middle towards costa; spot 
at end of cell merging with patch at costa (Fig. 9). Portion of aedeagus apical to hook small 
(Fig. 64) denotata (p. 22) 

3 Filament scarcely discernible (Fig. 60) montivaga (p. 18) 

Filament well developed (Figs 61, 62, 64-82) 4 

4 Uncus large, occasionally constricted at base but otherwise only slightly narrower than 

tegumen (Figs 61, 62, 66-72). Tegumen with pair of rounded lateral processes as in Fig. 31, or 

if absent, saccus extends far beyond tegumen anteriorly (Figs 70-72) 5 

- Uncus small, and usually considerably narrower than tegumen; tegumen without rounded 

lateral processes; saccus extends a comparatively short distance beyond tegumen anteriorly 

(as in Figs 76, 79-82) 13 

5 Tegumen without rounded lateral processes; saccus extends far beyond tegumen anteriorly 

(Figs 70-72) 6 

- Tegumen with pair of rounded lateral processes as in Fig. 31; saccus extends at most a short 

distance beyond tegumen anteriorly (Figs 61 , 62, 66-69) 8 

6 Filament strongly kinked near apex (Fig. 72) lentiginosella (p. 32) 

Filament straight (Figs 70, 71) 7 

7 Saccus of uniform width; basal half of gnathos increasingly very broad then narrowing sharply 

to hook-shaped apical half (Figs 46, 71) ocellinella (p. 29) 

Saccus spatulate, apex approximately twice width of base; gnathos slightly curved, basal half 
unmodified (Figs 47, 70) fasciata (p. 31) 

8 Filament not extending beyond apex of valva (Figs 61 , 68, 69) 9 

- Filament extends beyond apex of valva (Figs 62, 66, 67) 11 

9 Fore wing with zig-zag pattern of yellowish markings (Figs 11-14). Hind wing with veins Rs and 

M j separate , as in Fig. 3 10 

Fore wing without yellowish zig-zag markings. Hind wing with veins Rs and MI on common 
stalk, as in Fig. 4 scissella (p. 20) 

10 Sacculus with rounded, moderately broad apex (Fig. 68) fiavella (p. 26) 

Sacculus with pointed, moderately slender apex (Fig. 69) eburnella (p. 27) 

11 Fore wing without zig-zag pattern of yellowish markings. Saccus extremely short and very 

broad (Figs 35, 36, 62) rhodoptera(p.20) 

- Fore wing with zig-zag pattern of yellowish markings (Figs 11,12). Saccus moderately short and 

broad (Figs 39, 40, 66, 67) 12 



GELECHIID MOTHS 17 

12 Filament extends far beyond apex of valva posteriorly, and very far beyond tegumen anteriorly 

(Fig. 66) pallidipukhra (p. 24) 

- Filament extends short distance beyond apex of valva posteriorly, and moderately beyond 

tegumen anteriorly (Fig. 67) afiavella (p. 25) 

13 Sacculus uniformly slender (Figs 73, 80-82) 14 

- Sacculus broad, or broader apically than basally, club-shaped or spatulate; otherwise very 

small, hook-shaped (Figs 57, 59, 74-79) 17 

14 Uncus distinctly constricted at base; filament helical in apical half; sacculus gently S-curved 

(Fig. 73) constricta (p. 33) 

Uncus scarcely constricted at base; filament almost straight or curved dorsoventrally, not 
helical ; sacculus almost straight (Figs 80-82) 15 

15 Gnathos almost straight; sacculus does not reach gnathos arms (Fig. 81) ulicinella (p. 42) 

- Gnathos distinctly curved ; sacculus reaches or extends beyond gnathos arms (Figs 80 , 82) 16 

16 Filament stout, particularly basally, laterally compressed apically; median projection of hind 

edge of vinculum high (Fig. 82) mulinella (p. 43) 

- Filament very slender; median projection of hind edge of vinculum low (Fig. 80) cabezetta (p. 41) 

17 Sacculus stoutly club-shaped (Figs 57, 76, 78, 79) 18 

- Sacculus slender and spatulate (Fig. 77) or small and hook-shaped (Figs 74, 75) 20 

18 Gnathos extremely short ; saccus very slender; filament extends to hind edge of vinculum or just 

beyond (Figs 49, 76) monticolella (p. 37) 

- Gnathos developed normally; saccus comparatively broad; filament does not reach hind edge 

of vinculum (Figs 51, 52, 78, 79) 19 

19 Sacculus scarcely extends beyond median projection of hind edge of vinculum (Fig. 79) 

burdonella (p. 40) 

- Sacculus extends far beyond median projection of hind edge of vinculum (Fig. 59) 

fiavonigrella (p. 39) 

20 Gnathos very large, without median spine; sacculus long, slender, spatulate (Fig. 77) 

interuptella (p. 38) 

- Gnathos small, with median, sometimes minute, spine; sacculus small, hook-shaped (Figs 74, 

75) cytisella (p. 34) 

Females 

Note. The females ofscissella, monticolella and flavonigrella are unknown; constricta is also omitted since 

the female genitalia are unknown. 

1 Fore wing with large dark spots (Figs 9, 10) 2 

Fore wing with small dark spots or different markings 3 

2 Fore wing with large dark spot across fold narrowing gradually from middle towards costa; spot 

at end of cell comparatively distinct from patch at costa (Fig. 10). Signum present; antrum 
curved towards anterior, without indented anterior (Fig. 89) maculatella (p. 23) 

- Fore wing with large dark spot across fold narrowing sharply from middle towards costa ; spot at 

end of cell merging with patch at costa (Fig. 9). Signum absent or scarcely discernible; 
antrum straight , with indented anterior (Fig. 88) denotata (p. 22) 

3 Antrum same length as apophysis anterior or shorter (Figs 85-87 , 90-92 , 98) 4 

Antrum extends beyond apophysis anterior (Figs 94, 96-103) 10 

4 Antrum extremely short (Figs 85 , 98) 5 

- Antrum comparatively long (Figs 86, 87, 90-92) 6 

5 Signum a narrow ridge with faint surround (Fig. 98). Ductus bursae slightly longer than corpus 

bursae or of similar length cytisella (p. 34) 

- Signum elongate-oval to diamond-shaped with serrated edge (Fig. 85). Ductus bursae approx- 

imately four to six times length of corpus bursae montivaga (p. 18) 

6 Antrum almost straight, narrowing evenly towards anterior (Figs 92, 93). Fore wing with 

zig-zag pattern of yellowish markings (Figs 13,14) 7 

- Antrum curved or constricted towards anterior (Figs 86, 90, 91); if not curved and only very 

slightly constricted (Fig . 87) , forewing without zig-zag pattern of yellowish markings 8 

7 Antrum with rounded or scarcely indented anterior; abdomen with distinct, longitudinal, 

median band (Figs 93, 109) eburnella (p. 27) 

- Antrum with strongly indented anterior (Fig. 92); abdomen with, at most, very faint band 

ttavella (p. 26) 

8 Abdomen with pair of sclerites and membraneous sac at anterior margin of seventh segment 

(Fig. 107) pallidipulchra(p. 24) 



18 L. M. PITKIN 

Abdomen with unmodified anterior margin of seventh segment 9 

9 Signum with approximately three or four large spines projecting from one side, otherwise 

smooth; seventh abdominal segment comparatively broad (Figs 91, 108) attavella(p. 25) 

Signum covered with numerous spines of various sizes; seventh abdominal segment narrow 
(Figs 86, 87, 105,106) rhodoptera (p. 20) 

10 Antrum very long, coiled or strongly curved (Figs 94, 96, 97) 11 

Antrum moderately long, not coiled or strongly curved (Figs 98-103) 13 

11 Apophyses anteriores parallel, long; pair of membraneous sacs from sternite, between 

apophyses anteriores and antrum (Figs 94, 96) 12 

Apophyses anteriores diverging, short; without sacs between apophyses anteriores and antrum 
(Fig. 97) lentiginosella(p.32) 

12 Apophysis posterior three to four times length of apophysis anterior. Unsclerotized median 

area of eighth sternite at least as wide as each lateral sclerotized area (Fig. 94) ocellinella (p. 29) 
Apophysis posterior twice length of apophysis anterior. Unsclerotized median area of eighth 
sternite narrower than each lateral sclerotized area (Fig . 96) fasciata (p . 3 1 ) 

13 Apophysis anterior lobe-shaped; eighth sternite strongly sclerotized laterally, contrasting with 

weak sclerotization of median area (Figs 99, 100) 14 

Apophysis anterior rod-like (Figs 93, 101-103); eighth sternite slightly more sclerotized 
laterally than medially 15 

14 Signum present; median area of eighth sternite without horizontal striations (Fig. 99) 

interuptella (p. 38) 
Signum not discernible; median area of sternite with horizontal striations (Fig. 100) 

burdonella (p. 40) 

15 Antrum straight , narrowing evenly towards anterior ; abdomen with median longitudinal band , 

very distinct in seventh segment (Figs 93, 109) eburnella (p. 27) 

Antrum curved and constricted towards anterior (Figs 101-103); seventh abdominal segment 
without median longitudinal band 16 

16 Base of apophysis anterior with separate rounded lobe towards antrum (Fig. 101) cabezella (p. 41) 
Base of apophysis anterior without separate lobe (Figs 102, 103) 17 

17 Eighth tergite with pair of curved sclerotized lobes overlapping median longitudinal area 

(Fig. 102) ulidnella (p. 42) 

Eighth tergite without lobes (Fig. 103) nwlinclhi (p. 43) 

The montivaga-group 

Characters as described under montivaga. 

Mirificarma montivaga (Walsingham) comb. n. 
(Figs 1,2, 6, 32, 33, 60, 85, 104) 

Gelechia montivaga Walsingham, 1904: 221. LECTOTYPE cf, ALGERIA (BMNH), here designated 

[examined]. 
Gelechia pulverosella Zerny, 1936: 137, pi. 2, fig. 44. LECTOTYPE cf , MOROCCO (NM), here designated 

[examined]. Syn. n. 

O" , 7-5-8-5 mm. $ , 7-0-7-5 mm. Head cream. Labial palpus cream mottled with brown on outer surface. 
Thorax, tegula and fore wing (Fig. 6) cream to light brown mottled with brown. Hind wing with veins Rs 
and M\ separate as in Fig. 3. 

GENITALIA cf (Figs 32, 33, 60). Uncus large, only slightly narrower than tegumen. Gnathos a moderately 
large simple hook. Tegumen with pair of rounded lateral processes, sometimes very slight. Actual margin 
of tegumen slightly less emarginate than sclerotized margin anteriorly; sclerotized margin distinctly 
X-shaped medially. Sacculus moderately broad. Filament scarcely discernible; filament-supporting scler- 
ites not twisted around each other. Hind edge of vinculum a broad projection, scarcely emarginate 
medially. Vinculum short, with pair of very short faint sclerites converging from anterior edge. Saccus 
narrowing slightly towards rounded apex, extending short to moderate distance beyond tegumen anterior- 
ly. Aedeagus same length as tegumen plus uncus or slightly shorter. Aedeagus with bulbous basal half, 
small to moderate projection near apex, and long ductus ejaculatorius. 

GENITALIA $ (Figs 85, 104). Posterior margin of seventh abdominal segment with faint dorsal patch of 
sclerotization and pair of narrow longitudinal stripes ventrally. Very flimsy, broad, rounded invagination 



GELECHHD MOTHS 19 

of membrane between eighth tergite and papillae anales, directly opposite less flimsy, long, narrow 
invagination of membrane between sternite and papillae anales. Eighth segment with very weak sclejotiza- 
tion, longitudinally wrinkled, minutely spined anteriorly. Apophysis anterior rod-like, length 0-8-0-9 mm 
(2); apophysis posterior approximately three times length of apophysis anterior. Antrum slightly indented 
at anterior and extremely short. Ductus bursae gently coiled, narrow; coiled extent approximately four to 
six times length of oval or round corpus bursae. Signum very large, elongate oval to diamond shape, with 
deep longitudinal indentation edged with pair of serrated-edged projecting ridges. 

REMARKS. The frenulum was examined in the four females and consists of three setae in three of 
these and two setae in one, on both wings. The X-shaped anterior margin of the tegumen is 
particularly distinct in montivaga, although it occurs to a less degree in constricta and is very 
weak in several other Mirificarma species. 

M. montivaga is the only species of the genus in which the fore wing has no pattern other than 
the mottling of the ground colour, although the markings are extremely small in monticolella, 
lentiginosella and a few specimens of ocellinella. The ground colour is much darker in 
lentiginosella. M. montivaga differs from the rest of the genus in genitalic characters including 
the scarcely discernible male filament which is merely represented by a slight swelling between 
the supporting sclerites, the shape of the signum and the extremely long ductus bursae of the 
female. The long ductus bursae approaches that of rhodoptera, but this species has a much 
longer antrum than montivaga. 

M. montivaga was described from 3 cf , 3 9 from Algeria: El Kantara, all of which I have 
examined. I designate the specimen bearing Walsingham's number 96484 as the lectotype. This 
is referred to as the 'Type cf" on the specimen label and in the original description in which the 
date of capture is given as 3. v. 1903; however, 1 cf , 1 9 > bearing Walsingham's paratype labels, 
are labelled 9. v. 1903. 

G. pulverosella was described from 8 cf , 5 9 from Morocco, of which I have examined 3 cf , 
1 9- Of these 1 cf , 1 $ bear the labels Type cf' and Type 9' respectively and I designate the 
male as the lectotype. The 5 cf , 4 9 syntypes which I have not examined are from Tachdirt, 
middle and end vii (Zerny) . 

BIOLOGY. Host-plant unknown. Moths have been found from May to July. 
DISTRIBUTION. Morocco and Algeria. 

MATERIAL EXAMINED (including 6 cf , 3 9 genitalia preparations) 

Lectotype cf (montivaga), Algeria: [Constantine province,] El Kantara, 3. v. 1903 (Walsingham) (genita- 
lia slide no. 7124). Lectotype cf (pulverosella), Morocco: Haul ('Grosser') Atlas, c. 17 km SE. of Asm, in 
highest Iminene Valley, Tachdirt, 2200-2900 m, ll-19.vii. 1933 (Zerny) (genitalia slide no. 11223; NM). 

Morocco: 1 cf (pulverosella paralectotype), Haut Atlas, c. 70 km SW. of Marrakech, Goundafa area, 
3 km above Kasbah Goundafa, at junction of Agoundi Valley in Fis Valley, 'n Zala, above Ijjoukak, 
1500-1800 m, 21-29.vi.1933 (Zerny) (NM); 1 cf, Haul Atlas, Oukaimedene, 2500-2900 m, vii (coll. 
Burmann, Innsbruck); 1 cf, 1 9 (pulverosella paralectotypes), Haut Atlas, Tachdirt, 2200-2900 m, 
ll-19.vii.1933 (Zerny) (NM); 1 cf, Moyen Atlas, Val d'Ifrane, 1600-1700 m, vi (coll. Burmann, 
Innsbruck). Algeria: 2 cf , 3 9 (montivaga paralectotypes), El Kantara, 3, 9. v. 1903 (Walsingham). 

The maculatella-group 

Cf, 9- Fre wing sometimes with zig-zag pattern of yellowish markings; without median longitudinal 
stripe. Hind wing with veins Rs and M l separate or on common stalk. 

GENITALIA cf . Uncus large, only slightly narrower than tegumen. Gnathos a large simple hook. Tegumen 
with pair of rounded lateral processes. Filament-supporting sclerites usually crossed or spiralled round 
each other. Hind edge of vinculum a low, broad, sclerotized projection, usually with broad emargination, 
but without distinctly demarcated median projection. Vinculum without sclerites which extend from saccus 
and approach hind edge of vinculum. Projection near aedeagus apex usually large, angular and narrowing 
to point. 

GENITALIA 9- Invagination of flimsy membrane between eighth tergite and papillae anales present, 
immediately opposite invagination of slightly less flimsy membrane between sternite and papillae anales. 
Eighth segment weakly sclerotized. Apophysis anterior rod-like, long. Signum, if well developed, with 
large spines; small spines sometimes also present. 



20 L. M. PITKIN 

REMARKS. The filament-supporting sclerites of the male genitalia are more strongly spiralled 
round each other in the species in which they are long, particularly maculatella and rhodoptera. 
Although the vinculum does not have sclerites extending from the saccus and approaching the 
hind edge of the vinculum, in a few species the anterior margin of the vinculum is continued 
towards the median as faint, narrow, short sclerites on either side of the saccus. This is seen in 
rhodoptera and pallidipulchra and, even more faintly, in one specimen of aflavella; in scissella 
the anterior margin of the vinculum is sometimes continuous across the saccus posterior as a 
narrow sclerite. The sclerotization of the eighth segment is uniformly weak or limited to smaller 
areas than in most of the interuptella-group, but is always greater than in montivaga. 

BIOLOGY. Host-plants: Trifolieae (other than Ononis); Loteae; Coronilleae; Galegeae; Vicieae. 

Mirificarma scissella (Chretien) comb. n. 

(Figs 1,2, 7, 34, 61) 
Gelechia scissella Chretien, 1915: 319. LECTOTYPE cf , ALGERIA (MNHN), here designated [examined]. 

Cf , 7-0-7-5 mm. Head whitish cream. Labial palpus cream, mottled with brown at base. Thorax and tegula 
cream mottled with brown. Fore wing (Fig. 7) cream to pale brown, mottled with brown; slightly darker at 
extreme base and at costa in four-fifths. Dark brown spot across fold at one-third with trace at dorsal 
margin. Small dark brown spot at end of cell. Hind wing with veins Rs and MI on common stalk as in Fig. 4. 

GENITALIA cf (Figs 34, 61). Actual margin of tegumen less emarginate than sclerotized margin anteriorly. 
Sacculus broad at base, slightly wrinkled at narrower apex. Filament very slightly curved, slender, not 
reaching apex of valva posteriorly, extending short to moderate distance, beyond tegumen anteriorly. 
Filament-supporting sclerites not or scarcely twisted. Hind edge of vinculum with shallow U- or V-shaped 
emargination of sclerotization. Saccus usually moderately broad, scarcely narrower towards rounded 
apex, extending short distance beyond tegumen anteriorly. Aedeagus approximately same length as 
tegumen plus uncus, slender; apical projection moderately large, not distinctly pointed. 

GENITALIA $. Unknown. 

REMARKS. This species bears a superficial resemblance to rhodoptera, constricta and cabezella, 
although the head is paler and the spot in the fold of the fore wing scarcely extends to the dorsal 
margin; however, the only specimens known of scissella are in poor condition externally. It 
differs from rhodoptera in the hind wing venation, and the shorter filament and longer saccus of 
the male genitalia. It differs from constricta and cabezella in some genitalic characters, including 
the presence of a pair of rounded processes on the male tegumen, since these last two species are 
in the interuptella-group. 

M. scissella was described from an unspecified number of specimens of which I designate as 
lectotype the single type-specimen examined. The coating of debris on this specimen renders it 
almost unrecognizable externally. One of the rounded processes of the tegumen is absent in the 
lectotype. 

BIOLOGY. Host-plant unknown. Moths have been found in April and May. 
DISTRIBUTION. Algeria. 

MATERIAL EXAMINED (including 2 cf genitalia preparations) 

Lectotype cf , Algeria: [Constantine province,] Biskra, 7. v. 1907 (genitalia slide no. 43, LMP; MNHN). 
Algeria: 1 cf , M'zab ('Uzab') country, Oued Nsa ('Nza'), iv; 1 cf , Hamman-es-Salahine, iv. 

Mirificarma rhodoptera (Mann) 

(Figs 1, 2, 8, 35, 36, 62, 63, 86, 87, 105, 106) 

Gelechia rhodoptera Mann, 1866: 353, pi. 1 (B), fig. 10. Holotype cf, RUMANIA ('Turkey') (NM) 
[examined]. 

Cf, 5-0-7-5 mm, $,5-5-7-0 mm. (Typical form, cf , 6-5-7-5. $, 6-0-7-0. Small form, cf , 5-0-6-5 mm. <j>, 
5-5-6-0 mm.) Both forms. Head mid brown. Labial palpus mottled cream and brown; darker on outer 
surface. Thorax and tegula mottled mid brown. Fore wing (Fig. 8) mottled light brown, very faintly 



GELECHIID MOTHS 21 

pink-tinged, and dark brown; darker at base and in apical third; ochreous tinge near base. Small cream 
patch at costa at two-thirds extending diffusely to dorsal margin. Narrow, transverse dark brown spot 
across fold at one-third extending to dorsal margin. Smaller dark brown spot at end of cell, constricted 
medially, occasionally bisected. Both spots with ochreous cream surround. Very small diffuse dark brown 
spot in fold at one-fifth. Hind wing with veins Rs and M\ separate as in Fig. 3. 

GENITALIA cf (Figs 35, 36, 62, 63). Actual margin of tegumen less emarginate than sclerotized margin 
anteriorly. Sacculus short, moderately broad, wrinkled. Filament slightly helical, slender, extending 
posteriorly moderately beyond apex of valva, and very far beyond tegumen anteriorly. Hind edge of 
vinculum with broad, shallow U- or V-shaped emargination of sclerotization. Saccus very broad, usually 
with very shallow emargination at apex; extremely short, never reaching anterior margin of tegumen. 
Aedeagus approximately three-quarters length of tegumen plus uncus, basal half very slightly swollen; 
apical projection small. 

GENITALIA $ (Figs 86, 87, 105, 106). Posterior margin of seventh abdominal segment with area of weak 
sclerotization merging with progressively faint median longitudinal band. Seventh abdominal segment 
comparatively narrow, anterior margin unmodified. Invagination of membrane between eighth sternite 
and papillae anales indistinct, extremely long and narrow. Eighth sternite with lateral pair of weakly 
sclerotized strips. Tergite and rest of sternite very weakly sclerotized, with minute spines mainly on sternite 
extending into posterior of antrum. Apophysis anterior length in typical form 1-2-1-3 mm (3); small form 
0-6-0-7 mm (3); apophysis posterior twice length of apophysis anterior in each form. Antrum scarcely 
curved; anterior sometimes slightly indented, particularly in small form. Antrum of typical form very 
broad posteriorly, constricted strongly to narrow anterior half; shorter than apophysis anterior. Antrum of 
small form moderately broad posteriorly, constricted comparatively slightly for short distance at anterior; 
almost as long as apophysis anterior. Ductus bursae approximately two to four times length of oval corpus 
bursae. Signum covered with large and small spines; signum of typical form large, elongate kidney-shape, 
strongly curved inwards; signum of small form comparatively small, oval to broad kidney-shape, scarcely 
curved inwards. 

REMARKS. One specimen has veins Rs and MI on a very short common stalk in one hind wing. 
The frenulum, examined in seven females, consists of three setae in three of these, two setae in 
one of these, and in three it consists of three setae on one wing and two on the other. 

The specimens examined from Lakonia in Greece are smaller, on average, than the typical 
form, particularly in the genitalia. The small form is typical externally except that the fore wing 
markings tend to be relatively slightly smaller. Apart from the size, the female genitalia show 
some consistent differences in structure from the typical form. The eighth sternite has slightly 
longer, narrower sclerotized strips than in the typical form, the narrow anterior part of the 
antrum is considerably shorter and the signum differs in shape. The saccus is variable in length 
but tends to be more pronounced in the small form. It is possible that the Lakonia specimens 
may prove to be a distinct species; however, the differences are comparatively slight for the 
genus. There is insufficient material to determine whether the Lakonia population represents a 
distinct subspecies oirhodoptera. The few specimens I have seen from other Greek localities are 
of the typical form. A difference in the genitalia of one sex within a species is also discussed for 
cytisella (see p. 34). 

M. rhodoptera bears a superficial resemblance to scissella, constricta and cabezella. It differs 
from all these in the hind wing venation, and the very long filament and very short saccus of the 
male genitalia, and in addition differs from constricta and cabezella in many genitalic characters 
since these two species are in the interuptella-group. 

BIOLOGY. Host-plant unknown. Moths of the typical form have been found in June and July, 
those of the small form in May and June. 

DISTRIBUTION. Rumania, Greece, Turkey, Lebanon. 

MATERIAL EXAMINED 

Typical form (including 4 cf , 4 $ genitalia preparations) 

Holotype c? , Rumania (Turkey'): Dobrogea ('Dobrudscha'), far outside Tulcea ('Tultscha'), middle 
vi.1865 (Mann) (genitalia slide no. 11225; NM). 

Greece: 1 9 , Parnassfosl Mts, vii (MNHU); 1 cf , 3 $, Pelop[6nnisos], near Kalavfrita], Zakhlorou, vi 



22 L. M. PITKIN 

(Coll. Klimesch, Linz). Turkey: 1 <j>, Mara ('Marasch'), vii (MNHU). Lebanon: 2 cf, N., Bcharre 

('Becharre'), 1400 m, vi, vii (NM). 

Small form (including 3 cf , 4 9 genitalia preparations) 

Greece: 1 $ , Lakonia, 7 km SW. of Monemvasia, v; 4 cf , 5 $ , Lakonia, 5 km S. of Monemvasia, v, vi; 
1 cf , Lakonia, Mt Taygetos, 1000 m, vi (all ZM). 

Mirificarma denotata sp. n. 

(Figs 1,2, 9, 37, 64, 88) 

Cf, 7-5-8-5 mm. 9> 7-0-8-0 mm. Head light or occasionally mid brown. Labial palpus first and third 
segments dark brown and cream; second segment predominantly cream. Thorax and tegula brown; tegula 
base usually darker. Fore wing (Fig. 9) mottled brown and light pink-grey, ochreous tinge near base; apical 
quarter to third predominantly dark brown, edged with diffuse, basal, transverse pink and cream streak 
pronounced at costa; moderately dark brown patches slightly basal to streak at costa and dorsal margin, 
and at base. Large transverse dark brown spot across fold at one-third extending to dorsal margin, 
narrowing sharply from middle towards costa; dark brown spot at end of cell merging with patch at costa, 
comparatively distinct from patch at dorsal margin. Both spots with narrow yellow-ochre surround. Hind 
wing with veins Rs and MI on common stalk as in Fig. 4. 

GENITALIA cf (Figs 37, 64). Actual margin of tegumen less emarginate than sclerotized margin anteriorly. 
Sacculus short, moderately broad, flattened, slightly wrinkled at apex. Filament slightly helical, slender, 
extending posteriorly short distance beyond valva, and well beyond tegumen anteriorly. Hind edge of 
vinculum with moderately shallow U- or V-shaped emargination of sclerotization. Saccus broad basally, 
narrow apically, extending slightly beyond tegumen anteriorly. Aedeagus approximately same length as 
tegumen plus uncus. Aedeagus portion apical to hook small; apical hook moderately large. 

GENITALIA 9 (Fig- 88). Posterior margin of seventh abdominal segment with area of weak sclerotization 
mainly on tergite. Invagination of membrane between eighth sternite and papillae anales indistinct, long 
narrow tube without sac. Flimsy membrane between eighth tergite and papillae anales invaginated to 
broad rounded pouch. Eighth sternite with lateral pair of narrow, longitudinal sclerotized strips. Tergite 
and rest of sternite very weakly sclerotized, with faint longitudinal wrinkles. Apophysis anterior length 
0-9-1-1 mm (2); apophysis posterior approximately three times length of apophysis anterior. Antrum 
almost straight, narrowing considerably towards indented anterior; approximately same length as apo- 
physis anterior. Antrum with minute spines mainly in posterior half. Ductus bursae approximately twice 
length of oval corpus bursae. Signum absent or scarcely discernible. 

REMARKS. One specimen has the veins Rs and MI just separate in one hind wing. The frenulum 
was examined in one female and consists of three setae. During the preparation of the male 
genitalia, a tiny narrow projection was seen near the apex of the filament of one specimen, 
similar to that visible in maculatella and ocellinella, although not obviously tubular as in the 
latter. 

M. denotata is very similar to maculatella in the fore wing pattern, but differs very slightly in 
the form of the two large fore wing spots. The hind wing venation of denotata usually differs from 
that of maculatella. The male genitalia are very similar to those of maculatella although the shape 
of the extreme apex of the aedeagus differs; the female genitalia differ from those of maculatella 
in characters including the shape of the antrum and the absence of a signum. 

BIOLOGY. Host-plant: Galegeae: Astragalus lusitanicus Lamarck (type-specimens bred by 
Walsingham). The moths emerged in late April and May. 

DISTRIBUTION. Morocco. 

MATERIAL EXAMINED (including 3 cf , 2 $ genitalia preparations) 

Holotype cf, Morocco: [SE. of Tanger], El Fendek ('Fondak'), larva 28. iv. 1902, on Astragalus 
lusitanicus, moth emerged 27. v. 1902 (Walsingham) (genitalia slide no. 22027). 

Paratypes. 4 cf , 2 9 , same data as holotype, moths emerged 22-30. v. 1902. 



GELECHIID MOTHS 23 

Mirificarma maculatella (Hiibner) 
(Figs 1-3, 10, 38, 65, 89) 

Tinea maculatella Hiibner, 1796: 60, pi. 24, fig. 162 [legends to figs 161 and 162 are transposed]. 

LECTOTYPE cf , EUROPE (NM), here designated [examined]. 
Gelechia maculatella (Hiibner) Stainton, 1865: 228, pi. 7, fig. 3. 
[Gelechia vicinella Douglas; Bruand d'Uzelle, 1858: 481; Disque, 1908: 65. Misidentifications.] 

Cf , 7-0-9-5 mm. $, 7-5-9-0 mm. Head mottled mid brown. Labial palpus mottled cream or light brown, 
and dark brown. Thorax and tegula mottled mid brown. Fore wing (Fig. 10) mottled brown and light 
pink-grey; ochreous tinge near base; apical quarter to third predominantly dark brown, edged with diffuse, 
basal, transverse pink and cream streak pronounced at costa; moderately dark brown patches slightly basal 
to streak at costa and dorsal margin, and at base. Large transverse dark brown spot across fold at one-third 
extending to dorsal margin, narrowing gradually from middle towards costa; dark brown spot at end of cell 
comparatively distinct from patch at costa, usually merging with patch at dorsal margin. Both spots with 
narrow ochreous surround. Hind wing with veins Rs and MI separate (Fig. 3). 

GENITALIA d" (Figs 38, 65). Actual margin of tegumen less emarginate than sclerotized margin anteriorly. 
Sacculus short, broad, slightly flattened, strongly wrinkled at apex and inner edge. Filament slightly 
helical, slender, extending posteriorly well beyond apex of valva, and anteriorly, beyond tegumen. Hind 
edge of vinculum with broad, moderately shallow U- or V-shaped emargination of sclerotization. Saccus 
broad basally, narrow apically, extending a short distance beyond tegumen anteriorly. Aedeagus slightly 
longer than tegumen plus uncus. Aedeagus portion apical to hook large; apical hook large. 

GENITALIA $ (Fig. 89). Posterior margin of seventh abdominal segment with small area of weak 
sclerotization mainly on tergite. Invagination of membrane between eighth sternite and papillae anales 
very long narrow tube ending in pronounced, large oval sac. Eighth sternite with lateral pair of broad, 
moderately short sclerotized strips. Tergite and rest of sternite very weakly sclerotized; tergite with faint 
longitudinal wrinkles. Apophysis anterior length 0-9-1-0 mm (6); apophysis posterior approximately twice 
length of apophysis anterior. Membraneous area between antrum and each apophysis anterior produced 
into slight lobe, covered in minute spines. Antrum curved and narrower towards anterior; anterior not 
indented; approximately 1-5 times to less than twice length of apophysis anterior. Antrum with minute 
spines mainly in posterior half and extending into median area of sternite. Ductus bursae shorter than or of 
similar length to round corpus bursae. Signum moderately large, round or oval; with spines, large around 
edge. 

REMARKS. In approximately 7 per cent of the specimens examined the veins Rs and M l are on a 
common stalk in one or both hind wings. The frenulum of five out of six females examined 
consists of three setae; in the remaining female it consists of three setae on one wing and two on 
the other. During the preparation of the male genitalia, a tiny narrow projection was seen near 
the apex of the filament, similar to those in denotata and ocellinella, although not obviously 
tubular as in the latter. 

The fore wing pattern of maculatella clearly distinguishes it from all other species except 
denotata to which it is very similar. For differences from denotata see p. 22. 

M. maculatella was described from an unspecified number of specimens. In the NM there is 
one specimen from Mazzola's collection, here designated as lectotype, already labelled 'lecto- 
type' by Sattler. 

BIOLOGY. Host-plants: Coronilleae: Coronilla varia L. (moths bred by Miihlig, W. Germany; 
Stainton, 1865: 228). C. emerus L. (Bruand d'Uzelle, 1858: 482, as Gelechia vicinella Douglas; 
also subsequent authors). 

The larva occurs in May and June feeding in two opposite leaflets spun together. It pupates in 
a slight cocoon on the ground (Stainton, 1865: 228). Stainton records only one generation per 
year. Prose (1957: 111) records the larva in July. Moths have been collected from June to 
August. 

DISTRIBUTION. France; Germany; Austria; Czechoslovakia; Yugoslavia; Turkey; Syria. 

Additional records. Switzerland (Miiller-Rutz, 1914: 489); Italy (Hartig, 1964: 27); Poland 
(Schille, 1931: 179); Hungary (Gozmany, 1958: 227); Albania (Rebel, 1931: 146); U.S.S.R.: 
Ukraine (Piskunov, 1981: 674). 



24 L. M. PITKIN 

MATERIAL EXAMINED (including 8 cf , 6 $ genitalia preparations) 

Lectotype cf , Europe (Mazzola) (genitalia slide no. 11221; NM). 

France: 1 cf , 1 $, Paris; 1 cf , Basses-Alpes, viii. Germany: 1 cf , ; 1 cf , 1 $, 'N. Germany'. Germany 
(West): 1 cf, 4 $, Hessen, vii; 1 cf, 2 $, Bayern. Germany (East): 8 cf, 11 $, Gera. Austria: 1 cf, 
Nieder-Osterreich, vii. Czechoslovakia: 1 cf, Bohemia (NM). Yugoslavia: 1 cf, Croatia (NM); 1 $, 
Dalmatia (NM); vi. Turkey: 1 cf , Konya, vii (NM); 1 cf , Amasya (MNHU). Syria: 1 $, NW., vi (NM). No 
locality data: 15 ex. 

Mirificarma pallidipulchra (Walsingham) comb. n. 
(Figs 1,2, 11,39,66,90, 107) 

Rhinosia pallidipulchra Walsingham, 1904: 269. LECTOTYPE cf, ALGERIA (BMNH), here designated 

[examined]. 
Rhinosia striolella Turati, 1924: 166, pi. 6, fig. 11. LECTOTYPE cf , LIBYA (BMNH), here designated 

[examined]. Syn. n. 

Cf , 6-5-8-5 mm. $, 6-5-8-0 mm. Head cream to moderately light brown. Labial palpus cream frequently 
tinged with brown ventrally. Thorax and tegula as head. Fore wing (Fig. 11) with alternating transverse 
indistinct zig-zag patches of cream and ochre or occasionally deep yellowish brown; darker markings 
usually narrow. Several very narrow yellowish or brown striations radiating from wing base towards apex. 
Fore wing markings mostly weakly or moderately, occasionally strongly, contrasting. Hind wing with veins 
Rs and Af t , separate as in Fig. 3. 

GENITALIA cf (Figs 39, 66). Actual margin of tegumen usually coincides with sclerotized margin anteriorly. 
Sacculus moderately slender in apical half; apex wrinkled. Filament slightly helical, slender, extending 
posteriorly well beyond valva, and very far beyond tegumen anteriorly. Hind edge of vinculum with 
V-shaped emargination of sclerotization. Saccus moderately broad, apex truncate or very slightly 
emarginate; short, not quite reaching anterior of tegumen. Aedeagus approximately same length as 
tegumen plus uncus; basal half very slightly swollen, with moderately large apical hook. 

GENITALIA $ (Figs 90, 107). Abdomen with patchy, broad, median, longitudinal band constricted at 
anterior margin of sixth sternite. Seventh abdominal sternite with pair of sclerites and membraneous sac at 
anterior margin. Invagination of membrane between eighth sternite and papillae anales long, narrow, 
funnel-shaped. Eighth sternite with lateral pair of weakly sclerotized, broad strips. Tergite and rest of 
sternite very weakly sclerotized. Apophysis anterior length 0-9-1-0 mm (6); apophysis posterior approx- 
imately three times length of apophysis anterior. Antrum curved or constricted towards anterior, slightly 
shorter than or of similar length to apophysis anterior. Antrum with a few minute spines around posterior; 
anterior sometimes very slightly indented. Ductus bursae shorter than oval or pear-shaped corpus bursae. 
Signum large, spinose, particularly around edge, strongly curved inwards. 

REMARKS. In approximately 7 per cent of the specimens examined the veins Rs and MI are on a 
short common stalk in one hind wing. The frenulum was examined in five females and consists of 
three setae. In the male genitalia the sclerotized anterior margin of the vinculum does not 
coincide with the actual margin of the vinculum, although both are often indistinct. This occurs 
in a few other Mirificarma species but in these one of the margins is much more distinct than the 
other, whereas in pallidipulchra both margins appear equally faint. The constriction of the 
female abdominal band of this species and aflavella is seen to a much less degree in some other 
species (montivaga, rhodoptera, flavella, ocellinella and fasciatd) . 

M. pallidipulchra closely resembles eburnella, aflavella and flavella in the fore wing pattern, 
although its darker markings tend to be narrower, but it is distinguished from these species by 
the very long filament of the male genitalia and the pair of sclerites with the membraneous sac of 
the female abdomen. This last character is unique within the genus. 

M. pallidipulchra was described from 23 specimens from Algeria: Hamman-es-Salahine, 
18. iv, and El Kantara. I have examined 9 cf , 9 $ from El Kantara and designate as lectotype the 
specimen bearing Walsingham's number 96479. This is referred to as the 'Type cf' on the 
specimen label and in the original description. 1 cf , 1 $ of the remaining syntypes which I have 
not examined are in the NM. 

R. striolella was described from 4 cf from Libya, all of which I have examined. The specimen I 



GELECHIID MOTHS 25 

designate as lectotype, which bears Turati's 'Typus' label, was already labelled 'lectotype' by 
Sattler. 

BIOLOGY. Host-plant unknown. According to Walsingham (1904: 270) the moths are always 
found among stems and root-crowns of Teucrium (polium L. ?) (Labiatae), but it seems very 
unlikely that this is the host-plant. Moths have been found in March to May and July. 

DISTRIBUTION. Algeria, Tunisia, Libya. 
A record of this species from Egypt (Rebel, 1914: 268) is a misidentification of eburnella. 

MATERIAL EXAMINED (including 9 d", 6 $ genitalia preparations) 

Lectotype cf (pallidipulchra), Algeria: [Constantine province,] El Kantara, 5. v. 1903 (Walsingham) 
(genitalia slide no. 22491). Lectotype cf (striolella) , Libya: Cyrenaica, Benghazi, 20. Hi. 1922 (Kriiger) 
(genitalia slide no. 22490). 

Algeria: 1 $, Oran, Aflou, vii; 1 cf, 1 $, Hassi Bahbah, v (NM); 8 cf, 9 $ (pallidipulchra 
paralectotypes), El Kantara, 24.iv, 4, 5, 11. v. 1903 (Walsingham); 1 $, Constantine, Khenchela, v ? 
Tunisia: 1 $, Tozeur, iv (MNHN); 1 $, Hammamet, v (NM). Libya: 1 cf, 4 $, Tripolitania, Gharyan 
('Garian'),700m,iii,iv;5 $, Tripolitania, Tarhunah, 200 m,iv; 1 cf , 1 $?, Tripolitania, Bam WalId('Beni 
Ulid'), 300 m, iv; 5 cf , 1 $, Tripolitania, Al Khums ('Horns'), 10 m, iv; 2 cf (striolella paralectotypes), 
Cyrenaica, Benghazi, 18, 20.iii.1922 (Kriiger); 1 cf (striolella paralectotype), Cyrenaica, Benghazi, 
'Merg', 9.iv (Kriiger). 

Mirificarma afiavella (Amsel) stat. n. 

(Figs 1,2, 12,40,67,91,108) 
Rhinosiaflavella aflavella Amsel, 1935: 275. LECTOTYPE cf , ISRAEL (LN), here designated [examined]. 

Cf, 7-5-9-5 mm. $, 7-5-9-5 mm. Head moderately light golden-brown. Labial palpus mostly cream 
dorsally, golden-brown ventrally. Thorax and tegula as head. Fore wing (Fig. 12) with alternating 
transverse zig-zag patches of light golden-brown and mid brown, weakly contrasting. Hind wing with veins 
Rs and MI usually on common stalk as in Fig. 4. 

GENITALIA cf (Figs 40, 67). Actual margin of tegumen slightly less emarginate than sclerotized margin 
anteriorly. Sacculus moderately broad, apex wrinkled. Filament gently curved; moderately stout in basal 
half, tapering apically; extending posteriorly short distance beyond valva, and moderately far beyond 
tegumen anteriorly. Hind edge of vinculum with V-shaped emargination of sclerotization. Saccus 
moderately broad, narrowing slightly towards rounded apex; short, approximately reaching anterior of 
tegumen. Aedeagus almost as long as tegumen plus uncus; with moderately large apical hook. 

GENITALIA $ (Figs 91, 108). Abdomen with diffuse, broad, median, longitudinal band, slightly more 
distinct at posterior margin of seventh segment, strongly constricted at anterior margin of sixth sternite. 
Seventh abdominal segment comparatively broad, anterior margin unmodified. Invagination of membrane 
between eighth sternite and papillae anales long, narrow, funnel-shaped. Eighth sternite with lateral pair 
of broad sclerotized strips. Tergite and rest of sternite very weakly sclerotized. Apophysis anterior length 
0-7-0-9 mm (5); apophysis posterior three times length of apophysis anterior. Antrum curved or 
constricted anteriorly, of similar length to apophysis anterior; with minute spines posteriorly, extending 
into median area of sternite. Antrum anterior sometimes very slightly indented. Ductus bursae one- 
quarter to slightly more than one-third length of elongate pear-shaped corpus bursae. Signum with 
approximately three or four large spines arising from sclerotized ridge on one side, otherwise smooth. 

REMARKS. In approximately 25 per cent of the specimens examined the veins Rs and MI are 
separate in one or both hind wings. The frenulum consists of three setae (five females 
examined). In the female genitalia the ductus bursae merges gradually with the corpus bursae 
and is distinguished only by its paler appearance in stained preparations. 

This species resembles pallidipulchra, flavella and eburnella in fore wing pattern. It is 
particularly close to flavella in this respect although the fore wing pattern is slightly less 
contrasted. M. aflavella differs from these three species in the male genitalia, in which the 
filament extends a short distance beyond the apex of the valva, and in the form of the signum of 
the female genitalia. 

M. aflavella was originally described as a subspecies of flavella, with which it is allopatric. 
However, I consider that the genitalic differences between them are sufficient to separate them 



26 L. M. PITKIN 

as distinct species; moreover, aflavella appears to have even more features in common with 
pallidipulchra than with flavella. 

M. aflavella was described from a series collected by Amsel in Palestine, 1930: Tabgha, 12.iii; 
Jordan, Salt, 7.iv, and Waldheim (c. 15 km E. of Haifa), 9.v (Amsel, 1935: 265). I have 
examined 8 cf , 4 $ , from Tabgha, all of which probably belong to the type-series although some 
lack Amsel's type-labels and are dated IS.iii instead of 12.iii. The specimen I designate as 
lectotype, which bears Amsel's label 'Typus', was already labelled 'lectotype' by Sattler. 

BIOLOGY. Host-plant unknown. Moths have been found from March to May. 
DISTRIBUTION. Greece (Rodhos), Turkey, Israel (including Jordan west bank territories). 

MATERIAL EXAMINED (including 4 cf , 5 $ genitalia preparations) 

Lectotype cf , Israel: ('Palastina'), 10 km N. of Tiberias, En Sheva ('Tabgha'), 12.iii.1930 (Amsel) 
(genitalia slide no. 17, LMP.; LN). 

Greece: 1 $, Rodhos ('Rhodes'), Miramare [Hotel], iv. Turkey: 1 $, Gavur Daglari ('Amanus'), 
'Schechle', v (NM); 1 cf , Gavur Daglari, Tarib', v (NM); 3 cf , 1 $ , Haleb, Shar Deresy ('Shar Devesy'). 
Israel: 1 cf, 3 $, ('Palestine'), Haifa, iii-v; 3 cf, 3 $ (paralectotypes) , ('Palastina'), Tiberias, En Sheva 
('Tabgha'), iii.1930 (Amsel) (BMNH and LN); 4 cf , 1 $ (paralectotypes ?), Tiberias (including 3 cf , 1 $ , 
'See Genezareth'), En Sheva ('Tabgha'), iii, 15.iii.1930 (Amsel) (NM); 2 $, Palestine, plains of Jordan. 

Mirificarma ffavella (Duponchel) 

(Figs 1,2, 13,41,42,68,92) 

Acompsia flavella Duponchel, [1844]: 512, pi. 89, fig. 7. Type(s), FRANCE: Bondy Forest, end of vi 

(Begrand) [not traced]. 
Gelechia segetella Zeller, 1847: 847. LECTOTYPE cf, SICILY (BMNH), here designated [examined]. 

[Synonymized by Wocke, 1861: 115.] 

Cf , 7-0-9-0 mm. $ , 7-5-9-0 mm. Head yellowish. Labial palpus cream to yellowish, sometimes tinged with 
brown ventrally. Thorax yellowish, occasionally with faint brown median longitudinal stripe; tegula 
yellow-ochre to golden-brown. Fore wing (Fig. 13) with alternating transverse zig-zag patches of yellowish 
colour and deep yellow-tinged brown; moderately strongly contrasting. Hind wing with veins Rs and MI 
separate as in Fig. 3. 

GENITALIA cf (Figs 41, 42, 68). Actual margin of tegumen variable but usually slightly less emarginate than 
sclerotized margin anteriorly. Sacculus with rounded, moderately broad apex. Filament moderately stout 
at base, tapering and slightly curved towards apex; almost reaching apex of valva posteriorly, extending 
slightly or scarcely beyond tegumen anteriorly. Hind edge of vinculum with median U- or V-shaped 
emargination of sclerotization. Saccus broad to very broad, sometimes narrower at rounded apex, reaching 
or extending slightly beyond tegumen anteriorly. Aedeagus same length as tegumen plus uncus or slightly 
longer; with moderately large apical hook. 

GENITALIA $ (Fig. 92). Abdomen with at most faint median, longitudinal band. Eighth sternite and tergite 
very weakly sclerotized; sternite with faint lateral areas of sclerotization. Apophysis anterior length 
0-8-1-0 mm (8); apophysis posterior approximately three times length of apophysis anterior. Antrum 
almost straight, tapering, usually slightly, towards strongly indented anterior; slightly shorter than 
apophysis anterior; without minute spines. Ductus bursae slightly shorter to longer than oval or round 
corpus bursae. Signum small to medium-sized; spinose, particularly around edge. 

REMARKS. The fore wing pattern varies from predominantly yellowish to predominantly brown. 
The frenulum consists of three setae (five females examined). The sacculus of the male genitalia 
tends to be longer than that in eburnella or aflavella. The saccus is usually relatively broader than 
in eburnella although it is variable in both species. 

M. flavella resembles eburnella, aflavella and pallidipulchra in the fore wing pattern, 
particularly aflavella, although the fore wing pattern of flavella is usually more contrasted than in 
aflavella. The male genitalia of flavella differ from those of eburnella by the rounded apex of the 
sacculus, and from the other two species by the relatively short filament. The female genitalia 
differ in having a straight antrum together with an indented apex. 

M. flavella was described from an unspecified number of specimens from France: Bondy 



GELECHIID MOTHS 27 

Forest (near Paris). The type-specimens have not been traced and are not in the MNHN (Viette, 
pers. comm.). 

G. segetella was described from an unspecified number of specimens from Sicily: Siracusa. I 
have examined 1 Cf, 1 9 syntypes and designate as lectotype the male already labelled 
'lectotype' by Sattler. 

BIOLOGY. Host-plants: Trifolieae: Trifolium pratense L. (moth bred by Frey, France). T. minus 
Rehl. = procumbens G.G.' (the identity of this Trifolium species is not clear since minus is 
currently a synonym of T. dubium Sibthorp and procumbens is a synonym of T. campestre 
Schreber). Loteae: Lotus corniculatus L. 

These host-plants are referred to by Lhomme ([1948-1949] : 653) , who states that the larva has 
been found in May and June. Moths have been collected from April to July. 

G. segetella was described from moths collected on the edges of wheatfields and on 
chrysanthemum-like flowers (Compositae); however, there is no reason to suppose that the 
latter could be its host-plant. 

DISTRIBUTION. France, Corsica, Italy, Sardinia, Sicily, Greece, Crete, Cyprus, Algeria, Tunisia. 

Additional records. Spain (Agenjo, 1968: [4]); Belgium (Lhomme, [1948-1949]: 652); 
Greece: Attiki (Staudinger, 1871: 255). 

The following records should probably be referred to aflavella. Israel: Jerusalem (Caradja, 
1920: 112); 'Palestine'; Turkey ('Kleinasien') (Wocke, 1871: 300); Turkey ('Asia Minor'); Syria 
(Rebel, 1901: 158; Meess (in Spuler, 1910: 344); Meyrick, 1925: 142); Turkey ('Asia Minor') 
(Rebel, 1903:332). 

A record of the species from Bulgaria: Rilo (Rebel, 1903: 332) is based on a misidentification 
of Orophia ferrugella ([Denis & Schiffermiiller]) (Oecophoridae). A record oiflavella from 
Yugoslavia: Dalmatia (Rebel, 1903: 332) may also be a misidentification of O. ferrugella. 

MATERIAL EXAMINED (including 5 cf , 8 9 genitalia preparations) 
Lectotype cf (segetella), Sicily: Siracusa, v. (Zeller) (genitalia slide no. 7135). 
France: 1 $, ; 1 cf , Hie et Vilaine; 1 cf , Mayenne (MNHN); 1 cf, Calvados (MNHN); 1 $, Sarthe 

(MNHN); 1 cf , Essonne (MNHN); 2 cf , 5 $, Paris; 4 cf , 1 $, Alpes-Maritimes; vi, vii. Corsica: 7 cf , vi. 

Italy: 3 cf , 3 9 Toscana, vi, vii. Sardinia: 1 $, vi ? (NM). Sicily: 1 $ (segetella paralectotype), Siracusa, v. 

Crete: 2 $ , v, vi (NM). Cyprus: 7 cf , 8 $ , iv, v. Algeria: 2 cf , Algiers province (MNHN); 2 $ , Constantine 

province, vi. Tunisia: 1 $ . No locality data: 23 ex. 

Mirificarma eburnella ([Denis & Schiffermiiller]) comb. n. 
(Figs 1,2, 14,43,69,93, 109) 

Tinea eburnella [Denis & Schiffermiiller], 1775: 140. Syntypes, AUSTRIA: Wien area (lost). NEOTYPE cf , 

AUSTRIA (NM), here designated [examined]. 
Tinea formosella Hiibner, 1796: 62, pi. 23, fig. 160. Syntypes, SWITZERLAND: Genf ('Geneve') [not traced]. 

[Junior primary homonym of Tinea formosella [Denis & Schiffermiiller], 1775: 140 (Oecophoridae).] 
Carcina flammella Hiibner, [1825]: 410. [Objective replacement name for Tinea formosella Hiibner.] 

[Synonymized by Zeller, 1847: 848.] 
Gelechia rufeoformosella Bruand d'Uzelle, 1859: 652. [Objective replacement name for Tinea formosella 

Hiibner.] 
[Rhinosia palidipulchra Walsingham; Rebel, 1914: 268. Incorrect subsequent spelling of pallidipulchra 

Walsingham. Misidentification.] 
Mirificarma formosella (Hiibner) Capue, 1964: 17, figs 7, 8. 

Cf, 5-0-7-5 mm. 9> 5-5-7-5 mm. Head cream. Labial palpus cream, frequently tinged with brown 
ventrally. Thorax cream with faint, narrow, median, longitudinal, yellow-ochre stripe; tegula yellow- 
ochre. Fore wing (Fig. 14) with alternating transverse zig-zag patches of cream and yellowish to ochre, 
strongly contrasting. A few very narrow ochre striations radiating from wing base towards apex. Hind wing 
with veins Rs and MI separate as in Fig. 3. 

GENITALIA cf (Figs 43, 69). Actual margin of tegumen considerably less emarginate than sclerotized 
margin anteriorly. Sacculus with pointed, moderately slender apex. Filament moderately slender, almost 
straight, almost reaching apex of valva posteriorly, extending short to moderate distance beyond tegumen 



28 L. M. PITKIN 

anteriorly. Hind edge of vinculum with U- or V-shaped emargination of sclerotization. Saccus broad, 
slightly narrower at rounded apex, extending short distance beyond tegumen anteriorly. Aedeagus same 
length as tegumen plus uncus or slightly longer; apical hook usually moderately large. 

GENITALIA $ (Figs 93, 109). Abdomen with distinct broad, median, longitudinal band, most prominent in 
seventh segment. Eighth sternite and tergite very weakly sclerotized; sternite with faint lateral areas of 
sclerotization. Apophysis anterior length 0-6-0-9 mm (10); apophysis posterior three to four times length 
of apophysis anterior. Antrum almost straight, tapering, usually slightly, towards rounded or scarcely 
indented anterior; of similar length to apophysis anterior or slightly longer; without minute spines. Ductus 
bursae usually merging indistinguishably with elongate corpus bursae. Signum extremely small, sometimes 
apparently absent. 

REMARKS. In less than 2 per cent of the specimens examined the veins Rs and M l are on a short 
common stalk in one or both hind wings. The frenulum consists of three setae (five females 
examined). In the male genitalia, the uncus and tegumen posterior tend to be narrower than in 
flavella. The corpus bursae of the female genitalia is extremely flimsy and sometimes very small. 

M. eburnella closely resembles flavella, aflavella and pallidipulchra in the wing pattern, 
although it is usually slightly more contrasted in eburnella. It differs from these in the pointed 
apex of the sacculus together with the relatively short filament of the male genitalia, and the 
straight antrum without an indented apex in the female genitalia. 

M. eburnella was described from an unspecified number of specimens. Zeller (1847: 848) 
refers to two specimens of this species in the Schiffermuller collection, which is now lost. I 
designate a male neotype from the type-locality, Austria: Wien area. 

BIOLOGY. Host-plants: Trifolieae: Medicago saliva L., M. lupulina L. (Lhomme, [1948-1949]: 
652); M. polymorpha L. (Bur clover), Trifolium repens L. (variety - Ladino clover), T. hirtum 
Hall (Rose clover); Vicieae: Vicia americana Muhlenberg (Purple vetch) (California Depart- 
ment of Agriculture unpublished report); Coronilleae: Hippocrepis comosa L. (Lhomme, 
[1948-1949]: 652). 

The records of the California Department of Agriculture refer to the species in U.S.A.; the 
other records are European. I have seen a Walsingham specimen bred from 'Trifolium''; 
however, this name has been used broadly and may refer to Medicago. A record of the species on 
Populus (Salicaceae) (Hofmann, 1875: 118) is erroneous since it is based on a misidentification 
of the moth which was probably Schiffermuelleriaformosella [Denis & Schiffermuller] (Chretien 
in Lhomme, [1948-1949]: 652). 

The larva has been found in April and May, both in Europe (Lhomme, [1948-1949]: 652) and 
in U.S.A. (California Department of Agriculture unpublished report). This species has caused 
severe damage to clover in California, U.S.A., where the larva 'semiskeletonizes' leaves and 
lightly spins two leaves together, pupating inside the folded leaves (Anonymous, 1969: 69). 

Moths have been collected from March to July (also in August according to Piskunov, 1981: 
674); in May and June in U.S.A. Hartig (1964: 51) states that there are two generations in Italy, 
with the moths in June and August. 

DISTRIBUTION. M. eburnella occurs in western, central and southern Europe south of c. 50 N. 
and extends to North Africa, the Middle East and U.S.S.R.: Armeniya. It is also found in 
U.S.A.: California where it is presumed to have been introduced. 

According to the literature it has also been found in the following countries. Portugal 
(Zerkowitz, 1946: 132); Belgium (Lhomme, [1948-1949]: 652); Netherlands (Lempke, 1976: 
27); Sardinia (Hartig & Amsel, 1951: 86); Poland (Schille, 1931: 199); Czechoslovakia (Hruby, 
1964: 293); Bulgaria (Rebel, 1903: 332); Crete (specimen in NM, Sattler, pers. comm.); 
U.S.S.R.: European part (Piskunov, 1981: 674). 

MATERIAL EXAMINED (including 7 cf , 10 $ genitalia preparations) 

Neotype cf (eburnella), Austria: Nieder-Osterreich ('Austr. inf.'), [near Wien,] Klosterneuburg, 
2.vi.l918 ('Freiberg') genitalia slide no. 11230; NM). 

Spain: 6 cf , Granada; 1 cf , Mallorca; iv-vi. France: 1 cf , 1 $ , Sarthe (MNHN); 1 cf , SW. ; vii. Germany: 
3 cf . Switzerland: 2 cf , 3 $ , Valais. Corsica: 19 cf , 7 $, v-vii. Italy: 2 cf , 1 9, Toscana; 1 cf , Lazio; 1 cf , 
Campania; iv-vi. Sicily: 5 cf , 4 $, 5 ex., iii-v. Austria: 1 cf , 2 $?, ; 1 Cf , Wien area (NM); vi. Yugoslavia: 



GELECHIID MOTHS 29 

1 Cf, 1 $, Slovenija;3 cf, 1 $, Croatia; 1 cf, 1 $, Dalmatia; 2cf, Bosna i Hercegovina; vi, vii. Hungary: 
4 cf , Rumania: 6 cf , Timi, vi (BMNH; NM); 1 cf , Caras-Severin, vi? (NM). Albania: 2 cf , vii. Greece: 
1 $, Levkas, vi (NM); 1 cf, Lakonia, iv (ZM); 1 cf, 1 $, Rodhos I., iv. Cyprus: 4 cT, 3 $, 1 ex., iv, v. 
Turkey: 1 $ , Bursa (NM); 1 cf , Toros Daglari. Malta: 1 cf , 1 $ , iv. Morocco: 2 cf , 1 $ , iv, v. Algeria: 7 cf , 
3 $, Oran province; 2 $, Algiers province (BMNH; MNHN); 13 cf, 6 $, Constantine province; v, vi. 
Tunisia: 3 cf, 3 $, iv, v. Egypt: 2 cf , ; 1 $, Cairo area, iv (NM) (Rebel, 1914: 268, as Rhinosia 
palidipulchra Walsingham). Israel [including Jordan west bank territories]: 4 cf , 5 9, 'Palestine'; iv, v. 
Lebanon: 4 cf , v. Syria: 2 $, N.; 2 cf , 2 $, NW. U.S.S.R.: 1 $, Armeniya, vii (NM). U.S.A.: 6 cf , 4 $, 
California, v, vi. No locality data: 37 ex. 

The iJiterupte//a-group 

Cf, $ Fore wing without zig-zag pattern of yellowish markings; sometimes with median longitudinal 
stripe. Hind wing with veins Rs and MI on common stalk as in Fig. 4. 

GENITALIA cf . Uncus usually small or constricted at base; occasionally large. Gnathos a large or small 
simple hook or modified in shape. Tegumen without pair of lateral processes. Filament-supporting sclerites 
not crossed or spiralled round each other. Hind edge of vinculum a narrow projection or with distinctly 
demarcated, usually narrow, median projection. Vinculum with pair of sclerites or single sclerite extending 
from saccus and approaching hind edge of vinculum. 

GENITALIA $. One invagination of membrane between eighth segment and papillae anales present, 
comparatively sturdy, usually between tergite and papillae anales (between sternite and papillae anales in 
interuptelld) . Eighth segment usually moderately to strongly sclerotized. Apophysis anterior rod-like or 
very short, broad lobe; usually short. Signum, if present, with many small spines or without spines. 

REMARKS. In some species occasional specimens have the veins Rs and M\ separate in one or 
both hind wings. In mulinella the lateral edge of the male tegumen is sometimes wavy, producing 
slight projections but not the distinct rounded processes of the maculatella-group. The scler- 
otization of the female eighth segment is limited to small areas in fasciata and, particularly, 
ocellinella; the degree of sclerotization is greater in lentiginosella and cytisella but still slightly 
less than in the rest of the interuptella-group . The species in this group are more diverse in 
genitalia structure than those of the maculatella-group. 

BIOLOGY. Host-plants: Genisteae (cytisella also on Trifoliae: Ononis). 

Mirificarma ocellinella (Chretien) comb. n. 

(Figs 1, 2, 15, 16, 46, 71, 83, 94, 95) 

Gelechia ocellinella Chretien, 1915: 317. LECTOTYPE $, TUNISIA (MNHN), here designated [ex- 
amined]. 

Gelechia aurantiella Chretien, 1915: 317. LECTOTYPE , TUNISIA (MNHN), here designated [ex- 
amined]. Syn. n. 

Gelechia retamaeofoliella Dumont, 1931: 148. LECTOTYPE $, TUNISIA (MNHN), here designated 
[examined]. Syn. n. 

Cf , 9-0-11-0 mm. 9, 8-5-10-5 mm. Head cream to light brown. Labial palpus cream mottled with brown, 
sometimes dark brown, particularly on outer surface; second segment with slight to moderate brush below. 
Thorax and tegula cream to mid brown; tegula sometimes darker at base. Fore wing (Figs 15, 16) cream to 
light brown, sometimes mottled with darker brown; usually with narrow longitudinal dark brown stripes 
radiating to apical wing margin. Median stripe sometimes overlaid with broader median longitudinal band, 
occasionally very dark. Tiny spot sometimes present at one-third to half, on costal edge of cell, and 
occasionally at or near end of cell. 

GENITALIA cf (Figs 46, 71). Uncus large, two-thirds to nearly three-quarters width of tegumen. Gnathos 
basal half increasingly very broad then narrowing sharply to hook-shaped apical half. Actual margin of 
tegumen slightly less emarginate than sclerotized margin anteriorly. Sacculus very slightly S-curved, 
usually moderately slender, apex slightly wrinkled and with small irregular projections. Filament very 
moderately stout, almost straight; extending beyond apex of valva posteriorly, and very far beyond 
tegumen anteriorly. Tiny tube projecting from opening near filament apex. Hind edge of vinculum with 
sharply rectangular weakly sclerotized, scarcely emarginate, median projection. Vinculum short, with pair 



30 L. M. PITKIN 

of almost parallel sclerites. Saccus parallel-sided, moderately slender, apex not or scarcely wider than base; 
extremely long but not reaching anterior of filament. Aedeagus slightly less than twice length of tegumen 
plus uncus; apex with very narrow, distinctly sclerotized, slightly projecting structure. 

GENITALIA 9 (Figs 94, 95). Abdomen with at most faint median longitudinal band, scarcely more distinct at 
posterior margin of seventh segment. Invagination of membrane between eighth tergite and papillae 
anales funnel-shaped. Eighth sternite with lateral pair of relatively narrow longitudinal sclerotized strips; 
tergite and rest of sternite weakly sclerotized and faintly wrinkled longitudinally. Unsclerotized median 
area of sternite at least as wide as each sclerotized area. Apophyses anteriores parallel, rod-like, length 
1-0-1-3 mm (7); apophysis posterior three to four times length of apophysis anterior. Pair of membraneous 
sacs from sternite between apophyses anteriores and antrum, densely covered in minute spines. Antrum 
gently coiled, anterior not indented; extremely long, coiled length approximately twice length of apophysis 
anterior. Ductus bursae less than half length of oval or round corpus bursae. Signum strongly curved 
inwards, elongate-oval, distinctly covered with tiny spinules; with spiny-edged plate, usually medially 
indented, projecting obliquely inside bursa. 

REMARKS. The fore wing pattern of ocellinella varies by degrees from strongly contrasted 
markings to the comparatively uniform fore wing of the type-specimens of retamaeofoliella, 
which lacks stripes. The frenulum of six out of seven females examined consists of three setae; in 
the remaining female it consists of two setae. In the male genitalia the tiny tube near the apex of 
the filament was not visible in two specimens; however, it is easily lost during the preparation of 
the genitalia. The specimens examined show a wide but gradual range both in wing length and in 
the female sternite sclerotization. The lectotype of G. retamaeofoliella is the smallest specimen 
of ocellinella and has the shortest strips of sternite sclerotization. The signum of this specimen is 
more elongated than others examined and the projection of the signum has no median 
indentation. 

M. ocellinella is the largest species of the genus. It closely resembles fasciata in the male and 
female genitalia, but differs in the shape of the male gnathos and saccus, and to a less extent in 
the female sternite sclerotization and signum. 

M. ocellinella was described from an unspecified number of specimens from Tunisia. I 
designate the single type-specimen I have examined as the lectotype. The collector is likely to 
have been Chretien but may have been Oberthiir or Lucas (Chretien, 1915: 289). 

G. aurantiella was also described from an unspecified number of specimens from Tunisia. I 
have examined a single specimen labelled TYPE' [by Viette], 'Gelechia aurantiella'. It bears the 
locality label 'Mansour', not 'Gafsa' as cited in the original description; however, there is a 
locality of this name near Gafsa. Despite this discrepancy, I consider it to be a type-specimen 
and designate it as the lectotype. 

G. retamaeofoliella was described from a pair of specimens from Tunisia, both of which I have 
examined. They are labelled 'Coll. D. Lucas, 1952' and in Lucas's handwriting: 'Gelechia 
retamaeofoliella Dumont' on the female and 'Gelechia acupediella Frey' on the male. The 
specimens are conspecific and the label on the male is incorrect. According to Dr P. Viette (pers. 
comm.) the type-specimens, which are in the MNHN, belong to the Dumont Collection, not the 
Lucas Collection, and Lucas may have replaced their original labels. I designate the female as 
the lectotype. 

The type-specimens of retamaeofoliella represent the small, poorly-marked form of ocel- 
linella. 

BIOLOGY. Host-plant: Genisteae: Retama raetam (Forskal) Webb & Berthelot (R. 'retem Webb') 
(type-series of G. retamaeofoliella). The larva remains in a long silk tube at the base of the 
food-plant during the day and comes out to feed on the leafy branches at night (Dumont, 1931: 
149). It is fully grown at the beginning of March and the adults Dumont reared emerged in 
September and October. Moths have been found in January, March to May and September to 
November. A few females have been collected at light. 

DISTRIBUTION. Morocco, Algeria, Tunisia, Libya, Jordan. 

MATERIAL EXAMINED (including 7 d", 7 $ genitalia preparations) 
Lectotype $ (ocellinella), Tunisia: Tozeur, 31.x. 1904 (genitalia slide no. 28, LMP. ; MNHN). Lectotype 



GELECHIID MOTHS 31 

$ (aurantiella), Tunisia: [Gafsa], Mansour, 11. xi. 1908 (genitalia slide no. 29, LMP; MNHN). Lectotype $ 
(retamaeofoliella), Tunisia: Metlaoui, larva on Retama 'retem Webb', moth emerged 9.x 1921 (Dumonf) 
(genitalia slide no. 39, LMP; MNHN). 

Morocco: 1 cf , Agadir, 'Rocksin', xi (NM). Algeria: 1 cf , S. Oran, Aflou region, x; 3 cf , Lambese, x. 
Tunisia: 1 cf (retamaeofoliella paralectotype), Metlaoui, on Retama 'retem Webb' (MNHN); 1 $, 
Kasserine to Thelepte road, iv (MNHN); 1 cT, 1 $, 1 ex., Bu Hadmah ('Bou Hedma'), iii, x, (MNHN); 
6 Cf, 4 $, Maknassy, x, xi (MNHN). Libya: 1 $, W. ('occ.') Sirtica, Sawfajjin ('Sofeggin'), v; 1 cf , W. 
('occ.') Sirtica, Al Qaddahiyah ('Gheddahia'), ix. Jordan: 1 $, Jordan Valley, Zarqa ('Zerqa') R. Colony, 
c. 100 m below sea level, i. 

Mirificarma fasdata sp. n. 

(Figs 1,2, 17,47,70,96) 

Cf, $,8-5 mm. Head white mottled with grey. Labial palpus white mottled with grey; mainly grey on outer 
surface. Thorax mottled grey and white; tegula grey. Fore wing (Fig. 17) white and ochreous cream, 
mottled with dark brown scales mainly near wing margins; very faintly and diffusely forming longitudinal 
narrow stripes radiating from base towards apex. Brown median longitudinal band, very dark brown on 
costal side. Very narrow, broken, dark brown stripe along fold to dorsal margin. 

GENITALIA cf (Figs 47, 70). Uncus large, approximately two-thirds width of tegumen. Gnathos long and 
only slightly curved; basal half unmodified, extreme apex a very slight hook-shape. Actual margin of 
tegumen slightly less emarginate than sclerotized margin anteriorly. Sacculus straight, slender; extreme 
apex with small irregular projection. Filament straight, very moderately stout; reaching apex of valva 
posteriorly, extending far beyond tegumen anteriorly. Hind edge of vinculum with sharply rectangular, 
weakly sclerotized, scarcely emarginate median projection. Vinculum short, with pair of parallel sclerites. 
Saccus slender, parallel-sided except apex which is twice width of base; extremely long, extending beyond 
filament anteriorly. Aedeagus slightly less than twice length of tegumen plus uncus; apex with very narrow, 
distinctly sclerotized, slightly projecting structure. 

GENITALIA $ (Fig. 96). Posterior margin of seventh abdominal segment with area of weak dorsal 
sclerotization and pair of faint ventral patches merging with extremely faint median longitudinal band. 
Invagination of membrane between eighth tergite and papillae anales conical. Eighth sternite with lateral 
pair of broad longitudinal sclerotized strips; tergite and rest of sternite weakly sclerotized and very slightly 
wrinkled longitudinally. Less sclerotized median area of sternite narrower than each sclerotized area. 
Apophyses anteriores parallel, rod-like, length 0-7 mm (1); apophysis posterior twice length of apophysis 
anterior. Small pair of membraneous sacs from sternite between apophyses anteriores and antrum, densely 
covered in minute spines. Antrum coiled; anterior not indented; extremely long, actual length approx- 
imately three to four times length of apophysis anterior. Ductus bursae less than half length of oval corpus 
bursae. Signum moderately elongate-oval, with irregular surround and small inwards projection which is 
medially indented; covered with tiny spinules. 

REMARKS. The frenulum of the single female examined consists of two setae on one wing and 
three on the other. The longitudinal fore wing stripe of this species gives it a superficial 
resemblance to interuptella although the stripe is more diffuse than in the latter. However, it 
differs considerably in the genitalia, particularly in the shape of the male filament, saccus and 
sacculus, and in the female apophysis anterior and antrum. M. fasdata is also similar externally 
to specimens of ocellinella with a fore wing stripe and the two species appear to be closely 
related. It differs from ocellinella in the shape of the male gnathos and saccus and in the female, 
to a lesser degree, in the sternite sclerotization and the signum shape. 

BIOLOGY. Host-plant unknown. Moths have been found in December. 
DISTRIBUTION. Spain. 

MATERIAL EXAMINED (including 1 cf , 1 $ genitalia preparations) 

Holotype cf , Spain: [Malaga,] San Pedro de Alcantara, xii.1972 (Ffennelt) (genitalia slide no. 22083). 
Paratype. 1 $, same data as holotype. 



32 L. M. PITKIN 

Mirificarma lentiginosella (Zeller) 
(Figs 1,2, 18, 44, 72, 97) 

[Haemylis obscurella Hiibner; Treitschke, 1832: 240-241 (larva only). Misidentification.] 
Gelechia lentiginosella Zeller, 1839: 198; Stainton, 1865: 64, pi. 2, fig. 3. LECTOTYPE $, GERMANY 
(EAST) (BMNH), here designated [examined]. 

Cf , 6-5-8-5 mm. $ , 6-0-8-0 mm. Head mid to dark brown. Labial palpus dark brown with scattered cream 
scales; paler on dorsal or inner surface and at segment apices. Thorax and tegula mid to dark brown. Fore 
wing (Fig. 18) dark brown with scattered pinkish buff scales; small pinkish spot at costa, at two-thirds to 
three-quarters, sometimes extended to basal margin. Very small darker brown spots with ochreous 
surround; one in fold, one in cell, both approximately at one-third; one at end of cell. All spots indistinct. 

GENITALIA cf (Figs 44, 72). Uncus large, slightly constricted at base, at this point half to two-thirds width of 
tegumen, otherwise slightly narrower than tegumen. Gnathos a large simple hook. Actual margin of 
tegumen slightly less emarginate than sclerotized margin anteriorly. Sacculus moderately curved inwards, 
with evenly rounded apex; slender. Filament almost straight basally, without median lobe, with strong kink 
at apex, very moderately stout; extending posteriorly to hind edge of vinculum or slightly beyond; 
extending well beyond tegumen anteriorly. Hind edge of vinculum with low, weakly sclerotized median 
projection not emarginate ventrally. Vinculum with pair of sclerites, parallel or converging and undulating 
from saccus, almost reaching hind edge of vinculum. Saccus parallel-sided or broader at rounded apex, 
very long, extending far beyond tegumen anteriorly. Aedeagus approximately 1-5 times length of tegumen 
plus uncus, with moderately small apical projection. 

GENITALIA 9 (Fig- 97). Abdomen with at most faint, patchy, median longitudinal band scarcely more 
distinct at posterior margin of seventh segment. Invagination of membrane between eighth tergite and 
papillae anales long and narrow. Eighth segment not usually strongly sclerotized. Sternite with lateral pair 
of very broad, longitudinal sclerotized areas; median area of sternite less sclerotized, usually slightly, and 
narrower than each lateral area of sternite. Tergite with medially expanded sclerotized area at anterior and 
pair of tiny lateral patches at posterior. Apophyses anteriores diverging, rod-like, length 0-4-0-5 mm (8); 
apophysis posterior four to five times length of apophysis anterior; without sacs between apophyses 
anteriores and antrum. Antrum gently coiled or strongly curved, anterior not indented; extremely long, 
coiled extent two to four times length of apophysis anterior. Ductus bursae usually not more than half 
length of round corpus bursae. Signum small, oval, weakly sclerotized, slightly curved inwards, covered 
with tiny spinules. 

REMARKS. In six females examined, the frenulum consists of three setae, except for one female in 
which it consists of four on one wing. 

M. lentiginosella differs in the fore wing pattern of very small indistinct spots on a dark 
background, although this is very occasionally approached by extremely dark specimens of the 
externally variable mulinella. M. lentiginosella can be distinguished by the shape of the male 
filament, which is straight basally, kinked apically and without a lobe, and by the very long 
antrum together with the diverging apophyses anteriores in the female. This species appears to 
be close to constricta with which it has some common features of the uncus, filament and 
sacculus. However, the uncus and filament shape are similar in interuptella, although to a lesser 
degree. 

M. lentiginosella was described from a series of 25 specimens of which I designate as lectotype 
the single specimen I have examined, which was already labelled 'lectotype' by Sattler. The 
type-locality is neither stated in the original description nor on the lectotype label. The original 
description attributes the species name to Tischer, and I have seen a record, in Tischer's 
handwriting, of the species collected in Dresden (East Germany). It seems most likely that the 
specimens from Dresden referred to in Tischer's record were sent to Zeller and that Zeller based 
his description on these. 

BIOLOGY. Host-plants: Genisteae: Genista tinctoria L. (larvae and moths bred by Bankes and 
Ford, Great Britain; Stainton, 1865: 64); G. anglica L. (moths bred by Nielsen, Denmark; 
Sorhagen, 1886: 186); G. germanica L., G. sagittalis L. (Sorhagen, 1886: 186); Laburnum 
anagyroides Medicus (= Cytisus laburnum L.) (Miiller-Rutz, 1913-1914: 485). 

Records of Centaurium erythraea erythraea Rafn (= Erythraea centaurium Persoon) (Gen- 



GELECHIID MOTHS 33 

tianaceae) (Lhomme, [1946-1948]: 607, 'according to certain authors') and Salix repens L. ? 
(Salicaceae) (Sorhagen, 1886: 187) are extremely dubious as host-plants of this species. 
Sorhagen attributes his record to Hartm[ann] but I have not been able to trace the reference. 

The larva occurs in May and June, spinning together the young terminal shoots of the plant; it 
pupates in a cocoon amongst leaves on the ground (Stainton, 1865: 64), from June to July 
(Bradford, [1979]: 123). Moths have been collected from June to August (also in May according 
to Mariani, 1943:167). 

DISTRIBUTION. Great Britain, France, Denmark, Central Europe, Italy, Rumania. 

Additional records. Spain (Agenjo, 1968: [4]); Netherlands (Lempke, 1976: 26); Sweden 
(Krogerus et alii, 1971: 21); Poland (Schille, 1931: 176); Czechoslovakia (Nicked, 1908: 20); 
Yugoslavia (Mariani, 1943: 167); Hungary (Gozmany, 1958: 227); Turkey (Klimesch, 1961: 
648); U.S.S.R.: European part (Piskunov, 1981: 672); U.S.S.R.: Armeniya (Rebel, 1901: 14). 

MATERIAL EXAMINED (including 8 cf , 8 $ genitalia preparations) 

Lectotype $, [Germany (East): Dresden], larva on Genista tinctoria, ex viii (genitalia slide no. 22492). 

Great Britain (England): 67 ex., ; 2 ex., Worcester: 15 cf, 13 $, 3 ex., Dorset; 1 ex., New Forest; 
12 ex., Isle of Wight; 52 ex., Sussex; 8 ex., Kent; 7 ex., Essex; vii-viii. France: 1 cf, Basses Alpes; 
1 <j>, Alpes-Maritimes, viii. Denmark: 3 cf, 1 <j>, Jylland, vi (BMNH; ZM). Germany (West): 4 cf, 
1 $, Niedersachsen; 1 cf , Bayern, viii. Germany (East): 6 cf , 3 $, 1 ex., Gera. Switzerland: 1 cf , Zurich. 
Italy: 1 $ , Toscana, viii. Austria: 2 cf , 1 9 , Wien, viii (NM). Rumania: 1 cf , 1 $ , Cluj, viii (MINGA). No 
locality data: 34 ex. 

Mirificarma constricts sp. n. 

(Figs 1,2, 19,45,73) 

Cf , 5-5-6-0 mm. Head mid brown. Labial palpus mottled cream and brown. Thorax and tegula mid brown. 
Fore wing (Fig. 19) mottled mid brown, slightly darker at base. Dark brown spot across fold at one-third 
extending to dorsal margin. Small dark brown spot at end of cell. 

GENITALIA cf (Figs 45, 73). Uncus small, slightly more than half width of tegumen, strongly constricted at 
base. Gnathos large, only moderately curved; extreme apex a very slight hook-shape. Actual margin of 
tegumen slightly less emarginate than sclerotized margin anteriorly; sclerotized margin X-shaped medial- 
ly. Sacculus moderately S-curved; with evenly pointed apex, moderately slender. Filament basal half 
straight, very broad dorsoventrally and compressed laterally; median lobe projecting ventrally; apical half 
more slender, kinked; extending beyond hind edge of vinculum posteriorly, and moderately beyond 
tegumen anteriorly. Median projection of hind edge of vinculum with ventral V-shaped emargination; 
dorsally deeply U-emarginate. Vinculum with pair of sclerites converging from saccus, almost reaching 
hind edge of vinculum. Saccus moderately broad, parallel-sided or slightly bow-sided, with rounded apex; 
extending slightly beyond tegumen anteriorly. Aedeagus same length as tegumen plus uncus or slightly 
shorter; projection near apex slight. 

GENITALIA $. Unknown. 

REMARKS. M. constricta is very similar in fore wing pattern to cabezella and, to a less extent, 
scissella and rhodoptera but differs in the constricted uncus and the shape of the filament. M. 
constricta resembles lentiginosella in these male genitalic characters; however, the single known 
female of constricta lacks the abdomen and it is impossible to be certain of its relationships. 

BIOLOGY. Host-plant unknown. Moths have been found in August and October. 
DISTRIBUTION. Northern Morocco. 

MATERIAL EXAMINED (including 2 cf genitalia preparations) 

Holotype cf , Morocco: Tanger, 45 m, 30.x. 1934 (Querci) (genitalia slide no. 7477). 
Paratypes. Morocco: 1 cf , 1 $, Tanger, 45 m, 30. viii, 24.x. 1934 (Querci). 



34 L. M. PITKIN 

Mirificarma cytisella (Treitschke) 
(Figs 1, 2, 5, 20, 21, 48, 74, 75, 98, 110) 
Lita cytisella Treitschke, 1833: 99. 

Cf , 6-0-8-0 mm. $ , 6-0-7-5 mm. Head white to cream, eye socket edged with dark brown anteriorly. Labial 
palpus white to cream, with dark brown predominantly on outer surface of first segment and outer, basal 
two-thirds of second segment. Thorax and tegula white to cream with scattered brown scales. Fore wing 
(Figs 20, 21) white to cream, mottled, sometimes sparsely, with brown scales, usually slightly darker in 
apical fifth. Costa darker at base. Indistinct, narrow, transverse cream streak basal to apical fifth. Brown 
wedge-shaped spot, usually dark, across fold at one-third extending to dorsal margin. Smaller dark brown 
spot, sometimes bisected, at end of cell. Dark spots often with narrow ochreous surround. 

GENITALIA O" (Figs 48, 74, 75). Uncus small, narrowing progressively from tegumen, with tiny, pointed 
median projection at apex. Gnathos small, hook-shaped, with median spine on anterior surface. Actual 
margin of tegumen coincides with or slightly less emarginate than sclerotized margin anteriorly. Sacculus 
very small, hook-shaped. Filament slightly, sometimes irregularly, curved; moderately slender, gradually 
broader at base; almost reaching hind edge of vinculum or extending moderately short distance beyond, 
posteriorly; extending short distance beyond tegumen anteriorly. Hind edge of vinculum projecting 
slightly, medially; projection with truncate or irregular apex ventrally, slightly emarginate dorsally. 
Vinculum with pair of sclerites parallel or slightly diverging from saccus. Saccus almost parallel-sided, 
usually with truncate apex; almost reaching anterior of tegumen. Aedeagus of similar length to tegumen 
plus uncus; projection near apex small. 

GENITALIA $ (Figs 98, 110). Posterior margin of seventh abdominal tergite with well-defined crescent of 
sclerotization. Invagination of membrane between eighth tergite and papillae anales funnel-shaped. 
Eighth sternite sclerotized laterally; median area with sclerotization often weaker and patchy. Tergite very 
weakly sclerotized. Apophysis anterior rod-like, length 0-4 mm (7); apophysis posterior approximately 
three times length of apophysis anterior. Antrum extremely short. Ductus bursae usually of similar length 
to, or slightly longer than, pear-shaped or oval corpus bursae. Signum a narrow, inwards-projecting ridge, 
with faint surrounding area of sclerotization. 

REMARKS. The frenulum of five out of seven females examined consists of two setae; in the 
remaining two it consists of three setae on one wing and two on the other. 

Specimens from Portugal, the extreme west of the range, differ in fore wing pattern from 
specimens from France eastwards and were described as subspecies leonella. Examination of 
material from intermediate localities, particularly in Spain, is necessary to decide whether 
subspecific status is justified. 

M. cytisella has two forms of male genitalia. In the typical form of c. cytisella, the filament 
extends distinctly beyond the hind edge of the vinculum and is usually gently and evenly curved, 
or almost straight excluding the base. In other specimens of cytisella, including c. leonella, the 
filament scarcely extends beyond the hind edge of the vinculum and is usually more irregularly 
curved. The spine on the gnathos is usually shorter in the typical form than in the other 
specimens of c. cytisella, which I am referring to as the small form, or in leonella. The small form 
of c. cytisella, and leonella, have smaller male genitalia than the typical form of c. cytisella. In the 
female genitalia, the signum varies from small to large but without apparent correlation with the 
two forms of male genitalia. I have assigned the females to each form of c. cytisella on the basis of 
geographic association with the males. 

It is possible that the small form of c. cytisella could be a separate subspecies from the typical 
form, and might be more closely allied to leonella since the small form and leonella are similar in 
genitalia structure. However, the small form has a widely disjunct distribution, populations 
occurring both west and east of the typical form, thus it seems unsatisfactory to consider it a 
distinct subspecies. 

The fore wing spots of cytisella, other than in leonella, are usually more distinct than in the 
other species of Mirificarma, except maculatella and denotata which have larger spots. M. 
cytisella can be distinguished from the other species of the genus by the small hook-shaped 
sacculus, the spine of the gnathos, and the form of the signum. 



GELECHIID MOTHS 



35 



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1^ 



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T c 
E -E* J> 




J2 

~8J 
"x 

u 

.2 

I 
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u 



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O 






D 




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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 

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GELECHIID MOTHS 47 

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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, 
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Parts are published at irregular intervals as they become ready, each is complete in itself, 
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^v 

. /rt X* 

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World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.) 

1 J- 1LJ I__M 



Trustees of the British Museum (Natural History), 1984 



V 

^. . 







The Entomology series is produced under the general editorship of the 

Keeper of Entomology: Laurence A. Mound 

Assistant Editor: W. Gerald Tremewan 



ISBN 565 06001 5 
ISSN 0524-6431 

British Museum (Natural History) 
Cromwell Road 
London SW7 5BD 



Entomology series 
Vol48No2pp71-130 



Issued 23 February 1984 




Macronematine caddisflies of the genus 
Amphipsyche (Trichoptera: Hydropsychidae) 

N^P|, U\< 

P. C. Barnard 

Department of Entomology, British Museum (Natural History), Cromwell Road, London 
SW7 5BD 

Contents 

Synopsis 71 

Introduction 71 

Abbreviations of depositories 73 

Acknowledgements 73 

Taxonomic method 73 

Classification of the Macronematini 74 

Key to Old World genera of Macronematini 74 

Amphipsyche McLachlan 76 

Geographical distribution 78 

Biology 78 

Cladistic analysis 80 

Check-list of Amphipsyche species 85 

Key to species of Amphipsyche 85 

Thepro/wta-group 86 

The apicalis-group 94 

The meridiana-group 101 

References 127 

Index 130 

Synopsis 

In this revision of the genus Amphipsyche McLachlan 22 species are recognized, of which two are described 
as new. One new generic and ten new specific synonyms are established, and one species is transferred to 
Amphipsyche from Protomacronema Ulmer. Eight lectotypes are designated. Keys are given to the Old 
World genera of the tribe Macronematini and to the species of Amphipsyche. The classification of the 
species is based on a cladistic analysis, and some of the evolutionary and zoogeographical implications of 
the analysis are discussed. 

Introduction 

Amphipsyche McLachlan is an Old World genus of caddisflies having netspinning larvae that are 
frequently found in fast freshwater streams throughout the Afrotropical and Oriental regions. 
Some species have figured prominently in recent freshwater pollution and impoundment 
studies, and at least one species is a predator on larvae of Simulium Latreille (Diptera). 

Many of the recent ecological studies on tropical freshwater habitats are the result of the 
pressing need for knowledge of the effects of man's activities, the most obvious of which is direct 
pollution of the water by chemical or other agents. Although most Trichoptera are very sensitive 
to such pollutants and tend to disappear even at low levels of contamination, there is a selective 
response shown by different species of caddis. Resh & Unzicker (1975) have stressed that it is 
important to be able to identify the organisms at the specific level for this kind of study. Another 
important influence of man is the damming of rivers for hydroelectric or irrigation schemes. 
Although the ecology of such impounded water is usually studied, the effects on the regulated 
river itself are less well known and the few existing reports suggest that the natural watercourse 
may be altered for a considerable distance downstream of the impoundment. The release of 



Bull. Br. Mus. not. Hist. (Ent.) 48 (2): 71-130 Issued 23 February 1984 



72 P. C. BARNARD 

water rich in zooplankton from these dams leads to large populations of filter-feeding organisms 
such as Hydropsychidae, and among these Amphipsyche has often been reported as reaching 
pest proportions. However, it should be noted that the discharge of cold hypolimnial water from 
dams can suppress the populations of such organisms immediately below the impoundment 
(Stanford & Ward, 1981). Simulium, another filter-feeder, can also occur in large numbers in 
such habitats, and various insecticides such as DDT have been used to control populations of the 
S. damnosum complex, the vector of onchocerciasis. The effects of these control agents on 
non-target organisms is always monitored, and because Amphipsyche occurs at similar sites it 
has often figured prominently in such studies (Corbet, 1958; Statzner, 1981). Amphipsyche 
scottae is also known to be a predator of Simulium (Chutter, 1968). 

The identification of the organisms collected in all such freshwater studies, especially in the 
tropics, is always a major problem. Scott (1975) stated that the larval stages of less than 15 per 
cent of the African Trichoptera were known, and the corresponding figure for Asia must be 
considerably lower. Such identification relies on the correct association of larvae and adults, 
which often depends on long-term collecting programmes and rearing in the field; equally 
important is the provision of reliable keys for the identification of adults. The netspinning larvae 
of the Hydropsychidae are often one of the most abundant groups of macro-invertebrates in 
running water, and as part of a continuing study of the subfamily Macronematinae this paper 
deals with the adults of the genus Amphipsyche, in the tribe Macronematini. The species in this 
genus are superficially very similar to each other, and they also resemble species oiAethaloptera 
Brauer, in the Polymorphanisini (Barnard, 1980); I have frequently found these two genera 
confused in collections. Ulmer's (1907) monograph of the subfamily is still useful for some 
genera such as Macronema Pictet, but not for Amphipsyche; of the 22 species currently 
recognized only two were known to Ulmer. Kimmins (1962; 1963) described several African 
species, but new characters have been discovered in some of these. 

The keys here provided to the Old World genera of Macronematini, and to the species of 
Amphipsyche, are based on external characters as far as possible, but several species are known 
only from males and critical examination of the genitalia is often necessary. Using a cladistic 
analysis of the species of Amphipsyche the genus is divided into three main species-groups. 
Some of the evolutionary and zoogeographical implications of this classification are discussed, 
and it is intended to apply this approach to other genera of the Macronematinae and ultimately 
to test the current generic and tribal groupings within the whole subfamily. 

The methods of preparation and drawing of specimens are virtually the same as in the revision 
of the Polymorphanisini (Barnard, 1980). Temporary glycerine preparations of male and female 
genitalia were used for examination, and denuded wings were drawn from dry-mounted slide 
preparations wherever possible. 

The scale lines on the figures represent the following lengths: wings 1-0 mm; maxillary palps 
0-25 mm; legs 0-5 mm; genitalia 0-25 mm. All other features illustrated have their scale indicated 
on the figure. The arrows on some figures indicate features referred to in the keys or in the 
species descriptions. 

The nomenclature of wing veins and genitalia components follows Schmid's (1980) broadly 
based study. This means that some of the names previously used in the Polymorphanisini 
revision are now changed. Thus the aedeagus is here termed the phallotheca, and the gonopods 
are now called the inferior appendages. The wing venation terminology is unchanged, except 
that the apical forks are labelled I to V. Thus fork R 2 is now fork I, fork R 4 is fork II, fork M l is 
fork III, fork M 3 is fork IV, and fork C la is fork V. These forks are the same in both the fore and 
hind wing (except that fork IV never occurs in the hind wing of Trichoptera). 

No attempt has been made to homologize the endothecal spines of Amphipsyche males with 
those seen in some other genera. They are thus given the arbitrary names of dorsal, mid and 
ventral spines, according to their level of insertion on the apex of the phallotheca. The 
phallocrypt pocket may be homologous with the similar structure seen in some other families of 
Trichoptera (Nielsen, 1957), but its ontogeny is unknown. 

Under the heading 'Material examined' for each species are listed only the total numbers of 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 73 

each sex, the countries of collection and institutions holding the material. Full collection data are 
given only for type-specimens. Where there is further information on the distribution of a 
species which is not apparent from the list of material examined, this is noted in the correspond- 
ing 'Remarks' section. 

Abbreviations of depositories 

BMNH British Museum (Natural History), London, U.K. 

IP Institut fur Pflanzenschutzforschung, Eberswalde, D.D.R. 

IRSNB Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium 

MCZ Museum of Comparative Zoology, Harvard University, U.S.A. 

MNHN Museum National d'Histoire Naturelle, Paris, France 

MNHU Museum fur Naturkunde der Humboldt-Universitat, Berlin, D.D.R. 

MRAC Musee Royal de 1'Afrique Centrale, Tervuren, Belgium 

NAC Nanjing Agricultural College, Nanjing, China 

NM Naturhistorisches Museum, Vienna, Austria 

RNH Rijksmuseum van Natuurlijke Historic, Leiden, The Netherlands 

RSM Royal Scottish Museum, Edinburgh, U.K. 

USNM National Museum of Natural History, Smithsonian Institution, 

Washington D.C., U.S.A. 

ZI Zoological Institute, Lund, Sweden 

ZM Zoologisches Museum, Hamburg, B.R.D. 

ZSI Zoological Survey of India, Calcutta, India 

Acknowledgements 

I am very grateful to the following for the loan of specimens: Mr R. Danielsson, ZI; Dr J. 
Decelle, MRAC; Dr O. S. Flint, Jr, USNM; Dr D. C. Geijskes and Dr P. H. van Doesburg, 
RNH; Dr A. Kaltenbach, NM; Dr J. Legrand, MNHN; Dr G. Marlier, IRSNB; Mr A. F. 
Newton, MCZ; Prof. Dr H. Striimpel, ZM; Prof. Tian Li-xin, NAC; Frau H. Wendt, MNHU. I 
also thank the following for information: Dr S. K. Ghosh, ZSI; Ake Lilliestam, Kungliga 
Biblioteket, Stockholm; Dr Bo Tjeder, ZI; the Director, IP. Dr K. M. F. Scott kindly sent me 
the manuscript of the Amphipsyche section of her paper on the Hydropsychidae of southern 
Africa. 

Taxonomic method 

Although the cladistic method of classification is often taken to be equivalent to Hennig's (1966) 
phylogenetic systematics, Platnick (1979) has pointed out that there is no necessary connection 
between cladistics and the process of evolution: a cladogram can be constructed simply by 
studying the pattern of the distribution of characters in a group of organisms. Although this 
'transformed' cladistic approach has been criticized by several authors (e.g. Beatty, 1982) on the 
grounds that the claimed evolutionary neutrality is actually counter-productive, Platnick argued 
that cladistic methods are simply attempts to discover natural groups by analysing their 
characters, which is surely the aim of taxonomy in general. 

One of the difficulties with Hennig's phylogenetic method is that the taxonomist has to make a 
priori decisions about the polarity of character states, and to sort them into apomorphies and 
plesiomorphies on the basis of outgroup comparisons. This is a crucial step in the construction of 
a phylogeny, because groups can be recognized only on the basis of synapomorphies. Inevitably, 
some of these decisions on the polarity of character states are very hard to make, because the 
taxonomist has to assume at least some of the evolutionary history of the group before he starts. 
There is thus an element of circularity in the process, because one cannot make such 
assumptions about characters used to produce a phylogeny, and then use that phylogeny to draw 
independent conclusions about the evolution of the group. Platnick (1979) argued that the 
'plesiomorphic' state of a character is really the more general one, in that it is found in more 



74 P. C. BARNARD 

groups than the 'apomorphic', or less general, state. The group possessing the more specialized 
character state is therefore contained within the group showing the more general state, and this 
gives rise to the nested sets and subsets which form the hierarchical classification. This is an 
important concept, in that it avoids the idea that plesiomorphic and apomorphic states are 
alternatives: it also clearly shows why a group based on plesiomorphies alone cannot be a natural 
one because it would be recognized only by the absence of characters. Thus the production of a 
cladogram does not depend on the reconstruction of the evolutionary history of the group, but 
on the differentiation of more general characters from less general ones. The hierarchical 
structure of the cladogram is therefore a result of the inter-nested sets of unique characters, each 
delimiting a natural group. 

The test of whether the taxonomist has correctly identified the level of generality of a 
character is whether or not it is congruent with other characters at higher and lower levels. 
Instead of making decisions about the polarity of character states, one has only to distinguish the 
presence of a character from its absence, the latter being hypothesized as the more general 
condition. This highlights the problem of using the loss of a character to delimit a group. 
Phylogeneticists would decide that a loss character may be apomorphic by a priori outgroup 
reasoning, whereas transformed cladists would discover the level of generality of the 'loss' by its 
congruence with all the other characters examined. In practice, however, it is preferable to use 
presence characters to recognize groups, because without ontogenetic data it is hard to 
distinguish the secondary loss of a character from its absence at a more general level, unless 
there is a high degree of congruence. 

Having produced a cladistic classification without any assumptions of evolutionary history or 
speciation mechanisms, the taxonomist is then free to use the cladogram to infer something 
about the evolution of the group being studied, by hypothesizing a phylogenetic tree. Following 
the cladistic analysis of Amphipsyche I therefore discuss some of the phylogenetic and 
zoogeographic implications of the cladogram. The use of the transformed cladistic method and 
its application in biogeography are discussed in detail by Nelson & Platnick (1981). 

Classification of the Macronematini 

The current classification of the subfamily Macronematinae was discussed in a previous paper 
(Barnard, 1980). Of the two constituent tribes, the Polymorphanisini is almost certainly 
monophyletic, despite being delimited by loss characters. The adults are recognized by the loss 
of the mouthparts, and the larvae by the loss of the stridulatory organs on the head and fore legs 
(Scott, 1975). However, the tribe Macronematini lacks any diagnostic characters and is probably 
not monophyletic, although certain generic groups can be distinguished within it. For example, 
Macrostemum Kolenati, Amphipsyche and Protomacronema Ulmer can be grouped on both 
adult and larval characters (Scott, 1975), the most noticeable larval character being the raised 
carina on the head. The Neotropical genus Blepharopus Kolenati probably belongs here too 
(Flint & Wallace, 1980) although the carina is only poorly developed. On the other hand, the 
larvae of Leptonema Guerin-Meneville and Macronema s.str. (Flint & Bueno Soria, 1982) have 
no carina, but Leptonema and Macrostemum adults are often very similar superficially. More 
study is needed to clarify the validity of this tribe , but the group is retained here for convenience . 

Key to Old World genera of Macronematini 

1 Discoidal cell present in fore wing, but sometimes very small (Fig. 1) 2 

Discoidal cell absent in fore wing (Fig. 3) 5 

2(1) RI in hind wing ends on R 2 +i, joined to Sc by short cross-vein (Fig. 9) 3 

R\ in hind wing fuses with Sc (Fig. 8) 4 

3 (2) In fore wing, base of Rs entire (Fig. 1) PSEUDOLEPTONEMA Mosely 

In fore wing, base of Rs obsolete, joined to RI by cross-vein (Fig. 2) 

TRICHOMACRONEMA Schmid 

4 (2) Maxillary palp with second segment longer than third (Fig. 4) LEPTONEMA Guerin-Meneville 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



75 





III 



Figs 1-3 1, Pseudoleptonema sp. cf , fore wing; 2, Trichomacronema sp. cf , fore wing; 3, Leptopsyche 
gracills McLachlan cf , fore and hind wings. 



Maxillary palp with third segment longer than second (Fig. 5) MACROSTEMUMKolenati 

5 (1) Fork III in both wings with stalk (Fig. 3) LEPTOPSYCHE McLachlan 

Fork III in both wings sessile (Figs 6,7) 6 

6 (5) cf: anal area of fore wing strongly dilated (Fig. 12); $: Sc in hind wing ends on costal margin 

(Fig. 18) AMPHIPSYCHE McLachlan 

Cf : anal area of fore wing not dilated (Fig. 6); $ : Sc in hind wing fuses with R } to end on /? 7 + 3 

(Fig. 7) PROTOMACRONEMAUlmer 



76 



P. C. BARNARD 

AMPHIPSYCHE McLachlan 



Amphipsyche McLachlan, 1872: 68. Type-species: Amphipsyche proluta McLachlan, by monotypy. 
Phanostoma Brauer, 1875: 69. Type-species: Phanostoma senegalense Brauer, by monotypy. [Synony- 

mized by Martynov, 1935: 201.] 
Amphipsychella Martynov, 1935: 201. Type-species: Amphipsychella extrema Martynov, by original 

designation and monotypy. Syn. n. 




Figs 4-9 4, Leptonema sp. , maxillary palp; 5, Macrostemum sp. , maxillary palp; 6, Protomacronema sp. 
Cf, fore wing; 7, Protomacronema sp. $, hind wing; 8, Macrostemum sp. cf, hind wing; 9, 
Pseudoleptonema sp. cf , hind wing. 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



77 



Small to medium sized species, wing length cf 8-20 mm, $ 6-15 mm, yellowish or brownish in colour, 
rarely with markings on head or thorax. Antenna up to three and a half times wing length in cf , up to twice 
wing length in $ ; flagellar segments numerous (75-100 in cf , 45-70 in $), always elongate. Head with two 
pairs of setigerous warts in cf , hind pair indistinct, only one pair in $; genae in apicalis-group flat, with 
silverish pubescence. Maxillary palp with fifth segment usually very long and secondarily articulated, but 
sometimes reduced or even entirely fused with fourth segment. Spur formula basically 1.4.4, but often 
reduced to 0.4.4, 0.4.3, 0.4.2, 0.3.2 or 0.2.2. Tibia and tarsus of mid leg broad and flat in $ . Wing-coupling 
mechanism consists of single row of curved macrotrichia on costal margin of hind wing, enaging on anal 
fold of fore wing (Fig. 10). Discoidal cell absent in fore and hind wings ('false' discoidal cell formed by 
secondary fusion of R 4 and R 5 in fore wing of apicalis); median cell present in fore wing, usually absent in 
hind wing (present in magnd). In fore wing R\ and Rs often sinuous near anastomosis; fork I always stalked, 
fork II usually sessile, but stalked in apicalis-group. Sc in hind wing ends on costal margin, joined to RI by 
cross-vein. cf fore wing with strong dilated anal area. 

Cf genitalia with elongate two-segmented inferior appendages; phallocrypt pocket, associated with base 
of inferior appendages, and pre-anal appendages present in proluta-group only. Phallotheca usually with 
broad base, narrow stem and bulbous apex, with up to three pairs of endothecal spines. $ eighth sternite 
partially divided into two sclerites. 

REMARKS. Within the Macronematini, Amphipsyche seems most closely related to the African 
genus Protomacronema. Both genera have a very similar wing venation, although Protomac- 
ronema males do not have the dilated anal margin of the fore wing seen in Amphipsyche, and in 
the female hind wing Sc fuses with R^ to end on R 2 +i, instead of ending on the costal margin. The 
male genitalia are also superficially similar, Protomacronema having a pair of endothecal spines 
similar to those in the African species of Amphipsyche, but a detailed study of Protomacronema 
is needed in order to clarify the relationships of these two genera. 

Amphipsyche and Phanostoma Brauer have always been considered as being closely related, 
and have usually been separated on the spur formula. Martynov (1935: 201) synonymized them 
on the grounds that the species within Amphipsyche showed such variation in the number of 
spurs that the two genera were essentially the same. This was not accepted by all later authors 
(e.g. Ulmer, 1951) but eventually Kimmins (1962) showed that the spur formula of A. 
senegalensis (the type-species of Phanostoma) had been wrongly described, and that the 
distinction between the genera could no longer be maintained. Phanostoma is available as a 
subgeneric name for the meridiana-group recognized in the current study, but such a formal 
subdivision of the genus does not seem necessary. Kimmins also suspected that Amphipsychella 
Martynov was a synonym of Amphipsyche, and although I have seen no specimens of A. 
extrema, I am confident that this synonymy is correct. 




10 



0.5mm . 



Fig. 10 Amphipsyche berneri cf , wing coupling mechanism on costa of hind wing. 



78 P. C. BARNARD 

Geographical distribution 

Most species of Amphipsyche are restricted to the Old World tropics. The meridiana-group has 
representatives throughout the Afrotropical region, Madagascar, India and Sri Lanka, and 
through mainland South East Asia to Java, Borneo and the Philippines. The apicalis-group is 
restricted to S. India, Burma, Thailand, Vietnam, West Malaysia, Sumatra and Borneo, and the 
proluta-group occurs only in India, China and the Amur region of the U.S.S.R. Some 
zoogeographical implications of these distributions are discussed below (p. 84). 

A. senegalensis is the most widespread African species, being found throughout almost the 
whole of the Afrotropical region, whereas the other African species have very restricted 
distributions (Fig. 119). Similarly, meridiana is a very widespread species throughout India, Sri 
Lanka and South East Asia as far east as Java, although this distribution is apparently disjunct 
(Fig. 104). Most of the other species in the meridiana-group, and in the other groups, have more 
restricted distributions. The unique occurrence of proluta in central and northern China 
northwards to the Amur region of the U.S.S.R. shows an interesting parallel with Aethaloptera 
evanescens (McLachlan) in the Polymorphanisini (Barnard, 1980). There is a third species in the 
Macronematinae, Macrostemum radiatum (McLachlan), with a similar distribution, although 
this species extends into Siberia (like A. evanescens) and also occurs in Japan. 

Biology 

The first account of the immature stages of a species of Amphipsyche was by Hafiz (1937), who 
described the larva and pupa of meridiana (as indicd) from material collected near Calcutta. 
Ulmer (1957) gave detailed descriptions of Javan and Sumatran larvae and pupae of meridiana, 
but these vary in some features from Hafiz' account. Hafiz described the larval head as being 
uniformly dark brown, whereas Ulmer described (and figured) a pair of yellow flecks extending 
from the eyes onto the frontoclypeus. I have examined larvae recently collected from Java and 
they match Ulmer's figures of the head markings, so this feature may represent a genuine 
difference between the populations in India and Indonesia. The two descriptions also vary in the 
gill formula (allowing for the fact that the two authors used slightly different terminology for 
some gills) but here the recently collected Javan material matched exactly Hafiz' description of 
Indian specimens. Further information is needed to determine whether this species is polymor- 
phic or whether the two populations are perhaps subspecifically distinct. 

The larva of A. proluta was described by Lepneva (1947: redescribed, 1970). Despite the two 
species being in different species-groups it is apparent that the larvae of meridiana and proluta 
resemble each other very closely, the main difference being that in proluta the yellow head 
markings fuse to form a continuous transverse band. The gill formula of proluta matches that of 
the recently collected specimens of meridiana from Java. 

The larva of the African species senegalensis was first described from Ugandan material by 
Hickin (1955). Jacquemart (1957) gave a further detailed account of this species from Lake 
Edward (Zaire), but it should be noted that in his description the legends (and numbers) of the 
figures of the prothorax and mesothorax have been transposed, and the metathorax is figured 
upside-down. Ulmer (1963) described his Egyptian larval material as curvinerve, here con- 
sidered a synonym of senegalensis. Ulmer's description seems to differ slightly from those of 
Hickin and Jacquemart, but he made no direct comparisons with these earlier accounts, and 
without seeing material from these different areas one cannot draw any conclusions. Ulmer gave 
the gill formula for his 'curvinerve'' specimens, which is quite different from those of meridiana 
and proluta, but as neither Hickin nor Jacquemart described the gills of senegalensis, further 
comparison is impossible. Moreover, Ulmer's specimens may have represented ulmeri Kim- 
mins, and not 'curvinerve'. The pupa of senegalensis was first figured and briefly described by 
Gibbs (1973), with a detailed description by Marlier (1978). Several aspects of the biology of A. 
scottae have been described in papers by Chutter and Scott (see below) and a full description of 
the larva appears in Scott (in press). 

Although the larvae of only these few species have been described in any detail, there is 
sufficient in common between them to recognize some generic characters. This has been done by 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 79 

Lepneva (1970), Gibbs (1973) and Scott (1975; in press), all of whom give characters sufficient to 
distinguish Amphipsyche larvae from those of other macronematine genera, especially Macros- 
temum (as Macronema), Leptonema and Protomacronema . Scott (1975) has demonstrated that 
the larvae of Protomacronema and Amphipsyche seem to show a close relationship between 
these two genera, thus confirming the evidence suggested by the adult characters (see 'Remarks' 
p. 77). 

Ulmer (1957) separated the larvae of Amphipsyche and Phanostoma on the form of the hind 
tarsal claw. This was described as half the length of the tarsus and pointed in Amphipsyche 
meridiana, and only one-third the tarsal length and blunt in Phanostoma. This character was 
later figured in 'Phanostoma curvinerve' (Ulmer, 1963). However, if Hickin's (1955) and 
Jacquemart's (1957) figures of senegalensis are accurate, the claw is also half the tarsal length and 
pointed in this species. It is possible that the short, blunt claw in Ulmer 's specimens is due to 
excessive abrasion on a rocky substrate (which is known to affect both the anal and tarsal claws in 
other species of Trichoptera). 

Habitats 

Larvae of Amphipsyche are generally found in fast-flowing rivers on a stony substrate. Hickin 
(1955) also recorded A. senegalensis in Lake Victoria, but the larvae were near the outfall of the 
Nile and were therefore still in fast water. Scott (1970) found the same species in Lake Kariba, in 
a deep bay at the mouth of a stream, and Seshadri (1955) described the mass occurrence of A. 
meridiana in the very rapid water near the sluice gates of a reservoir. 

Chutter (1963) described the ecological requirements of A. scottae in some detail. The species 
was found on the Vaal River in South Africa, immediately below the man-made Vaal Barrage. 
Here the larval population dropped in winter and built up again in September-November, 
presumably in direct response to the increase in zooplankton populations, although some larvae 
were present all the year round. Further down the same river Chutter (1968) found that most 
adults of this species were caught in January, when the larval populations were again low. The 
gut contents of some larvae showed that they are apparently omnivorous, feeding on algae as 
well as on insects such as Simulium larvae. 

Boon (1979) discovered populations of A. meridiana below the artificial Lake Rawapening on 
the River Tuntang in central Java. Many organisms have difficulty in living in such a regulated 
river which is subject to sudden large changes in both water level and current speed. Parts of the 
substrate of this river are formed from vesicular volcanic lava, and large numbers of meridiana 
larvae live in the vesicles in the rock. Boon has suggested four advantages of this habitat: (1) the 
spacing of the vesicles enforces the spacing of the larvae, both within the same species and 
between the other two hydropsychid species in the same river, thus preventing overcrowding; 
(2) the fairly deep vesicles give protection against predation; (3) the larvae are protected from 
being dislodged during high water levels; (4) they are protected from desiccation during 
low water levels. Moreover, meridiana larvae also construct very tough feeding nets, which are 
more resistant to damage than those of most Hydropsychidae and also do not collapse in low 
current speeds or even when exposed at low water. Boon also showed that larvae apparently 
co-operate in building large communal nets, which is unusual in this family. It there- 
fore seems that meridiana is a particularly adaptable species, and this may be linked 
to its widespread distribution through India, South East Asia and Indonesia. The fact that 
senegalensis has been found in both rivers and lakes (albeit always in fast water) suggests 
that it too may be an adaptable species, possibly accounting for its widespread Afrotropical 
distribution. 

Corbet (1958) studied the fauna of the Victoria Nile below the Owen Falls Dam, subsequent 
to the use of DDT to eliminate populations of the Simulium damnosum complex in an effort to 
control onchocerciasis. Trichoptera in general are very sensitive to DDT, and the previously 
large populations of A. senegalensis disappeared entirely from the treated stretch of river 
immediately after the addition of the insecticide. Over a year later the populations of 
senegalensis were still very small, despite the chance of recolonization from unaffected popula- 



80 P. C. BARNARD 

tions immediately upstream. Corbet showed that this kind of insecticidal treatment can have 
long-term effects on many such macroinvertebrates as well as on the fish which rely on them for 
food. 

'Pest' species 

Where Amphipsyche larvae have colonized the fast, zooplankton-rich water immediately below 
man-made reservoirs and impoundments, either the larvae or the adults have sometimes 
reached 'pest' proportions. Seshadri (1955) gave an account ofmeridiana larvae occurring below 
the sluice gates of a reservoir in India. Here the adults were the problem, flying in enormous 
numbers every night between September and November, swarming around the street-lights and 
causing a great nuisance to people living nearby. Seshadri vividly describes how 'By about 8 P.M. 
it was a remarkable sight to see these insects in their millions dashing against lamps, and 
dropping to the ground so as to cause considerable annoyance to passers-by and vehicles. This 
went on throughout the night and every morning, to the Town Sanitary Staff fell the task of 
cleaning up the streets and removing basket loads of dead insects, especially from under the 
fluorescent lamps, where they formed shallow heaps several inches thick and many square feet in 
extent.' The larval nets were found encrusting the rocks for a few hundred yards downstream of 
the sluice gates and at times of low water the decaying stranded larvae were 'emanating a foul 
stench all over the entire locality'. Hickin (1955) described the 'peculiar sickly odour' of the dead 
bodies of vast numbers of adults of senegalensis which, together with a species of Cheumatop- 
syche Wallengren, had been swarming around a light at the Ripon Falls, Lake Victoria. Adults 
of senegalensis, together with a mayfly, were the main insect nuisance at the lights of the Owen 
Falls Dam (Uganda) according to Corbet (19580), and he also (19586) reported that larvae of 
senegalensis and two Cheumatopsyche species occasionally occurred in such numbers as to 
obstruct filters in the same dam. 

Flight activity 

The flight period of Amphipsyche species, like that of most Trichoptera, is virtually continuous 
in the tropics, but more noticeably seasonal in the more temperate regions. For example, A. 
meridiana adults are found in virtually every month of the year, whereas scottae adults, in South 
Africa, have been captured mainly from December to March. 

Corbet & Tj0nneland (1955) studied the flight activity of different species of Trichoptera 
throughout the night. The general pattern was of two peaks, one at dusk and one at dawn, 
although not every species exhibited both peaks of activity. A. senegalensis was exceptional in 
flying throughout the night, with no recognizable peaks; this species also flies in daylight. Only 
females of senegalensis were caught, which led Corbet (1966) to postulate that this species may 
be parthenogenetic. However, light-trap catches often show abnormal sex ratios because of 
differential attraction to light, and the material examined in the present study contains 
appreciable numbers of males of this species, many caught at light. Corbet's claim therefore 
seems unjustified, although facultative parthenogenesis cannot be ruled out. 

Cladistic analysis 

The list of characters used in the analysis is given below, grouped under broad morphological 
divisions. They are all 'presence' characters (see p. 74) and the state of the absence of the 
character is given in parentheses after each one. In the data matrix (Table 1) their presence is 
indicated by a plus sign and their absence by a dash, and the order of characters is re-arranged to 
show how the groupings were constructed to produce the cladogram (Fig. 11). Two apomorphic 
loss characters could have been used in this analysis, but are unnecessary. The meridiana-group 
can be recognized by the loss of the fore tibial spurs and by the hind spurs being reduced to two or 
three. The spur formula of 1.4.4 in theproluta- and apicalis-groups is demonstrably plesiomor- 
phic for the genus (by outgroup comparison with the other genera in the tribe) but the presence 
of character 21 is sufficient to distinguish the meridiana-group, making such phylogenetic 
reasoning unnecessary. The list of characters used is as follows. 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 81 

Head 

1 Vertex with dark brown markings (no markings) 

2 Genae flat with silverish pubescence (rounded with no pubescence) 

3 Fifth segment of maxillary palp simple (annulated) 

4 Fourth and fifth segments of maxillary palp fused (separate) 

Thorax 

5 Mesoscutellum with pair of dark markings (no markings) 

Fore wing 

6 Anal margin dilated in male (margin straight) 

7 Fork II stalked (sessile) 

8 'False' discoidal cell present (absent) 

9 Fork I with dark marking (no marking) 

10 Series of dark spots at wing apex (no spots at apex) 

11 Sc-Ri cross-vein with dark marking (no marking) 

12 Diagonal marking proximal to anastomosis (no marking) 

Hind wing 

13 Af 3+ 4-Cw la cross-vein present (absent) 

Male genitalia 

14 Ninth segment with lateral row of setae (only dorsal row present) 

15 Phallocrypt pocket present (absent) 

16 Basal segment of inferior appendage broad distally (narrow distally) 

17 Basal segment of inferior appendage entirely broad (entirely narrow) 

18 Inferior appendage with median setigerous projection on inner side (no setigerous projection) 

19 Ventral apex of phallotheca produced (apex rounded) 

20 Ventral apex of phallotheca pointed (apex rounded) 

21 Phallotheca with ventral median groove meeting gonopore (no groove) 

22 Eversible endotheca present (no endotheca) 

23 Base of phallotheca flattened dorso-ventrally (base rounded) 

24 Base of phallotheca extended into two pointed lobes (base rounded) 

25 Base of phallotheca broadly triangular (base rounded) 

26 Stem of phallotheca thickened in lateral view (stem narrow) 

27 Ventral endothecal spines present (absent) 

28 Mid endothecal spines present (absent) 

29 Dorsal endothecal spines present (absent) 

30 Dorsal leaf-like lobes on phallotheca (lobes absent) 

31 Mid endothecal spines blunt, rod-like (spines pointed) 

32 Mid endothecal spines very long and thickened (spines short and narrow) 

33 Mid endothecal spines fused (spines paired) 

34 Mid endothecal spines sharply up-turned (spines straight or only slightly curved) 

Three species were not included in the cladistic analysis. A. bengalensis and extrema were 
omitted because I was unable to examine material, and delicata because males of this species are 
unknown (male genitalic characters constitute over half the characters used in the analysis). A. 
bengalensis and extrema belong to the meridiana-gioup on the basis of their spur formulae (that 
of bengalensis probably being wrongly quoted - see p. 112). The male genitalia of bengalensis, 
which seem to have only the mid endothecal spines present, modified into blunt rods, also place 
the species in this group, presumably near to sinhala. Little can be surmised about extrema as it is 
known only from Martynov's figures of the female, but the highly reduced spur formula and 
shortened maxillary palps would suggest that it may also be closely related to sinhala. Although 
specimens of delicata have been examined, the affinities of the species are doubtful as it 
possesses none of the non-genitalic characters used in the analysis; it also has the intermediate 
spur formula of 0.4.4. I suspect that it belongs in the proluta-group; it cannot belong in the 
apicalis-group because it lacks characters 2, 7 and 9, and its Chinese distribution makes its 
inclusion in the meridiana-group unlikely, though not impossible. 



82 



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P. C. BARNARD 



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MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



83 



Incongruencies in the cladogram 

Although the cladogram shown is the most parsimonious one that can be constructed from the 
available data, there are a number of apparent incongruencies in the data matrix (Table 1) to 
which attention is drawn. Character 11 delimits a distinct group of African species (Fig. 11) but 
this wing-marking also occurs in senegalensis and exsiliens. However, its appearance in 
senegalensis is not consistent, and in exsiliens it is part of the broader stripe across the 
anastomosis. Character 19, the produced ventral apex of the phallotheca, delimits a large 
section of the meridiana-group, and also occurs in apicalis, but the phallotheca of this species is 
different in all other respects. Character 30, the presence of leaf-like lobes on the phallotheca, 
occurs in both meridiana and gratiosa, but each lobe in gratiosa is distinctive in bearing a spine at 
its tip. Thus each of these apparent incongruencies probably arises from the non-homology of 
the character, and no real doubt is cast on the validity of the groups suggested by the cladogram. 

Character 13, the M 3+4 -CMi a cross-vein in the hind wing, occurs independently in instabilis 
and proluta, and can thus be considered convergent for these two species. 

The remaining three incongruencies are of more interest in that they may highlight some real 
difficulties. Character 14, the presence of a row of lateral setae on the ninth abdominal segment 
of the male, is used to combine the proluta- and apicalis-groups. However, the character is 
absent in gratiosa and distincta, yet it occurs independently in pellucida of the meridiana-group. 
Only the discovery of further characters at the same level of generality can test the validity of this 



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Fig. 11 Cladogram of Amphipsyche species. 



+ African spp. 



84 P. C. BARNARD 

group. Character 4, the fusion of the 4th and 5th segments of the maxillary palp, delimits the 
berneri-corbeti-fuscata subgroup, yet this character is also seen in instabilis. Similarly, character 
3, the lack of secondary articulation on the 5th segment of the maxillary palp, appears to delimit 
the 'African' subgroup of the meridiana-group (excluding senegalensis)\ however, this character 
does not occur in ulmeri but is seen in magna. Again, these incongruencies may indicate 
incorrect groupings in the cladogram, or else possible homoplasy ; it seems reasonable to suggest 
that character 4 in instabilis and character 3 in magna have each arisen independently, as this 
tendency for simplification and reduction of the maxillary palps is well known in other genera of 
the Macronematinae. 

Thus there is still scope for further testing of the groups hypothesized in the cladogram, and 
the discovery of more characters, perhaps in the larvae, is needed to confirm or modify these 
groupings. Such extra data will show whether the incongruencies arise from non-homology of a 
character, from the usage of the character at the wrong level of generality, or whether 
homoplasy can be demonstrated in this genus. 

It will be noted that several of the individual species do not have autapomorphies indicated on 
the cladogram. All of these species are easily recognized, as is demonstrated in the key to 
species, but they do not have easily described unique features which will differentiate them from 
all other species in the genus rather than merely from their nearest neighbour. Thus berneri and 
corbeti can be distinguished from each other by the lobes of the male tenth segment, which are 
broad and divergent in berneri but narrow and sub-parallel in corbeti. Neither of these character 
states would distinguish one or other species from all others in the genus, and their use would 
generate confusion and repetition in the cladogram. 

Evolutionary and zoogeographical considerations 

The geographical distributions of the main species-groups and some of their components have 
been outlined earlier (p. 78). In brief, the three species-groups have noticeably different 
distributions, although all three overlap to some degree. It is interesting to try and deduce 
something of the evolutionary history of the genus from these data, coupled with the informa- 
tion from the cladogram (Fig. 11). For these purposes the cladogram can be considered as a 
phylogram, showing degrees of common ancestry (Nelson & Platnick, 1981: 171), although 
branching points do not denote actual speciation events or real ancestors. 

The current distribution of the genus in Africa, Madagascar, India and South East Asia 
suggests that it arose when the African, Malagasy and Indian land-masses were still closely 
associated, namely before the end of the Cretaceous (Smith, Hurley & Briden, 1981). Each of 
these three components carried its own fragmented group of species, and the further dispersal 
and evolution of the genus would have occurred when India became linked with South East Asia 
(Late Eocene/Oligocene). 

This kind of model seems more useful than postulating the origin of the genus in one of the 
three main areas of its distribution (Africa, India, South East Asia) with subsequent long-range 
dispersal to the other two. Several other macronematine genera have similar widespread 
distributions, such as Polymorphanisus Walker, Aethaloptera and Macrostemum (the latter also 
occurring in the New World), and their long-range dispersal seems similarly unlikely in view of 
their relatively limited powers of flight and their restriction to certain freshwater habitats. Some 
other genera of the subfamily are restricted to one continent; Protomacronema, which may be 
the sister-group of Amphipsyche is found only in Africa, and future studies on these genera 
should indicate whether or not this model is satisfactory. 

The meridiana-group of Amphipsyche has representatives in Africa, Madagascar, India and 
South East Asia, but it is important to note that not all the African species form a monophyletic 
group. A. senegalensis was apparently an early coloniser of Africa, where it is now the most 
widespread species (Fig. 119), but all the other African species (including pellucida from 
Madagascar) form a monophyletic group with allopatric distributions. Each member of this 
sub-group is individually sympatric with senegalensis, which tends to confirm their more distant 
relationship with that species (Nelson & Platnick, 1981: 384). The apicalis- and proluta-groups 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 85 

have no African representatives but both occur in India and South East Asia. It thus seems likely 
that both arose from a common Indian ancestral species, and that each group later dispersed and 
speciated throughout South East Asia, along with some members of the meridiana-group. 

Check-list of Amphipsyche species 

Although most of the species in this list are arranged according to the relationships suggested by 
the cladistic analysis (Fig. 11), they are not phyletically sequenced (Wiley, 1981: 211). This is 
partly because three species, bengalensis, delicata and extrema, were omitted from the clado- 
gram owing to lack of suitable material; these species would be sedis mutabilis sensu Wiley. 
Moreover, the cladogram is not of the simple asymmetrical 'pectinate' type which lends itself to 
this sequencing convention without the proliferation of formal subgroup names. 

AMPHIPSYCHE McLachlan parva Banks 

Phanostoma Brauer meridiana Ulmer 
Amphipsychella Martynov syn. n. nirvana Banks syn. n. 

pro/ute-group vedana Banks syn. n. 

pro/ufa McLachlan propinqua Ulmer syn. n. 

paraproluta Hwang syn. n. indica Martynov syn. n. 

bifasciata Navas tricalcarata Martynov 

distincta Martynov sigmosa Navas syn. n. 

delicata Banks sinhala sp. n. 

ap/ca/is-group bengalensis Martynov 

apicalis Banks extrema (Martynov) comb. n. 

exsiliens sp. n. pellucida (Navas) comb. n. 

gratiosa Navas instabilis Kimmins 
petiolata Ulmer plicata (Jacquemart) syn. n. 

minima Banks syn. n. ulmeri Kimmins 

pubescens Kimmins syn. n. scottae Kimmins 

meridiana-group fuscata Kimmins 

senegalensis (Brauer) corbeti Kimmins 

curvinerve (Navas) syn. n. berneri Kimmins 
magna Banks 

Key to species of Amphipsyche 

Of necessity this key is based largely on features of the male genitalia, which are often the only reliable way 
of distinguishing species in this genus; moreover, the females of nine of the species are unknown. However, 
geographical distribution and external characters such as wing venation and spur formulae are used where 
feasible, so that isolated female specimens can be identified as far as possible. 

1 Spur on fore tibia absent 

Spur on fore tibia present 16 

2 (1) Four spurs on hind tibia; $ ; RI in fore wing ends on Sc (Fig. 39) (O" unknown) delicata (p. 93) 

Two or three spurs on hind tibia; R^ in fore wing ends on wingmargin 3 

3 (2) c?:phallotheca with three pairs of endothecal spines (Fig. 82) 4 

Cf : phallotheca with only two pairs of endothecal spines or less 5 

4 (3) Very large species, fore wing 15-20 mm; pair of round markings on mesoscutellum(Fig. 

87) (Philippines) magna (p. 102) 

Small species, fore wing 8 mm; no markings on mesoscutellum ($ unknown) 
(Borneo) parva (p. 105) 

5 (3) Indian or Sri Lankan species 6 

African or Malagasy species 

6 (5) 9 : maxillary palps very short (Fig. 116); only one or two spurs on mid tibia (cf 

unknown) extrema (p. 112) 

Maxillary palps unmodified; four spurs on mid tibia 

7 (6) SpursO.4.2 8 

Spurs 0.4. 3 or 0.4. 4 meridiana (p. 106) 



86 P. C. BARNARD 

8 (7) cf : endothecal spines rod-like, longer than breadth of phallotheca stem (Fig. 117) (9 

unknown) (India) bengalensis (p. Ill) 

Cf : endothecal spines much shorter than breadth of phallotheca stem (Fig. 98) (Sri 
Lanka) sinhala (p. 105) 

9 (5) Hind tibia with three spurs; cf : base of phallotheca extended into two pointed lobes 

(Fig. 134) (Malagasy species) pellucida (p. 1 15) 

Hind tibia with two spurs ; cf : phallotheca of other shape ( Af rotropical species) 10 

10 (9) cf: endothecal spines absent (Fig. 124) senegalensis (p. 112) 

Cf : at least one endothecal spine present 11 

11 (10) Cross-vein present between M 3+4 and Cw la in hind wing (Figs 140, 147); cf: mid 

endothecal spines fused into single structure (Fig. 145) instabilis (p. 1 17) 

No such cross-vein in hind wing; cf : mid endothecal spines paired 12 

12 (11) cf : fifth segment of maxillary palp fused with fourth (line of fusion visible infuscata, Fig. 

167); apex of phallotheca rounded (Fig. 175) 13 

Cf : fifth segment of maxillary palp distinct (sometimes reduced in length); apex of 
phallotheca pointed (Fig. 159) 15 

13 (12) cf : diagonal fuscous marking on fore wing (Fig. 165); base of phallotheca narrow (Fig. 

169) (9 unknown) fuscata (p. 123) 

Cf : no diagonal wing marking; base of phallotheca broadly triangular (Fig. 175) 14 

14 (13) cf : lobes of tenth segment broad and divergent in dorso-ventral view (Fig. 180) (9 

unknown) (Ghana) berneri (p. 127) 

Cf: lobes of tenth segment narrow, subparallel in dorso-ventral view (Fig. 176) ($ 
unknown) (Uganda) corbeti (p. 125) 

15 (12) Cf : mid endothecal spines gently curved dorsally (Fig. 159) (South Africa) . . scottae (p. 121) 

Cf: mid endothecal spines turned abruptly dorsally (Fig. 154) (9 unknown) 
(Sudan) ulmeri (p. 120) 

16 (1) Genae flat with silverish pubescence; fork II stalked in fore wing (Fig. 54) 17 

Genae rounded with no pubescence ; fork II sessile in fore wing (Fig . 1 2) 20 

17 (16) 'False' discoidal cell in fore wing (enclosing corneous spot) (Fig. 43) (India) . . . apicalis (p. 94) 

No 'false' discoidal cell 18 

18 (17) cf : phallotheca with eversible endotheca (Figs 59, 60) (9 unknown) (Burma) exsiliens (p. 96) 

Cf : phallotheca without eversible endotheca 19 

19 (18) Fore wing with striking pattern of five dark brown spots with other paler brown 

markings (Fig. 61); cf: phallotheca with large dorsal leaf-like lobes (Fig. 65) 

( 9 unknown) gratiosa (p. 98) 

Fore wing with only one dark brown spot in fork I (Fig. 68); cf : phallotheca with no 
dorsal lobes (Fig. 72) petiolata (p. 99) 

20 (16) Cross-vein present between M 3+4 and Cw la in hind wing (Fig. 12) proluta (p. 86) 

No such cross-vein in hind wing 21 

21 (20) Vertex of head, antennal scape and pedicel with dark brown markings (Fig. 30) 

distincta (p. 89) 
No markings on head ($ unknown) bifasciata (p. 89) 

The pro/ufa-group 

Genae rounded, with no pubescence. Fork II sessile in fore wing. Spurs usually 1.4.4 but subject to 
reduction. Fifth segment of maxillary palp long and secondarily annulated. cf interior appendages with 
basal segment broad, at least distally; phallocrypt pocket present; pre-anal appendages present though 
small (absent in distincta). Phallotheca lacking endothecal spines. Ninth segment with two rows of setae 
(dorsal and lateral) in lateral view. 

India, China and U.S.S.R. (Amur region). 

Amphipsyche proluta McLachlan 
(Figs 12-21) 

Amphipsyche proluta McLachlan, 1872: 70. Lectotype cf, U.S.S.R. (BMNH), designated by Kimmins, 

1957ft: 105 [examined]. 
Amphipsyche paraproluta Hwang, 1957: 387. Holotype cf, CHINA: Jiangsu Prov., Nanjing, 15.viii.1956 

(Hwang) (NAC) [not examined]. Syn. n. 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



115 








129 



Figs 126-129 Amphipsyche senegalensis $. 126, wing venation; 127, mid and hind legs; 128, eighth 
sternites; 129, maxillary palp. 

and Ulmer (1963) (as curvinerve); the pupa was described by Gibbs (1973) and Marlier (1978). 
In addition to the distribution records below, Navas (1923) also recorded this species from 
Madagascar, but this is unconfirmed. 

MATERIAL EXAMINED 

Lectotype d" of senegalensis , Senegal: 1869 (Steindachner) (NM). Lectotype $ oi curvinerve, Egypt: 
Cairo, 20.vii.1916 (Alfieri) (USNM). 

79 cf, 194 $, 3 larvae, 3 pupae, Chad, Sudan, Ethiopia, Ghana, Nigeria, Cameroun, Zaire, Uganda, 
Tanzania, Zambia, Malawi, Zimbabwe, South Africa (Transvaal) (BMNH, IRSNB, MRAC, RSM, 
USNM, ZI). 

Amphipsyche pellucida (Navas) comb. n. 

(Figs 130-139; distribution, Fig. 119) 
Protomacronema pelluddum Navas, 1923: 26. Holotype $, MADAGASCAR (MNHN) [examined]. 

Cf . Antenna 45 mm, with c. 80 segments. Fore wing 15-16 mm. Body yellowish brown; basal antennal 
segments annulated with dark brown, apical segments fuscous. Fore wing very pale yellow, with faint 
darker stripe from R { to M in line with M-Cu\ cross-vein. Venation as in Fig. 130. Spurs 0.4.3 (Fig. 131). 
Maxillary palp 5-segmented, 5th segment not secondarily annulated, shorter than segments 1-3 combined 
(Fig. 132). 

$. Antenna 15 mm, with c. 60 segments. Fore wing 11-13 mm. Coloration as in cf . Fore wing with no 
markings, venation as in Fig. 136. Spurs 0.4.3 (Fig. 137). Maxillary palp similar to that of cf (Fig. 139). 

GENITALIA cf (Figs 133-135). Ninth segment broadly rounded laterally. Base of phallotheca narrow, with 



116 



P. C. BARNARD 




Figs 130-135 Amphipsyche pellucida cf . 130, wing venation; 131, mid and hind legs; 132, maxillary palp; 
133, genitalia, lateral view; 134, phallotheca, lateral view; 135, genitalia, ventral view. 

basal corners produced into pointed lobes. Stem of phallotheca narrow, apex produced into an elongate 
lobe. Only mid endothecal spines present, very short and blunt. Inferior appendage slender and sinuous, 
terminal segment clearly differentiated. 

GENITALIA $ (Fig. 138). Eighth sternites oval, each sclerite almost symmetrical, with all corners rounded. 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



87 




. \ 



16 



Figs 12-17 Amphipsyche proluta cf . 12, wing venation; 13, legs; 14, maxillary palp; 15, genitalia, lateral 
view; 16, phallotheca, lateral view; 17, genitalia, ventral view. 

O". Antenna c. 30 mm, with c. 80 segments. Fore wing 11-14 mm. Body pale yellowish brown, antenna pale 
yellow, narrowly annulated with brown, becoming more fuscous towards antennal apex. Fore wing very 
pale yellow, with slightly darker marking around Sc-R\ cross- vein, sometimes extending further onto 
anastomosis. Venation as in Fig. 12; cross-vein present between M 3+4 and Cwi a in hind wing. Spurs 1.4.4; 



P. C. BARNARD 



pre-apical spurs on hind tibia very short (Fig. 13). Maxillary palp 5-segmented, 5th segment secondarily 
annulated, over 1-5 times length of segments 1-4 combined (Fig. 14). 

$. Antenna c. 12 mm, with c. 50 segments. Fore wing 8-10 mm. General coloration as in cf , fore wing 
with no dark markings. Venation as in Fig. 18; M 3+4 _CM la cross-vein in hind wing, as in cf. Spurs 
1.4.4 (Fig. 19). Maxillary palp similar to that of cf, but 5th segment approximately same length as 
segments 1-4 combined (Fig. 21). 

GENITALIA cf (Figs 15-17). Ninth segment narrow laterally, pre-anal appendages present as small 
setigerous projections on tenth segment (Fig. 15). Phallocrypt pocket relatively broad and rounded(Fig. 
15). Basal segment of inferior appendage broad apically only, terminal segment partly differentiated. 
Phallotheca slender with narrow base; apex truncate, with slightly pointed lobe dorsally. 

GENITALIA $ (Fig. 20). Eighth sternites broad, squarish; thickened inner margins wide, extending far down 
inner edges. 

REMARKS. Although easily identified by the male genitalia, both sexes of this species can be 
distinguished from the rest of the proluta-group by the M 3+4 _Cw la cross-vein in the hind wing. 
The only other species to show this character is instabilis, in the meridiana-group. 

Hwang (1957) described paraproluta as differing from proluta in the male genitalia. However, 
it seems that he had only McLachlan's original description on which to base his comparison, and 
both McLachlan's description and figure of proluta are very poor. McLachlan saw only dried 
material, and the pointed valves, which he thought were the intermediate appendages, are 
apparently the tenth segment. McLachlan's (1878: 352) redescription of the species was rather 
better, with a clearer figure; here he noted the setigerous pre-anal appendage as 'a distinct 
tooth'. McLachlan's type-series of three males and two females is extant in the BMNH, although 
Kimmins (19576) did not mention this in his lectotype designation. 







21 



Figs 18-21 Amphipsyche proluta $ . 18, wing venation; 19, legs; 20, eighth sternites; 21, maxillary palp. 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 89 

Although Hwang's (1957) description was apparently based on one male holotype, I was lent a 
male specimen from NAC labelled as paratype, which had identical data to the type. Whether or 
not this specimen has any type-status, it is an important 'topotypic' specimen, and examination 
of it confirmed the synonymy of paraproluta with proluta. 

Three specimens examined (MNHN, MCZ) are labelled 'Hanlseon', apparently from China, 
according to Navas (1914). However, this is an impossible combination of letters for a Chinese 
place-name, and the label data must have been mis-copied from another, presumably hand- 
written, source. I tentatively suggest that the 'Is' is a misreading of the single (handwritten) letter 
'k', and the final 'n' a misreading of 'u'. This would give the more plausible transliteration of 
'Hankeou', the French spelling of Han-kow, for material collected by the Frenchman, de 
Guerne. 

The larva of proluta was described by Lepneva (1947; 1970). 

MATERIAL EXAMINED 

Lectotype cf of proluta, U.S.S.R.: 'Amur Land' (Maack) (BMNH). 

19 Cf, 8 <j>, U.S.S.R.: Amurskaya (2 cf, 2 $ paralectotypes of proluta), China (1 cf 'paratype' of 
paraproluta) (BMNH, MCZ, MNHN, NAC). 

Amphipsyche bifasciata Navas 
(Figs 22-27) 

Amphipsyche bifasciata Navas, 1931a: 7. Holotype cf , CHINA: 'meridionale' (Bris) (lost). 
[Amphipsyche proluta McLachlan; Banks, 1940: 207; Mosely, 1942: 361. Misidentifications.] 

Cf . Antennal length unknown (both specimens damaged). Fore wing 10-15 mm. Antennal segments pale 
yellow, with golden brown annulations. Head, thorax and abdomen yellowish brown. Fore wing pale 
yellow, shaded pale brown at apex and with darker brown stripe across anastomosis. Venation as in Fig. 22; 
fork I in fore wing approximately equal in length to its stalk. Spurs 1.4.4 (Fig. 24). Maxillary palp 
5-segmented, 5th segment secondarily annulated, longer than segments 1-4 combined (Fig. 23). 
$. Unknown. 

GENITALIA cf (Figs 25-27). Ninth segment relatively narrow laterally; pre-anal appendages present as 
small setigerous projections on tenth segment. Phallocrypt pocket elongate and sac-like in lateral view, 
shield-shaped in ventral view (Fig. 27). Basal segment of inferior appendage very broad, viewed laterally 
and ventrally; terminal segment clearly differentiated. Phallotheca slender, apex truncate, with pair of 
pointed unsclerotized lobes (superficially resembling endothecal spines); dorsally a single pointed lobe. 

REMARKS. Well-marked examples of this species are easily recognized by the wing markings, but 
the single male examined from Szechwan has lost virtually all these markings, and is easily 
confused with proluta, hence Banks's (1940) misidentification. 

According to the original description the holotype of bifasciata should be in Navas's 
collection, now in Barcelona, but when this was examined in 1979 the type was apparently 
missing (T. R. New, pers. comm.). However, Navas's illustration of the wing is sufficient to 
identify the species; it is obvious from his figure that the type is a male, although Navas does not 
mention this. 

MATERIAL EXAMINED 
2 cf , China (BMNH, USNM). 

Amphipsyche distincta Martynov 

(Figs 28-38) 

Amphipsyche distincta Martynov, 1935: 196. 6 cf syntypes, INDIA: Madhya Pradesh, river at Mandla, 
Nerbudda Survey (Pruthi) (lost from ZSI). 

Cf. Antenna up to 25mm, with c. 80 segments. Fore wing 8-10 mm. Body pale yellowish brown; 
setigerous warts on vertex of head dark greyish brown; antennal scape and pedicel with dark brown 
longitudinal stripe on dorsal surface (Fig. 30); front femur dark brown. Fore wing pale yellow, no dark 
markings. Venation as in Fig. 28; in fore wing fork IV with short stalk; in hind wing Sc not reaching wing 



90 



P. C. BARNARD 









27 



Figs 22-27 Amphipsyche bifasciata of. 22, wing venation; 23, maxillary palp; 24, legs; 25, genitalia, 
lateral view; 26, phallotheca, lateral view; 27, genitalia, ventral view. 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



91 




30 



31 



32 






Figs 28-34 Amphipsyche distincta d". 28, wing venation; 29, mid and hind tibiae; 30, head, dorsal view; 
31, maxillary palp; 32, genitalia, lateral view; 33, phallotheca, lateral view; 34, genitalia, ventral view. 



92 



P. C. BARNARD 



margin, fork V very narrow. Spurs 0.3.2 (Fig. 29). Maxillary palp 5-segmented, 5th segment secondarily 
annulated, longer than segments 1-4 combined (Fig. 31). 

$. Antenna up to 10 mm, with c. 50 segments. Fore wing 6-8 mm. Coloration as in cf . Venation as in 
Fig. 35; in fore wing R\ ends on Sc, fork IV sessile. In hind wing Cu\ not forked, 2A absent. Spurs (Fig. 36) 
and maxillary palp (Fig. 37) as in cf . 

GENITALIA cf (Figs 32-34). Ninth segment broad laterally, pre-anal appendages absent. Phallocrypt pocket 
rounded in lateral view (Fig. 32), broadly triangular in ventral view. Inferior appendage broad and sinuous 
in ventral view, terminal segment not differentiated. Phallotheca broad basally, with narrow stem; apex 
triangular with no obvious appendages or lobes. 

GENITALIA 9 (Fig. 38). Eighth sternites broad, with squarish corners; thickened inner margins extending 
barely half length of sclerites. 

REMARKS. This aptly named species is so distinctive that it is easily recognized; no other species 
has markings on the head and antennae (the only other species with any body markings is 
magna, with a pair of spots on the mesoscutellum). 

Banks (1939) redescribed distincta, drawing attention to the dark front femur in both sexes. 
Martynov had described all the legs as being pale, but since all six syntypes are apparently lost 
(Ghosh, in lift. ) this character cannot be checked on Martynov's material. Banks also stated that 
fresh specimens were 'plainly greenish'. 

Martynov listed as the syntypes '4 cf ' followed by '2 cf ' with identical data; one of these may 




y 



Figs 35-38 Amphipsyche distincta $ . 35, wing venation; 36, mid and hind tibiae; 37, maxillary palp; 38, 
eighth sternites. 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



93 



be an error for '9'- No descriptions of the female are given in his text, but this is also true of his 
description ofindica (= meridiana), where both males and females make up the type-series. 
A neotype designation does not seem necessary for such an easily recognizable species. 

MATERIAL EXAMINED 

19 cf , 28 9, India (MCZ, USNM). 

Amphipsyche delicata Banks 

(Figs 39-42) 
Amphipsyche delicata Banks, 1939: 58. LECTOTYPE $ , CHINA (MCZ), here designated [examined]. 

Cf . Unknown. 

$. Antenna over 12 mm, with more than 50 segments (all specimens damaged). Fore wing 6-8 mm. 
Antennal segments pale yellow, slightly annulated with brown. Head, thorax and abdomen yellowish 
brown. Fore wing very pale yellow with no markings, venation as in Fig. 39. RI in fore wing ends on Sc. 
Spurs 0.4.4 (Fig. 40). Maxillary palp 5-segmented, 5th segment secondarily annulated, longer than 
segments 1-4 combined (Fig. 41). 




Figs 39-42 Amphipsyche delicata <j>. 39, wing venation; 40, mid and hind tibiae; 41, maxillary palp; 42, 
eighth sternites. 



94 p. c. BARNARD 

GENITALIA $ (Fig. 42). Eighth sternites rounded, bluntly pointed posteriorly; thickened inner edge 
relatively narrow. 

REMARKS. The relationships of this species are in some doubt, mainly because no males have yet 
been discovered. Banks (1939) compared it with minima (= petiolata), from which it differs in 
venation, and with distincta, which he distinguished by the dark fore femora. It seems to be most 
closely related to distincta, with which it shares the venational feature of /?! ending on Sc in the 
fore wing; this may be a synapomorphy for this pair of species. However, the placing of this 
species in the proluta-group is complicated by its unique spur formula of 0.4.4. It cannot belong 
in the apicalis-group because of the sessile fork II in the fore wing, and its Chinese distribution 
and unmodified maxillary palps make its inclusion in the meridiana-group unlikely. 
The specimen here designated as lectotype was labelled 'type' by Banks, but not so published. 

MATERIAL EXAMINED 

Lectotype $, China: Hainan Tao I., Chung Kon, 18.vii.1935 (Gressitt) (type no. 23470, MCZ). 
6 $ (paralectotypes), China (MCZ). 

The apicalis-group 

Genae flat, with silverish pubescence. Fore wing with fork II stalked, dark marking in fork I. Spurs always 
1.4.4. Fifth segment of maxillary palp long and secondarily annulated (except $ apicalis). cf inferior 
appendages slender; phallocrypt pocket and pre-anal appendages absent. Phallotheca lacking endothecal 
spines. Ninth segment with two rows of setae as mproluta-group. 

India, Burma, Thailand, Vietnam, West Malaysia, Sumatra and Borneo (Sarawak). 



Amphipsyche apicalis Banks 

(Figs 43-53) 
Amphipsyche apicalis Banks, 1939: 56. LECTOTYPE cf , INDIA (MCZ), here designated [examined]. 

Cf . Antenna over 20 mm, with more than 50 segments (broken in both specimens examined). Fore wing 
12-13 mm. Antennal segments pale golden brown, becoming more fuscous towards apex, slightly 
annulated with brown. Head, thorax and abdomen yellowish brown. Fore wing golden yellow, with dark 
brown spot in fork I, glabrous brown streaks at wing apex proximal to dark spot and across anastomosis 
(Fig. 43). In fore wing 'false' discoidal cell enclosing corneous spot at base of R 4+5 , fork I with short stalk. 
R 2 +3 fused in hind wing. Spurs 1.4.4, pre-apical spurs on hind tibia short (Fig. 44). Maxillary palp 
5-segmented, 5th segment secondarily annulated, longer than segments 1-4 combined (Fig. 45). 

$ . Antenna over 12 mm, with more than 50 segments (broken in both specimens examined). Fore wing 
9-10 mm. Body coloration as in cf . Fore wing yellowish brown with pale brown spot on wing margin in fork 
I and another at anterior end of anastomosis (Fig. 50). Fore wing with 'false' discoidal cell as in cf , but not 
so clearly defined. Spurs 1.4.4, one pre-apical spur on hind tibia very small (Fig. 51). Maxillary palp 
5-segmented, 5th segment not annulated, approximately equal in length to segments 1 and 2 combined 
(Fig. 52). 

GENITALIA cf (Figs. 46-49). Ninth segment with enlarged rounded side-pieces. Phallotheca with slender 
stem, enlarged apically to form two ventrally directed subtriangular processes; apical process with fine 
teeth at ventral apex, and hingeing dorsally to form a simple eversible endotheca (Figs 48, 49). Inferior 
appendage with pronounced setigerous projection mid-dorsally; terminal segment not clearly differenti- 
ated. 

GENITALIA $ (Fig. 53). Eighth sternites with sharp indentation in posterior edge. 

REMARKS. Superficially this species is very similar to exsiliens and petiolata, but is easily 
distinguished by the characteristic shape of the male phallotheca (Fig. 48). 

Banks (1939) was unsure whether the two females from Coimbatore belonged to this species, 
but after examination of his material there seems little doubt that they are correctly placed here. 
Banks apparently overlooked the 'false' discoidal cell in the females (which is admittedly less 
obvious than in the males), and he also did not notice the very small pre-apical spur on the hind 
tibia. The faint wing markings in the female are a reduced form of the pronounced male pattern; 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



95 







Figs 43-49 Amphipsyche apicalis cf . 43, wing venation; 44, mid and hind tibiae; 45, maxillary palp; 46, 
genitalia, lateral view; 47, genitalia, ventral view; 48, phallotheca, lateral view; 49, phallotheca with 
endotheca everted. 



96 



P. C. BARNARD 





Figs 50-53 Amphipsyche apicalis 9 . 50, wing venation; 51, mid and hind tibiae; 52, maxillary palp; 53, 
eighth sternites. 

this is also seen \npetiolata, and probably also in the closely related gratiosa and exsiliens, but the 
females of the last two species have yet to be discovered. 
The specimen designated as lectotype was labelled 'type' by Banks, but not so published. 

MATERIAL EXAMINED 

Lectotype Cf , India: Mysore, Shimoga, R. Tunga, 1865' [560 m], at light, lO.vi. [not iv as stated by 
Banks] [year unknown] (Nathan) (type no. 22677, MCZ). 

1 Cf (paralectotype), 2 <j>, India (MCZ). 



Amphipsyche exsiliens sp. n. 

(Figs 54-60) 

Cf. Antennal length unknown (broken in all specimens). Fore wing 12-14 mm. Body yellowish brown, 
back of head and dorsal surface of thorax brown; antennal segments pale yellow, narrowly annulated with 
brown. Fore wing pale yellow with brown spot in fork I and another centred on Sc-Ri cross-vein; pale 
brown stripe across anastomosis and very pale shading at wing apex. Venation as in Fig. 54. Spurs 1.4.4 
(Fig. 55). Maxillary palp 5-segmented, 5th segment secondarily annulated, longer than segments 1-4 
combined (Fig. 56). 
$. Unknown. 

GENITALIA cf (Figs 57-60). Ninth segment broad laterally. Base of phallotheca broadly triangular, stem 
slender with triangular apex. Eversible sac-like endotheca present, connective membrane bearing many 
small spines (Fig. 60). Inferior appendage slender, terminal segment scarcely differentiated. 

REMARKS. This species is easily distinguished from the other members of the apicalis-group by 
the rounded sac-like endotheca, which is more mobile than that of apicalis. In the latter species it 
hinges through only a few degrees, but in exsiliens it can be invaginated almost entirely inside the 
phallotheca apex, or hinged through almost 180 to lie dorsal to the apex (Figs 59, 60). 

MATERIAL EXAMINED 

Holotype cf , Burma: Tenasserim Valley (Doherty) (BMNH). 
Paratypes. 2 cf , data as holotype (BMNH). 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



97 




Figs 54-60 Amphipsyche exsiliens cf . 54, wing venation; 55, legs; 56, maxillary palp; 57, genitalia, lateral 
view; 58, genitalia, ventral view; 59, phallotheca, lateral view; 60, phallotheca with endotheca everted. 



98 



P. C. BARNARD 

Amphipsyche gratiosa Navas 



(Figs 61-67) 
Amphipsyche gratiosa Navas, 1922: 62. Holotype cf , VIETNAM: Tonkin, Hag Song, vii.1918 (lost). 

C?. Antenna 25 mm, with c. 85 segments. Fore wing 10-12 mm. Body yellowish brown, antennal segments 
narrowly annulated with brown. Fore wing pale yellow with striking pattern; pale brown streak across 
whole width of wing proximal to anastomosis; five dark brown spots in apical area, partially linked to pale 
brown areas in apical forks; venation as in Fig. 61. Spurs 1.4.4 (Fig. 62). Maxillary palp 5-segmented, 5th 
segment secondarily annulated, much longer than segments 1-4 combined (Fig. 63). 
. Unknown. 





63 







66 





67 



Figs 61-67 Amphipsyche gratiosa cf. 61, wing venation; 62, legs; 63, maxillary palp; 64, genitalia, lateral 
view; 65, phallotheca, lateral view; 66, phallotheca, dorsal view; 67, genitalia, ventral view. 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 99 

GENITALIA cf (Figs 64-67). Ninth segment relatively narrow laterally. Base of phallotheca elongate, 
flattened; stem narrow; apex with pair of leaf-like lobes, each bearing single spine on inner surface; pointed 
dorsal lobe proximal to these apical lobes. Inferior appendage slender; setigerous projection midway on 
inner surface in ventral view; terminal segment not differentiated. 

REMARKS. This species is easily recognized by its prominent wing pattern. The unusual genitalia 
are also diagnostic, and suggest no close affinities with the other species in the apicalis-group. I 
have assumed that the dorsal lobes are not modified endothecal spines, despite their superficial 
similarity to those structures, because no other species in the genus has spines with lobate bases. 
There is certainly a close superficial similarity between the phallotheca of this species and that of 
meridiana for example, although all other critical characters of this species definitely place it in 
the apicalis-group. One male examined has a 'false' discoidal cell in the left fore wing only, a 
character otherwise seen only in apicalis. 

The type of this species should be in the Navas collection (now in Barcelona) but is apparently 
missing (T. R. New, pers. comm.); however the species is easily recognizable from Navas's 
figure. In addition to the distribution records below, the type was collected in Vietnam. 

MATERIAL EXAMINED 
4 cf , Burma, Thailand (BMNH). 

Amphipsyche petiolata Ulmer 
(Figs 68-77) 

[Amphipsyche proluta McLachlan; Ulmer, 1910: 55; 1913: 79. Misidentifications.] 

Amphipsyche petiolata Ulmer, 1930: 434. Lectotype $ [listed as 'holotype' by Weidner, 1964: 67], JAVA 

(ZM), designated by Ulmer, 1951: 197 [examined]. 
Amphipsyche minima Banks, 1931: 395. LECTOTYPE $, WEST MALAYSIA (BMNH), here designated 

[examined]. Syn. n. 
Amphipsyche pubescens Kimmins, 1955: 387. Holotype cf, BORNEO (BMNH) [examined]. Syn. n. 

Cf . Antenna c. 35 mm, with c. 80 segments. Fore wing 9-11 mm. Body yellowish brown; posterior part of 
vertex and dorsal surface of mesothorax brown. Antenna pale yellow, annulated with brown, segments 
becoming more fuscous towards apex. Fore wing pale golden yellow, with pale brown apex, brownish 
stripe across anastomosis and dark brown spot in fork I; venation as in Fig. 68. Spurs 1.4.4 (Fig. 69). 
Maxillary palp 5-segmented, 5th segment secondarily annulated, about three times length of segments 1-4 
combined (Fig. 70). 

$ . Antenna 10 mm, with c. 45 segments. Fore wing 7-8 mm. General coloration as in cf . Fore wing very 
pale yellow, pale brown stripe across anastomosis, sometimes slight brown marking in fork I; venation as in 
Fig. 74. Spurs 1.4.4 (Fig. 75). Maxillary palp 5-segmented, 5th segment secondarily annulated, about twice 
length of segments 1-4 combined (Fig. 77). 

GENITALIA cf (Figs 71-73). Ninth segment broadly rounded laterally. Base of phallotheca narrow, 
rectangular, narrow stem abruptly right-angled. Tip of phallotheca globose with bifurcate membranous 
process arising from apical depression, abruptly up-turned at apex (Fig. 72). Inferior appendage narrow 
with setigerous median projection on inner surface, terminal segment not differentiated. 

GENITALIA 9 (Fig. 76). Eighth sternites narrow, inner thickened edges slightly incurved; posterior margin 
produced as rounded point. 

REMARKS. The male of this species can be distinguished from others in the apicalis-group by the 
form of the phallotheca. The apical rod-like process superficially resembles an endothecal spine 
but it is a membranous median structure. Within this species-group the only other known female 
is that of apicalis, which has differently shaped eighth sternites and a short apical segment of the 
maxillary palp. 

I have taken Ulmer's subsequent (1951) listing of the Javan syntype as 'type' as a lectotype 
designation. Weidner (1964) listed this specimen as the holotype, but this is incorrect as Ulmer's 
original description was based on three syntypes. 

It is not clear from Banks's (1931) description of minima how many specimens constituted the 
type-series. Of the two extant syntypes (with identical data, and both labelled 'type' by Banks) 



100 



P. C. BARNARD 




70 





72 



Figs 68-73 Amphipsyche petiolata cf . 68, wing venation; 69, legs; 70, maxillary palp; 71 , genitalia, lateral 
view; 72, phallotheca, lateral view; 73, genitalia, ventral view. 



the MCZ specimen was labelled 'paratype' by H. H. Ross in 1965. 1 have therefore designated 
the BMNH syntype as lectotype. Ulmer (1951) remarked on the similarity of minima to 
petiolata. 

Mosely identified the type-material of pubescens as petiolata, but Kimmins decided that it 
represented a distinct species on the grounds that Ulmer had not mentioned an apical wing spot 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 

-I 



101 




Figs 74-77 Amphipsyche petiolata $ . 74. wing venation; 75, legs; 76, eighth sternites; 77, maxillary palp. 



in the description of petiolata. However, Ulmer's species was described from females only, and 
the wing markings are very faint in this sex. 

In addition to the distribution records below, Ulmer (1930) also recorded this species from 
Sumatra. 

MATERIAL EXAMINED 

Lectotype $ of petiolata, Java: Wonosobo, iv.1909 (Jacobson) (ZM). Lectotype $ of minima, West 
Malaysia: Kedah, nr Jitra, catchment area, 9.iv.l928 (Pendlebury) (BMNH). Holotype d" oipubescens, 
Borneo: Sarawak, foot of Mt Dulit, junction of Rivers Tinjar and Lejok, 20.viii.1932 (Hobby & Moore) 
(BMNH). 

3 cf, 5 $, West Malaysia (1 $ paralectotype of minima); Borneo: Sarawak (1 cf, 2 $ paratypes of 
pubescens) (BMNH, MCZ). 

The meridiana-group 

Genae rounded, with no pubescence. Fork II sessile in fore wing. spurs on fore leg; mid spurs sometimes 
reduced to 3 or 2; hind spurs always reduced to 3 or 2. Fifth segment of maxillary palp often reduced, or 
fused with 4th segment, c? inferior appendages slender; phallocrypt pocket and pre-anal appendages 
absent. Phallotheca with up to three pairs of endothecal spines. Ninth segment with single (dorsal) row of 
setae in lateral view. 

Africa, Madagascar, India, Sri Lanka, Nepal, Cambodia, West Malaysia, Sumatra, Java, 
Borneo, Philippines. 



102 



P. C. BARNARD 



Amphipsyche magna Banks 

(Figs 78-88) 
Amphipsyche magna Banks, 1939: 58. Holotype cf , PHILIPPINES (MCZ) [examined]. 

Cf (holotype only). Antennae missing, described by Banks (1939) as 'pale, tips of joints dark'. Fore wing 
20mm. Head, thorax and abdomen pale yellowish brown, mesoscutellum with two round dark brown 




Figs 78-83 Amphipsyche magna cf . 78, wing venation; 79, mid and hind tibiae; 80, maxillary palp; 81. 
genitalia, lateral view; 82, phallotheca, lateral view; 83, genitalia, ventral view. 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



103 




87 



Figs 84-88 Amphlpsyche magna. 84. 9 wing venation; 85, $ mid tibia, 86, $ maxillary palp; 87, 
thorax, dorsal view; 88, $ eighth sternites. 



markings (Fig. 87). Fore wing elongate, yellowish brown with no markings. Venation as in Fig. 78; closed 
median cell in hind wing formed by M 2 -M 3+4 cross-vein. Spurs 0.4.2 (Fig. 79), not 1 .4.2 as stated by Banks. 
Maxillary palp 5-segmented, 5th segment short, not secondarily annulated (Fig. 80). 

9 (single example). Antennal length unknown (specimen damaged). Fore wing 15 mm. Coloration as in 
Cf, with similar round markings on mesoscutellum. Basal antennal segments pale yellow, narrowly 
annulated with brown. Venation as in Fig. 84; closed median cell in hind wing as in cf Spurs 0.4. [? 2] (hind 
legs missing) (Fig. 85). Maxillary palp 5-segmented, 5th segment shorter than in cf (Fig. 86). 

GENITALIA cf (Figs 81-83). Ninth segment broadly rounded laterally. Base of phallotheca strongly 
flattened dorso-ventrally, apex rounded. Three pairs of endothecal spines present; dorsal pair directed 
ventrally, mid and ventral pairs curved dorsally. Inferior appendage thin and strongly sinuous; terminal 
segment moderately clearly differentiated. 

GENITALIA $ (Fig. 88). Eighth sternites subrectangular, much longer than broad; inner thickened margins 
broad. 



104 



P. C. BARNARD 



REMARKS. Despite the unlikely sounding combination of names, it seems that magna, the largest 
species in the genus, andparva, one of the smallest, are sister species. They are, of course, easily 
separable by the great disparity in size, and magna is particularly easy to recognize by the 
mesoscutellar markings and hind wing median cell in both sexes. Both magna andparva have the 
full complement of three pairs of endothecal spines and both have the unusually shaped 
phallothecal base, which is elongate and strongly flattened in lateral view. 

This is the first description of the female of magna. The two striking external characters of the 
species, the mesoscutellar markings and the hind wing median cell, are the same in each sex, as is 
the reduced condition of the maxillary palps. 

MATERIAL EXAMINED 

Holotype cf , Philippines: Luzon, Del Carmen, 15. xi. 1927 (Uichanco) (type no. 23471, MCZ). 
1 $, Philippines: Luzon (USNM). 




91 



92 



Figs 89-93 Amphipsyche parva c?. 89, wing venation; 90, mid and hind tibiae; 91, genitalia, lateral view; 
92, phallotheca, lateral view; 93, genitalia, ventral view. 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 105 

Amphipsyche parva Banks 

(Figs 89-93) 
Amphipsyche parva Banks, 1920: 354. Holotype d", BORNEO (MCZ) [examined]. 

Cf (holotype only). Antenna c. 25 mm with c. 80 segments. Fore wing 8 mm. Antennal segments pale 
yellow, slightly annulated with brown. Head, thorax and abdomen yellowish brown. Fore wing very pale 
yellow, almost colourless, with no markings. Venation as in Fig. 89: RI in fore wing strongly sinuous, fork I 
in hind wing clearly stalked. Spurs 0.4.2 (Fig. 90). Maxillary palps missing. 
$. Unknown. 

GENITALIA cf (Figs 91-93). Ninth segment only slightly broadened laterally. Base of phallotheca strongly 
flattened dorso-ventrally, apex rounded. Three pairs of endothecal spines present; dorsal pair short, 
curved latero-ventrally; mid pair strongly curved dorsally; ventral pair long and almost straight, slightly 
directed dorsally. Inferior appendage thin and sinuous, terminal segment scarcely differentiated. 

REMARKS. A. parva and magna are the only two species in the genus to possess three pairs of 
endothecal spines. A. parva is easily distinguished from magna (its sister species) by its small size 
and lack of thoracic markings. Little can be surmised about the maxillary palps of this species 
(which are missing in the holotype): although these are often reduced in the meridiana-group, 
this is not invariably the case, and the shortened apical segment in the sister species magna may 
be a unique character within this species-pair. 

MATERIAL EXAMINED 
Holotype cf , Borneo: Mindai, vi.1882 (Grabowsky) (type no. 10886, MCZ). 



Amphipsyche sinhala sp. n. 

(Figs 94-103) 

Cf . Antenna c. 35 mm, with c. 85 segments. Fore wing 10-12 mm. Body pale yellowish brown; basal 
antennal segments narrowly annulated with pale brown, apical segments fuscous. Fore wing very pale 
yellow with no markings; venation as in Fig. 94. Spurs 0.4.2 (Fig. 95). Maxillary palp 5-segmented; 5th 
segment secondarily annulated, approximately equal in length to segments 1-4 combined (Fig. 96). 

$. Antenna c. 14 mm, with c. 65 segments. Fore wing 7-8 mm. Coloration as in cf . Venation as in Fig. 
100. Spurs 0.4.2 (Fig. 101). Maxillary palp as in cf , but 5th segment slightly shorter than segments 1-4 
combined (Fig. 103). 

GENITALIA cf (Figs 97-99). Ninth segment broad laterally. Phallotheca with moderately narrow stem, 
broadly truncate at apex. Mid endothecal spines very short, rod-like (Fig. 98); dorsal and ventral 
endothecal spines absent. Inferior appendage moderately narrow, slightly sinuous; terminal segment 
clearly differentiated. 

GENITALIA $ (Fig. 102). Eighth sternites narrow, outer borders strongly sloping, posterior border broadly 
pointed. 

REMARKS. A. sinhala is apparently restricted to Sri Lanka, and the only other species reported 
from that country is meridiana. They can be separated easily on genitalic differences in both 
sexes, as well as on the spur formula; meridiana always has at least three spurs on the hind tibia in 
both sexes. Moreover, there is little overlap in size, sinhala being a noticeably small species. A. 
sinhala also resembles bengalensis, but is distinguished by the much shorter endothecal spines. 

MATERIAL EXAMINED 

Holotype cf , Sri Lanka: Panamure, 15-21. x. 1970 (Flint) (USNM). 

Paratypes. Sri Lanka: 16 cf , 36 $ , data as holotype (all in USNM except 2 cf , 2 $ in BMNH) ; 2 cf , 18 $ , 
Sella Kataragama, Menik Ganga, 24.x. 1970 (Flint) (USNM). 



106 



P. C. BARNARD 




Figs 94-99 Amphipsyche sinhala cf . 94, wing venation; 95, mid and hind tibiae; 96, maxillary palp; 97, 
genitalia, lateral view; 98, phallotheca, lateral view; 99, genitalia, ventral view. 



Amphipsyche meridiana Ulmer 
(Figs 104-1 14) 

Amphipsyche meridiana Ulmer, 1909: 134. LECTOTYPE $ , JAVA (RNH), here designated [examined]. 

[Phanostoma sp. Betten, 1909: 234.] 

Amphipsyche nirvana Banks, 1913: 236. Holotype cf , INDIA (MCZ) [examined]. Syn. n. 

Amphipsyche vedana Banks, 1913: 235. Holotype 9, INDIA (MCZ) [examined]. Syn. n. 

Ampsipsyche [sic] propinqua Ulmer, 1927: 177. LECTOTYPE cf, CAMBODIA (MNHU), here designated 

[examined]. Syn. n. 

[Amphipsyche proluta McLachlan; Navas, 1931ft: 91; 1934: 227. Misidentifications.] 
Amphipsyche indica Martynov, 1935: 199. 8 syntypes, INDIA: 1 d", Bihar, Mokameh, at light; 1 cf , Bihar, 

Dinapore, at light (Annandale); 2 CT, 2 $>, Bihar, Pusa, 5-10.xi.1915 (Gravely); 2 cf, E. Bengal, 

Damukdia Ghat, at light on board steamer, 30. vi. 1908 (2 syntypes in ZSI, the other 6 lost) [not 

examined]. Syn. n. 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



107 




Figs 100-103 Amphlpsyche sinhala $ . 100, wing venation; 101, mid and hind legs; 102, eighth sternites; 
103, maxillary palp. 



Amphipsyche tricalcarata Martynov, 1935: 197. Holotype $, INDIA: Orissa, Puri district, Bhubaneswar, 
4-6.xi.1912 (Gravely) (lost from ZSI). [Synonymized with indica by Schmid, 1958: 107.] 

Amphipsyche sigmosa Navas, 1935: 105. LECTOTYPE d", INDIA (MNHN), here designated [examined]. 
Syn. n. 

C?. Antenna c. 40 mm with up to 100 segments. Fore wing 13-15 mm. Body pale yellowish brown, antennal 
segments pale golden brown. Fore wing pale golden yellow, sometimes with pale brown marking behind 
R\-Rs cross-vein. Venation as in Fig. 105; RI in fore wing strongly sinuous both proximal and distal to 
anastomosis. Spurs 0.4.3 or 0.4.4 (pre-apical spurs on hind tibia always very small) (Fig. 106). Maxillary 
palp 5-segmented, 5th segment secondarily annulated, approximately equal in length to segments 1-4 
combined (Fig. 107). 

$. Antenna c. 18 mm, with c. 70 segments. Fore wing 8-12 mm. Coloration as in cf ; dark marking on 
fore wing always absent. Venation as in Fig. Ill; fork IV in fore wing occasionally stalked. Spurs (Fig. 112) 
and maxillary palp (Fig. 114) as in cf . 

GENITALIA cf (Figs 104, 108-109). Ninth segment moderately broad laterally. Phallotheca elongate, with 
narrow stem. Dorsal endothecal spines absent, in their place a pair of semi-membranous leaf-like lobes, 
variable in shape (Fig. 109). Mid and ventral endothecal spines short, varying in relative length; ventral 
pair occasionally lost. Inferior appendage narrow and sinuous, terminal segment moderately well 
differentiated. 

GENITALIA 9 (Fig. 113). Eighth sternites broad and squarish with broadly rounded corners. 



108 



P. C. BARNARD 




-0 



70 



Fig. 104 Variation in cf genitalia of Amphipsyche meridiana throughout its range. 

REMARKS. A. meridiana is the most common and widespread of the Asian species in the genus. 
The male is easily recognized by the dorsal leaf-like lobes on the phallotheca, but the female may 
be confused with sinhala unless it is examined closely; the hind pre-apical spurs are always 
extremely small, thus the spur formula may be taken erroneously as 0.4.2. 

The large number of synonyms of this species is partly a result of its morphological variability 
over a wide geographical range. Banks (1913) said that Betten's (1909) "Phanostoma sp.' was the 
same as, or very similar to, nirvana, and Martynov (1935) said that it was identical with his new 
species indica. Banks (1939) apparently regarded indica as a synonym of nirvana, as he placed 
the name indica in parentheses after nirvana. Meanwhile, Ulmer (1927) had compared 
propinqua with nirvana (but not with his own species meridiana) but later (1951) noted that 
nirvana, propinqua and meridiana were all very similar. Thus the Indian species were long 
considered as being synonymous, but the synonymy with meridiana was not suspected, partly 
because of the geographical separation (meridiana being described from Java) and partly 
because the male of meridiana was not described until 1951. 

The extent of variation in the male phallotheca is shown semi-diagrammatically in Fig. 104. 
The mid endothecal spines are moderately consistent in size and form throughout the whole 
range, although in the single known male from Bombay they are very short and broad, and bent 
abruptly upwards. However, the ventral endothecal spines vary greatly, and there seems to be a 
correlation with distribution, such that in the most eastern specimens they are much longer than 
the mid spines, whereas in the western (Indian) populations they are usually much shorter, and 
even lost in specimens from west India (and also occasionally Sri Lanka). There is in fact a 
discontinuity in the distribution of this species, with no specimens known from countries 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



109 




Figs 105-110 Amphipsyche meridiana d" 105, wing venation; 106, mid and hind tibiae; 107, maxillary 
palp; 108, genitalia, lateral view; 109, phallotheca, lateral view; 110, genitalia, ventral view. 

between India and Cambodia. There may be justification for considering the two populations as 
subspecifically distinct, in which case the Indian subspecies would have to be named meridiana 
nirvana, with the nominate subspecies in South East Asia, but I do not propose such a formal 
division at present. 



110 



P. C. BARNARD 




113 





Figs 111-114 Amphipsyche meridiana 
sternites; 114, maxillary palp. 



Ill, wing venation; 112, mid and hind tibiae; 113, eighth 



I do not believe that the 'paratype' of meridiana mentioned by Weidner (1964) has any 
type-status. Although from the type-locality, it bears a printed label with the date 'Dec. 1908'. A 
'Paratype' label has also been attached, bearing the hand-written date '8.1907', not in Ulmer's 
hand, to conform to the published type-data. However, the other labels do not match those on 
the two remaining syntypes in Leiden; of the original three syntypes mentioned by Ulmer in the 
RNH, one has apparently been lost (Geijskes, in lift.). 

Of the three syntypes oipropinqua described by Ulmer (1927) I have examined the two males 
in MNHU and designate as lectotype the one labelled 'type' by Ulmer. The third male syntype, 
now a paralectotype (IP, not examined), lacks its abdomen (Director, IP, in lift.). 

I was informed by Ghosh (in litt.) of the apparent loss of six syntypes of indica, and of the 
holotype of tricalcarata from the ZSI. The female syntypes oisigmosa (from Khandala) are also 
apparently lost, so the sole remaining male syntype in the MNHN is here designated as 
lectotype. 

The larva of this species was described by Hafiz (1937, as indica), and by Ulmer (1957). There 
are some differences between these two descriptions, both in the gill formulae and in the head 
markings. Specimens from Java that I have examined differ slightly in gill counts from Ulmer's 
description, even though his material was also from Java (and Sumatra). Some aspects of the life 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



111 



history are described by Seshadri (1955) and Boon (1979) - see p. 79. Some specimens in the 
BMNH, received via the Commonwealth Institute of Entomology, were captured on paddy- 
fields in India, but any economic significance of this is unknown. 

MATERIAL EXAMINED 

Lectotype $ of meridiana, Java: Batavia, viii.1907 (Jacobson) (RNH). Holotype cf of nirvana, India: 
Bengal, Pusa, at light, 23.iii.1908 (type no. 11755, MCZ). Holotype $ of vedana, India: Bengal, Pusa, 
15. ix. 1907 (type no. 11757, MCZ). Lectotype cf of propinqua, Cambodia: Mekong, Pnom-Pech, i.1914 
(Friederichs) (MNHU). Lectotype cf of sigmosa, India: Bombay, Lonawla [= Lonavla, = Lonauli], 
9.x. 1934 (Benavent) (MNHN). 

229 Cf, 332 9, c. 75 larvae, 2 pupae, India, Sri Lanka, Nepal, Cambodia (1 cf paralectotype of 
propinqua), West Malaysia, Sumatra, Java (1 9 paralectotype and 1 $ as 'paratype' of meridiana; see 
'Remarks' above) (BMNH, MCZ, MNHU, RNH, USNM, ZM). 

Amphipsyche bengalensis Martynov 
(Figs 117, 118) 

Amphipsyche bengalensis Martynov, 1935: 201. 2 cf syntypes, INDIA: Bengal, Calcutta, at light, 19. vi. 1907 
(Hodgart) (ZSI) [not examined]. 

Cf (from Martynov, 1935). 'Body pale yellow. Antennae yellow, with narrow dark annulations. Anterior 
wings pale. R[/?i] curved in its apical portion even more strongly than in A. indicum [= meridiana] ... In 
posterior wings first (apparently false) fork sessile, not pedicellate; . . . formula of spurs 1.4.2 [probably 
0.4.2]. Length of body 6mm.' 
9- Unknown. 

GENITALIA cf (Figs 117, 118) (from Martynov, 1935). '10th [9th] segment as in [meridiana], but its 
side-lobes appear to be somewhat broader. Lower end-lobe of the penis [phallotheca] broader, distinctly 
excised in the middle and curved upwards (if seen from side); upper leaf-like lobes lacking, in their place is 
an oval elevation, behind which are situated two stick-shaped appendages [mid endothecal spines]; 
underside of the penis thickened before its lower end-lobe.' 

REMARKS. This species seems to be most closely related to sinhala, but the male genitalia are 
different, assuming Martynov's figures to be accurate. Although Martynov gave no wing-length 
for bengalensis, this would also seem to be a larger species than sinhala, whose body length is 




116 



117 




118 



Figs 115-118 115, 116, Amphipsyche extrema ?, (115) wing venation; (116) palps. 117, 118, Amphip- 
syche bengalensis cf , (117) phallotheca, lateral view; (118), phallotheca, dorsal view. (After Martynov.) 



112 P. C. BARNARD 

only 4-5 mm. The 6 mm body length of bengalensis suggests that it is of a similar size to 
meridiana. The spur formula of this species should almost certainly be 0.4.2; Martynov probably 
mistook an apical seta on the fore tibia for a spur. 

The two syntypes in the ZSI are damaged (Ghosh, in litt.) and I was unable to examine them. 
The species is known only from these two specimens from Bengal. 

Amphipsyche extrema (Martynov) comb. n. 
(Figs 115, 116) 

Amphipsychella extrema Martynov, 1935: 202. 2 syntypes, INDIA: 1 $ , Bengal, Calcutta, Eden Garden, at 
light, 26.V.1912 (Gravely); 1 $, Bengal, Calcutta, v.1915 (Gravely) (both lost from ZSI). 

Cf . Unknown. 

$ (from Martynov, 1935). Tale yellow. Antennae very slender, yellowish, with narrow darker 
annulations. Maxillary palpi very short, not reaching eyes; 2-4 joints subequal; 5th joint shorter than 3rd 
and 4th combined, its distal half slender and but very indistinctly annulated; labial palpi also very short 
[Fig. 116] . . .In anterior wings [Fig. 115] three false veinlets are seen between C and Sc; 1st apical fork a 
little longer than its pedicel and somewhat approximated to R [R\]', R long, slightly arcuate; . . . 4th apical 
fork with a short pedicel. RS 1+2 in posterior wings simple, not united at its base with RS 3 . Abdomen pale. 
Length of body 5-5 mm.' [From generic diagnosis of Amphipsychella] 'Spurs 0.2(1). 2, the outer spur on the 
median legs reduced, indistinct.' 

REMARKS. It is difficult to comment on the relationships of this species, as it is known only from 
the female which I have not examined; apparently both syntypes are lost from the ZSI (Ghosh, 
in litt.). However, the highly reduced spur formula, and the shortened maxillary palp place it in 
the meridiana-group. It would seem most closely related to bengalensis, meridiana and sinhala, 
but can be distinguished from these other Indian species by the spur formula and the maxillary 
palp. 

Amphipsyche senegalensis (Brauer) 
(Figs 120-129; distribution, Fig. 119) 

Phanostoma senegalense Brauer, 1875: 71. Lectotype cf , SENEGAL (NM), designated by Kimmins, 1962: 86 

[examined]. 
Phanostoma curvinerve Navas, 1927: 214. LECTOTYPE $ , EGYPT (USNM), here designated [examined]. 

Syn. n. 
Amphipsyche senegalensis (Brauer) Kimmins, 1962: 85. 

Cf . Antenna c. 40 mm, with c. 75 segments. Fore wing 11-17 mm. Body pale yellowish brown, antenna 
narrowly annulated with brown. Fore wing pale yellow, often with brownish wedge-shaped pterostigmal 
marking; venation as in Fig. 120. Spurs 0.4.2 (Fig. 121). Maxillary palp 5-segmented; 5th segment- 
secondarily annulated, slightly longer than segments 1 and 2 combined (Fig. 122). 

$. Antenna c. 15 mm, with c. 65 segments. Fore wing 9-12 mm. General coloration as in cf Fore wing 
usually unmarked, rarely with pale brown pterostigmal marking; venation as in Fig. 126; Rs in fore wing 
strongly sinuous. Spurs 0.2.2, 0.3.2 (Fig. 127) or 0.4.2. Maxillary palp similar to that of cf ; 5th segment 
approximately equal to 1 and 2 combined (Fig. 129). 

GENITALIA cf (Figs 123-125). Ninth segment broadly rounded laterally. Base and stem of phallotheca 
narrow, apex bluntly rounded; no endothecal spines present. Inferior appendage slender; terminal 
segment moderately well differentiated. 

GENITALIA $ (Fig. 128). Eighth sternites broadly rounded; slight indentation in middle of posterior edge, 
and outer posterior corners produced. 

REMARKS. This species is easily distinguished from the other African species by the complete 
absence of endothecal spines in the male. The female can probably be distinguished by the very 
sinuous Rs in the fore wing, but this cannot be confirmed until the females of all the African 
species have been discovered. A. senegalensis has a very wide distribution, being found in 
virtually every country in the Afrotropical region except in the south-west and along the east 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



113 



berneri 



..-...- v '"'.' 'Vv ulmeri 0\ 



i. ';..'; senegalensis '-..' '.;.': .'.'.' '.''.'.' 



(.'': '''.'.'.'.':':'. ''.''' corbeti '.; 




scottae 



Fig. 119 Distribution of the African species oiAmphipsyche. 

coast (Fig. 119); this distribution coincides closely with the distribution of permanent waters in 
Africa (Gourou, 1970). The species is often caught in very large numbers, especially at light. 

The synonymy of curvinerve with senegalensis was suspected by Kimmins (1962); the only 
remaining syntype of curvinerve, from the Alfieri collection, now in the USNM, has genitalia 
indistinguishable from those of typical senegalensis. Ulmer (1963) retained the name curvinerve 
when describing the larva of this species, but he admitted that he could not separate the two 
species. Ulmer had seen no females from West Africa to compare with his Egyptian examples, 
and he rightly suspected that his Sudanese specimens represented a different species: this was 
described as ulmeri by Kimmins (1962). 

The female figured by Savigny (1813) from Egypt is certainly this species, though not named; 
this was the first published figure of a species olAmphipsyche. 

The larva of senegalensis was described by Hickin (1955), Jacquemart (1957), Marlier (1962) 



114 



P. C. BARNARD 




Figs 120-125 Amphipsyche senegalensis cf . 120, wing venation; 121, mid and hind legs; 122, maxillary 
palp; 123, genitalia, lateral view; 124, phallotheca, lateral view; 125, genitalia, dorsal view. 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



111 




Figs 136-139 Amphipsyche pellucida $ . 136, wing venation; 137, mid and hind legs; 138, eighth sternites; 
139, maxillary palp. 



REMARKS. Although this species clearly belongs in the 'African' section of the meridiana-group, 
it has no close affinities with any other species. It is the only species found in Madagascar, and 
morphologically the form of the phallotheca renders it easily identifiable. This is the first time 
that the male has been described. 

Navas (1923) mis-read the type-locality of pellucida as 'Maeratanana'; this also applies to 
other species described in the same paper. 

MATERIAL EXAMINED 

Holotype 9> Madagascar: Maevatanana [no further data] (MNHN). 
3 cf , 13 $. Madagascar (BMNH, USNM). 

Amphipsyche instabilis Kimmins 
(Figs 140-150; distribution, Fig. 119) 

Amphipsyche instabilis Kimmins, 1963: 126. Holotype cf , ETHIOPIA (BMNH) [examined]. 
Phanostoma plicata Jacquemart, 1963: 363. LECTOTYPE cf , ZIMBABWE (ZI), here designated [ex- 
amined]. Syn. n. 

O" . Antenna up to 33 mm, with c. 90 segments. Fore wing 11-14 mm. Body pale yellowish brown, antennal 



118 



P. C. BARNARD 




U2 




U5 





Figs 140-146 Amphipsyche instabilis C". 140, wing venation; 141, mid and hind tibiae; 142, maxillary 
palp; 143, genitalia, lateral view; 144, genitalia, ventral view; 145, phallotheca, lateral view; 146, 
phallotheca, dorsal view. 

segments becoming gradually more fuscous towards apex, narrowly annulated with brown. Fore wing pale 
yellow with no markings; venation as in Fig. 140; cross-vein present between M 3+4 and Cw la in hind wing. 
Apical venation often irregular in both fore and hind wings. Spurs 0.4.2 (Fig. 141). Maxillary palp 
4-segmented, apical segment short and not secondarily annulated (Fig. 142). 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



119 



9 (allotype only). Antennal length unknown (specimen damaged). Fore wing 9 mm. Coloration as in cf , 
venation as in Fig. 147; cross-vein between A/ 3+4 and Cu la in hind wing as in cf . Spurs 0.2.2 (Fig. 148). 
Maxillary palp 4-segmented (Fig. 149), similar to that of cf . 

GENITALIA cf (Figs 143-146). Ninth segment only moderately broad laterally. Base of phallotheca very 
large and rounded, stem short and greatly thickened; dorsal clavate process present, with curved stem, 
apex covered with fine spines (Fig. 145); on either side of process a similarly spinose triangular lobe. Mid 
endothecal spines fused to form single median spine, very thick and curved dorsally. Lower apex of 
phallotheca elongate, curved dorsally, extreme apex globular. Inferior appendage narrow, slightly sinuate; 
terminal segment moderately well differentiated. 

GENITALIA 9 (Fig- 150). Eighth sternites broad and squarish, with rounded indentation in posterior edge; 
inner thickened edges narrow. 

REMARKS. Although this species is closely related to the other African species of the genus, it is 
easily recognized. Externally, both sexes are easily identified by the extra cross-vein between 
M 3+4 and Cw la in the hind wing which is unique amongst the African species, though paralleled 
inproluta, the type-species of the genus. The male genitalia are highly distinctive; the elongate 
dorsal process and single median endothecal spine are not found in any other species. The only 
known female closely resembles the female of senegalensis , which is found in the same locality, 
but apart from the venational character already mentioned, it can easily be distinguished by the 
4-segmented maxillary palp and the squarish eighth sternites. 

The specimen here designated as lectotype ofplicata was labelled 'type' by Jacquemart, and 
the paralectotypes as 'paratypes', but these designations were not pubished. 

Although the descriptions ofinstabilis andplicata were published in the same year, there is no 
doubt that Kimmins's appeared first. His paper was officially published on 20th February 1963 
and, according to data in the BMNH Entomology Department Library, it was definitely 
available before the end of that month. The book in which Jacquemart's paper appeared has no 
exact date of publication, but the copy in the BMNH Zoology Department Library was received 




U8 



Figs 147-150 Amphipsyche instabilis $. 147, wing venation; 148, mid and hind tibiae; 149, maxillary 
palp; 150, eighth sternites. 



120 



P. C. BARNARD 



on llth October 1963. Further enquiries to the ZI revealed that their copy was received on 25th 
November 1963 (Tjeder, in litt.), and the copy in the Kungliga Biblioteket, Stockholm (the 
Swedish copyright library) was not received until February 27th 1964 (Lilliestam, in litt). It 
seems certain, therefore, that the description oiplicata was not published until at least October 
1963, and that the name instabilis has priority. 

MATERIAL EXAMINED 

Holotype cf of instabilis, Ethiopia: Dawa River, 12 km N. of Hudat, 12. iv. 1961 (Tj0nneland) (slide 
preparation, BMNH). Lectotype cf of plicata, Zimbabwe: Victoria Falls, 16. v. 1951 (slide preparation, 
ZI). 

38 cf , 1 $ , Ethiopia (18 cf paratypes of instabilis and 1 $ allotype inadvertently labelled as 'cf paratype' 
by Kimmins), Zimbabwe (5 d" paralectotypes of plicata), Zambia (BMNH, IRSNB, USNM, ZI). 

Amphipsyche ulmeri Kimmins 
(Figs 151-154; distribution, Fig. 119) 

[Phanostoma senegalense Brauer; Ulmer, 1923: 19 (partim - specimens from Sennar only); 1924: 2. 

Misidentifications . ] 
Amphipsyche ulmeri Kimmins, 1962: 89. Holotype cf , SUDAN (NM) [examined]. 



151 




153 



Figs 151-154 Amphipsyche ulmeri cf . 151, wing venation; 152, genitalia, ventral view; 153, genitalia, 
lateral view; 154, phallotheca, lateral view. 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 121 

Cf (holotype only). Antenna 32 mm, with c. 70 segments. Fore wing 14 mm. Antennal segments pale 
golden brown, annulated with dark brown. Head, thorax and abdomen yellowish brown. Fore wing pale 
yellow with indistinct brownish shading across anastomosis; venation as in Fig. 151. Spurs 0.4.2. Maxillary 
palp as in senegalensis (cf. Fig. 122). 

9- No specimens seen, but see 'Remarks' below. 

GENITALIA cf (Figs 152-154). Lateral part of ninth segment broad and squarish. Base of phallotheca broad, 
extreme apex bluntly pointed; mid endothecal spines long and stout, curved abruptly dorsally (Fig. 154). 
Inferior appendage strongly sinuous in ventral view, terminal segment moderately well differentiated. 

REMARKS. A. ulmeri and scottae are closely related, despite their widely separate distributions 
(Fig. 119) , both species having very similar male genitalia. However, ulmeri can be distinguished 
by the sharply up-turned mid endothecal spines. 

Kimmins (1962) mentioned the existence of females from the type-locality, Sennar, in 
Ulmer's collection. Subsequently Ulmer (1963) described these females as being distinguishable 
from Egyptian females of curvinerve (= senegalensis) by the less sinuous Rs in the fore wing and 
the spur formula of 0.3.2. It is quite probable that these are females of ulmeri, but the spur 
formula is not significant, as Kimmins (1963) showed that there is great variation in the spurs of 
female senegalensis, 0.3.2 occurring in that species also. 

MATERIAL EXAMINED 
Holotype cf , Sudan: Sennar, 18-27. ii. 1914 (Ebner) (NM). 



Amphipsyche scottae Kimmins 
(Figs 155-164; distribution, Fig. 119) 
Amphipsyche scottae Kimmins, 1962: 93. Holotype cf , SOUTH AFRICA (BMNH) [examined]. 

Cf . Antenna c. 45 mm, with c. 95 segments. Fore wing 16-19 mm. Body yellowish brown, antenna narrowly 
annulated with brown, segments becoming more fuscous towards apex. Fore wing pale yellow, with slightly 
darker area along costa and near Sc-Ri cross-vein, often indistinct. Venation as in Fig. 155. Spurs 0.4.2 
(Fig. 156). Maxillary palp 5-segmented, 5th segment approximately equal in length to segments 1 and 2 
combined, not secondarily annulated (Fig. 157). 

$. Antenna c. 20 mm, with c. 70 segments. Fore wing 14-15 mm. General coloration as in cf ; fore wing 
with no darker markings. Venation as in Fig. 161. Spurs 0.4.2 (Fig. 162). Maxillary palp similar to that of 
Cf , but 5th segment shorter (Fig. 164). 

GENITALIA cf (Figs 158-160). Ninth segment broadly rounded laterally. Base of phallotheca broadly 
triangular, with pronounced corner on dorsal side. Ventral apex forming pair of rounded lobes; extreme 
apex bluntly pointed (Fig. 159). Mid endothecal spines long, curved dorsally. Inferior appendage slender, 
terminal segment moderately well differentiated. 

GENITALIA $ (Fig. 163). Eighth sternites broad and squarish, with slight indentation in middle of posterior 
edge. Inner thickened margin extending far towards anterior edge. 

REMARKS. A. scottae most closely resembles ulmeri in that the males of both species have the tip 
of the phallotheca bluntly pointed in lateral view, but scottae can be distinguished by the gently 
curved mid endothecal spines, which are sharply up-turned in ulmeri. 

The larva of this species was described by Scott (in press), and some aspects of its biology are 
mentioned by Chutter (1963; 1968); see p. 79. 

MATERIAL EXAMINED 

Holotype cf, South Africa: Natal, Wilge R., 5 miles [8 km] below Harrismith, 10.ii.1959 (slide 
preparation, BMNH). 

12 Cf , 2 $ , South Africa (12 cf , 1 $ paratypes) (BMNH). 



122 



P. C. BARNARD 




Figs 155-160 Amphipsyche scottae C? . 155, wing venation; 156, mid and hind tibiae; 157, maxillary palp; 
158, genitalia, lateral view; 159, phallotheca, lateral view; 160, genitalia, ventral view. 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



123 




Figs 161-164 Amphipsyche scottae $. 161, wing venation; 162, mid and hind legs; 163, eighth sternites; 
164, maxillary palp. 



Amphipsyche fuscata Kimmins 
(Figs 165-170; distribution, Fig. 119) 
Amphipsyche fuscata Kimmins, 1963: 128. Holotype cf , ETHIOPIA (BMNH) [examined]. 

Cf . Antenna up to 35 mm, with c. 85 segments. Fore wing 12-17 mm. Body pale yellowish brown; basal 
12-15 segments of antenna yellowish brown, remainder of flagellum fuscous. Fore wing pale yellow, with 
fuscous streak running obliquely from costal margin proximal to anastomosis; hind margin fuscous (these 
markings may be very faint). Venation as in Fig. 165. Spurs 0.4.2 (Fig. 166). Maxillary palp short, 4th and 
5th segments imperfectly separated, 5th segment narrow apically, not secondarily annulated (Fig. 167). 
$. Unknown. 

GENITALIA cf (Figs 168-170). Lateral lobe of ninth segment somewhat squarish. Base of phallotheca 
narrow and rounded, apex elongate, produced into a bifid lobe. Mid endothecal spines long, curved 
upwards. 

REMARKS. Well-marked specimens of this species can be recognized by the oblique wing- 
marking, but identification of specimens with faint markings depends on the male genitalia, and 
here the close similarity with several other African species is apparent. However , fuscata differs 



124 



P. C. BARNARD 




Figs 165-170 Amphipsyche fuscata d". 165, wing venation; 166, mid and hind tibiae; 167, maxillary palp; 
168, genitalia, lateral view; 169, phallotheca, lateral view; 170, genitalia, ventral view. 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 



125 



from scottae and ulmeri in having the apex of the phallotheca rounded, not pointed, and from 
berneri and corbeti in having a narrow base to the phallotheca. A.fuscata is further distinguished 
by the unique maxillary palps, with the 4th and 5th segments only partly fused. Kimmins (1963) 
noted the colour of the palps, but did not notice their unusal form. 

MATERIAL EXAMINED 

Holotype d", Ethiopia: Koka Dam, 29.iii.1964 (Tj0nneland) (slide preparation, BMNH). 
62 cf , Ethiopia (60 cf paratypes), Sudan (BMNH). 

Amphipsyche corbeti Kimmins 
(Figs 171-176; distribution, Fig. 119) 
Amphipsyche corbeti Kimmins, 1962: 89. Holotype cf , UGANDA (BMNH) [examined]. 

Cf. Antenna up to 30 mm, with c. 85 segments. Fore wing 11-12 mm. Body pale yellowish brown, thorax 
pale brown dorsally, antenna pale greyish brown. Fore wing pale yellowish brown, usually with pale brown 
shading around Ri~Rs cross-vein. Venation as in Fig. 171. Spurs 0.4.2 (Fig. 172). Maxillary palp 
4-segmented, terminal segment narrow and slightly elongate, but not secondarily annulated (Fig. 173). 
$. Unknown. 




Figs 171-176 Amphipsyche corbeti cf . 171, wing venation; 172, mid and hind tibiae; 173, maxillary palp; 
174, genitalia, lateral view; 175, phallotheca, lateral view; 176, genitalia, ventral view. 



126 



P. C. BARNARD 



GENITALIA cf (Figs 174-176). Base of phallotheca broadly triangular, stem thickened, apex forming pair of 
lobes ventrally; mid endothecal spines long, curved dorsally. Inferior appendage slender, terminal 
segment scarcely differentiated. 

REMARKS. This species closely resembles berneri and, to a lesser extent, ulmeri. It differs from 
ulmeri in the thicker stem of the phallotheca and the longer endothecal spines, and from berneri 
in the lobes of the tenth segment. These lobes are narrower in dorsal view in corbeti, and do not 
diverge. There are also slight differences in the apex of the phallotheca, best seen in ventral 
view. 




Figs 177-182 Amphipsyche berneri cf . 177, wing venation; 178, mid and hind legs; 179, maxillary palp; 
180, genitalia, ventral view; 181, phallotheca, ventral view; 182, genitalia, lateral view. 



MACRONEMATINE CADDISFLIES OF THE GENUS AMPHIPSYCHE 127 

MATERIAL EXAMINED 

Holotype cf, Uganda: Northern Province, Victoria Nile, Karuma Falls [no date] (Corbet) (slide 
preparation, BMNH). 

33 Cf paratypes, Uganda (BMNH). 



Amphipsyche berneri Kimmins 
(Figs 177-182; distribution, Fig. 119) 

[Phanostoma senegalense Brauer; Kimmins, 1957a: 13 (partim - specimens from Gold Coast [Ghana] 

only). Misidentification.] 
Amphipsyche berneri Kimmins, 1962: 91. Holotype cf , GHANA (BMNH) [examined]. 

Cf . Antenna c. 25 mm, with c. 85 segments. Fore wing 11-12 mm. Body coloration uncertain (both 
specimens originally preserved in alcohol). Fore wing yellowish brown, with darker marking centred on 
R\-Rs cross-vein; venation as in Fig. 177. Spurs 0.4.2 (Fig. 178). Maxillary palp 4-segmented, terminal 
segment scarcely longer than 3rd (Fig. 179), not secondarily annulated. 
$. Unknown. 

GENITALIA cf (Figs. 180-182). Base of phallotheca broadly triangular; ventral apex forming pair of lobes; 
mid endothecal spines long and curved dorsally. Inferior appendage slender, terminal segment not clearly 
differentiated. 

REMARKS. This species is very similar to corbeti, both externally and in the form of the genitalia. 
It can be distinguished by slight differences in the shape of the apex of the phallotheca and by the 
lobes of the tenth segment. These are broader and more divergent in dorsal view in berneri, 
although it should be noted that in the paratype (figured here) the lobes are less divergent than in 
the holotype figured by Kimmins (1962: 91, fig. 26). The holotype is now mounted laterally as a 
permanent slide preparation. 

Kimmins did not comment on the 4-segmented maxillary palps, although these are clearly 
visible in his slide preparation of the holotype. 

MATERIAL EXAMINED 

Holotype cf , Ghana: Volta R., Senchi, l.viii.1950 (Berner) (slide preparation, BMNH). 
1 cf paratype, Ghana (BMNH). 

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Index 

Synonyms are in italics; principal references are in bold. References to Table 1 (p. 82), the cladogram 
(p. 83) and the check-list (p. 85) are not included. 



Aethaloptera72,78,84 
Amphipsyche 71, 75, 76 
Amphipsychella 76, 77, 112 
apicalis77,83,94,96,99 
apicalis-group 77, 78, 80, 81, 83, 
84, 94, 96, 99 

bengalensis 81, 105, 111,112 
berneri 77, 84, 125,126, 127 
bifasciata 89 
Blepharopus 74 

Cheumatopsyche 80 
corbeti 84, 125, 127 
curvinervelS, 79, 112, 113, 115, 
121 

delicata81,93 
distincta83,86,89,94 

exsiliens83,94,96 
extrema76, 81, 112 

fuscata 84, 123 
gratiosa83,96, 98 



indica 78, 93, 106, 108, 110, 111 
instabilis83,84,88, 117 

Leptonema 74, 79 
Leptopsyche 75 

Macronema 72, 74, 79 
Macronematinae 72, 74 
Macronematini 72, 74 
Macrostemum 74, 75, 78, 79, 84 
magna 77, 84, 92, 102 
meridiana 78, 79, 80, 83, 93, 99, 

105, 106, 111, 112 
meridiana-group 77, 78, 80, 81, 

83,84,85,88,94,101,105, 

112,117 
minima 94, 99 

nirvana 106, 108, 109 

paraproluta 86, 88 
parva 104, 105 
pellucida84, 115 
petiolata 94, 96, 99 
Phanostoma 76, 77, 79, 108 
plicata 117, 119, 120 



Polymorphanisini 72, 74 
Polymorphanisus 84 
proluta76,78,83,86,89,99, 

106, 119 
proluta-group 77, 78, 80, 81, 83, 

84, 86, 88, 94 
propinqualQ6,10S, 110 
Protomacronema 74, 75, 77, 79, 

84 

Pseudoleptonema 74 
pubescens 99, 100 

scottae 72, 79, 80, 121,125 
senegalensis 76, 78, 79, 80, 83, 

84, 112, 119, 121 
sigmosa 107, 110 
Simulium71,72, 79 
sinhala81,105, 108, 111,112 

tricalcarata 107, 110 
Trichomacronema 74 

ulmeri78,84, 113,120, 121, 
125, 126 

vedana 106 



British Museum (Natural History) 

Blue Butterflies of the Lycaenopsis-group 

J. N. Eliot and A. Kawazoe 

The most wide-spread member of the Lycaenopsis-group is known and loved in Britain as the 
Holly Blue, in North America as the Spring Azure and in Japan as Ruri Shijimi (the Small 
Lapis Lazuli). In appearance and behaviour it is typical of the group, which attains its 
maximum diversity and abundance in the mountains of South East Asia and New Guinea. 
Hitherto the systematics of the group have been in a state of confusion. In this work 112 
species are recognised divided among 21 genera. These are defined mainly on characters of the 
genitalia, which are figured for the males of each, and the females of most, species. There are 
keys to, and descriptions of, the genera, subgenera, species and subspecies, including 8 new 
genera and 27 new species. The new species and those not previously figured are illustrated in 
the plates at the rear of the book, and references are given to published figures of the 
remaining species. Finally, there is a complete bibliography, enabling the original descriptions 
of all the taxa to be traced. 

John Eliot is the author of many papers of a taxonomic nature and was the reviser of the 
third edition of Corbet & Pendlebury's classic work "The Butterflies of the Malay Peninsula" 
first published half a century ago. His contribution to zoology has been recognised by the 
presentation of the Stamford Raffles Award of the Zoological Society of London and the 
J. H. Bloomer Award of the Linnean Society of London. 

Akito Kawazoe has published many taxonomic papers, mainly in Japanese journals, and is 
best known in Europe as the senior author of "Coloured Illustrations of the Butterflies of 
Japan", a work notable not only for its superb coloured plates but also for numerous black and 
white drawings illustrating structural characters. His skill as artist and microscopist is again 
demonstrated in this book by more than two hundred figures of genitalia which will prove 
indispensable to a proper understanding of the Lycaenopsis-group. 

Blue butterflies of the Lycaenopsis-group. J. N. Eliot & A. Kawazoe 
1983, 296pp, 18 plates include 6 in colour, 560 text figures. (0 565 00860 9) 
28.00 



Titles to be published in Volume 48 



Gelechiid moths of the genus Mirificarma. 

By Linda M.Pitkin. 

Macronematine caddisflies of the genus Amphipsyche (Trichoptera: Hydropsychidae) 

By P. C. Barnard. 

A review of the genera oflndo- Pacific Encyrtidae (Hymenoptera: Chalcidoidea). 

By John S. Noyes and M. Hayat 



Photoset by Rowland Phototypesetting Ltd, Bury St Edmunds, Suffolk 
Printed in Great Britain by Henry Ling Ltd, Dorchester 






Bulletin of the 

British Museum (Natural History) 



A review of the genera 
of Indo-Pacific Encyrtidae 
(Hymenoptera: Chalcidoidea) 



John S. Noyes & M. Hay at 



Entomology series 

Vol 48 No 3 28 June 1984 



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 
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World List abbreviation: Bull. Br. Mas. nat. Hist. (Ent.) 



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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